To refer to this article use this url: http://www.zoologischemededelingen.nl/83/nr04/a34


Zoologische Mededelingen, 83 (September 2009)

The nephtheid soft coral genus Gersemia Marenzeller, 1878, with the description of a new species from the northeast Pacific and a review of two additional species (Octocorallia: Alcyonacea)

G.C. Williams, L. Lundsten

Gary C. Williams, Department of Invertebrate Zoology and Geology, California Academy of Sciences, 55 Music Concourse Drive, Golden Gate Park, San Francisco, California 94118-4503, U.S.A., gwilliams@cal-academy.org

Lonny Lundsten, Monterey Bay Aquarium Research Institute, 7700 Sandholdt Road, Moss Landing, California 95039-9644, U.S.A., lonny@mbari.org

Keywords: Cnidaria; Anthozoa; Octocorallia: Alcyonacea; Nephtheidae: Gersemia; new species; eastern Pacific.

Abstract


A new species of nephtheid soft coral inhabiting the northeast Pacific Ocean is described from samples collected using remotely operated vehicles (ROVs) and a benthic trawl. Two hundred thirty ROV video observations provide additional information about the biogeographical distribution and habitat characteristics of this new species and are used to supplement the information ascertained from collected specimens. The species described herein is found through a broad range of depths (519-2034 m), has been observed through a latitudinal range of approximately 1581 km (33.10472°N; 47° 51.412’N), and has primarily been observed living upon hard-rock substrate. In addition, we revise the generic designation of the South African nephtheid Litophyton liltvedi and we address the status of the Northeast Pacific nephtheid previously referred to as Gersemia rubiformis.

Introduction


Soft corals of the family Nephtheidae have received limited attention in recent literature. Since morphological characters of several genera have been shown to overlap, revisionary systematics of much of the family is necessary (L.P. van Ofwegen, pers. comm., and Ofwegen & Groenenberg, 2007). Three nephtheid genera have recently been described, Pacifiphyton Williams, 1997, Leptophyton Ofwegen & Schleyer, 1997, and Chromonephthea Ofwegen, 2005.

Species in the genus Gersemia are cold-temperate to polar taxa, ranging from 21 m to depths over 2000 m (present paper). The descriptions of G. antarctica and G. marenzelleri (Kükenthal 1906a and Kükenthal 1906b, respectively) are the most recent descriptions of species belonging to this genus, with the exception of the new and revised species described here. The remaining nine species were described between 1806 and 1900.

Throughout the past twenty years, the Monterey Bay Aquarium Research Institute (MBARI) has been using remotely operated vehicles (ROVs) to explore the deep in the Monterey Bay and other locations in the northeast Pacific Ocean. ROV surveys at nearby seamounts and dive sites within Monterey Canyon have yielded many new species descriptions recently, including several gorgonian corals (Cairns, 2007) and an unusual predatory sponge, Cladorhiza pteron (Reiswig & Lee, 2007).

The new species of Gersemia described in this paper was first discovered and collected at Rodriguez Seamount in 2004 and has since been observed and collected at other seamounts off the coast of California, in Monterey Canyon, and on the continental slope off northern California, central Oregon, and northern Washington (fig. 1).

FIG2

Fig. 1. Distribution of Gersemia juliepackardae spec. nov. Light-blue hexagon represents locations of collected specimens. Red cross represents specimens observed in ROV video only. Contours are 500 m.

Material and methods


Most of the type material was collected using ROV manipulators and is preserved in 75% ethanol. One specimen (CAS 175885) was originally fixed in 10% seawater formalin and later transferred to 75% ethanol. In addition to the specimens collected by ROVs, approximately twelve specimens representing one lot were recently found in the invertebrate zoology collection of California Academy of Sciences (CAS). These specimens were collected by benthic trawl off the Oregon coast in 1973 and were preserved in 75% ethanol. We have found this material to be conspecific with the new taxon. Microscopes used in this study include a Nikon SMZ-10 dissecting microscope, an Olympus CH-2 compound microscope, and a Leo 1400 series scanning electron microscope.

In situ observations of living colonies were made using high-resolution video (Ikegami HDL-40 and Panasonic WV-E550) and digital still (Nikon Cooplpix 990) cameras (figs. 2, 3). Over 250 hours of video recordings from San Juan, Rodriguez, Davidson and Pioneer Seamounts, Monterey Canyon, and the continental slope off northern California were reviewed using MBARI’s Video Annotation Reference System, VARS (Schlining & Jacobsen Stout, 2006). Living Gersemia colonies and associated habitats were identified in video and added to the searchable VARS database. Within VARS, these video observations were merged with ancillary data (latitude, longitude, depth, temperature, and oxygen concentration) that were collected by the ROV at the time of deployment. The VARS query was used to export video observation data for analysis and the data were mapped using ArcGIS 9.3. In addition, six video transects were collected to estimate organism density (Monterey Canyon n=5; Pioneer Seamount n=1). Two parallel red lasers (640 nm) positioned 29 cm apart were used to estimate transect width. Transect length was calculated in ArcView® 3.2 using the Animal Movement Analysis Extension, Version 2, which was used to calculate successive distance between the start and end points of each transect (Hooge & Eichenlaub, 1997).

Systematics


Class Anthozoa Ehrenberg, 1831

Subclass Octocorallia Haeckel, 1866

Order Alcyonacea Lamouroux, 1812

Family Nephtheidae Gray, 1862

Gersemia Marenzeller, 1878

Description.― Colonies erect, arborescent from one main stalk. Polyps relatively congested at the ends of short and narrow terminal branches, an effect that is observed more readily in wet preserved specimens. Polyps non-retractile with calyces, supporting bundles of polyps, and polyp crowns absent. Polyp points present: the polyp walls and tentacle rachises usually with numerous sclerites. Points of the polyp walls sometimes arranged more-or-less en chevron. Stalk sclerite complement commonly includes modified or irregularly-shaped capstan-like structures. Sclerites often coloured. Species azooxanthellate. Worldwide distribution in temperate to polar regions, mostly in colder waters, up to over 1200 meters in depth.

Type species.― Gersemia loricata Marenzeller, 1878: 377.

FIG2

Fig. 2. In situ digital still images of the Gersemia juliepackardae spec. nov.; A. Image of the holotype growing on a dead sponge. Numerous ophiuroids are also seen living upon the sponge; other organisms visible in the image include the white sponge, Sclerothamnopsis compressa (left, background), the soft coral, Anthomastus ritteri (at base of dead sponge), and numerous crinoids, Florometra serratissima. B. Close-up image of the holotype. C. Detail of the holotype. Scale bar in 2A: distance between red laser points = 29 cm.

Gersemia juliepackardae spec. nov.

(figs 2, 3, 4A-B, 5, 6, 7)

Material examined.― Holotype CASIZ 175883; Sample Number T1101-A4; U.S.A., central California, Pioneer Seamount (37.37255 N 123.403252 W); 1004.9 m; 19.vi.2007; MBARI - David A. Clague; one whole specimen attached to a fragment of dead sponge. Paratype CASIZ 175884; Sample Number V3191-A1; U.S.A., central California, Monterey Canyon (36.720495° N 122.005016° W); 1186 m; 10.iv.2008; MBARI - James P. Barry; one whole specimen removed from granitic rock substratum. Paratype RMNH Coel. 39007; Sample Number T1100-A12; U.S.A., central California, Pioneer Seamount (37.368573° N 123.39311° W); 1253 m; 18.vi.2007; MBARI - David A. Clague; one fragment sampled from volcanic lava substratum.
Other material: CASIZ 175885; Sample Number T661-A2; U.S.A., southern California, Rodriguez Seamount (34.055796° N 121.084348° W); 889 m; 28.iv.2004; MBARI - David A. Clague; one fragment sampled from volcanic lava substratum. CASIZ 134171; Field Number OTB-500; U.S.A., central Oregon, off shore (43.00° 21.60’ N; 125.00° 10.20’ W); 1600 m; 4.iv.1973; A. Carey Jr. on board R/V Cayuse via trawl; ca. 12 whole specimens. CASIZ 179446; Container Number 779; U.S.A., northern Washington, Olympic Coast National Marine Sanctuary (47° 51.412’ N 125° 28.363’ W); 879 m; 11.vii.2008; National Oceanic and Atmospheric Administration; four whole specimens attached to a rock.

Description of the holotype growth form.― The colony is attached to a fragment of dead hexactinellid sponge. The preserved and contracted holotype is 88 mm in length and 41 mm in width. Basal stalk width at proximal portion of holdfast approximately 40 mm. Lateral branching begins immediately above the holdfast into two mostly opposite rows of primary lateral branches. The primary branches each give rise to several smaller or secondary branches toward their distal ends.

Polyps.― Polyps mostly concentrated on the outer edges of ultimate branches. Some polyps occur on the primary branches in groups of two or more. Polyps are not retractile but tentacles are retractable into the body of each polyp. Polyps are tubular/cylindrical, mostly 4.5-5.5 mm long by 1.2-1.5 mm wide. Some of the smallest (presumably younger) polyps are approximately 1.5 mm in length. The distal half of polyps that contains the anthocodia are bulbous, more or less oval in shape, salmon pink in colour, and 2.3-3.0 mm long. Distal half of each polyp with eight en chevron points composed of salmon pink sclerites, distinct crown not developed (fig. 4A). The basal halves of polyps are white and mostly tubular. Tentacles with rods and spindles arranged en chevron along the longitudinal median (fig. 4B).

Sclerites.― Sclerites include spindles, club-shaped spindles, needles, branched needles; branched club-shaped spindles; and branched spindles. Polyp body wall sclerites are needles and spindles 0.09-0.51 mm (figs 5B, 6A-K). Tentacle sclerites are mostly irregularly-shaped rods and stout spindles 0.08-0.22 mm (figs 5A, 6L-N). Branched spindles of the tentacles are rare to infrequent in abundance. One end of branching sclerites are weakly bifurcate or trifurcate. Surface of the upper stalk contains eight radiates with some irregular radiates; rarely seven radiates 0.06-0.10 mm (figs 5C, 7A-H). Sclerites of the interior of the middle to upper portions of the stalk absent. Surface of the basal portion of the stalk with eight radiates 0.10-0.12 mm (figs 5D, 7I-P). Rod-like sclerites of the interior of the base of the stalk at the holdfast level extremely sparse.

Colour.― Coloration of the tentacles and the distal regions of the polyps (anthocodiae) is salmon pink; basal portion of polyps white; the holdfast and lower region of the stalk are white; the branches of the stalk and the middle to upper parts of stalk are pale pink. The coloration is permanent and was observed in situ (figs. 2, 3), and in all preserved specimens as well, including those collected in 1973.

FIG2

Fig. 3. In situ images of Gersemia juliepackardae spec. nov., non-types; A. Living colony at Pioneer Seamount on ROV Tiburon dive 892 (digital still image, depth = 977 m). B. Living colony from Rodriguez Seamount on Tiburon dive 661 (digital still image, depth = 971 m). C. Living colony at Pioneer Seamount on Tiburon dive 1100 (digital still image, depth = 1,253 m, specimen was collected as T1100-A12).

Etymology.― The new species is named in honour of Julie Packard, executive director of the Monterey Bay Aquarium, Monterey, California, for her dedication to ocean stewardship and conservation, and for elevating public awareness about the ocean environment.

Distribution of collected specimens.— Rodriguez Seamount (southern California) to northern Washington (fig. 1); 888-1600 meters depth. Of the five specimens collected using ROVs, four were collected from hard rock substrata. One other specimen, the holotype, was collected from where it was attached to a dead hexactinellid sponge. The trawled material, CASIZ 134171, came from unknown habitat type on the continental shelf off central Oregon.

Distribution of material from video images presumed to be of the new species.― San Juan Seamount (southern California) to northern California (fig. 1); 520-2034 meters depth. Of the 230 individuals that were observed in video only (i.e., no specimen was collected) and identified as Gersemia juliepackardae by the second author, the vast majority (95%) were observed on hard-rock substrate including volcanic lavas, outcropping granite, outcropping bedded sandstone, and talus material from volcanic and granitic sources. In addition, several individuals of G. juliepackardae (4.4%) were observed upon either dead or living hexactinellid sponge. Only one organism was observed living upon soft sediment, but it is assumed that this organism was attached to a small rock below the sediment surface as has been observed for other deep-sea corals. Ancillary data that were collected by the ROV during video observations indicate that this species may occupy a temperature range of 1.98 – 6.13 °C (mean 3.86 ± 0.025 SE, n=235), a salinity range of 34.17 – 34.59 psu (mean 34.44 ± 0.008 SE, n=235), and oxygen concentrations of 0.21 – 1.42 mL/L (mean 0.52 ± 0.009 SE, n=235). Video transect length (mean 143.8 m ± 21.75 SE, n=6) and width (mean 0.91 m ± 0.14 SE, n=6) varied. Organism density ranged from 0.0 – 0.28 individuals m-2 (mean 0.09 individuals m-2 ± 0.05 SE, n=6).

Variability.― One Oregon specimen (CAS 134171) has numerous sclerites in the interior of the base of the stalk (the region just above the holdfast). These sclerites are rod-like with robust, thornlike, elongated tubercles.

Remarks.― Until now only three species have originally been described in the genus Gersemia: Gersemia loricata Marenzeller, 1879 (type species) from the Arctic Ocean, G. marenzelleri Kükenthal, 1906b, from the Sea of Japan, and G. subtilis Tixier-Durivault, 1961, from the tropical Atlantic Ocean. Other species that are possibly assignable to the genus Gersemia have been described in various genera including Alcyonium, Eunephthya, Nephthea, Paraspongodes, and Voeringia. Gersemia juliepackardae spec. nov. differs from the previously described species of Gersemia as follows. Polyps of the new species have eight points but no crown development (fig. 4A), whereas the polyps of G. subtilis have well-developed crown and points (Tixier-Durivault, 1961: 246, fig. 11f). Stalk sclerites of the new species are mostly eight radiates and 0.07-0.12 mm long, while stalk sclerites of G. marenzelleri are spindles and clubs 0.12-0.20 in length (Kükenthal, 1906b: 287), and those of G. loricata are robust spindles and rods (Marenzeller, 1878: pl. 3, fig. 3A). Eunephthya antarctica Kükenthal, 1906a superficially resembles Gersemia juliepackardae spec. nov. (Kükenthal, 1906a: pl. 3, figs. 14, 15), but the new species differs in having eight radiates in the stalk, whereas the sclerites of E. antarctica are mostly rods and robust spindles with pronounced tubercles (Kükenthal, 1906a: pl. 12, figs. 73, 74).

Gersemia, Litophyton, and Alcyonium

(figs 4C-D)

Gersemia liltvedi (Verseveldt & Williams, 1988), comb. nov.

Material examined.— CASIZ 108791; Litophyton sp.; Japan, Ryukyu Islands, Okinawa, Seragaki, 1.3 km ENE of Maekizaki (26.00˚ 30.40’N 127.00˚ 52.60’E); 58 m; 09.xi.1996; coll. R.F. Bolland, SCUBA; one whole specimen. CASIZ 167912; Gersemia liltvedi; South Africa, Cape Province, Port Elizabeth, Thunderbolt Reef, 21-24 m; 16.ii.1999; J. Starmer, SCUBA; one whole specimen.
FIG2

Fig. 4. A-B. Gersemia juliepackardae spec. nov. A. Polyp armature, paratype. Scale bar = 1.5 mm. B. Tentacle armature, paratype; Scale bar = 1.0 mm C. Litophyton sp. (CASIZ 108791), stalk sclerite; Scale bar = 0.1 mm D. Gersemia liltvedi (CASIZ 167912), stalk sclerites; Scale bar = 0.1 mm.

A soft coral from South Africa was described as Litophyton liltvedi by Verseveldt & Williams (1988: 321; see also Williams, 1992: 337). Subsequent investigation has shown that the proper generic designation should be Gersemia. Species of Litophyton Forskǻl, 1775, have some sclerites of the branches and stalk that are spindles, often with spine ornamentation more pronounced on one side (= caterpillars) (fig. 4C; see also Fabricius & Alderslade, 2001: 106). Gersemia liltvedi comb. nov., differs in having stalk sclerites that are modified capstans (fig. 4D). Species of both genera do not have supporting bundles or crowns in the polyps. However, they differ as follows: Gersemia species are azooxanthellate, circumglobal temperate to polar, colder water taxa, that do not inhabit coral reefs; in mostly deeper water 21-1253 m (with video images depth is increased to 2034 m); sclerites are often coloured; the stalk usually has modified or irregularly-shaped capstan-like sclerites < 1 mm. On the other hand, Litophyton species are zooxanthellate, tropical, warmer water taxa that inhabit coral reef regions of the Indo-Pacific; in shallow water < 50 m; sclerites are colourless; some stalk sclerites are spindles > 0.1 mm. Taking this information into consideration, we here regard Gersemia liltvedi as the correct binomial designation for this taxon.

FIG2

Fig. 5. Gersemia juliepackardae spec. nov., paratype; A. Sclerites from a tentacle rachis, 0.10-0.21 mm in length. B. Sclerites from an anthocodia, 0.36-0.46 mm in length. C. Sclerites from the upper portion of the stalk, 0.06-0.10 mm. D. Sclerites from the base of the stalk, 0.11-0.13 mm. Scale bar for A-D = 0.1 mm.

FIG2

Fig. 6. Gersemia juliepackardae spec. nov., holotype. Scanning electron micrographs of polyp sclerites. A-K. Sclerites from the body of a polyp. A. 0.11 mm. B. 0.31 mm. C. 0.17 mm. D. 0.20 mm. E. 0.21 mm. F. 0.28 mm. G. 0.10 mm. H. 0.09 mm. I. 0.11 mm. J. 0.10 mm. K. Detail of a tubercle from the sclerite shown in G (white arrow); scale bar = 0.004 mm. L-N. Tentacle sclerites. L. 0.15 mm. M. 0.16 mm. N. 0.11 mm.

FIG2

Fig. 7. Gersemia julieparkardae spec. nov., holotype. Scanning electron micrographs of stalk sclerites. A-H. Sclerites from upper portion of stalk. A. 0.10 mm. B. 0.09 mm. C. 0.07 mm. D. 0.09 mm. E. 0.08 mm. F. 0.07 mm. G. 0.10 mm. H. 0.08 mm. I-P. Sclerites from basal portion of stalk. I. 0.10 mm. J. 0.04 mm. K. 0.09 mm. L. 0.08 mm. M. 0.08 mm. N. 0.08 mm. O. 0.08 mm. P. 0.09 mm.

In addition, a species previously referred to by several authors as ‘‘Gersemia rubiformis’’ and the common name of ‘‘Sea Strawberry‘‘ from the Pacific northwest coast of North America (southern Alaska to northern California) is more correctly designated as an unidentified species of Alcyonium (Williams, 2007: 184-185, 188), and will be dealt with in a subsequent publication. The species can be aligned with other species in the genus Alcyonium due to its lobate growth form, the possession of crowded rounded lobules, small retractile polyps that can completely withdraw into the coenenchyme of the lobules and sclerites of densely warted spindles, radiates, and ovoid or clubbed forms similar to other species in the genus, such as Alcyonium gruveli Tixier-Durivault, 1955, A. monodi Tixier-Durivault, 1955, A. grandis Casas et al., 1997, A. southgeorgiensis Casas et al., 1997, and A. valdiviae Kükenthal, 1906 (Verseveldt & Ofwegen, 1992: 161-167; Casas et al., 1997: 304-310; Verseveldt & Williams, 1988: 316-321; Williams, 1992: 292-295).

Discussion


The reliance upon video and still images to supplement the ecological and biogeographical information of single species (Harbison et al., 2001; Matsumoto et al., 2003; Haddock, 2004; Cairns, 2007; Reiswig & Lee, 2007) and, also, to characterize whole faunas (Braby et al., 2007; Lundsten et al., 2009) has become more accepted recently, particularly when describing habitats such as the deep-sea. Here we have used video observations to add to our knowledge of the biogeography and ecology of this new species, with the aim of more accurately characterizing this octocoral.

The known depth and latitudinal ranges of Gersemia juliepackardae is large. Mean depth for collected specimens and video observations, when combined, is 1039 m (± 8.36 SE, n = 235 range 520-2034 m). Video observations also expand the latitudinal range of collected specimens by approximately 100 km to 1686 km (33.10472°N - 47.85686°N). We are confident that the combined latitudinal range of collected specimens and those observed in ROV video only are accurate, however, we suggest that the true latitudinal range may be larger than our data currently show, as this trend has been observed for other organisms found upon the continental slope and at seamounts along the west coast of the USA (Lundsten et al., 2009). Our estimate of organism density was relatively low, however, the video transects were quite long and this tends to lower the estimate for patchily distributed organisms. Localized density was often greater than our estimates indicate, as we did observe G. juliepackardae in densely clustered groups of four to eight individuals.

Surprisingly, the new species described here was often observed living upon dead and live sponges. The second author has observed zoanthid anemones and hydrozoan’s living upon both dead and living corals and sponges, however, in a recent analysis of invertebrate community composition at three seamounts off California (Lundsten et al., 2009), no octocorals, aside from this species, were seen living upon any other organisms. This may represent the first account of a deep-sea octocoral living upon sponges. Other individuals of the new species were in this study were found living upon hard-rock substrata.

Acknowledgements


This work was partially supported by the David and Lucile Packard Foundation contributions to MBARI. We thank principal investigators James P. Barry, David A. Clague, the crew of the research vessels Western Flyer and Point Lobos, the pilots of ROVs Tiburon and Ventana, and Mary Sue Brancato of the Olympic Coast National Marine Sanctuary. We also thank Jei-Ying Chen for her help in the making of scanning electron micrographs and Judith Connor, Kyra Schlining, Nancy Jacobsen Stout, and Susan von Thun for proofreading the manuscript. We also thank the anonymous referees for reviewing the manuscript.

Received: 12.iii.2009

Accepted: 6.v.2009

Edited: L.P. van Ofwegen

References


Braby, C.E., G.W. Rouse, S.B. Johnson, W.J. Jones, & R.C. Vrijenhoek, 2007. Bathymetric and temporal variation among Osedax boneworms and associated megafauna on whale-falls in Monterey Bay, California.― Deep-Sea Research I., 54: 1773-1791.

Cairns, S.D., 2007. Calcaxonian Octocorals (Cnidaria; Anthozoa) from Eastern Pacific Seamounts.― Proceedings of the California Academy of Sciences 58(25): 511-541.

Casas, C., F. Ramil & L.P. van Ofwegen, 1997. Octocorallia (Cnidaria: Anthozoa) from the Scotia Arc, South Atlantic Ocean. I. The genus Alcyonium Linnaeus, 1758.― Zoologische Mededelingen Leiden 71(26): 299-311.

Danielssen, D.C., 1887. Alcyonida. Den Norske Nordhavs-Expedition 1876-1878, 17.― Zoologi, Alcyonida: 1-169.

Ehrenberg, C.G., 1834. Beiträge zur physiologischen Kenntniss der Corallenthiere im allgemeinen, und besonders des rothen Meeres, nebst einem Versuche zur physiologischen Systematik derselben.― Abhandlungen der Königlichen [preussischen] Akademie der Wissenschaften zu Berlin. Aus dem Jahre 1832. Erster Theil: 225-380.

Fabricius, K. & P. Alderslade, 2001. Soft corals and sea fans - a comprehensive guide to the tropical shallow-water genera of the Central-West-Pacific, the Indian Ocean and the Red Sea.― Australian Institute of Marine Science, Townsville. 264 pp.

Harbison, G.R., G.I. Matsumoto, & B.H. Robison, 2001. Lampocteis cruentiventer gen. nov., sp. nov.: A new mesopelagic lobate ctenophore (Phylum Ctenophora, Class Tentaculata, Order Lobata, Family Lampoctenidae).― Bulletin of Marine Science. 68(2): 299-311.

Haddock, S.H.D., 2004. A golden age of gelata: past and future research on planktonic cnidarians and ctenophores.— Hydrobiologia 530/531:549-556.

Hooge P.N. & B. Eichenlaub, 1997. Animal movement extension to ArcView. ver. 1.1.― Alaska Science Center, Biological Science Office, U.S. Geological Survey, Anchorage, AK, USA.

Koren, J. & D.C. Danielssen (eds), 1877. Description of some new Norwegian coelenterates (pages 77-81).― Fauna littoralis Norvegiae 3: 1-163.

Kükenthal, W., 1906a. Alcyonacea.― Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf dem Dampfer “Valdivia” 1898-1899, 13(1) Lieferung 1: 1-111, pls. 1-12.

Kükenthal, W., 1906b. Diagnosen neuer japanischer Alcyonaceen.― Zoologische Anzeiger 30 (8/9): 280-289.

Lundsten L., J.P. Barry, G.M. Cailliet, D.A. Clague, A.P. DeVogelaere, & J.B. Geller, 2009. Benthic Invertebrate Communities on Three Seamounts off Southern and Central California, USA.― Marine Ecology Progress Series 374: 23-32.

Marenzeller, E. von, 1878. Die Coelenteraten, Echinodermen und Wurmer der K.K. Österreichisch-Ungarischen Nordpol-Expedition.― Denkschriften der Kaiserlichen Akademi der Wissenschaften, Mathematisch-Naturwissenschaftliche Classe 35: 357-398.

Matsumoto, G.I., K.A. Raskoff, & D.J. Lindsay, 2003. Tiburonia granrojo n. sp., a mesopelagic scyphomedusa from the Pacific Ocean representing the type of a new subfamily (class Scyphozoa: order Semaeostomeae: family Ulmaridae: subfamily Tiburoniinae subfam. nov.).― Marine Biology 143 (1): 73-77.

May, W.,1900a. Beitrage zur Systematik und Chorologie der Alcyonaceen.― Jenaische Zeitschrift Naturwissenschaft 33 (Neue Folge 26): 1-180.

May, W., 1900b. Die arktische, subarktische und subantarktische Alcyonaceenfauna.― Fauna Arctica 1: 379-408.

Ofwegen, L.P. van, 2005. A new genus of nephtheid soft corals (Octocorallia: Alcyonacea: Nephtheidae) from the Indo-Pacific.― Zoologische Mededelingen Leiden 79(4): 1-236.

Ofwegen, L.P. van & D.S.J. Groenenberg, 2007. A centuries old problem in nephtheid taxonomy approached using DNA data (Coelenterata: Alcyonacea).― Contributions to Zoology 76(3): 153-178.

Ofwegen, L.P. van & M.H. Schleyer, 1997. Corals of the South-west Indian Ocean V. Leptophyton benayahui gen. nov. & spec. nov. (Cnidaria Alcyonacea) from deep reefs off Durban and on the KwaZulu-Natal coast, South Africa.― South African Association of Marine Biological Research, Oceanographic Research Institue Investigational Report 71: 1-12.

Rathke, J., 1806. In: Müller, O.F., 1788-1806. Zoologia danica, seve Animalivm Daniae et Norvegiae rariorum ac minvs notorvm descriptiones et historia … Avetore Othone Friderico Müller … Ad formam tabvlarvm denvo edidit frater avctoris. Vol. 4: Descripserunt et tabvlas dedervnt P.C. Abildgaard, M. Vahl, J.S. Holten, J. Rathke (Page 20).― Havniae, typis N. Mölleri aulae regiae typographi et filii 1788-1806.

Reiswig, H.M. & W.L. Lee, 2007. A new species of Cladorhiza (Porifera: Cladorhizidae) from S California (USA). pp. 517-523 in: Custodio, M.R., G. Lobo-Hajdu, E. Hajdu, G. Muricy (eds.), Porifera Research: Biodiversity, Innovation & Sustainability.― Proceedings of the 7th International Sponge Symposium, Buzios, Brazil, May 2006.

Sars, M., 1860. Om nogle nye eller lidet bekjendte norske Coelenterater.― Forhandlinger i Videnskabs-selskabet i Christiania, 1860: 140-151.

Schlining, B. & N. Jacobsen Stout, 2006. MBARI’s Video Annotation and Reference System. In: Proceedings of the Marine Technology Society / Institute of Electrical and Electronics Engineers Oceans Conference.― Boston, Massachusetts: 1-5.

Studer, T., 1891. Note preliminaire sur les alcyonaires provenant des campagnes du yacht “l’Hirondelle” 1886, 1887, 1888. Part 2. Alcyonacea and Pennatulacea.― Memoires de la Societe zool. de France 4(2): 86-95.

Tixier-Durivault, A., 1961. Les octocoralliaires du Golfe de Guinee et des Iles du Cap-Vert (Alcyonacea, Pennatulacea). Campagne del la “Calypso”: Golfe de Guinee.― Annales de l’Institut Océanographique 39: 237-262.

Verseveldt, J. & G.C. Williams, 1988. A redescription of the soft coral Alcyonium valdiviae Kükenthal, 1906, with the description of a new species of Litophyton.― Annals of the South African Museum 97(12): 315-328.

Verseveldt, J. & L.P. van Ofwegen, 1992. New and described species of Alcyonium Linnaeus, 1758 (Anthozoa: Alcyonacea).― Zoologische Mededelingen 66(7): 155-181.

Williams, G.C., 1992. The Alcyonacea of southern Africa. Stoloniferous octocorals and soft corals (Coelenterata, Anthozoa).― Annals of the South African Museum 100 (3): 249-358.

Williams, G.C., 1997. A new genus and species of nephtheid soft coral (Octocorallia: Alcyonacea) from the western Pacific Ocean, and a discussion of convergence with several deep-sea benthic organisms.― Proceedings of the California Academy of Sciences 49(12): 423-437.

Williams, G.C., 2007. Anthozoa: Octocorallia. Pages 184-188 in J.T. Carlton (editor), Intertidal invertebrates from central California to Oregon.― University of California Press, Berkeley, California, U.S.A. 1001 pp.