Research Article |
Corresponding author: Odalisca Breedy ( odaliscab@gmail.com ) Academic editor: Bert W. Hoeksema
© 2021 Odalisca Breedy, Leen van Ofwegen, Catherine S. McFadden, Catalina Murillo-Cruz.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Breedy O, van Ofwegen L, McFadden CS, Murillo-Cruz C (2021) Rhodolitica on rhodoliths: a new stoloniferan genus (Anthozoa, Octocorallia, Alcyonacea). ZooKeys 1032: 63-77. https://doi.org/10.3897/zookeys.1032.63431
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Rhodolitica occulta gen. nov. et sp. nov. (Clavulariidae) is described from Cocos Island National Park, Pacific Ocean, Costa Rica. The species was found at various islets and rocky outcrops around the island, 20−55 m in depth. The genus is characterised by tubular, single, erect anthosteles interconnected by thin basal ribbon-like stolons on the surfaces of living rhodoliths. The anthosteles are devoid of fused sclerites, which are only present in the stolons. Coenenchymal sclerites are mostly spindles of various shapes, with a characteristic cylindrical warty type in the outer layer, crosses and radiates. Anthocodiae are armed with points, lacking collarets. Colonies and sclerites are red. Using an integrative taxonomic approach, we separate the new genus from similar genera through both morphological comparison and a molecular phylogenetic analysis. This research is a contribution to the knowledge of the octocoral biodiversity in Cocos Island and marine biodiversity in the eastern tropical Pacific.
Biodiversity, Cocos Island, new species, oceanic island, soft corals, taxonomy
Cocos Island, Costa Rica, an oceanic eastern tropical Pacific (ETP) island, has been considered a biodiversity and endemism hot-spot for marine organisms (
In 2007, during a CIMAR expedition aboard the R/V Proteus, while dredging at depths of 40–45 m in Chatham Bay, scientists collected rhodoliths with a red stoloniferous octocoral. Rhodoliths are found at several points around Cocos Island (
The Stolonifera include a group of octocorals that consist of individual tubular polyps that arise separately from ribbon-like stolons and present a series of transitional forms, from solitary separated polyps to united polyps joined at their bases in a common extended coenenchyme (
The specimens were collected by scuba diving down to 30 m depth, by dredging, and by the submersible HOV ‘DeepSee’ of the M/V ‘Argo’ to 55 m depth. Cocos Island National Park is an oceanic island located between 5°30'–5°34'N and 87°01'–87°06'W in the eastern Tropical Pacific (ETP) approximately 500 km southwest of Costa Rica and more than 600 km northeast of the Galápagos Islands, Ecuador (
Preserved specimens were morphologically analysed and photographed. For taxonomic identification, sclerites from different parts of the colonies (polyp, coenenchyme, base and stolons) were obtained by dissolving the tissue in 5% sodium hypochlorite; dissociated sclerites were washed several times in distilled water until organic matter was completely removed, dehydrated with 100% ethanol, and subsequently dried in an oven. Sclerites were prepared for light microscopy, mounted in glycerine, and photographed with an Olympus LX 51 inverted microscope. For scanning electron microscopy (SEM), sclerites were mounted on SEM stubs by double stick carbon tape and silver paint, then sputter-coated with gold, 30–60 nm layer, in an Eiko IB-5 Ion Coater; the images were obtained using a Hitachi SEM S-3700N (at 15kV). All specimens were preserved with the anthocodiae partially or totally retracted therefore the description is based on polyp dissection. Measurements of the sclerites were obtained from the SEM images. The holotype and paratypes are deposited at the Zoology Museum, University of Costa Rica, Costa Rica (MZUCR).
DNA was extracted from ethanol-preserved tissues with the NucleoSpin® Tissue kit (Macherey-Nagel, Germany) according to the manufacturer’s instructions, and kept at −20 °C until further processing. A partial region of the mitochondrial mismatch repair gene (mtMutS) was amplified with ND42599F (5'-GCCATTATGGTTAACTATTAC-3';
Sequences obtained from MZUCR 2513 were aligned with reference sequences from a wide range of octocoral taxa (Suppl. material
Subclass Octocorallia Haeckel, 1866
Order Alcyonacea Lamouroux, 1812
Family Clavulariidae Hickson, 1894
Colonies composed of tubular, single, erect anthosteles up to 8 mm tall and closely spaced, 1–3 mm apart forming interconnected groups of 2–15 anthosteles. Anthosteles arise directly from thin basal ribbon-like stolons that encrust irregular surfaces of living rhodoliths. Stolons composed of conspicuous red brittle fused sclerites not extending into anthostelar walls. Bright red coenenchymal sclerites compose the anthostelar walls, including long warty spindles, up to 0.27 mm in length; complexly warted cylindrical spindles up to 0.32 mm long; bent spindles and smaller crosses and radiates. Polyps whitish to transparent. Anthocodiae retractile, armed with eight interseptal points of slender warty spindles, up to 0.24 mm long, and small orange biscuit-like rods along the tentacles. Collaret absent. Collected specimens were preserved with the anthocodiae partially or totally retracted therefore a full description of polyps was not possible.
Colour of colonies bright red. Azooxanthellate.
Rhodolitica occulta spec. nov. by original designation.
The generic name is in reference to the substratum to which the colonies were attached: rhodolith rocks.
Holotype. MZUCR 2514, lot 1, ethanol preserved, Cocos Island, Chatham Bay, 05°33.347'N, 87°02.336'W, dredging, 45 m depth, J. Cortés, J. Sibaja-Cordero, Proteus-CIMAR Expedition, R/V Proteus, 13 January 2007. Paratypes. MZUCR 2514, lots 2, 3, ethanol preserved, same geographical data as the holotype. MZUCR 2734, lot ethanol preserved, Roca Chatham Bay, Langosta, Punta Ulloa, 05°33.222'N, 87°02.053'W, dredging 39–44 m depth, J. Cortés, J. Sibaja-Cordero, Proteus-CIMAR Expedition, R/V Proteus, 17 January 2007. MZUCR 2513 (GenBank accession numbers MW491885 (28S rDNA), MW491886 (mtMutS) and MW491887 (COI)), ethanol preserved, Chatham Bay, Punta Ulloa, 05°33.222'N, 87°02.063'W dredging 45 m depth, J. Sibaja-Cordero, 14 April 2008. MZUCR 3301, ethanol preserved, Bahía Iglesias, Muela Rock, 23 m depth, J. Cortés, 14 January 2007. MZUCR 3302, lot preserved in ethanol, Chatham Bay, 05°32'43.6"N, 87°01'41.7"W, 20 m depth, J. Cortés, 3 April 2009. MZUCR 3303, lot ethanol preserved, off Cocos Island, Everest pinnacle, 55 m depth, HOV DeepSee Dive 2375, A. Klapfer, 25 October 2015.
Chatham Bay, Cocos Island, Costa Rica, 45 m depth.
The holotype is composed of several bright red tubular, single anthosteles extending up to 8 mm above the surface of a 4 cm-diameter rhodolith (from one lot of five rhodoliths, 4–7 cm diameter) (Fig.
The paratypes were found on rhodoliths of 3–9 cm in diameter. Some variation in colour was found in a few paratypes. They present red and orange anthosteles in the same colony. All other characteristics were consistent with those of the holotype.
The colonies were found only associated with living rhodoliths, from 20 to 55 m depth. The rhodoliths were in aggregations, forming extensive beds or dispersed on sandy bottoms (Fig.
The species is presently only known from Cocos Island.
An adjective (L), occultus, meaning hidden, secret, mysterious. Historically, numerous expeditions to Cocos Island seeking pirate treasures buried somewhere in the mysterious island were completely unsuccessful. The new species, out of sight, hidden amongst rhodoliths, shows that the real treasure of the island is its biodiversity.
All phylogenetic analyses placed MZUCR 2513 in a well-supported clade (ML bootstrap (bs) = 95%; Bayesian posterior probability (pp) = 1.0) that included families Tubiporidae, Acrossotidae, Arulidae, and several genera of Clavulariidae (Fig.
Maximum likelihood reconstruction (2417 nt of concatenated mtMutS, COI, 28S rDNA) of the Holaxonia-Alcyoniina clade of Octocorallia. Two stoloniferous taxa belonging to the Calcaxonia-Pennatulacea clade (Cornularia, Telestula) were included to root the tree. Clades that did not include any stoloniferans have been collapsed to triangles to facilitate readability. Solid circles on nodes indicate strong support from both maximum likelihood and Bayesian analyses (bootstrap > 70%, pp > 0.95)
From the morphological point of view, Rhodolitica gen. nov. is related to Rhodelinda, Paratelesto, and Stragulum Ofwegen & Haddad, 2011, all within the family Clavulariidae, a conclusion that is supported by the phylogenetic analysis. Morphologically, Paratelesto grows in bushy clusters, the colony branches many times, up to four times with secondary polyps (
The morphological distinction of Rhodolitica from Rhodelinda, Stragulum, and Tubipora is also supported by phylogenetic analyses. Although all analyses strongly supported the placement of Rhodolitica in a clade with Rhodelinda, Paratelesto, Stragulum, and Tubipora, the relationships among those genera remained poorly resolved. Lack of clear evidence for a sister relationship between Rhodolitica and any one of these other related but morphologically distinct genera supports the decision to establish a new genus for R. occulta sp. nov., and highlights the importance of the integrative approach to octocoral taxonomic work.
We are grateful to Bert Hoeksema (Naturalis Biodiversity Center, Leiden, the Netherlands) and the anonymous reviewers for their suggestions and comments that definitely improved our publication. We thank the Servicio Nacional de Áreas Protegidas de Costa Rica and the personnel of the Parque Nacional Isla del Coco for allowing our research at the Island. We express our appreciation to the crew of the R/V Proteus, 2007 trip, and the submersible DeepSee and M/V Argo (UnderSea Hunter Group). We thank Jeffrey Sibaja and Jorge Cortés for collecting the samples. We appreciate Luis Solís and Andrea Moya (UCR) for their contribution in this manuscript. The research was funded by Vicerrectoría de Investigación, Universidad de Costa Rica (projects B5172, B6773 and B5159), Marviva Foundation and Conservation International.
Table S1. Sequence data used in phylogenetic analysis
Data type: Phylogenetic
Explanation note: Table S1 contains the sequence data of Octocorallia used in the phylogenetic analysis for a new genus and species: Rhodolitica occulta gen. nov. sp. nov.
Fig. S1
Data type: Phylogenetic
Explanation note: Maximum likelihood tree for mtMutS (738 nt). Only the stoloniferan clade that includes Rhodolitica occulta gen. nov. sp. nov. is shown, rooted to Clavularia inflata. Numbers on branches are bootstrap values (% of 100 replicates)
Fig. S2
Data type: Phylogenetic
Explanation note: Maximum likelihood tree for COI (693 nt). Only the stoloniferan clade that includes Rhodolitica occulta gen. nov. sp. nov. is shown, rooted to Clavularia inflata. Numbers on branches are bootstrap values (% of 100 replicates).
Fig. S3
Data type: Phylogenetic
Explanation note: Maximum likelihood tree for 28S rDNA (813 nt). Only the stoloniferan clade that includes Rhodolitica occulta gen. nov. sp. nov. is shown, rooted to Clavularia inflata. Numbers on branches are bootstrap values (% of 100 replicates).