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Revision of the Hawaiian Stylasteridae (Cnidaria: Hydrozoa: Athecata)1

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Revision of the Hawaiian Stylasteridae (Cnidaria: Hydrozoa: Athecata)1 Revision of the Hawaiian Stylasteridae (Cnidaria: Hydrozoa: Athecata)1 Stephen D. Cairns2 Abstract: Four species of stylasterids are described from the Northwestern Ha- waiian Islands (Kure to Kaua‘i) at depths of 293–583 m, including three new species: Distichopora asulcata, Stylaster griggi, and S. infundibuliferus. In addition, specimens of Distichopora anceps were observed and collected, showing it to be the most common macroinvertebrate on the northwestern slope of Laysan Island but not known from any other Hawaiian locality. Its description was amended to include branching colonies with up to 20 lobes; a suggested ontogeny of these growth forms is illustrated. Also, unique sexually dimorphic features of both male and female ampullae of D. anceps are described. Although there had been several undoc- environments, preferably living on seamounts umented reports of stylasterid corals from or on the slopes of volcanic islands having a the Hawaiian Islands before 1978, Boschma landmass less than 36,000 km2. Such oceanic (1959:134, 1964:186) surmised that they regions usually have a constant salinity, a low were all based on inaccurate reports or suspended sediment load, hard substrate, and concerned specimens that were bought in low nutrient levels, all factors that seem to fa- Hawai‘i but probably transported from an- vor stylasterids and all factors that are com- other island group, such as the Marquesas, mon throughout the Hawaiian Islands. The to be sold to tourists in O‘ahu. Boschma largest of the Hawaiian Islands, Hawai‘i, is (1959:134) finally concluded: ‘‘Taking these about 10,400 km2 in area. Thus, one might data into consideration it seems fairly con- expect stylasterids to occur throughout the clusive that Stylasterine corals do not occur archipelago, and when the deepwater envi- in the Hawaiian Islands.’’ However, the first ronment is better known I expect that many and, until now, only documented record of a more species will be reported. stylasterid from the Hawaiian Islands was the report of four specimens of Distichopora anceps materials and methods Cairns, 1978, from 360 to 402 m (corrected depth) off Laysan. In this report three addi- Most of the specimens reported herein and tional species are reported for the Northwest- the impetus for this study resulted from an ern Hawaiian Islands, bringing the total for oceanographic expedition to the Northwest- Hawai‘i to four species, but the paucity of ern Hawaiian Islands during October and stylasterids from this archipelago is still sur- November 2003, in which I participated. prising for a region of this size and character- The support vessel, the R/V Ka‘imikai-o- istics. Cairns (1992) mapped the worldwide Kanaloa of the Hawaiian Underseas Research distribution of all reported stylasterids, show- Laboratory, was the base for dives of the ing that they occur predominantly in insular three-person submersible Pisces V, rated to 2,000 m, and the RCV-150, a tethered re- motely operated vehicle rated to 914 m. Specimens were also obtained from a series 1 Manuscript accepted 8 August 2004. 2 Department of Invertebrate Zoology, MRC 163, of cruises of the R/V Townsend Cromwell dur- National Museum of Natural History, Smithsonian Insti- ing 1970–1972, called the SANGO expedi- tution, P.O. Box 27012, Washington, D.C. 20013-7012 tions. Several specimens were also obtained (e-mail: [email protected]). from the private collections of Beatrice and Tom Burch. Pacific Science (2005), vol. 59, no. 3:439–451 The scanning electron photomicrographs : 2005 by University of Hawai‘i Press were made by me on an AMRAY 1810 scan- All rights reserved ning electron microscope. Most specimens 439 440 PACIFIC SCIENCE . July 2005 are deposited at the National Museum of RCV-150, dive 252, 25 50.49 0 N, 171 Natural History, Smithsonian Institution; 57.822 0 W, 396–397 m, 15 October 2003, some duplicates are also deposited at the observations only (no specimens collected); Bishop Museum. RCV-150, dive 253, 25 50.5190 N, 171 The following abbreviations are used in 57.668 0 W, 362–381 m, 15 October 2003, the text: BPBM, Bernice Pauahi Bishop observations only (no specimens collected); Museum, Honolulu, Hawai‘i; H :W, height- RCV-150, dive 254, 25 50.4700 N, 171 to-width ratio of a gastrostyle; NMNH, Na- 57.655 0 W, 353–388 m, 15 October 2003, 1 tional Museum of Natural History (formerly gender indet., usnm 1021951. USNM), Smithsonian Institution, Washing- remarks. This uniquely shaped stylas- ton, D.C.; RCV, a tethered remotely con- terid was described and redescribed by Cairns trolled vehicle; SANGO, a series of cruises (1978, 1983), who included aspects of its of the R/V Townsend Cromwell (sango is the morphology and microstructure (SEM), zo- Japanese word for coral); USNM, United oid structure, and nematocysts. Cairns and States National Museum (now the National Macintyre (1992) later added observations on Museum of Natural History, Smithsonian its mineralogy, but all of these observations Institution). were based on only four type specimens, two of which were dead and somewhat worn species account when collected. The 46 additional colonies and fragments reported herein allow substan- Class Hydrozoa tial amplification of the original description Order Filifera of several characters, which are described Superfamily Hydractinoidea Bouillon, 1978 here. Family Stylasteridae Gray, 1847 Originally the species was thought to con- sist of a stem that supported two flabellate Distichopora anceps Cairns, 1978 lobes and was thus called anceps, meaning Figures 1A–H, 2A–I ‘‘two headed.’’ The collected specimens and observation of thousands more in situ show Distichopora (Haplomerismos) anceps Cairns, that, although the two-headed form is the 1978:84–86, figs. 1–6; 1983:474–476, fig. most common shape of this species, constitut- 16A–G, 24C, 25F, 28E (histology); ing probably more than 90% of the observed 1991:17 (deposition of types).—Cairns colonies, the species is capable of forming and Macintyre, 1992:98 (mineralogy). larger colonies (up to 12 cm in height and a Distichopora anceps.—Cairns, Hoeksema, and basal diameter of 12.7 mm [Figure 1A]) hav- van der Land, 1999:42 (listed). ing many more lobes (e.g., more than 20). Figure 2A–E show diagrammatically some of material examined. Pisces V-533a, 25 the growth forms observed and a suggested 52.041 0 N, 171 57.8480 W, 501–577 m, 19 growth transition for colonies having one, October 2003, 19 female colonies or frag- two, three, four, and eight lobes. Some colo- ments, 16 male colonies or fragments, SEM nies exhibit asymmetric growth (Figure 2F– stubs 1101–1103, usnm 1021949, 1 male and I ), which is most likely the result of envi- 1 female colony, bpbm D1071; Pisces V-533b, ronmental conditions. All specimens were 25 52.046 0 N, 171 57.839 0 W, 481–485 m, uniformly white. 19 October 2003, 4 female colonies, 2 male As noted in the original description, the colonies, 2 gender indet., usnm 1021950; flabellar faces and pedicel are covered with RCV-150, dive 250, 25 53.90 0 N, 171 closely spaced, parallel ridges. These ridges 55.280 W, 429–495 m, 14 October 2003, are 0.3–0.7 mm apart, a new one intercalating observations only (no specimens collected); as the flabellum increases in width; the ridges RCV-150, dive 251, 25 50.53 0 N, 171 themselves are prominent, up to 0.40 mm in 58.098 0 W, 370–403 m, 14 October 2003, height, and sharp-edged. Most colonies also observations only (no specimens collected); bear numerous, small, homogeneously placed Figure 1. Distichopora anceps from Pisces V-533a: A, a large colony with multiple lobes, Â0:55; B, surface depressions of 15 male ampullae, Â17; C, longitudinal fracture through two male ampullae, Â38; D, several conical nematopores between two flabellar ridges, Â90; E–F, several female ampullae nestled between flabellar ridges, some efferent pores visible, Â11, Â25, respectively; G, stereo pair of longitudinal fracture through two female ampullae showing incipient efferent pore from within ampullae, also cross section of pore row showing dactylopores and gastropores, Â17; H, higher magnification of two female ampullae of G, Â35. 442 PACIFIC SCIENCE . July 2005 A C B D E F G H I Figure 2. Diagrams of colonial forms of Distichopora anceps, synthesized from in situ RCV observations: A–E, sug- gested ontogeny from two- to eight-lobed colony form; F, an asymmetrical one-lobed colony; G, an asymmetrical three-lobed colony; H, an asymmetrical four-lobed colony; I, a large, well-developed, asymmetrical 20-lobe colony. Diagrams not drawn to scale. papillae that cover the flabellar faces, each often contains one yellow egg. Circular effer- conical mound about 0.10 mm in diameter ent pores, measuring 0.10–0.15 mm in diam- and 0.06 mm in height, each bearing a small eter, are often found at both the distal and slit at its apex, the slit about 25 mm in length proximal ends of the ampulla, flush with the and 11 mm wide. These are presumed to be coenosteal surface, implying that two eggs/ nematopores (Figure 1D). larvae may develop simultaneously. This is Sexual dimorphism and differentiation was unique in the Stylasteridae. Often a new fla- unclear in the type specimens, due to their bellar ridge will originate on top of a female immature size and worn condition, but of ampulla and continue to the flabellar edge. the 46 newly collected specimens, 24 were The holotype can now be determined to be a female, 19 male, and three of indeterminate female colony. Male ampullae are completely gender, making it possible to fully describe internal and much smaller but also occur lin- the sexual dimorphism of this species, which early between the flabellar ridges.
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