Eunicea clavigera, BAYER, 1961

Sánchez, Juan Armando, 2009, Systematics of the candelabrum gorgonian corals (Eunicea Lamouroux; Plexauridae; Octocorallia; Cnidaria), Zoological Journal of the Linnean Society 157 (2), pp. 237-263 : 248-253

EUNICEA CLAVIGERA BAYER, 1961 ( FIGS 4–7 , 8H )

Eunicea clavigera Bayer, 1961: 147 (Holotype: USNM 50265 ).

Eunicea sp. Sánchez & Lasker, 2003: 2040.

Eunicea sp. 1 Sánchez & Wirshing, 2005: 515.

Description: Colonies up to 1.8 m in height, long branches, irregular branching ( Fig. 4 ). Terminal branches 5–15 mm without the calyx. Calyx long, paler than the surrounding tissue in dry samples. Middle layer sclerites robust spindles between 0.58 and 2.66 mm in length and 0.08 and 0.72 in diameter ( Fig. 5 ). Axial layer sclerites with ornate spindles, lavender or purple, or irregular bodies up to 0.25 mm ( Fig 7D–F ). Surface layer with small foliate and irregular clubs between 0.07 and 0.19 mm in length ( Fig. 6 ). Polyp with anthocodial armature consisting of robust rods between 0.15 and 0.57 mm and also small ones between 0.08 and 0.18 mm ( Fig. 7A–C ); little sclerite bodies and octoradiate forms in the tentacles. Longer tentacles when expanded, usually pale brown probably because of content of zooxanthellae.

There were two morphotypes in this species amongst all the examined material except for the type material. The ‘slender form’ is about 20 cm in height with branches up to 12 cm long. It has often few club sclerites mostly of small sizes ( Fig. 6A ) and larger polyp rods ( Fig. 7C ). Middle layer sclerites are robust ( Fig. 7A ) and visible at the surface, which has a brown appearance in dry specimens. The ‘thick form’ has colonies about 40 cm in height ( Fig. 5C, D ) with thick branches up to 2 cm in diameter near the base. Black coloured coenemchyme with the calyx aperture pale and about three orders of branching but most branches are longer than 20 cm. Both morphotypes have tubular and prominent calyces up to 0.5 mm in length with a semihermetic aperture and with the lower lip longer than the upper one as an enlarged lobule. Calyx tube whiter than the rest of the colony ( Fig. 4 ). The type material is not well preserved and most of these features were not observed but they are mentioned in Bayer (1961). Another interesting feature of E. clavigera is the size of the extended polyps in comparison with other Euniceopsis species (e.g. Fig. 8E–I ).

Distribution and habitat: Antilles, Central America, and the Caribbean coast of South America, probably widespread in Caribbean. Semi-exposed reef terraces and slopes between 10 to 30 m depths.

Material examined: USNM 50265 , holotype, (70% ETOH), Caribbean Sea, Netherlands Antilles, Curacao, Caracas Baai, col. Zaneveld J.S. & P.W. Hummelinck, 22.iv.1955, on chain buoy for 15+ years, submerged several metres deep . USNM 50076 , Paratype, Bermuda, North Rock , col. diver For E. Deichmann, 23.vii.1951, donor : MCZ, Harvard . USNM 100644 , Riding Rock Reef , Leeward terrace, 12 m , San Salvador, Bahamas, 15.viii.1999, col. J.A. Sánchez. USNM 97700 (32), Tesoro island, Rosario islands , Caribbean coast of Colombia, col. J.A. Sánchez & A. Ramirez, 16.ix.1992; fore-reef terrace 20 m ; ICN-MHN-CO-097 (B-38), Isla Fuerte, Banco Bushnell, Colombian Caribbean, Col. J. A. Sánchez, 4.x.1995, mixed coral zone 18 m. INV-CNID-385, (F-14), isla Fuerte, bajo ‘el bobito’, Caribbean coast of Colombia, 3.x.1995, col. J.A. Sánchez, marginal terrace 10 m. USNM 100646 , Riding Rock Reef, Leeward terrace, slope edge, 17 m , San Salvador, Bahamas, 15.viii.1999, col. J.A. Sánchez. ICN-MHN- CO-088 (U-60), Isla Narsa, Capurganá, Gulf of Urabá , Caribbean coast of Colombia, col. J. A. Sánchez, terrace edge 15 m. (30 cm in height) . ICN-MHN-CO- 075 (E-162), Albuquerque Cays, San Andrés and Providencia Archipelago, Colombian Caribbean, col. J.A. Sánchez, vi.1994, outer slope 20 m. INV-CNID- 386, (E-135), Courtown Cays (12°27′ 21.2″N, 81° 27′ 57.2″W), San Andrés and Providencia Archipelago, Colombian Caribbean, col. J.A. Sánchez, 23.vi.1994; fore-reef terrace 20 m. USNM 54910 , (70% ETOH), North Atlantic Ocean, Caribbean Sea; Jamaica, Portland Point (17° 35′ 54″N, 077° 21′ 48″W), St. No. 1220, 24– 27 m , 6.vii.1970, Pillsbury R / V, University of Miami . USNM 97692 (24), isla Tesoro, islas del Rosario, Caribbean coast of Colombia, col. J.A.Sánchez & A.Ramirez, xii.1992; fore-reef terrace slope 25 m. USNM 92593 , Bahamas, Crooked island, 18 m , 30.ix.1994, col. Jenkins K.M., W. Fenical ( No. 6) .

EUNICEA CALYCULATA ELLIS & SOLANDER, 1786

( FIGS 2M, S, W , 8A, G )

Gorgonia calyculata Ellis & Solander, 1786: 95 (holotype: British Museum).

Eunicea calyculata: Bayer, 1961: 154 .

Description: Tall colonies up to 1.8 m in height with thick and long branches ( Fig. 8A, G ). Branches cylindrical and flexible ranging between 6 to 20 mm thick. High polymorphism in branch thickness and calyx. Calyx either present or not, some with only projecting aperture borders, others prominent with hermetic seals or conic walls. Several colours: reddish ochre, beige, and dark ochre. Middle sclerite layer composed of transparent spindle sclerites of 0.6–2 mm in length sometimes curved and widened at middle. Axial layer sclerites pale violet spindles up to 0.25 mm length with a few but notable ornamentations ( Fig. 2S ). Anthocodial sclerites numerous; wide collar with rod sclerites of about 0.1–0.5 mm length ( Fig. 2W ).

Diagnostic features: The surface sclerites contain two discrete kinds of club sclerites. There are small and enlarged clubs with multiple and radial rows in the handle ending in a sharp head, reaching a length up to 0.12 mm ( Fig. 2M ). In addition, torch clubs are present with just one row of ornamentations in the handle.

Comments: Because the different calyx morphotypes described above usually live in the same habitat, it is possible that this taxon is a complex of several species. The morphotype E. coronata described by Bayer (1961) is very likely to be a valid and different species. Notwithstanding the huge variation of Eunicea calyculata , the morphotype E. coronata does not match either the sclerites or the external variation of the former. Morphotype E. coronata ’s branches are significantly thinner than E. calyculata . The sclerites are quite different with larger anthocodial rods as well as club sclerites. This morphotype deserves a careful revision, hopefully including some sort of ecological information. I reviewed the type of the morphotype but new observations and material are needed from its geographical range (Florida, both Atlantic and Gulf Mexico).

Distribution and habitat: Caribbean province excluding some places from the West coast of the Gulf of Mexico. Fore-reef terraces and seaward reefs with moderate exposition to wave and currents between 10 and 30 m in depth.

Examined material: ICN-MHN-CO-090 (E-83), Albuquerque Cays (12° 12′ 07″N, 81° 50′ 43″W), Colombian Caribbean, 5.vi.1995, col. J.A. Sánchez, fore-reef terrace 18 m. USNM 97694 (26), Tesoro island , Rosario islands , Colombian Caribbean, col. J.A. Sánchez & A. Ramirez, 16.ix.1992; fore-reef terrace edge, mixed coral zone 15 m. USNM 97695 (27) Tesoro island , Rosario islands , Colombian Caribbean, col. J.A. Sánchez & A. Ramirez, 16.ix.1992; fore-reef terrace edge, mixed coral zone 15 m. INV-CNID-370 (E-85), Bolivar Cays (Courtown) (12° 24′ 25″N, 81° 29′ 24″W), Colombian Caribbean, 24.v.1995, leeward terrace; col. J.A. Sánchez. INV-CNID-369 (F-1), Isla Fuerte, bajo ‘el bobito’, Colombian Caribbean, 3.x.1995, col. J.A. Sánchez, marginal terrace 10 m. USNM 51741 , Hipotype, Florida, Soldier Key, col. Bayer F.M., 10.x.1960. USNM 87107 , Gulf Of Mexico , United States, Florida, Cape Sable, (25° 17′ 32″N, 81° 39′ 49″W) 13 m, col. LGL Ecological Research Associates , 15.v.1984, Suncoaster R / V, Expedition Southwest Florida Shelf , Minerals Management. USNM 50079 , Bermuda, col. Deichmann E., 1951, donor: Museum Comp. Zool. Harvard. USNM 51981 , Bermuda, Bermuda Island, Southampton, South Shore, 24–27 m, col. Tucker T., 3.ix.1960, donor: Peterson L.W .

Figures

Figure 2. Scanning electron micrograph images of sclerites of Eunicea spp. and Plexaura spp. A–D, sclerites (ornamented spindles) from the middle layer: A, Eunicea flexuosa (Lamouroux, 1821) (scale bar = 0.2 mm); B, Eunicea laciniata Duchassaing & Michelotti (scale bar = 100 Mm); C, Plexaura kuna Lasker et al. (scale bar = 0.1 mm); D, Eunicea clavigera (scale bar = 0.2 mm). E–P, club sclerites from the coenenchyme surface: E, P. kuna (scale bar = 0.1 mm); F, stereopair, Eunicea laxispica (Lamarck) (scale bar = 0.02 mm); G–H, E. laxispica (scale bar = 0.05 mm); I, Eunicea mammosa Lamouroux (scale bar = 0.02 mm); J, E. flexuosa (scale bar = 0.04 mm); K, E. clavigera Bayer (scale bar = 0.01 mm); L, Eunicea pallida Garcia-Parrado & Alcolado (scale bar = 0.02 mm); M, Eunicea calyculata (Ellis & Solander) (scale bar = 0.02 mm); N, Eunicea tourneforti Milne Edwards & Haime (scale bar = 0.02 mm); O, Eunicea fusca Duchassaing & Michelotti (scale bar = 0.01 mm); P, Eunicea tayrona sp. nov. (scale bar = 0.01 mm). Q–U, axial cortex layer sclerites: Q, stereopair, E. laxispica (scale bar = 0.02 mm); R, E. tourneforti (scale bars = 0.06 mm, respectively); S, E. calyculata (scale bar = 0.05 mm); T, Eunicea pallida (scale bars = 0.02 mm); U, E. clavigera. V–Z, polyp sclerites: V, anthocodial rods, E. tourneforti (scale bar = 0.06 mm); W, E. calyculata (scale bar = 0.06 mm); X, E. clavigera (scale bars = 0.01, 0.05, and 0.02 mm); Y, E. mammosa (scale bar = 0.06 mm); Z, octoradiates from the tentacles (stereopair), E. asperula (scale bar = 0.01 mm).

Figure 4. Photographs of Eunicea clavigera. A–B, ‘thick’ morphotype (USNM 100644), A, underwater view of the colony alive, Riding Rock Reef, Leeward terrace, 17 m, San Salvador, Bahamas, 15 August 1999 (diver L. Kauffman); B, dry specimen (calliper open at 2 cm). C–D, ‘slender’ morphotype specimen (USNM 100645). C, dry specimen (calliper open at 2 cm); D, underwater views of the colony alive, Riding Rock Reef, Leeward terrace, 12 m, San Salvador, Bahamas, 15 August 1999. Right photos show the same branch with different levels of polyp retraction.

Figure 5. Scanning electron microscope images from sclerites of Eunicea clavigera ‘slender’ or typical morphotype (USNM 100645). A, sclerite (ornamented spindle) from the middle layer (scale bar = 0.1 mm) with detail of wart ornamentation (scale bars = approximately 0.5 and 0.05 mm, respectively); B, club sclerites (foliate) from the surface layer (scale bars = approximately 0.005 mm); C, rod sclerites from the anthocodial crown/armature of the polyps (scale bars = approximately 0.015 mm); D, rod sclerites from the anthocodial crown/armature of the polyps (scale bars = approximately 0.005 mm).

Figure 6. Scanning electron microscope images from Eunicea clavigera thick morphotype (USNM 100644). A–B, sclerites (ornamented spindles) from the middle layer (scale bars = 0.1 mm) with detail of wart ornamentation (scale bars = 0.01 mm); C–D, club sclerites (foliate) from the surface layer (scale bars = 0.01 mm); E, rod sclerites from the anthocodial crown/armature of the polyps (scale bars = 0.01 mm).

Figure 7. Scanning electron microscope stereopairs of Eunicera clavigera sclerites (‘slender’ morphotype USNM 100645). A, sclerite, spindle, from the middle layer (scale bar = 0.2 mm); B, detail from A, wart ornamentation (scale bar = 0.02 mm); C, axial layer sclerites (scale bar = 0.02 mm); D, warts from spindles of the middle layer (scale bar = 0.01 mm); E, axial layer sclerites (scale bar = 0.01 mm); F, rod sclerites from the anthocodial crown/armature of the polyps (scale bar = 0.02 mm); G–H, club sclerites (foliate) from the surface layer (scale bar = 0.01 mm).

Figure 8. Photographs of Caribbean candelabrum corals in situ. A, Eunicea calyculata (Ellis & Solander), Serrana Bank, Colombia, 18 m; B, Eunicea knighti Bayer, Bocas del Toro Panama, 15 m; C, Eunicea pallida García-Parrado & Alcolado, Bocas del Toro, Panamá, 15 m; D, Eunicea flexuosa (Lamouroux), Carrie Bow Cay, Belize, 12 m. Polyp details (Bocas del Toro, Panamá): E, Eunicea tourneforti; F, Eunicea laciniata; G, E. calyculata; H, Eunicea clavigera; and I, Eunicea tayrona sp. nov.