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Black Corals of the Flower Garden Banks National Marine Sanctuary

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Black Corals of the Flower Garden Banks National Marine Sanctuary Gulf of Mexico Science, 2016(1), pp. 47–67 Black Corals of the Flower Garden Banks National Marine Sanctuary DENNIS M. OPRESKO,MARISSA F. NUTTALL, AND EMMA L. HICKERSON Black corals (Cnidaria: Anthozoa: Antipatharia) of the Flower Garden Banks National Marine Sanctuary (FGBNMS) are described. A key to the species is provided and supplemented with in situ and laboratory photos. The microscopic skeletal features of the species are illustrated from photographs taken with a scanning electron microscope. Thirteen species of black corals are found in the FGBNMS, belonging to the families Antipathidae (Antipathes furcata Gray 1857; Antipathes atlantica Gray 1857, Stichopathes luetkeni Brook 1889; Stichopathes pourtalesi Brook 1889, and Stichopathes sp.); Aphanipathidae [Aphanipathes pedata (Gray 1857), Elatopathes abietina (Pourtale`s 1874), Acanthopathes thyoides (Pourtale`s 1880) and Phanopathes expansa (Opresko and Cairns 1992)]; and Myriopathidae [Plumapathes pennacea (Pallas 1766), Tanacetipathes tanacetum (Pourtale`s 1880), Tanacetipathes hirta (Gray 1857), and Tanacetipathes cf. thamnea (Warner 1981)]. INTRODUCTION about 20 m. The larvae (planulae) of black corals are hypothesized to be negatively photo- lack corals (Order Antipatharia) are colonial tactic. This has been shown to be the case for one B organisms that produce a branched or species studied in the laboratory (Miller, 1996), unbranched skeletal axis that is similar in general and the same may be true for other shallow- appearance to that of gorgonian corals; species water species, as colonies are often found under of both groups are commonly referred to as overhangs, in crevices, and in underwater caves sea fans, sea bushes, sea feathers, or sea whips. or in areas with high levels of suspended Each colony consists of hundreds to thousands of particulate matter where light intensity is re- individual polyps. Antipatharian polyps are gen- duced (Pe´re`s and Piccart, 1949; Grigg, 1965). erally no more than a few millimeters in diameter The Flower Garden Banks National Marine and each polyp has six simple (unbranched) Sanctuary (FGBNMS) encompasses three geo- tentacles. In contrast, the polyps of gorgonians graphically separate locations south of the Texas have eight tentacles and each tentacle is pinnately and Louisiana coasts on the continental shelf branched like a feather. margin in the Gulf of Mexico (Fig. 1). The East The skeleton of black corals, referred to as the corallum, is organic, and is made of concentric and West Flower Garden Banks are 19.5 km apart layers of protein and chitin (Goldberg, 1978, 1991). and are located approximately 160 km south of Unlike gorgonians, black corals do not produce the Texas/Louisiana border. Stetson Bank is any type of calcium carbonate skeletal structures, located 48 km northwest of the West FGB, and is such as sclerites, in either the skeletal axis or in 85 miles southeast of Galveston, TX. Antipathar- their soft tissues. All black corals do, however, have ians are relatively common in some areas of the skeletal spines, although in some species the spines FGBNMS; 13 species have now been recorded are present only on the smallest branches. The from the sanctuary. They occur at depths ranging spines, rarely more than 0.5 mm in height, range in from 50 to 150 m. shape from being compressed and triangular This paper includes: (1) a key to the FGBNMS (when viewed from the side), to conical, thorn- black coral species; (2) descriptions of the major like, blade-like, needle-like, or even bister-like. In morphological features of the species; (3) in situ some taxa they become forked, multilobed at the photographs to aid in field identification; (4) tip, or even antler-shaped in appearance (Opresko, laboratory photos of the microscopic skeletal 2001, fig 17b). The spines may be smooth or features, which are the key characters used to covered with minute surface features such as identify species; and (5) information on the relative papillae, tubercles, or fine striations, details that abundance of the species within the FGBNMS. may only be observable under high magnification. Regarding the last item, reported abundance for Although a few species of antipatharian corals each species is based on the use of in situ imagery are found in waters as shallow as 5 m, most in deep-reef habitat (Schmahl et al., 2008). species prefer deeper waters, and some live in Imagery was collected over four cruises between the deepest parts of the ocean. In the Atlantic, 2009 and 2012 where surveys were conducted black corals are only rarely found shallower than randomly over the reef, but biasedly restricted to E 2016 by the Marine Environmental Sciences Consortium of Alabama 48 GULF OF MEXICO SCIENCE, 2016, VOL. 33(1) Fig. 1. Location of the Flower Garden Banks National Marine Sanctuary. hard bottom structure. For genera where in situ Abpolypar: Refers to the side of a branch identification to species level is difficult (Sticho- opposite to the side on which the polyps occur pathes spp., Tanacetipathes spp., Acanthopathes (in most species the polyps occur in a single row thyoides, Phanopathes expansa,andAntipathes only on one side of the branch). atlantica), remarks on associated black coral fauna were not included and the distribution of Actinopharynx: A tubular involution of ectoder- species among collected specimens is used to mal tissue that extends down from the mouth infer species abundance within the community, into the polyp. as in situ identifications are not possible. Anastomosing: When adjacent branches or Abundance groups are defined as: branchlets fuse together to form a net-like Abundant: Comprises $20% of observed Anti- pattern. patharia species in in situ imagery, and $40% of Anisomorphic: Not of equal size—specifically collected specimens when in situ identifications referring to the spines on the polypar side of the made to genus level or higher. axis, in which case the spines around the edges Moderate: Comprises 5–20% of observed Anti- of the polyp are longer than those between the patharia species in in situ imagery, and 20–40% of polyps, and sometimes the spines directly below collected specimens when in situ identifications the mouth of the polyp are very small or even made to genus level or higher. absent. Rare: Comprises ,5% of observed Antipatharia Axis: The skeletal material of the corallum, species in in situ imagery, and ,20% of collected a so-called scleroprotein lacking any calcareous specimens when in situ identifications made to elements. genus level or higher. Bottlebrush: A type of branching pattern in which the smallest branchlets, referred to as TERMINOLOGY pinnules, extend out horizontally on all sides of the branches. In some cases the colony forms Some specialized terms are used in the a single upright stem with the pinnules extend- scientific descriptions of antipatharian corals; ing out on all sides and often at right angles to these are briefly described here: the stem. Acicular: Needle-like, used primarily in reference Branchlets: The smallest branches of a colony, to the shape of the spines—long and cylindrical, not of uniform size and arrangement (see with a sharp or rounded tip. pinnules). OPRESKO ET AL.—BLACK CORALS OF THE FLOWER GARDEN BANKS 49 Central canal: a chambered tube (that may mouth (at each end of the sagittal axis of the become hollow or secondarily filled) running polyp). down the center of the axis of the stem and all branches and pinnules. Subpinnulate: Branching pattern in which the stem and branches of a colony have pinnules and Circumpolypar spines: Referring to the spines the pinnules themselves have smaller secondary located around the perimeter of a polyp. and possibly tertiary pinnules, which are referred to as subpinnules. Corallum: A general term referring to the entire colony and particularly to the skeletal part of the Terminal branchlets: The smallest unbranched colony. branchlets on the corallum. Distal: In the direction of the free end of Transverse axis of a polyp: A line extending a branch, branchlet, or pinnule. through the center of the polyp in the direction of the branch, branchlet, or pinnule. Flabellate: Fan shaped, in reference to the shape of the corallum. Transverse diameter of polyp: This is a measure- ment of the size of the polyp in the direction of Hypostomal: Area on the skeletal axis directly the branch (from the distal side of the base of below the mouth of the polyp. the distal tentacles to the proximal side of the Interpolypar: Area on the colony surface be- base of the proximal lateral tentacles). tween adjacent polyps. Tuberculate: Referring to a type of skeletal spine Lateral tentacles: Two pairs of tentacles; one pair that has distinct conical protuberances on its located at the distal end of the polyp, and one surface. pair located at the proximal end of the polyp. TAXONOMIC CLASSIFICATION Mesentery: Partitions of tissue that extend from the interior wall of the polyp to the wall of the Seven families are currently recognized in the actinopharynx. Mesenteries divide the upper order Antipatharia: Antipathidae, Schizopathi- interior part of the polyp into pie-like sections. dae, Cladopathidae, Stylopathidae, Leiopathi- The number of mesenteries is usually 10 (6 dae, Myriopathidae, and Aphanipathidae (Daly primary ones and 4 secondary ones); however, et al., 2007). Representatives of the families in one family the polyps have only 6, and in Antipathidae, Aphanipathidae, and Myriopathi- another family there are 12. dae are found in the FGBNMS. Families are differentiated on the basis of several taxonomic Papillose: Referring to microscopic bumps or characters including the number of mesenteries protuberances present on the surface of the in the polyps, size and shape of the polyps, size spines in some species. and appearance of the tentacles of the polyps, Peristomal: Area surrounding the mouth of the and the morphology of the spines. Within the polyp.
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