Abstract
The systematics of bamboo corals of the Family Keratoisididae are evaluated using both DNA sequences and morphological data. Sequence data were obtained from 398 specimens, from which 77 unique haplotypes representing the mtMutS and 18S gene regions were identified. These were aligned with sequences downloaded from GenBank from an additional 12 keratoisids and 6 octocoral outgroups. Phylogenetic analyses recovered seven well-supported major clades, the most recently derived of which consists of several subclades. Each clade and subclade can be characterized by a suite of morphological characters that include axis construction, branching pattern, polyp form, and sclerite type and arrangement. This analysis also shows that keratoisid genera described >100 years ago are paraphyletic and need revision and that a large number of new genera will need to be described.
References
Alderslade, P. & McFadden, C.S. (2012) A new genus and species of the family Isididae (Coelenterata: Octocorallia) from a CMAR Biodiversity study, and a discussion on the subfamilial placement of some nominal isidid genera. Zootaxa, 3154 (1), 21–39. https://doi.org/10.11646/zootaxa.3154.1.2
Apakupakul, K., Siddall, M. E. & Burreson, E.M. (1999) Higher level relationships of leeches (Annelida: Clitellata: Euhirudinea) based on morphology and gene sequences. Molecular phylogenetics and evolution, 12, 350–359. https://doi.org/10.1006/mpev.1999.0639
Bayer, F.M. (1990) A new isidid octocoral (Anthozoa: Gorgonacea) from New Caledonia, with descriptions of other new species from elsewhere in the Pacific Ocean. Proceedings of the Biological Society of Washington, 103, 205–228.
Bayer, F.M., Grasshoff, . & Verseveldt, J. (1983) Illustrated Trilingual Glossary. E.J. Brill, Leiden, 75 pp.
Berntson, E.A., Bayer, F.M., McArthur, A.G. & France, S.C. (2001) Phylogenetic relationships within the Octocorallia (Cnidaria: Anthozoa) based on nuclear 18S rRNA sequences. Marine Biology, 138, 235–246. https://doi.org/10.1007/s002270000457
Berntson, E.A. & France, S.C. (2001) Generating DNA sequence information from museum collections of octocoral specimens (Phylum Cnidaria: Class Anthozoa). Bulletin of the Biological Society of Washington, 10, 119–129.
Bilewitch, J.P. & Degnan, S.M. (2011) A unique horizontal gene transfer event has provided the octocoral mitochondrial genome with an active mismatch repair gene that has potential for an unusual self-contained function. BMC Evolutionary Biology, 11, 228. https://doi.org/10.1186/1471-2148-11-228
Brockman, S.A. & McFadden, C.S. (2012) The mitochondrial genome of Paraminabea aldersladei (Cnidaria: Anthozoa: Octocorallia) supports intramolecular recombination as the primary mechanism of gene rearrangement in octocoral mitochondrial genomes. Genome Biology and Evolution, 4, 994–1006. https://doi.org/10.1093/gbe/evs074
Brugler, M.R. & France, S.C. (2008) The mitochondrial genome of a deep-sea bamboo coral (Cnidaria, Anthozoa, Octocorallia, Isididae): genome structure and putative origins of replication are not conserved among octocorals. Journal of Molecular Evolution, 67, 125–36. https://doi.org/10.1007/s00239-008-9116-2
Cordeiro, R., McFadden, C., van Ofwegen, L., & Williams, G. (2021). World List of Octocorallia. Keratoisididae Gray, 1870. Accessed through: World Register of Marine Species. Available from: http://www.marinespecies.org/aphia.php?p=taxdetails&id=1544106 (accessed 11 January 2022)
Dueñas, L. & Sánchez, J. (2009) Character lability in deep-sea bamboo corals (Octocorallia, Isididae, Keratoisidinae). Marine Ecology Progress Series, 397, 11–23. https://doi.org/10.3354/meps08307
Dueñas, L.F., Alderslade, P. & Sánchez, J.A. (2014) Molecular systematics of the deep-sea bamboo corals (Octocorallia: Isididae: Keratoisidinae) from New Zealand with descriptions of two new species of Keratoisis. Molecular Phylogenetics and Evolution, 74, 15–28. https://doi.org/10.1016/j.ympev.2014.01.031
Farris, J.S., Kallersjö, M., Kluge, A.G. & Bult, C. (1994) Testing significance of incongruence. Cladistics, 10, 315–319. https://doi.org/10.1111/j.1096-0031.1994.tb00181.x
France, S.C. (2007) Genetic analysis of bamboo corals (Cnidaria: Octocorallia: Isididae): Does lack of colony branching distinguish Lepidisis from Keratoisis? Bulletin of Marine Science, 81, 323–333.
France, S.C. & Hoover, L.L. (2001) Analysis of variation in mitochondrial DNA sequences (ND3, ND4L, MSH) among Octocorallia (=Alcyonaria) (Cnidaria: Anthozoa). Bulletin of the Biological Society of Washington, 10, 110–118.
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, 149–155. https://doi.org/10.1023/A:1016517724749
Grant, R. (1976) The marine fauna of New Zealand: Isididae (Octocorallia: Gorgonacea) from New Zealand and the Antarctic. New Zealand Oceangraphic Institute Memoir, 66,1–54.
van der Ham, J.L., Brugler, M.R. & France, S.C. (2009) Exploring the utility of an indel-rich mitochondrial region as a molecular barcode for bamboo corals (Octocorallia:Isididae). Marine Genomics, 2, 183–192. https://doi.org/10.1016/j.margen.2009.10.002
Heestand Saucier, E., France, S.C. & Watling, L. (2021) Toward a revision of the bamboo corals: Part 3, deconstructing the Family Isididae. Zootaxa, 5047 (3), 247–272. https://doi.org/10.11646/zootaxa.5047.3.2
Heestand Saucier, E., Sajjadi, A. & France, S.C. (2017) A taxonomic review of the genus Acanella (Cnidaria: Octocorallia: Isididae) in the North Atlantic Ocean, with descriptions of two new species. Zootaxa, 4323 (3), 359–390. https://doi.org/10.11646/zootaxa.4323.3.2
Hellberg, M.E. (2007) Footprints on water: the genetic wake of dispersal among reefs. Coral Reefs, 26, 463–473. https://doi.org/10.1007/s00338-007-0205-2
Hogan, R.I., Hopkins, K., Wheeler, A.J., Allcock, A.L. & Yesson, C. (2019) Novel diversity in mitochondrial genomes of deep-sea Pennatulacea (Cnidaria: Anthozoa: Octocorallia). Mitochondrial DNA Part A, 30, 764–777. https://doi.org/10.1080/24701394.2019.1634699
Huelsenbeck, J.P. & Ronquist, F. (2001) MrBayes: Bayesian inference of phylogeny. Bioinformatics, 17, 754–755. https://doi.org/10.1093/bioinformatics/17.8.754
Katoh, K., Misawa, K., Kuma, K. & Miyata, T. (2002) MAFFT: a novel method for rapid multiple sequence alignment based on Fast Fourier transform. Nucleic Acids Research, 30, 3059–3066. https://doi.org/10.1093/nar/gkf436
Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution, 30, 772–780. https://doi.org/10.1093/molbev/mst010
Lapointe, A. & Watling, L. (2015) Bamboo corals from the abyssal Pacific: Bathygorgia. Proceedings of the Biological Society of Washington, 128, 125–136. https://doi.org/10.2988/0006-324X-128.2.125
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, 19–31. https://doi.org/10.1111/j.1755-0998.2010.02875.x
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, 513–27. 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, 389–410. https://doi.org/10.1093/icb/icq056
Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In: Proceedings of the Gateway Computing Environments Workshop. New Orleans, Louisiana, pp. 1–8. https://doi.org/10.1109/GCE.2010.5676129
Muzik, K. (1978) A bioluminescent gorgonian, Lepidisis olapa, new species (Coelenterata: Octocorallia), from Hawaii. Bulletin of Marine Science, 28, 735–741.
Pante, E., France, S. C., Couloux, A., Cruaud, C., McFadden, C. S., Samadi, S., & Watling, L. (2012) Deep-sea origins and in-situ diversification of chrysogorgiid octocorals. PLoS ONE, 7, e38357. https://doi.org/10.1371/journal.pone.0038357
Pante, E., Heestand Saucier, E., & France, S. C. (2013) Molecular and morphological data support reclassification of the octocoral genus Isidoides. Invertebrate Systematics, 27, 365–378. https://doi.org/10.1071/IS12053
Pante, E., Abdelkrim, J., Viricel, A., Gey, D., France, S.C., Boisselier, M. & Samadi, S. (2015) Use of RAD sequencing for delimiting species. Heredity, 114, 450–459. https://doi.org/10.1038/hdy.2014.105
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
Smith, P.J., McVeagh, S.M., Mingoia, J.T. & France, S.C. (2004) Mitochondrial DNA sequence variation in deep-sea bamboo coral (Keratoisidinae) species in the southwest and northwest Pacific Ocean. Marine Biology, 144, 253–261. https://doi.org/10.1007/s00227-003-1206-5
Stamatakis, A. (2006) RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22, 2688–2690. https://doi.org/10.1093/bioinformatics/btl446
Stamatakis, A., Hoover, P. & Rougemont, J. (2008) A fast bootstrapping algorithm for the RAxML web-servers. Systematic Biology, 57, 758–771. https://doi.org/10.1080/10635150802429642
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, 11, 1–72.
Watling, L. (2020) The revision of the bamboo corals: The Subfamily Muricellisdinae (Octocorallia: Isididae). Zootaxa, 4881 (2), 361–371. https://doi.org/10.11646/zootaxa.4881.2.9
Watling, L. & France, S.C. (2011) A new genus and species of bamboo coral (Octocorallia: Isididae: Keratoisidinae) from the New England seamounts. Bulletin of the Peabody Museum of Natural History, 52, 209–220. https://doi.org/10.3374/014.052.0202
Watling, L. & France, S.C. (2021) Towards a revision of the bamboo corals: Part 2, untangling the genus Lepidisis (Octocorallia: Isididae). Bulletin of the Peabody Museum of Natural History, 62, 97–110. https://doi.org/10.3374/014.062.0202