Abstract
The family Plexauridae Gray, 1859 is characterized by grouping octocorals that have thick branches and coenenchyme. However, due to their relatively simple body plan, the establishment of homologous and, therefore, systematically informative morphological characters is a challenge for the systematics of Octocorallia. During the last decade, molecular studies suggested that Plexauridae is polyphyletic, with representatives grouping with Acanthogorgiidae Gray, 1859 and also Gorgoniidae Lamouroux, 1812. Represented by 22 extant species, mostly of which occur in the Atlantic Ocean, the plexaurid genus Thesea Duchassaing & Michelotti, 1860 has also been purported to be polyphyletic, with a Pacific lineage related to the former “Paramuriceidae” and an Atlantic lineage more closely related to Gorgoniidae. Thus, aiming to further improve our understanding of the evolutionary position of the Southwestern Atlantic Thesea, sequencing of the extended Octocorallia barcode reinforces the need for a re-evaluation of the position of the genus within plexaurids. Molecular and macro and micromorphological analyzes indicate the occurrence of an undescribed species distributed from Rio de Janeiro to Santa Catarina, here named Thesea pyrrha sp. nov. Results presented herein also suggest that T. pyrrha is close related to Adelogorgia and Psammogorgia, both genera exclusive to the Pacific Ocean.
References
Bayer, F.M. (1956) Octocorallia. In: Moore, R.C. (Ed.), Treatise on Invertebrate Paleontology Part F. Coelenterata. Geological Society of America and University of Kansas Press. Lawrence, Kansas, 498 pp.
Bayer, F.M. (1958) Les Octocoralliaires plexaurides des côtes occidentales d’Amerique. Mémoires du Muséum National D’Histoire Naturelle, Nouvelle Série, (A), 16 (2), 41–56, 6 pls.
Bayer, F.M. (1959) Octocorals from Surinam and the adjacent coast of South America. Studies on the Fauna of Suriname and other Guyanas, 3 (1), 1–43.
Bayer, F.M. (1961) The shallow-water Octocorallia of the West Indian region. Studies on the Fauna of Curacao and other Caribbean Islands, 12 (1), 1–373.
Bayer, F.M. (1981) Key to the genera of Octocorallia exclusive of Pennatulacea (Coelenterata: Anthozoa), with diagnosis of new taxa. Proceedings of the Biological Society of Washington, 94, 902–947.
Bayer, F.M., Grasshoff, M. & Verseveldt, J. (1983) Illustrated trilingual glossary of morphological terms applied to Octocorallia. E.J. Brill, Leiden, 74 pp.
Bayer, F.M. & Macintyre, I.G. (2001) The mineral component of the axis and holdfast of some gorgonacean octocorals (Coelenterata: Anthozoa), with special reference to the family Gorgoniidae. Proceedings of the Biological Society of Washington, 114 (1), 309–345.
Breedy, O. & Guzman, H.M. (2018a) A new alcyonacean species (Cnidaria: Anthozoa: Octocorallia) from a seamount in the tropical Pacific Ocean. Bulletin of Marine Science, 94 (4), 1515–1524. https://doi.org/10.5343/bms.2018.0027
Breedy, O. & Guzman, H.M. (2018b) Revision of the genus Adelogorgia Bayer, 1958 (Cnidaria: Anthozoa: Octocorallia) with the description of three new species. Zootaxa, 4369 (3), 327–348. https://doi.org/10.11646/zootaxa.4369.3.2
Breedy, O. & Guzman, H.M. (2020) A revision of the genus Psammogorgia Verrill, 1868 (Cnidaria, Anthozoa, Octocorallia) in the tropical eastern Pacific Ocean. ZooKeys, 961, 1–30. https://doi.org/10.3897/zookeys.961.54846
Breedy, O., Rouse, G.W., Stabbins, A., Cortés, J. & Cordes, E. (2019) New records of Swiftia (Cnidaria, Anthozoa, Octocorallia) from off the Pacific Costa Rican margin, including a new species from methane seeps. Zootaxa, 4671 (3), 407–419. https://doi.org/10.11646/zootaxa.4671.3.6
Castro, C.B., Pires, D.O., Medeiros, M.S., Loiola, L.L., Arantes, R.C.M., Thiago, C.M. & Berman, E. (2006) Filo Cnidaria. Corais. In: Lavrado, H.P. & Ignacio, B.L. (Eds.), Biodiversidade bentônica da região central da Zona Econômica Exclusiva brasileira. Série Livros. No. 18. Museu Nacional, Rio de Janeiro, pp. 147–192.
Cordeiro, R., McFadden, C., van Ofwegen, L. & Williams, G. (2021) World List of Octocorallia. Thesea Duchassaing & Michelotti, 1860. Accessed through: World Register of Marine Species at: https://www.marinespecies.org/aphia.php?p=taxdetails&id=125315 (accessed 23 August 2021)
Daly, M., Brugler, M.R., Cartwright, P., Collins, A.G., Dawson, M.N., France, S.C., McFadden, C.S., Opresko, D.M., Rodriguez, E., Romano, S. & Stake, J. (2007) The phylum Cnidaria: A review of phylogenetic patterns and diversity 300 years after Linnaeus. In: Zhang, Z.-Q. & Shear, W.A. (Eds.), Linnaeus Tercentenary: Progress in Invertebrate Taxonomy. Zootaxa, 1668, pp. 127–182. https://doi.org/10.11646/zootaxa.1668.1.11
Darriba, D., Taboada, G. L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature methods, 9 (8), 772. https://doi.org/10.1038/nmeth.2109
Deichmann, E. (1936) The Alcyonaria of the western part of the Atlantic Ocean. Memoirs of the Museum of Comparative Zoology at Harvard, 53, 1–317. https://doi.org/10.5962/bhl.title.49348
Duchassaing, P. & Michelotti, G. (1860) Mémoire sur les Coralliaires des Antilles. Memorie della Reale Accademia delle Scienze di Torino, Serie Seconda, 19, 56–87. https://doi.org/10.5962/bhl.title.11388
Ehrenberg, C.G. (1834) Beitrage zur physiologischen Kenntniss der Corallenthiere im allgemeinen, und besonders des rothen Meeres, nebst einem Versuche zur physiologischen Systematik derselben. Abhandlungen der Koniglich Preussische Akademie Wissenschaften zu Berlin. Aus dem Jahre 1832. Erster Theil, Berlin, 156 pp. [pp. 225–380]
France, S.C. & Hoover, L.L. (2002) DNA sequences of the mitochondrial COI gene have low levels of divergence among deep-sea octocorals (Cnidaria: Anthozoa). Hydrobiologia, 471 (1), 149–155. https://doi.org/10.1023/A:1016517724749
France, S.C., Rosel, P.E. & Ewann, J. (1996) DNA sequence variation of mitochondrial large-subunit rRNA. Mol. Mar. Biol. Biotechnology, 5 (1), 15–28.
Frometa, J., Etnoyer, P.J., Quattrini, A.M., Herrera, S. & Greig, T.W. (2021) Genetic divergence and polyphyly in the octocoral genus Swiftia [Cnidaria: Octocorallia], including a species impacted by the DWH oil spill. Diversity, 13 (4), 172. https://doi.org/10.3390/d13040172
Gray, J.E. (1859) On the arrangement of zoophytes with pinnated tentacles. Annals and Magazine of Natural History, 3(4), 439–444.
Gray, J.E. (1868) Descriptions of some new genera and species of alcyonoid corals in the British Museum. Annals and Magazine of Natural History, 2 (12), 441–445. https://doi.org/10.1080/00222936808695849
Haeckel, E. (1866) Generelle morphologie der Organismen, vol. 2. Verlag von Georg Reimer, Berlin.
Halàsz, A., Reynolds, A.M., McFadden, C.S., Toonen, R.J. & Benayahu, Y. (2015) Could polyp pulsation be the key to species boundaries in the genus Ovabunda (Octocorallia: Alcyonacea: Xeniidae)? Hydrobiologia, 759 (1), 95–107. https://doi.org/10.1007/s10750-014-2106-z
Herrera, S., Baco, A. & Sánchez, J.A. (2010) Molecular systematics of the bubblegum coral genera (Paragorgiidae, Octocorallia) and description of a new deep-sea species. Molecular phylogenetics and evolution, 55 (1), 123–135. https://doi.org/10.1016/j.ympev.2009.12.007
Hickson, S.J. (1928) The Gorgonacea of Panama Bay together with a description of one species from the Galapágos Islands and one of Trinidad. Videnskavelige Meddelelser fra den naturhistoriske Forening i Kovenhavn for Aarene, 85, 325–422.
Jaramillo, C., Montes, C., Cardona, A., Silvestro, D., Antonelli, A. & Bacon, C.D. (2017) Comment (1) on “Formation of the Isthmus of Panama” by O’Dea et al. Science Advances, 3 (6), e1602321. https://doi.org/10.1126/sciadv.1602321
Johnson, S.K. & Hallock, P. (2020) A review of symbiotic gorgonian research in the western Atlantic and Caribbean with recommendations for future work. Coral Reefs, 39 (2), 1–20. https://doi.org/10.1007/s00338-020-01891-0
Joseph, E.O., Carlo, J.M. & Lasker, H.R. (2015) Plasticity and conservatism in sclerites of a Caribbean octocoral. Hydrobiologia, 759 (1), 109–118. https://doi.org/10.1007/s10750-014-2153-5
Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular biology and evolution, 30 (4), 772–780. https://doi.org/10.1093/molbev/mst010
Kim, E., Lasker, H.R., Coffroth, M.A. & Kim, K. (2004) Morphological and genetic variation across reef habitats in a broadcast-spawning octocoral. Hydrobiologia, 530 (1), 423–432. https://doi.org/10.1007/s10750-004-2646-8
Kluge, A.G. (1989) A concern for evidence and a phylogenetic hypothesis of relationships among Epicrates (Boidae, Serpentes). Systematic Biology, 38 (1), 7–25. https://doi.org/10.1093/sysbio/38.1.7
Kluge, A.G. (1998) Total evidence or taxonomic congruence: cladistics or consensus classification. Cladistics, 14 (2), 151–158. https://doi.org/10.1111/j.1096-0031.1998.tb00328.x
Kükenthal, W. (1919) Gorgonaria Wissenschaftliche Ergebnisse der deutschen Tiefsee–Expedition auf dem Dampfer “Valdivia”, 1898–1899, 1–946.
Kükenthal, W. (1924) Gorgonaria. Das Tierreich. Vol. 47. Walter de Gruyter and Co., Berlin und Leipzig, 478 pp.
Kumar, J.S.Y., Geetha, S., Raghunathan, C. & Sornaraj, R. (2018) A note on shallow water Octocorallia from Nicobar Islands, India. Indian Journal of Geo-Marine Sciences, 47 (1), 60–66.
Lamouroux, J.V.F. (1812) Extrait d’un mémoire sur la classification des polypiers coralligènes non entièrement piérreux. Nouveau Bulletin des Sciences, Société Philomathique de Paris, 3 (63), 18l–188
Madeira, F., Park, Y.M., Lee, J., Buso, N., Gur, T., Madhusoodanan, N., Basutkar, P., Tivey, A.R., Potter, S.C. & Finn, R.D. (2019) The EMBL-EBI search and sequence analysis tools APIs in 2019. Nucleic acids research, 47 (W1), W636–W641. https://doi.org/10.1093/nar/gkz268
Matsumoto, A.K. & van Ofwegen, L.P. (2016) Species of Elasmogorgia and Euplexaura (Cnidaria, Octocorallia) from Japan with a discussion about the genus Filigella. ZooKeys, 589, 1–21. https://doi.org/10.3897/zookeys.589.8361
McFadden, C.S., France, S.C., Sánchez, J.A. & Alderslade, P. (2006) A molecular phylogenetic analysis of the Octocorallia (Cnidaria: Anthozoa) based on mitochondrial protein-coding sequences. Molecular phylogenetics and evolution, 41 (3), 513–527. https://doi.org/10.1016/j.ympev.2006.06.010
McFadden, C.S., Sánchez, J.A. & France, S.C. (2010) Molecular phylogenetic insights into the evolution of Octocorallia: a review. Integrative and Comparative Biology, 50 (3), 389–410. https://doi.org/10.1093/icb/icq056
McFadden, C.S., Benayahu, Y., Pante, E., Thoma, J.N., Nevarez, P.A. & France, S.C. (2011) Limitations of mitochondrial gene barcoding in Octocorallia. Molecular ecology resources, 11 (1), 19–31. https://doi.org/10.1111/j.1755-0998.2010.02875.x
McFadden, C.S. & van Ofwegen, L.P. (2013) A second, cryptic species of the soft coral genus Incrustatus (Anthozoa: Octocorallia: Clavulariidae) from Tierra del Fuego, Argentina, revealed by DNA barcoding. Helgoland Marine Research, 67 (1), 137–147. https://doi.org/10.1007/s10152-012-0310-7
McFadden, C.S., Brown, A.S., Brayton, C., Hunt, C.B. & Van Ofwegen, L.P. (2014) Application of DNA barcoding in biodiversity studies of shallow-water octocorals: molecular proxies agree with morphological estimates of species richness in Palau. Coral Reefs, 33 (2), 275–286. https://doi.org/10.1007/s00338-013-1123-0
Nutting, C.C. (1910) The Gorgonacea of the Siboga Expedition III. The Muriceidae. Siboga-Expedition Monograph, 13b, 1–108, pls. 1–22.
O’Dea, A., Lessios, H.A., Coates, A.G., Eytan, R.I., Restrepo-Moreno, S.A., Cione, A.L., Collins, L.S., De Queiroz, A., Farris, D.W. & Norris, R.D. (2016) Formation of the Isthmus of Panama. Science advances 2, 1–11. https://doi.org/10.1126/sciadv.1600883
Pérez, C.D., de Moura Neves, B., Cordeiro, R.T., Williams, G.C. & Cairns, S.D. (2016) Diversity and distribution of Octocorallia. In: The Cnidaria, Past, Present and Future. Springer, Cham, pp. 109–123. https://doi.org/10.1007/978-3-319-31305-4_8
Rambaut, A. (2014) FigTree. Version 1.4.3. Institute of Evolutionary Biology, University of Edinburgh, Edinburgh. Available from: https://tree.bio.ed.ac.uk/software/figtree/ (accessed 22 February 2022)
Riess, M. (1929) Die Gorgonarien Westindies. Kap. 8. Die Familie Muriceidae. Zoologische Jahrbücher, Jena, 16, 377–420.
Roberts, J.M., Wheeler, A.J. & Freiwald, A. (2006) Reefs of the deep: the biology and geology of cold-water coral ecosystems. Science, 312 (5773), 543–547. https://doi.org/10.1126/science.1119861
Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19 (12), 1572–1574. https://doi.org/10.1093/bioinformatics/btg180
Sánchez, J., McFadden, C., France, S. & Lasker, H. (2003) Molecular phylogenetic analyzes of shallowwater Caribbean octocorals. Marine Biology, 142 (5), 975–987. https://doi.org/10.1007/s00227-003-1018-7
Sánchez, J.A. & Wirshing, H.H. (2005) A field key to the Identification of Tropical Western Atlantic Zooxanthellate Octocorals (Octocorallia: Cnidaria). Caribbean Journal of Science, 41 (3), 508–522.
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
Tixier-Durivault, A. (1970) Octocoralliaires. Campagne de la “Calypso” au large des côtes atlantiques de l’Amérique du Sud (1961–1962). Annales de l’Institut Océanographique, 47, 145–169.
Vargas, S., Eitel, M., Breedy, O. & Schierwater, B. (2010) Molecules match morphology: mitochondrial DNA supports Bayer's Lytreia-Bebryce-Heterogorgia (Alcyonacea: Octocorallia) clade hypothesis. Invertebrate Systematics, 24 (1), 23–31. https://doi.org/10.1071/IS09033
Verrill, A.E. (1868) Notes on the Radiata in the Museum of Yale College, with descriptions of new genera and species. 4. Notice of corals echinoderms collected by Prof. C.F. Hartt, at the Abrolhos Reefs, Province of Bahia, Brazil, 1867. Transactions of the Connecticut Academy of Arts and Sciences, 1, 351–371.
Verrill, A.E. (1869) Critical remarks on the halcyonoid polyps with descriptions of new species in the Museum of Yale College, No. 4. American Journal of Science and Arts, 2 (48), 419–429. https://doi.org/10.2475/ajs.s2-48.143.244
Verrill, A.E. (1883) Report on the Anthozoa, and on some additional species dredged by the “Blake” in 1877-1879, and by the U.S. Fish Commission steamer “Fish Hawk” in 1880-82. Bulletin of the Museum of Comparative Zoology at Harvard, 11, 1–72.
Villesen, P. (2007) FaBox: an online toolbox for fasta sequences. Molecular Ecology Notes, 7 (6), 965–968. https://doi.org/10.1111/j.1471-8286.2007.01821.x
Waterhouse, A.M., Procter, J.B., Martin, D.M., Clamp, M. & Barton, G.J. (2009) Jalview Version 2—a multiple sequence alignment editor and analysis workbench. Bioinformatics, 25 (9), 1189–1191. https://doi.org/10.1093/bioinformatics/btp033
West, J.M., Harvell, C.D. & Wall, A.-M. (1993) Morphological plasticity in a gorgonian coral (Briareum asbestinum) over a depth cline. Marine Ecology Progress Series, 4, 61–69. https://doi.org/10.3354/meps094061
Wirshing, H., Messing, C.G., Douady, C., Reed, J., Stanhope, M. & Shivji, M.S. (2005) Molecular evidence for multiple lineages in the gorgonian family Plexauridae (Anthozoa: Octocorallia). Marine Biology, 147 (2), 497–508. https://doi.org/10.1007/s00227-005-1592-y