Primnoidae (Octocorallia: Calcaxonia) from the Emperor Seamounts, with Notes on Callogorgia Elegans (Gray, 1870)

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Primnoidae (Octocorallia: Calcaxonia) from the Emperor Seamounts, with Notes on Callogorgia elegans (Gray, 1870)

Stephen D. Cairns, Robert P. Stone, Hye-Won Moon, Jong Hee Lee

Pacific Science, Volume 72, Number 1, January 2018, pp. 125-142 (Article)

Published by University of Hawai'i Press

For additional information about this article https://muse.jhu.edu/article/683173

[ This content has been declared free to read by the pubisher during the COVID-19 pandemic. ] Primnoidae (Octocorallia: Calcaxonia) from the Emperor Seamounts, with Notes on Callogorgia elegans (Gray, 1870)1

Stephen D. Cairns,2,6 Robert P. Stone,3 Hye-Won Moon,4 and Jong Hee Lee 5

Abstract: Six primnoid species are reported from depths of 280 – 480 m from the southern Emperor Seamounts, including two new species (Callogorgia imperialis and Thouarella taylorae). Only the new species are fully described and illustrated. Also, Callogorgia elegans, which has a confused taxonomic history, is discussed and illustrated. Not unexpectedly, the Emperor Seamount primnoids have a strong affinity with the fauna of the Hawaiian Islands, an affinity that is expected to increase as more collecting is done in the region.

The United Nations General Assembly nations around the world are developing pro- approved Resolution 61/105 in December tocol and policy on fishing encounters with 2006 ( United Nations General Assembly 2007) the sensitive biota (Durán Muñoz et al. 2012). that calls on States to directly, or through Here we report on collections made on fish- Regional Fisheries Management Organiza- ing vessels in the Emperor Seamounts, North tions, apply a precautionary ecosystem ap- Pacific Ocean, as part of a joint project be- proach to sustainably manage fish stocks and tween the United States and the Republic of protect vulnerable marine ecosystems ( VMEs) Korea. in deep-sea fisheries on the high seas (FAO The Emperor Seamounts are a series of 2009). Some deep-sea corals are indicator very old and heavily eroded guyots (flat- species of VMEs, and accordingly fishing topped) and seamounts stretching approximately 6,700 km from the Aleutian Trench to the Northwestern Hawaiian Islands (Figure 1 This research was supported by a joint project 1). Several nations, including the Republic of agreement between the U.S. National Marine Fisheries Korea and Japan, conduct fishery operations Service (Alaska Fisheries Science Center) and the Repub- lic of Korea National Institute of Fisheries Science (proj- for two important fish species, North Pacific ect no. R2017027). The findings and conclusions in this armorhead (Pseudopentaceros wheeleri ) and article are those of the authors and do not necessarily splendid alfonsino (Beryx splendens), with gear represent the views of the National Marine Fisheries Ser- that occasionally contacts the seafloor. vice, NOAA. Manuscript accepted 6 June 2017. 2 Department of Invertebrate Zoology, National Mu- seum of Natural History, Smithsonian Institution, P.O. materials and methods Box 37012, Washington, D.C. 20013-7012. 3 NOAA, National Marine Fisheries Service, Alaska Coral specimens were collected during fish- Fisheries Science Center, Juneau, Alaska 99801. ing operations aboard the FV Oryong501 in 4 Department of Marine Bio-Resources, National Marine Biodiversity Institute of Korea, Seocheon 33662, 2014 and the FV Oyang96 in 2014, 2015, and Republic of Korea. 2016 (see Table 1). Specimens were frozen 5 National Institute of Fisheries Science, Busan, Re- (−20°C) onboard the fishing vessels and trans- public of Korea. 6 ferred to the National Institute of Fisheries Corresponding author (e-mail: [email protected]). Science (Busan, Republic of Korea) where they were preliminarily identified to family. Pacific Science (2018), vol. 72, no. 1:125 – 142 Whole specimens or samples of primnoid doi:10.2984/72.1.8 octocorals were shipped to the U.S. National This article was created by a U.S. government employee Museum of Natural History ( NMNH ), and is in the public domain. Public domain information Smithsonian Institution ( Washington D.C.), may be freely distributed and copied, but it is requested that in any subsequent use the Smithsonian Institution where they were ultimately identified to spe- and the journal, Pacific Science, be given appropriate cies. Specimens are deposited primarily at the acknowledgment. National Marine Biodiversity Institute of

125 126 PACIFIC SCIENCE · January 2018

Figure 1. Map of the Emperor Seamounts in the North Pacific Ocean showing the location of six species of primnoid corals collected during this study. Taxa are as follows: Callogorgia imperialis, n. sp. (green circles), Callogorgia formosa (gray circles), Faniella tuberculata ( black circles), Candidella helminthophora ( brown circles), Narella vermifera (orange circle), and Thouarella taylorae, n. sp. (white circles).

Korea, and voucher fragments and scanning BM British Museum ( Natural His electron microscopy (SEM) stubs of most tory), now the Natural History species are also deposited at the NMNH. Museum, London The morphological terminology used here L:W Length to width ratio of a sclerite follows that of Bayer et al. (1983). The scan- MZW Museum of Natural History, ning electron photomicrographs were taken Wrocław University, Poland by S.D.C. using a scanning electron micro- MABIK National Marine Biodiversity In- scope (Zeiss EVO MA15). Authorship of the stitute of Korea, Seocheon 33662, two new species is attributed to the first au- Republic of Korea thor (Cairns) alone. NMNH National Museum of Natural His- Abbreviations used in the text include the tory, Smithsonian following: SEM Scanning electron microscopy Primnoidae from the Emperor Seamounts · Cairns et al. 127

USNM United States National Museum 152-5, colony, mabik CN00076508, branch (now the NMNH, but acronym fragment, usnm 1422521; Oyang 2014-96- used for cataloging purposes) 153-1, colony, mabik CN00076513, branch ZMA Zo¨ologische Museum, Amster‑ fragment, usnm 1422510; Oyang 2014-96- dam 153-5, colony, mabik CN00076514, branch fragment, usnm 1422518; Oyang 2014-96- results 158-2, colony, mabik CN00079269, branch fragment, usnm 1422513; Oyang 2014-96- systematic treatment 181-5, colony, mabik CN00076544, branch Subclass Octocorallia fragment, usnm 14225217; Oyang 2015-96-4- Order Alcyonacea 1, colony, mabik CN00076567, branch frag- Suborder Calcaxonia ment, usnm 1422520; Oyang 2016-96-37-1, Family Primnoidae Milne Edwards, 1857 colony, mabik CN00076578, fragments, usnm Genus Callogorgia Gray, 1858 1422519; Oryong 2014-501-13-1, branch fragment, mabik CN00076125, fragments, usnm Callogorgia Gray, 1858:286. — Bayer, 1982:119 – 1422522; Oryong 2014-501-131-3, branch 123 ( key to Indo-Pacific species). — Cairns fragments, mabik CN00076288, fragments, and Bayer, 2009:40, fig. 12A – D (more usnm 1422523. complete synonymy). — Cairns, 2016:58. type locality: Oyang 2015-96-138-1: 35.6217° N, 171.3167° E ( Kökö Guyot, Em- type species: Gorgonia verticillata Pallas, peror Seamounts), 347 m. 1766, by monotypy. distribution: Emperor Seamounts diagnosis: Colonies uniplanar, pinnately ( Kökö Guyot), 284 – 380 m. Elsewhere: Saga- or dichotomously branched. Polyps cylindri- mi Bay, Japan (depth unknown). cal to clavate, arranged in whorls of up to 12, description: Colonies are uniplanar and all polyps facing upward. Polyps covered with taller than wide, the largest specimen (the ho- eight longitudinal rows of body wall scales, lotype, Figure 2A) measuring 42 cm in height the number of scales per row decreasing from and 22 cm in width, having a broken basal ab- to adaxial polyp side. Body wall scales branch diameter of 7.2 mm. Branching is ir- granular, smooth, pitted, or covered with tall regularly dichotomous (not pinnate) and the ridges. Inner side of opercular scales covered branches are rather stiff, the axis being longi- with serrated multiple keels. tudinally ridged. The polyps are arranged in remarks: Including the new species de- whorls of 6 or 7 (Figure 3B) on distal branches scribed herein there are 27 valid species in the (8 or 9 on proximal stems); 5 to 6 whorls oc- genus. A key to the Indo-Pacific Callogorgia is cur per cm branch length; the whorl diameter in preparation. ranges from 2.8 to 3.5 mm. The polyps are distribution: Indo-Pacific, North At- 1.5 – 1.9 mm in length, curved, and slightly lantic, 37 – 2,472 m. clavate (Figures 3A, C ). The color of the colony is light orange. Callogorgia imperialis Cairns, n. sp. There are eight longitudinal rows of body Figures 2A, 3A – D, 4A – I wall scales, decreasing in number from the ab- to adaxial polyp side; the body wall scler- Caligorgia elegans: Kinoshita, 1908:40 – 42, ite formula is 12 – 16: 6 – 10: 2 – 5: 2 – 3. Most pl. 3, figs. 17 – 18; pl. 6, fig. 48. body wall scales are wider than taller, slightly curved to accommodate the curvature of the material examined (types): Holotype: polyp, and rather thick (e.g., about 0.08 mm). Oyang 2015-96-138-1, mabik CN00076574, The abaxial body wall scales (Figure 4A) branch fragments, SEM stubs 2359 – 2363, range from 0.33 to 0.36 mm in width, the usnm 1422524. Paratypes: Oyang 2014-96- outer laterals (Figure 4B) 0.37 – 0.42 mm, and 13-2, colony, mabik CN00076352, branch the inner laterals (Figure 4C ) 0.14 – 0.16 mm. fragments usnm 1422525; Oyang 2014-96- The abaxial and outer laterals often have a 128 PACIFIC SCIENCE · January 2018

Figure 2. Colony images of: A, Callogorgia imperialis, holotype; B, Callogorgia formosa, Oryong 2014-501-33-1; C, Fanel- lia tuberculata, Oyang 2014-96-37-6; D, Candidella helminthophora, Oryong 2014-501-110-6; E, Narella vermifera, Oyang 2016-96-20-1; F, Thouarella taylorae, holotype. serrate distal edge. The adaxial body wall type scales (Figures 3C, 4E ), or this region scales (Figure 4D) are squarish, ranging from may be covered with polygonal scales similar 0.19 to 0.21 mm on a side, and thinner, al- to those of the coenenchyme. The outer face though even smaller adaxials occur lower on of the body wall scales is fairly smooth, the polyp. Proximal to the adaxial scales the covered with very small (5 µm in diameter) polyp is naked (Figure 3D). At the proximal granules. The opercular scales (Figure 4F ) margin of the polyp, where the polyp scales range in length from 0.23 to 0.49 mm (L:W = transition to the coenenchymal scales, there 2.1 – 2.2), decreasing in length from the ab- may be 2 wide (up to 0.6 mm wide) infrabasal- to adaxial polyp side, forming a low opercu- Figure 3. Stereo views of the holotype of Callogorgia imperialis: A, lateral view of two polyp whorls; B, apical view of a polyp whorl; C, lateral view of a polyp; D, adaxial-oral view of a polyp. Figure 4. Holotype of Callogorgia imperialis: A, outer surface of an abaxial body wall scale; B, outer lateral body wall scales; C, outer surface of an inner lateral body wall scale; D, inner surface of an adaxial body wall scale; E, curved infrabasal scale; F, opercular scales; G, stereo view of coenenchymal scales in situ and a juvenile polyp; H, coenen chymal scales in situ; I, thick coenenchymal scales. Primnoidae from the Emperor Seamounts · Cairns et al. 131 lum. They are lanceolate in shape with a blunt is dichotomous. Without having additional tip. Their outer surface is covered with low material or seeing the type, Versluys (1906) granules, loosely arranged in several rows dis- doubted the synonymy with C. flabellum but tally; their inner surface is covered with tu- suggested that the type be re-examined. bercles proximally and distally bears serrated Kinoshita (1908) provisionally reported a ridges. specimen from Japan as C. elegans; that speci- The coenenchymal scales are polygonal men has a flabellate, dichotomously branched (Figure 4I ) and quite thick (up to 0.12 mm), corallum, a sclerite formula of 12 – 13:7:3:1 – 3, fitting together in a closely adjacent mosaic four to six polyps per whorl, a polyp length of (tessellate) pattern (Figures 4G – H ). These 1.0 – 1.1 mm, and 8 – 8.5 whorls per cm. From scales range from 0.35 to 1.02 mm in greater this description, Kü kenthal (1919, 1924), and length, which is aligned with the branch axis. later Bayer (1982), defined and provided keys These scales are flat to slightly concave and to C. elegans based on this characterization. essentially smooth ( bearing tiny granules Unfortunately, the Japanese specimen is not about 5 µm in diameter) and have serrate C. elegans, as indicated by the unpublished edges, such that the projecting points of one SEM (Figures 5, 6A – D) and notes of the dry scale often interdigitate with the recessed bm holotype ( bm 1965-12-15-006 and 008), region of an adjacent scale, locking them in both made by F. M. Bayer, who showed this position. Below the coenenchymals is a thin species to have a sclerite formula of 7 – 8:2+ layer of small (35 – 45 µm in diameter) tuber- 1:0:1, four polyps per whorl, a polyp length of culate spheroids. 0.78 – 0.93 mm, and about nine polyps per cm. comparisons: Using the keys of (The 2+1 convention indicates that the outer Kü kenthal (1919, 1924) and Bayer (1982), this lateral sclerites are disjunct). Kinoshita’s C. species keys closest to C. elegans (Gray, 1870), elegans, however, does seem to be the same as but as explained in Remarks that species has C. imperialis, as described herein. Using the been incorrectly diagnosed since its descrip- key provided by Bayer (1982) and Kü kenthal tion. Callogorgia imperialis is unique among (1919, 1924), C. elegans would key closest to the Indo-Pacific species by having a high C. indica Versluys in Thomson and Hender- number of abaxial body wall scales and tessel- son (1906), known only from the Andaman late coenosteal scales. It appears to be the spe- Islands at 82 – 490 m. They both have a similar cies Kinoshita (1908) described as “? Caligor- sclerite formula and polyp size, differing only gia elegans.” in that C. indica has only two or three polyps etymology: Named imperialis ( Latin per whorl and a more delicate colony than for “of the empire” or “emperor”), an allu- that of C. elegans. sion to its type locality on the Emperor ecological remarks: Several colonies Seamounts. of C. imperialis hosted one or more large remarks: As mentioned earlier, C. impe- ophiuroids, Family Asteroschematidae and rialis keys closest to C. elegans (Gray, 1870) in likely Asteroschema sp. (Gordon Hendler, Los the classic keys of Kü kenthal (1919) and Bayer Angeles County Museum of Natural His (1982), but that species is poorly known. The tory, pers. comm.) when collected. The brittle seven-word original description and two illus- stars are quite large, up to 24 cm in total trations of C. elegans (Gray, 1870) of a speci- length (arm tip to arm tip), and apparently use men from Formosa (= Taiwan) do not help the corals as elevated feeding or spawning to distinguish it from other Callogorgia with platforms. dichotomous branching. Studer’s (1878) ref- erence to C. elegans simply transferred the Callogorgia formosa ( Kü kenthal, 1907) species from Callicella to Calligorgia. Wright Figure 2B and Studer (1889) synonymized it with C. flabellum (Ehrenberg, 1834), but as Versluys Caligorgia formosa Kü kenthal, 1907:208 – 209; (1906) and Bayer (1982) later showed, C. fla- 1919:366 – 369, text figs. 155 – 159; pl. 30, bellum has pinnate branching, and C. elegans fig. 1; pl. 40, fig. 47. 132 PACIFIC SCIENCE · January 2018

Figure 5. Holotype of Callogorgia elegans, bm 1965-12-15-006.

Callogorgia formosa: Bayer, 1982:130 – 134, from opposite sides of the main branch (op- figs. 9 – 10. — Cairns, 2010:427 – 429, figs. posite pinnate), the undivided branchlets up 1E, 12A; 2016:61 – 65, figs. 31A, 32, 33. to 15 cm in length (Figure 2B). Polyps arranged in whorls of 3 to 5; polyps slightly cla- material examined: Oryong 2014-501- vate, 1.4 – 1.7 mm in length. Body wall sclerite 58-3, fragments, mabik CN00076181, 2 branch formula: 8 – 12: 7 – 9: 2 – 5: 1 – 3; the proximal fragments, usnm 1422534; Oryong 2014-501- adaxial polyp side is unprotected. Body wall 33-1, fragments, mabik CN00079275, 2 scales rectangular with a coarse serrate distal branch fragments, usnm 1422535. margin. Opercular scales unique in that both types and type locality: Syntype de- outer and distal inner surfaces covered with posited as mzw 25. Type locality: Southwest coarse blunt spines, producing a thick, blunt of Great Nicobar Islands, Indian Ocean, apex. Coenenchymal scales elongate and 362 m. granular. distribution: Emperor Seamounts: remarks: Of the 27 species in the genus, Kökö Guyot, 280 – 289 m. Elsewhere: Off C. formosa is the only one to have opposite Great Nicobar, Nicobar Islands, northern pinnate branching and is thus probably the New Zealand, Hawaiian Islands, 296 – 750 m. most easily recognized species in the genus. It diagnosis: Colony uniplanar, usually has been more than adequately described and consisting of one straight main branch from figured at least four times (see synonymy) and which pairs of undivided branchlets diverge thus is not redescribed herein. Figure 6. Stereo views of the holotype of Callogorgia elegans, bm 1965-12-15-006. A, lateral view of a whorl of polyps; B – D, lateral views of individual polyps. 134 PACIFIC SCIENCE · January 2018

Genus Fanellia Gray, 1870 ment, usnm 1422508; Oyang 2014-96-37-6, colony, mabik CN00076378, branch frag- Fanellia Gray, 1870:46. — Bayer, 1982:134 – ment, usnm 1422506; Oyang 2014-96-39-3, 135 ( key to species). — Bayer and Stefani, colony, mabik CN00076383, branch frag- 1989:470 – 471 ( key to species). — Cairns ment, usnm 1422505; Oyang 2014-96-40-3, and Bayer, 2009:40 – 41, fig. 12E – M. — colony, mabik CN00076389, branch frag- Cairns, 2016:72 – 73. ment, usnm 1422503. types and type locality: Holotype: type species: Primnoa compressa Verrill, formerly zma Coel. 2286, now at Centre for 1865, by monotypy. Biodiversity Naturalis, Leiden. Type locality: diagnosis: Colonies uniplanar, branches 5.717° N, 119.667° E (Sulu Sea), 522 m. alternate-pinnate or dichotomously arranged. distribution: Emperor Seamounts: Polyps often clavate, arranged in pairs or Kammu, Yüryaku, and Kökö Guyots, 343 – 480 whorls of up to 16, all polyps facing up- m. Elsewhere: northern New Zealand and ward. Polyps arranged in 4 to 8 longitudinal Kermadec Ridge, Sulu Sea, Bikini, Japan, rows of body wall scales, the number of scales Hawaiian Islands, 128 – 522 m. in each row decreasing from ab- to adaxial diagnosis: Colony uniplanar and side. Outer surface of body wall and coenen- alternate-pinnate in branching (Figure 2C ), chymal scales covered with closely spaced, the terminal branchlets rarely more than 4 cm complex, granular tubercles, sometimes fus- in length. Polyps arranged in pairs or whorls ing into low, flat-topped radial ridges. Inner of 3, often lacking from the terminal cm of face of operculars bears a keel composed each branchlet; polyps 0.9 – 1.4 mm in length. of multiple ridges. Coenenchymal scales or Body wall scale formula: 4 – 9: 1 – 3: 0 – 3: 1 – 2; plates thick, and imbricate or tessellate in the short adaxial side of the polyp is unpro- arrangement. tected. Body wall scales rectangular, with a remarks: Nine species are known in the straight distal edge (not pointed). Coenen- genus, most of them keyed by Bayer and chymal scales (plates) thick, trapezoidal to rect- Stefani (1989). The genus is primarily differ- angular, and arranged in a tessellate pattern. entiated from Callogorgia by its unique tuber- comparisons: Fanellia tuberculata is com- culate surface sculpturing; however, based on pared to other species using tabular keys molecular sequencing evidence, Taylor and (Cairns 2010, 2016) and dichotomous keys Rogers (2015) suggested that Fanellia is prob- (Bayer 1982, Bayer and Stefani 1989) and is ably synonymous with Callogorgia. fully described and illustrated by Bayer (1982) distribution: Western and Central and Cairns (2016), and thus only a diagnosis is Pacific, from New Zealand to Alaska, 82 – provided herein. 1,341 m. Genus Candidella Bayer, 1954 Fanellia tuberculata ( Versluys, 1906) Figure 2C Candidella Bayer, 1954:296 (nom. nov.). — Cairns and Bayer, 2004:476 – 477; 2009:46, Caligorgia tuberculata Versluys, 1906:80 – 81, fig. 16H – N (synonymy and discussion). — text-figs. 95– 96. Cairns, 2009:440; 2016:102. Fanellia tuberculata: Bayer, 1982:144 – 154, figs. 18 – 26. — Bayer and Stefani, 1989:471. — type species: Primnoa imbricata Johnson, Cairns, 2010:433 – 434, table 3; Taylor and 1862, by monotypy. Rogers, 2015:fig. 1. — Cairns, 2016:73 – 74, diagnosis: Colonies unbranched (flagel- figs. 31E, 40, 41, table 4. liform) or uniplanar, with dichotomous branching. Polyps arranged in whorls, usually material examined: Oyang 2014-96- standing perpendicular to branch. Each polyp 15-1, colony, mabik CN00076354, branch ringed distally with 4 large marginal scales, fragment, usnm 1422507; Oyang 2014-96-20- which form a cowl around operculum; body 1, colony, mabik CN00079270, branch frag- wall scales much smaller, occurring in 2 to 4 Primnoidae from the Emperor Seamounts · Cairns et al. 135 submarginal tiers. Opercular scales with an chymal scales elliptical to irregular in shape, inner keel. Coenenchymal scales elliptical and having a flat to slightly concave outer surface. highly concave. remarks: This species has been fully de- remarks: Four species are known in the scribed and figured recently (Cairns 2009, genus, discussed and figured by Cairns (2009) 2016), and thus only a diagnosis is given here. and Cairns and Bayer (2009). It is distinguished from congenerics by Cairns distribution: Amphi-Atlantic, Fiji, Ha- (2009). waiian Islands, New Zealand, Emperor Sea- mounts, 378 – 2,165 m. Genus Narella Gray, 1870

Candidella helminthophora ( Nutting, 1908) Narella Gray, 1870:49. — Cairns and Bayer, Figure 2D 2003:618 – 619; 2007:84 – 86 ( list of species); 2009:43, fig. 14A – G. — Cairns and Stenella helminthophora Nutting, 1908:575 – Baco, 2007:392 – 393. — Cairns, 2012:14. — 576, pl. 44, figs. 6– 9; pl. 47, fig. 5. Taylor and Rogers, 2015:190 ( list of Candidella helminthophora: Grigg and Bayer, species). 1976:171 ( listed). — Cairns, 2009:440 – 443, pl. 1A, H, 19 – 20 (complete synonymy); type species: Primnoa regularis Duchas- 2016:102 – 104, figs. G,46 60 – 62. saing & Michelotti, 1860, by monotypy. diagnosis: Colonies dichotomously or material examined: Oyang 2014-96- pinnately branched, or unbranched. Polyps 20-2, colony, mabik CN00079271, 2 branch arranged in whorls, the polyps always facing fragments, usnm 1422541; Oryong 2014-501- downward. Polyps covered by 3 pairs of large F-110-6, colony, mabik CN00079272, 2 body wall scales ( basal, medial, and buccal); branch fragments, usnm 1422542. smaller adaxial body wall scales may be pres- types and type locality: The holo- ent, although proximal portion of adaxial type is deposited at the NMNH (usnm polyp usually naked. Opercular scales usu 23385). Type locality: 23.786° N, 166.415° W ally prominently keeled. Coenenchymal scales (off French Frigate Shoals, Hawaiian Islands), often quite thick (plates) and ridged, ar- 722 – 726 m. ranged in an imbricate or tessellate (mosaic) distribution: Emperor Seamounts fashion. ( Yüryaku and Kökö Guyots), 300 – 470 m. remarks: Narella is one of the most Elsewhere: Southern Kermadec Ridge to species-rich among the 43 primnoid genera, North Cape, and Kermadec Ridge, New containing 41 valid species (Cairns 2012, Zealand, throughout Hawaiian Islands, 417 – Taylor and Rogers 2015). More extensive dis- 1,801 m. cussions of the genus may be found in Cairns diagnosis: Colonies uniplanar, equally and Bayer (2009) and Cairns (2012). and dichotomously branched (Figure 2D). distribution: All ocean basins, 128 – Polyps occur in whorls of 2 – 4, individual 4,594 m (Cairns 2012). polyps strongly flared and up to 3.8 mm in length, standing perpendicular to branch. Of Narella vermifera Cairns & Bayer, 2007 the 4 marginal scales, the 2 abaxial marginals Figure 2E are wider; 4 submarginals occur below marginals. Below submarginals the body wall Narella vermifera Cairns & Bayer, 2007:98 – 99, scales are irregular but usually consist of at figs. 1E, 8A – D, 9A – H. least 2 more tiers. Two tall basal scales firmly attach polyp to coenenchymal scales, some- material examined: Oyang 2016-96- times becoming greatly enlarged and reflexed 20-1, 1 colony, mabik CN00076585, branch in shape, forming a tube for a commensal fragment, usnm 1422530. polynoid polychaete, Gorgoniapolynoe muzikae . types and type locality: The types Opercular scales triangular, each bearing a are deposited at the NMNH. Type locality: prominent keel on their inner side. Coenen- 23° 02′ N, 162° 05′ W (off Ka‘ula Rock, near 136 PACIFIC SCIENCE · January 2018

Ni‘ihau), northwest Hawaiian Islands, 275 – or ornate longitudinal keel. Opercular scales 527 m. usually keeled as well. distribution: Emperor Seamounts: remarks: The genus Thouarella was com- Kökö Guyot, 296 m. Elsewhere: Northwest- prehensively revised by Taylor et al. (2013). ern Hawaiian Islands (off Ni‘ihau and Nïhoa), They included 25 species in the genus, 271 – 527 m. which were divided into two groups (see Di- diagnosis: Colonies uniplanar and diagnosis). Two species have been described chotomously branched, up to 17 cm in height since (Cairns 2011) and one herein, result- and 19 cm in width, with a basal axis diameter ing in 28 species in the genus, making it of 5.8 mm (specimen reported herein, Figure one of the more species-rich among the 2E ). Polyps arranged in whorls of 3 to 5; 5 primnoids. The species key in Taylor et al. whorls per cm branch length; whorl diameter (2013) makes it relatively easy to identify the about 3.9 mm. Polyps 1.8 – 2.0 mm in length. species, which, until then, had been a difficult Basal scales have a rounded dorsolateral edge process. and often have greatly enlarged, recurved distribution: Worldwide from the outer edges that form a tube for a commensal Aleutian Islands to Antarctica, 11 – 2,100 m. polynoid polychaete of the genus Gorgonia polynoe. Medial scales quite short; both basal Thouarella taylorae Cairns, n. sp. and buccal pairs of scales form complete but Figures 2F, 7A – D, 8A – E unfused rings. Coenenchymal scales thick, flat, and polygonal in shape, arranged in a material examined: Types. mosaic pattern. types and type locality: Holotype: remarks: This is the first report of this Oryong 2014-501-178-2, colony, mabik species subsequent to its original description. CN00076337, several branch fragments, Its characteristics were tabulated by Cairns usnm 1422540. Paratypes: Oryong 2014-501- and Bayer (2007: table 2), who also compared 93-1, colony fragment, mabik CN00079274, it to N. biannulata ( Kinoshita, 1907), a similar branch fragment, usnm 1422536; Oryong species known from off Japan. 2014-501-94-2, colony fragment, mabik CN00076228, 2 branch fragments, usnm Genus Thouarella Gray, 1870 1422539; Oryong 2014-501-110-1, 1 colony, mabik CN00079273, polyps, usnm 1422538; Thouarella Gray, 1870:45. — Cairns and Bayer, Oyang 2014-96-15-2, colony fragment, mabik 2009:33 – 34 (more complete synonymy). — CN00076355, branch fragment, usnm Cairns, 2011:4 – 5. — Taylor et al., 2013:20 – 1422537. Type locality: 34° 33.3′ N, 171° 21 (revision of genus, including complete 13.7′ E, 297 m. synonymy and discussion). distribution: Emperor Seamounts: Kammu and Kökö Guyots, 280 – 372 m. type species: Primnoa antarctica Valen description: The colonies are essen ciennes, 1846, by monotypy. tially uniplanar, consisting of relatively few diagnosis: Branching of colonies as bot- main branches from which long (up to 8 cm in tlebrush, uniplanar pinnate, or dichotomous, length) branchlets originate in an irregularly the main branch itself sparsely divided. Polyps pinnate manner, although some branchlets isolated (i.e., occurring on all sides of branch). diverge slightly out of the plane, producing Group 1 sensu Taylor et al. (2013), or ar- a slightly bushy aspect but not considered to ranged in pairs and /or whorls (Group 2 sensu be bottlebrush branching (Figure 2F ). The Taylor et al. 2013). Each polyp covered with holotype is 17 cm in height and 2 mm in basal 6 or 8 longitudinal rows of body wall scales, axis diameter. The axis is bronze and finely those of the adaxial side reduced in size and longitudinally striate. number. Marginal scales often arranged in 2 The white polyps occur individually on the circlets that alternate with one another, the main branches and in pairs and whorls of 3 or inner face of each marginal bearing a simple 4 on the branchlets, paired polyps most com- Figure 7. Stereo views of the holotype of Thouarella taylorae: A, distal polyps arranged in whorls of three and pairs; B, lateral view of a polyp; C, adaxial view of a polyp; D, opercular view of a polyp (o, 5 opercular scales; m, the 8 marginal scales). 138 PACIFIC SCIENCE · January 2018 mon at branchlet tips (Figure 7A), the whorls T. laxa Versluys, 1906; and T. moseleyi Wright more developed proximally. The polyps are & Studer, 1889. Thouarella taylorae can be dis- 0.8 – 1.1 mm in length and slightly flared distinguished from both T. tydemani and T. laxa tally, curved upward along the branchlet axis by having pointed (not spinose) marginal (Figures 7B – C ); about 10 pairs of whorls oc- scales (and thus a much lower L:W ), smaller cur per cm branchlet length. polyps (those of the other two species are up Each polyp is covered with 8 longitudinal to 1.5 mm in length), and more abaxial body rows of body wall scales having the body wall scales (those of the other two have only wall sclerite formula of: 8 – 9:4 – 7:4 – 5:3 – 5. four to six abaxial body wall scales per row). The adaxial body wall scales (Figure 7C ) are Thouarella taylorae differs from T. moseleyi in slightly fewer and smaller than the others but having longer branchlets, shorter polyps, still cover the entire adaxial side. The mar- whorled polyps (only pairs occur in T. mose- ginal scales (Figures 7D, 8A) are quite broad leyi ), and more body wall scales in its abaxial (up to 0.3 mm, L:W = 0.75 – 1.1), slightly rows. curved to fit the circumference of the polyp, etymology: This species is named in and arranged in an inner and outer overlap- honor of Michelle Taylor, who published a ping quartet; all marginals are the same size seminal paper consisting of a revision of all and shape. The sculpturing of their outer and species of the genus Thouarella (Taylor et al. inner surfaces is similar to that of the opercu- 2013). lars. Their medial keel folds over and fits into ecological remarks: At the one site on the concavity on the outer side of the adjacent Kammu Guyot where specimen 2014-96-15- opercular scale. Some, but usually not all, 2 was collected, approximately six additional of the 8 submarginals also have keeled inner specimens were collected, possibly indicating surfaces. Otherwise the body wall scales (Fig- that this species is found in patches. ure 8B) are elliptical, fairly flat, up to 0.25 mm in width, granular above and tuberculate be- discussion low, and lack keels. The opercular scales (Fig- ures 7D, 8C) also occur in two quartets of 4, Of the six primnoid species reported herein, but alternate in size and shape. The opercu- two (Callogorgia imperialis and Thouarella lars of the outer quartet are lanceolate, up to taylorae ) are thus far known only from the 0.28 mm in length, and have an L:W of Emperor Seamounts, although a broader dis- 1.6 – 2.2. The operculars of the inner quartet, tribution off the Hawaiian Islands is expected; which are usually overshadowed by the larger one ( Narella vermifera) is known elsewhere quartet and thus not visible, are only up to only in the Hawaiian Islands, this record 0.21 mm in length and much more slender being the first subsequent record since its de- (L:W = 2.5 – 3.1). All opercular scales have a scription; and the remaining three species highly concave outer surface that is covered (Callogorgia formosa, Fanellia tuberculata, and with granules and short radiating ridges, Candidella helminthophora) are known from whereas their inner faces are tuberculate throughout the western Pacific, including the basally and prominently keeled distally. The Hawaiian Islands. Thus, four of the six species coenenchymal scales (Figures 8D – E ) are are known from the Hawaiian Islands, and it elongate elliptical, about 0.11 – 0.15 mm in is expected that the remaining two will even- width and 0.25 mm in length, but otherwise tually be found there as well. Twenty-eight similar to the body wall scales. primnoid species are known from the Hawai- comparisons: Thouarella taylorae belongs ian Islands (Cairns 2010), a much more heav- to “species group 2” sensu Taylor et al. (2013), ily collected and studied faunistic region, and a group of 9 of the 28 known species in this it is expected that the number of primnoids genus characterized by having their polyps from the Emperor Seamounts will increase, arranged in pairs and /or whorls. Only three including some of these Hawaiian species, as species in this group have pinnate branching, more collecting is done. The key to the gen- as does T. taylorae: T. tydemani Versluys, 1906; era of Hawaiian primnoids given by Cairns Figure 8. Holotype of Thouarella taylorae: A, marginal body wall scales; B, body wall scales; C, opercular scales; D, coenenchymal scales; E, coenenchymal scales in situ. 140 PACIFIC SCIENCE · January 2018

TABLE 1 Station Data for the 26 Primnoid Specimens Collected during This Study

Specimen No. Date Latitude ( N ) Longitude (E) Depth (m)

2015-96-138-1 10 May 2015 35.6217° 171.3167° 347 2014-96-13-2 14 March 2014 35.0550° 171.7100° 380 2014-96-152-5 21 May 2014 34.9617° 171.8700° 343 2014-96-153-1 21 May 2014 34.9567° 171.8200° 348 2014-96-153-5 21 May 2014 34.9567° 171.8200° 348 2014-96-158-2 23 May 2014 34.9817° 171.8550° 344 2014-96-181-5 31 May 2014 35.5533° 171.3850° 364 2015-96-4-1 8 April 2015 34.9050° 171.8100° 328 2016-96-37-1 24 March 2016 35.6000° 171.3333° 284 2014-501-13-1 26 March 2014 35.1117° 171.7000° 296 2014-501-131-3 25 May 2014 35.4817° 171.4350° 285 2014-501-58-3 25 April 2014 35.5800° 171.3967° 280 2014-501-33-1 11 April 2014 35.2900° 171.5083° 289 2014-96-15-1 15 March 2014 32.2733° 172.8650° 370 2014-96-20-1 17 March 2014 32.6967° 172.2567° 470 2014-96-37-6 28 March 2014 32.6383° 172.2850° 480 2014-96-39-3 29 March 2014 32.6150° 172.2833° ~450 2014-96-40-3 31 March 2014 32.0100° 173.1300° 343 2014-96-20-2 17 March 2014 32.6967° 172.2567° 470 2014-501-110-6 15 May 2014 35.4533° 171.5617° 300 2016-96-20-1 24 March 2016 35.5167° 171.3000° 296 2014-501-178-2 15 June 2014 35.4800° 171.5583° 297 2014-501-110-1 15 May 2014 35.4533° 171.5617° 300 2014-501-93-1 9 May 2014 35.5467° 171.3617° 280 2014-501-94-2 9 May 2014 35.5667° 171.5650° 372 2014-96-15-2 15 March 2014 32.2733° 172.8650° 370

(2010) also serves to identify the genera from the biological specimens. Michele Masuda the Emperor Seamounts. provided Figure 1, and Justin Heard assisted The Aleutian Islands and the seamounts with Figures 1 and 2. We are grateful to F. B. of the Gulf of Alaska, only a slightly greater Bayer for the use of his illustrations that form distance to the north than the Hawaiian Is- Figures 5 and 6. Gordon Hendler is acknowl- lands are to the southeast, also have a rich edged for having identified the commensal primnoid fauna [e.g., 19 species off the Aleu- ophiuroids. tians (Cairns 2011) and five different species from the deep seamounts in the Gulf of Literature Cited Alaska (Cairns and Baco 2007)]. However, no species are held in common between the Bayer, F. M. 1954. New names for two genera Emperor Seamounts and this more northerly of Octocorallia. J. Wash. Acad. Sci. 44:296. region, a finding recently reported by Miya- ——— . 1982. Some new and old species of moto et al. (2017). the primnoid genus Callogorgia Gray, with a revalidation of the related genus Fanellia Gray (Coelenterata: Anthozoa). Proc. Biol. acknowledgments Soc. Wash. 95:116 – 160. We thank Loh-Lee Low, Seok-Gwan Choi, Bayer, F. M., M. Grasshoff, and J. Verseveldt, Inja Yeon, and Eunjung Kim for project sup- eds. 1983. Illustrated trilingual glossary of port. We thank Chanho Hwang, Sungsu Kim, morphological and anatomical terms ap- Yungil Lee, and Seochul Kim for collecting plied to Octocorallia. E. J. Brill, Leiden. Primnoidae from the Emperor Seamounts · Cairns et al. 141

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