Abstract
Caymanostellidae is a group of rarely collected and morphologically unusual sea stars that have been exclusively encountered on wood falls in the deep sea. There are currently three genera and seven species described, occurring in the Atlantic, Pacific and Indian Oceans with a depth range between 418 and 6780 m. Three new species are here described from specimens collected from wood falls in multiple localities across the Pacific margin of Costa Rica and near the Gulf of California (Mexico): Caymanostella scrippscognaticausa sp. nov., Caymanostella davidalani sp. nov. and Caymanostella loresae sp. nov. These records expand the known geographical distribution of caymanostellids and constitute their first report from wood falls found at methane seeps. This study also includes the first descriptions of early-stage juvenile caymanostellids and reveals that traits previously considered useful for diagnosis might represent intraspecific and ontogenetic variability, with important consequences for caymanostellid taxonomy.
References
- Altschul, S.F., Gish, W., Miller, W., Myers, E.W. & Lipman, D.J. (1990) Basic local alignment search tool. Journal of Molecular Biology, 215, 403–410. https://doi.org/10.1016/S0022-2836(05)80360-2
- Alvarado, J.J., Chacón-Monge, J.L., Azofeifa-Solano, J.C. & Cortés, J. (2022) Diversity of deep-sea echinoderms from Costa Rica. Frontiers in Marine Science, 9, 918878. https://doi.org/10.3389/fmars.2022.918878
- Arndt, A., Marquez, C., Lambert, P. & Smith, M.J. (1996) Molecular phylogeny of eastern Pacific sea cucumbers (Echinodermata: Holothuroidea) based on mitochondrial DNA sequence. Molecular Phylogenetics and Evolution, 6, 425–437. https://doi.org/10.1006/mpev.1996.0091
- Aziz, A. & Jangoux, M. (1984) Description de quatre nouvelles espèces d’astérides profonds (Echinodermata) de la région Indo-Malaise. Indo-Malayan Zoology, 1, 187–194.
- Baker, A.N., Rowe, F.W.E. & Clark, H.E.S. (1986) A new class of Echinodermata from New Zealand. Nature, 321, 862–864. https://doi.org/10.1038/321862a0
- Barroso, R., Kudenov, J.D., Halanych, K.M., Saeedi, H., Sumida, P.Y.G. & Bernardino, A.F. (2018) A new species of xylophilic fireworm (Annelida: Amphinomidae: Cryptonome) from deep-sea wood falls in the SW Atlantic. Deep Sea Research Part I: Oceanographic Research Papers, 137, 66–75. https://doi.org/10.1016/j.dsr.2018.05.005
- Becker, P., Samadi, S., Zbinden, M., Hoyoux, C., Compère, P. & De Ridder, C. (2009) First insights into the gut microflora associated with an echinoid from wood falls environments. Cahiers de Biologie Marine, 50, 343–352.
- Belyaev, G.M. (1974) A new family of abyssal starfishes. Zoologischeskii Zhurnal, 53, 1502–1508.
- Belyaev, G.M. (1990) Is it valid to isolate the genus Xyloplax as an independent class of echinoderms? Zoologicheskii Zhurnal, 69, 83–96.
- Bernardino, A.F., Levin, L.A., Thurber, A.R. & Smith, C.R. (2012) Comparative composition, diversity and trophic ecology of sediment macrofauna at vents, seeps and organic falls. PLoS ONE, 7, e33515. https://doi.org/10.1371/journal.pone.0033515
- Bienhold, C., Pop Ristova, P., Wenzhöfer, F., Dittmar, T. & Boetius, A. (2013) How deep-sea wood falls sustain chemosynthetic life. PLoS ONE, 8, e53590. https://doi.org/10.1371/journal.pone.0053590
- Blake, D.B. (1987) A classification and phylogeny of post-Palaeozoic sea stars (Asteroidea: Echinodermata). Journal of Natural History, 21, 481–528. https://doi.org/10.1080/00222938700771141
- Brett, C.H. (2017) Evolutionary biology of wood-eating sea urchins (Temnopleuridea: Trigonocidaridae). Master of Science Thesis, San Francisco State University. Available from: http://hdl.handle.net/10211.3/199487 (accessed 7 October 2024)
- Clement, M., Posada, D. & Crandall, K.A. (2000) TCS: a computer program to estimate gene genealogies. Molecular Ecology, 9, 1657–1659. https://doi.org/10.1046/j.1365-294x.2000.01020.x
- Colgan, D.J., Ponder, W.F., Beacham, E., & Macaranas, J. (2003) Gastropod phylogeny based on six segments from four genes representing coding or non-coding and mitochondrial or nuclear DNA. Molluscan Research, 23, 123–148. https://doi.org/10.1071/MR03002
- Darriba, D., Posada, D., Kozlov, A.M., Stamatakis, A., Morel, B. & Flouri, T. (2019) ModelTest-NG: A new and scalable tool for the selection of DNA and protein evolutionary models. Molecular Biology and Evolution, 37, 291–294. https://doi.org/10.1093/molbev/msz189
- Dilman, A.B., Minin, K.V. & Petrov, N.B. (2022) New record of the wood-associated sea star Caymanostella, with notes on the phylogenetic position of the family Caymanostellidae (Asteroidea). Zoological Journal of the Linnean Society, 194, 14–35. https://doi.org/10.1093/zoolinnean/zlab060
- Distel, D.L., Baco, A.R., Chuang, E., Morrill, W., Cavanaugh, C. & Smith, C.R. (2000) Do mussels take wooden steps to deep-sea vents? Nature, 403, 725–726. https://doi.org/10.1038/35001667
- Duperron, S., Lorion, J., Samadi, S., Gros, O. & Gaill, F. (2009) Symbioses between deep-sea mussels (Mytilidae: Bathymodiolinae) and chemosynthetic bacteria: diversity, function and evolution. Comptes Rendus Biologies, 332, 298–310. https://doi.org/10.1016/j.crvi.2008.08.003
- Edler, D., Klein, J., Antonelli, A. & Silvestro, D. (2021) raxmlGUI 2.0: A graphical interface and toolkit for phylogenetic analyses using RAxML. Methods in Ecology and Evolution, 12, 373–377. https://doi.org/10.1111/2041-210X.13512.
- Fujita, T., Stampanato, S. & Jangoux, M. (1994) Beyaevostella hyugaenis: a new species of deep-sea asteroid (Asteroidea, Caymanostellidae) found on a sunken wood from off southern Japan. Bulletin of the National Science Museum, Tokyo, 20, 183–188.
- Gale, A.S. (2011) The phylogeny of post-palaeozoic Asteroidea (Neoasteroidea, Echinodermata). Special Papers in Palaeontology, 85, 1–112.
- Hatch, A.S., Liew, H., Hourdez, S. & Rouse, G.W. (2020) Hungry scale worms: Phylogenetics of Peinaleopolynoe (Polynoidae, Annelida), with four new species. ZooKeys, 932, 27–74. https://doi.org/10.3897/zookeys.932.48532
- Hookabe, N., Moritaki, T., Jimi, N. & Ueshima, R. (2022) A new oerstediid discovered from wood falls in the Sea of Kumano, Japan: Description of Rhombonemertes rublinea gen. et sp. nov. (Nemertea: Eumonostilifera). Zoologischer Anzeiger, 301, 154–162. https://doi.org/10.1016/j.jcz.2022.10.003
- Janies, D.A., Voight, J.R. & Daly, M. (2011) Echinoderm phylogeny including Xyloplax, a progenetic asteroid. Systematic Biology, 60, 420–438. https://doi.org/10.1093/sysbio/syr044
- 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
- Kozlov, A.M., Darriba, D., Flouri, T., Morel, B. & Stamatakis, A. (2019) RAxML-NG: a fast, scalable and user-friendly tool for maximum likelihood phylogenetic inference. Bioinformatics, 35, 4453–4455. https://doi.org/10.1093/bioinformatics/btz305
- Leigh, J.W. & Bryant, D. (2015) PopART: Full‐feature software for haplotype network construction. Methods in Ecology and Evolution/British Ecological Society, 6, 1110–1116. https://doi.org/10.1111/2041-210x.12410
- Limaye, A. (2012) Drishti: a volume exploration and presentation tool. In: Developments in X-Ray Tomography VIII. SPIE, Bellingham, Washington, pp. 191–199. https://doi.org/10.1117/12.935640
- Linchangco, G.V., Foltz, D.W., Reid, R., Williams, J., Nodzak, C., Kerr, A.M., Miller, A.K., Hunter, R., Wilson, N.G., Nielsen, W.J., Mah, C.L., Rouse, G.W., Wray, G.A. & Janies, D.A. (2017) The phylogeny of extant starfish (Asteroidea: Echinodermata) including Xyloplax, based on comparative transcriptomics. Molecular Phylogenetics and Evolution, 115, 161–170. https://doi.org/10.1016/j.ympev.2017.07.022
- Maddison, W.P. & Maddison, D.R. (2023) Mesquite: a modular system for evolutionary analysis. Version 3.70. Available from: http://www.mesquiteproject.org (accessed 27 June 2023)
- Magalhães, W.F., Linse, K. & Wiklund, H. (2017) A new species of Raricirrus (Annelida: Cirratuliformia) from deep-water sunken wood off California. Zootaxa, 4353 (1), 51–68. https://doi.org/10.11646/zootaxa.4353.1.3
- Magalhães, W.F. & Hilliard, J. (2022) Two new deep-sea species of Capitella (Annelida: Capitellidae) from sunken wood in the Northeast Pacific. Zootaxa, 5125 (2), 229–240. https://doi.org/10.11646/zootaxa.5125.2.7
- Mah, C.L. (2006) A new species of Xyloplax (Echinodermata: Asteroidea: Concentricycloidea) from the northeast Pacific: comparative morphology and a reassessment of phylogeny. Invertebrate Biology, 125, 136–153. https://doi.org/10.1111/j.1744-7410.2006.00048.x
- Mah, C.L. & Blake, D.B. (2012) Global diversity and phylogeny of the Asteroidea (Echinodermata). PLoS ONE, 7, e35644. https://doi.org/10.1371/journal.pone.0035644
- Martin-Cao-Romero, C., Solís-Marín, F.A. & Bribiesca-Contreras, G. (2021) Crinitostella laguardai, new genus and species of wood-dwelling deep-sea sea-star (Asteroidea: Caymanostellidae) from the Gulf of Mexico. Journal of the Marine Biological Association of the United Kingdom, 101, 591–597. https://doi.org/10.1017/S0025315421000448
- Okanishi, M., Kato, M., Watanabe, H.K., Chen, C. & Fujita, T. (2020) Large populations of two new species of Ophiambix (Echinodermata, Ophiuroidea) discovered on Japanese hot vents and cold seeps. Raffles Bulletin of Zoology, 68, 196–213. https://doi.org/10.26107/RBZ-2020-0017
- Palumbi, S.R. (1996) Nucleic acids II: the polymerase chain reaction. In: Hillis, D.M., Moritz, G. & Mable, B.K. (Eds.), Molecular Systematics. 2nd Edition. Sinauer, Sunderland, Massachusetts, pp. 205–247.
- Payne, C.Y., Tilic, E., Boschen-Rose, R.E., Gannon, A., Stiller, J., Hiley, A.S., Grupe, B.M., Mah, C.L. & Rouse, G.W. (2023) Xyloplax princealberti (Asteroidea, Echinodermata): A new species that is not always associated with wood falls. Diversity, 15, 1212. https://doi.org/10.3390/d15121212
- Pereira, O.S., Gonzalez, J., Mendoza, G., Le, J., McNeill, M., Ontiveros, J., Lee, R.W., Rouse, G.W., Cortés, J. & Levin, L.A. (2022) Does substrate matter in the deep sea? A comparison of bone, wood, and carbonate rock colonizers. PLoS ONE, 17, e0271635. https://doi.org/10.1371/journal.pone.0271635
- Pop Ristova, P., Bienhold, C., Wenzhöfer, F., Rossel, P.E. & Boetius, A. (2017) Temporal and spatial variations of bacterial and faunal communities associated with deep-sea wood falls. PLoS ONE, 12, e0169906. https://doi.org/10.1371/journal.pone.0169906
- Romano, C., Nunes-Jorge, A., Le Bris, N., Rouse, G.W., Martin, D. & Borowski, C. (2020) Wooden stepping stones: Diversity and biogeography of deep-sea wood boring Xylophagaidae (Mollusca: Bivalvia) in the North-East Atlantic Ocean, with the description of a new genus. Frontiers in Marine Science, 7. https://doi.org/10.3389/fmars.2020.579959
- Rowe, F.W.E. (1988) Review of the extant class Concentricycloidea and reinterpretation of the fossil class Cyclocystoidea. In: Burke, R.D., Mladenov, P.V., Lambert, P. & Parsley, R.L. (Eds.), Echinoderm Biology. Balkema, Rotterdam, pp. 3–15.
- Rowe, F.W.E. (1989) A review of the family Caymanostellidae (Echinodermata: Asteroidea) with the description of a new species of Caymanostella Belyaev and a new genus. Proceedings of the Linnean Society of New South Wales. Linnean Society of New South Wales, 111, 293–307.
- Saeedi, H., Bernardino, A.F., Shimabukuro, M., Falchetto, G. & Sumida, P.Y.G. (2019) Macrofaunal community structure and biodiversity patterns based on a wood-fall experiment in the deep South-west Atlantic. Deep Sea Research, Part I: Oceanographic Research Papers, 145, 73–82. https://doi.org/10.1016/j.dsr.2019.01.008
- Sagorny, C., von Döhren, J., Rouse, G.W. & Tilic, E. (2022) Cutting the ribbon: bathyal Nemertea from seeps along the Costa Rica margin, with descriptions of 2 new genera and 9 new species. European Journal of Taxonomy, 845, 132–174. https://doi.org/10.5852/ejt.2022.845.1959
- Smith, A.B. (1988) To group or not to group: the taxonomic position of Xyloplax. In: Burke, R.D., Mladenov, P.V., Lambert, P. & Parsley, R.L. (Eds.), Echinoderm Biology: Proceedings of the Sixth International Echinoderm Conference. Balkema, Rotterdam, pp. 17–23.
- Smith, C.R., Glover, A.G., Treude, T., Higgs, N.D. & Amon, D.J. (2015) Whale-fall ecosystems: recent insights into ecology, paleoecology, and evolution. Annual review of marine science 7, 571–596. https://doi.org/10.1146/annurev-marine-010213-135144
- Swofford, D.L. (2002) PAUP* Phylogenetic Analysis Using Parsimony (and other methods). Available from: https://paup.phylosolutions.com (accessed 3 May 2024)
- Tandberg, A.H.S., Rapp, H.T., Schander, C. & Vader, W. (2013) A new species of Exitomelita (Amphipoda: Melitidae) from a deep-water wood fall in the northern Norwegian Sea. Journal of natural history, 47, 1875–1889. https://doi.org/10.1080/00222933.2012.725778
- Vaidya, G., Lohman, D.J. & Meier, R. (2011) SequenceMatrix: concatenation software for the fast assembly of multi-gene datasets with character set and codon information. Cladistics: the international journal of the Willi Hennig Society, 27, 171–180. https://doi.org/10.1111/j.1096-0031.2010.00329.x.
- Voight, J.R. (2015) Xylotrophic bivalves: aspects of their biology and the impacts of humans. The Journal of molluscan studies, 81, 175–186. https://doi.org/10.1093/mollus/eyv008
- Voight, J.R., Marshall, B.A., Judge, J., Halanych, K.M., Li, Y., Bernardino, A.F., Grewe, F. & Maddox, J.D. (2019) Life in wood: preliminary phylogeny of deep-sea wood-boring bivalves (Xylophagaidae), with descriptions of three new genera and one new species. The Journal of molluscan studies, 85, 232–243. https://doi.org/10.1093/mollus/eyz003
- Wolff, T. (1979) Macrofaunal utilization of plant remains in the deep sea. Sarsia, 64, 117–143. https://doi.org/10.1080/00364827.1979.10411373