Branches of colonies slender. Polyps monomorphic, with prominent calyces, anthocodiae usually exsert. Axis not jointed, without a cross-chambered central core. Medulla surrounded by longitudinal boundary canals (of roughly equal size) separating it from cortex; medulla only rarely perforated by gastrodermal solenia in smaller branches and even then, not as extensively as in lower parts of colony; in larger branches, medulla perforated by solenia. Generally, medulla with separable sclerites; medullar sclerites stout spindles (not needle-like), thorny, ornamented with warts, spines or branching processes, that may link sclerites together. Sclerites of coenenchyme longer fusiform spindles, sometimes clavate or bent, occasionally with radiate bodies and capstans (rarely).

Within the Subclass Octocorallia, taxonomic placement of this family reflects the changeable history the Order Alcyonacea has experienced since its inception. Currently, Alcyonacea is one of three orders in the subclass (Williams and Cairns 2014). The current Order Alcyonacea was, however, originally divided into four orders (Alcyonacea, Gorgonacea, Stolonifera, and Telestacea). Current coral taxonomy now divides Order Alcyonacea into five nominal groups: Calcaxonia, Holaxonia, Scleraxonia, Alcyoniina and Stolonifera (Bayer 1981c, Fabricius and Alderslade 2001). While the Family Anthothelidae is today recognized as valid in the Order Alcyonacea [Scleraxonia], a number of species in the family were originally placed in the older Order Stolonifera, often within the Family Clavulariidae. A few researchers may still group some of the families of soft corals in an Order Stolonifera, but since then, a number of genera and several species have been moved out of Stolonifera. Fabricius and Alderslade (2001) noted that the “decision whether to categorize a particular genus as a stoloniferan becomes so subjective that the name plainly has limited classificatory value . . . . .” Use of the Order Stolonifera, and placement of the Family Anthothelidae in it, or the nominal group, Scleraxonia (as opposed to Stolonifera), has had a continued, tumultuous history (Hickson 1915, Molander 1918, Kükenthal 1919, 1924, Madsen 1944, Bayer 1961, Bayer 1981a, b, c, Hochberg 1979). Currently, the classification of species in the family is determined by the presence of a membranous colony form, presence, or absence of coenenchyme layers surrounding an axis, and the way in which polyps arise from the membranous base.

Any membranous octocoral colony currently held in the SBMNH collection (few in number, small in size, deteriorated due to early formalin storage) could be a member of either the genus Clavularia or the genus Anthothela (the latter, a genus within the Order Alcyonacea, Scleraxonia). A detailed examination of the few colonies held in the SBMNH collection, in comparison with material housed elsewhere is needed; that necessitates a separate project, to be undertaken at some future date. Most of the material in the SBMNH collection with membranous colony form is present in a very fragile state; a more detailed description for each will not be easy, in some instances, not possible at all. It is likely that even with a more thorough examination of the material held at SBMNH, the results will necessarily be inconclusive. A complete revision of the genus Anthothela was recently completed by Moore et al. (2017), utilized here.

Briareum grandiflorum Sars, 1856b (by subsequent designation).

With slender, rounded, tortuous, commonly upright, abundant anastomosing branches producing tangled colonies. Branches always solid; no main stems developed, branches grading upwards from broadened membranous base. Polyps present on base as well as on branches, widely scattered on all sides, sometimes clustered into large masses. Polyps elongated in expansion arising from distinctly projecting, short yet elevated cylindrical calyces. Polyps partially retractile, seldom entirely retractile; large anthocodiae commonly preserved exsert, arising from either extended, rather thin, slightly sharp, spiculose, but spongy, basal membrane (encrusting) or from slender irregular stems (branched). Axis spiculose, well differentiated, not firm. Long, strongly warted, often irregular spindles and short, girdled rods in coenenchyme. Sclerites of axis more irregular; bear fewer, larger warts, knobs or lobes. Spongy base filled with thin spindles and rods, permeating tissue. Bright rosy-red or brownish in life, but other color forms likely.

USA, California, China Point, (?) San Clemente Island, SW tip, 50 fm (91 m).

Location of type unknown.

One specimen in SBMNH collection was identified as this species (see Appendix 1: List of material examined).

Colony (Figure 1A) of thin, flattened branches, crooked, tortuous, somewhat anastomosed; no large main stem development. Rather low-lying, prostrate form; height 0.5 mm-7.0 mm; “stolon” thin, membranous; arrangement of medulla and “cortex” as described for family. Calyces (Figure 1B) prominent, projecting cylinders up to 2.5–3.0 mm tall; polyps stout, with highly retractile portion up to 2.5 mm in length; total calyx/polyp size 5.0 mm tall by 2.0 mm wide. Calyces monomorphic, with eight deep, longitudinal grooves, delineating eight ridges or ribs with sharp edges. Distal part of polyp usually exsert, bearing eight double-rows of spindles. Polyps large, rather widely spaced. Color of living colony described as light yellow or straw-colored to grayish white; in alcohol, creamy white. Sclerites (Figure 2) of coenenchyme large, elongate, thin, pointed spindles, often with prominent projections on edges; generally, fusiform (0.25–0.3 mm long), not capstans; often appearing bent. Older descriptions indicate that sometimes these are strongly clavate at terminal end, appearing as clubbed spindles; this condition not seen in material examined. Anthocodial armature strong spindles, often clavate or bent, only rarely as radiate bodies and capstans. Spindles at base of tentacles (collaret), 0.18 mm long, those of calyx 0.22 mm long, none of these club-shaped. Sclerites widely spaced, showing transverse disposition at base of tentacles; sclerites of medulla strong thorny spindles. Colony surface rough to the touch due to projecting sclerites.

Figure 1. 

Anthothela pacifica, SBMNH 265939. A Large portion of colony fragment; several rocks accompanied specimen (presumed substrate); colony unattached, ~7.0 cm in length B close-up of polyps, each polyp ~0.5 mm tall.

Figure 2. 

Anthothela pacifica, SBMNH 265939, SEM image. Coenenchymal sclerites, colored a very pale yellow to white. Length of largest sclerites examined ~400 µm.

Presumably named for type locality, northern Pacific Ocean.

Dwarf white gorgonian.

Not definitively known at this time for coastal western United States; potentially from southern California: USA, California, San Diego, Point Loma, 200 m (Nutting 1909) and China Point, (?) San Clemente Island, SW tip, 91 m, to at least northern California, Monterey Bay, 900 m (Nutting 1909); possibly as far north as southern British Columbia, Canada (Lamb and Hanby 2005).

According to Kozloff (1987), a shallow subtidal form. Lamb and Hanby (2005) state it as “subtidal, below 40 m (133 ft).”

Kükenthal (1913a) indicated species is not equal to Sympodium armatum Wright and Studer as cited in Nutting (1909). However, an interesting aspect presented itself through an unexpected source. A letter, found in SBMNH archives, dated 1978, from Dr Frederick Bayer to Dr Eric Hochberg, indicated the suggestion of a possible synonymy between Anthothela argentea Studer, 1894 and Anthothela pacifica. In that letter Dr Bayer made two significant statements, the first being that the two lots reported and identified (and misidentified) by Nutting (1909: 686) as “Sympodium armatum Wright & Studer, 1889,” subsequently identified by Kükenthal (1913) as Clavularia pacifica, are in actuality gorgonians of the genus Anthothela. Bayer (1961) made mention of this in relation to the Atlantic Ocean species Anthothela tropicalis Bayer, 1961. Secondly (key point of the letter), Bayer believed that it was “possible that ‘Clavularia’ pacifica is identical with Anthothela argentea Studer, in which case Studer’s name takes precedence.” Further reading in Bayer (1961) showed that there was indeed a genus name change. Bayer, discussing family Anthothelidae, genus Anthothela, made the following statement: “A distinctly different species has now been recorded from the Gulf of Mexico, also in deep water, which proves to be a new species closely related to the eastern Pacific Anthothela pacifica (Kükenthal), forming with it an eastern Pacific-western Atlantic twin-pair of species.” Note the change in genus name (from Clavularia to Anthothela). Regarding the distributional range for A. argentea, location records noted on distribution maps posted on the Encyclopedia Of Life website showed it occurring throughout the southeastern Pacific Ocean, extending into the northeastern Pacific, to at least central California (USA; USNM 94428 was collected in proximity of the western edge of the Bight). This indicated a possible geographic overlap with (or the possibility that it was synonymous with) A. pacifica. However, the work of Moore et al. (2017) puts the suggested synonymy of Bayer in question, having reassigned Anthothela argentea to the genus Victorgorgia López-González & Briand, 2002 and the new family Victorgorgiidae, as they indicated that there are clear morphological and genetic differences from the genus Anthothela.

From the World List of Octocorallia, the World Register of Marine Species (WoRMS), Anthothela pacifica is an accepted scientific name, while Anthothela argentea has been accepted as Victorgorgia argentea, and from that listing it is clear that these two are considered separate species (Cordeiro et al. 2018a).

Identification of specimen relied on notes made by Dr Hochberg, with a description given by Kükenthal; fragility of specimen did not permit an extensive examination, but as far as it could be done, one was done with the specimen to hand.

Hickson (1915) lamented: the “widely distributed and very variable genus Clavularia is badly in need of revision. It is probable that such a revision would lead to a considerable reduction in the numbers of species, many of which have been founded on very in-adequate characters.” He went on to say that the genus Clavularia is well represented in the North Pacific Ocean; how many other species belong in the genus Clavularia, and how many may be members of the genus Anthothela, or some other genus, remains to be seen. Bayer (1961:70) affirmed that Nutting’s and Kükenthal’s specimens are gorgonians of the genus Anthothela. Hochberg (1979) stated, “comparison with the type is needed to identify what has been called Clavularia pacifica from this area in the past. The generic change (to Anthothela) is an obscure reference, being only a few lines in Bayer’s work (1961) on Caribbean octocorals.”

Location of type is at issue; someone (unknown) noted: “China Point, ‘San Clemente Island (SW tip).’ ” There was no way to confirm this statement, and there is the added problem of a China Point on Santa Catalina Island (SW side of island), as well. There was no means to identify which “China Point” was the correct collection location.

Robust, profusely branched colonies with dimorphic polyps (feeding autozooids, reproductive siphonozooids). Axial skeletal structure solely a continuous medulla, containing separable sclerites. Medulla perforated by gastrodermal canals all the way to branch tips, not separated from cortex by a ring of boundary canals.

Alcyonium arboreum Linné, 1758; [= by subsequent designation, Paragorgia arborea (Linnaeus, 1758), by monotypy].

Massive, tree-like colonies with thick branches, measuring up to 7.0 meters tall, perhaps as much as 6.0–7.0 meters wide. Sclerites in axial medulla, long, ornate rods (spindles) with branching processes, derived from capstan type, up to 0.6–0.8 mm in length, colorless or pink; elsewhere (coenenchyme, tentacles, etc.) sclerites small (less than 0.15 mm in length), differing shapes, commonly pink or red. Surface sclerites six-, seven-, and eight-radiate capstans, always less than 0.1 mm long, with globular, smooth, grooved or lobulated ornamentation. Sclerites in subsurface/outer medulla of intermediate form, ranging between radiates and spindles. Autozooid polyp tentacles have distinctively blunt, stubby rods or ovals, less than 0.1 mm.

Canada, British Columbia, Vancouver Island, Jervis Inlet, ~20 m.

Holotype YPM-5373 [dry].

~1–2 lots (see Appendix 1: List of material examined).

Collection lot studied contains one branch fragment (Figure 3); in most respects, examination of fragment revealed characters that align with the description given in Sánchez (2005, pages 15–20). The branch is distinctive in its knobby aspect, but sclerites fall well within the parameters of morphology as discussed and shown in Sánchez (2005).

Figure 3. 

Paragorgia arborea var. pacifica, SBMNH 422977. Single branch fragment, ~5.0 cm long, displaying knobby condition of branches seen in some specimens.

The variety name pacifica was presumably proposed in reference to the location/distribution of the species in the Pacific Ocean.

Referred to frequently as “Bubblegum coral.” Cairns et al. (2003) referred to Breeze et al. (1997) where this genus/species is also listed as “Rubber Trees” and “Strawberry Plants.” Specifically, it could be called the “Pacific bubblegum tree.”

Recorded from Alaskan waters, Bering Sea, ‘Albatross’, 54°02'40"N, 166°42'00"W, at a depth of 504 m; USNM 3315. Also, recorded from Unalaska to Kodiak, ‘Albatross’, 54°19'00"N, 159°40'00"W, taken by dredge, 114 m; USNM 3338. Bayer indicated (unpublished ms 2, Cairns 2009) that known distribution of this species was from the Bering Sea south to Vancouver Island (specimen collected by Mr Wm Spreadborough, at Ucluelet, Vancouver Island, BC, at a depth of 16 m, June, 1909. [Col. No. 51, Coelenterates, Victoria Memorial Museum, Ottawa]) on the east side of the Pacific, and likely over to Japan on the west; southern limit (at the time Bayer was writing) was unknown. Based on more recent work, including the review of systematics for the family by Sánchez (2005), this species, in the Pacific Ocean (a possible variant of P. arborea), extends to at least the northern limit of the California Bight on the eastern side (at question, further south, at depth), and down into New Zealand waters on the Pacific Ocean’s western side.

Specimens collected or photographed in the Monterey Bay Marine Sanctuary have harbored polychaete worms, purple in color (species identification not determined), with the worms wound around the branches of the colony (Langstroth and Langstroth 2000). On a specimen examined from Mexico, Gulf of California, Baja, Bahia de Los Angeles, there was a complete over-covering of what may be some sort of grey colonial or zoanthid-type organism (the specimen was not included in the list of material examined, as the overgrowth of the zoanthids precluded any clear examination of the host gorgonian itself). A MBARI video clip, viewed on a visit to MBARI, had an excellent segment of this species heavily colonized by numerous basket stars, so many in fact, that the entire, large tree-like colony displayed a heavy growth of “hair.” According to Langstroth and Langstroth (2000), on a Paragorgia specimen, a feather star, Florometra serratissima, was seen clutching the gorgonian with its leg-like cirri. While this was seen in a lab setting, the feather star likely may have come with the gorgonian during the collection process. If so, other filter-feeding echinoderms might be seen living on/with these gorgonians in situ. Evidence from recent OCNMS expeditions, as well as numerous MBARI and NOAA video clips support this. Colonies living in deeper water grow very slowly in some areas and could be several hundred years old (Andrews et al. 2005, Sherwood et al. 2005), reaching heights of several meters (DeVogelaere et al. 2005). It is speculated (Brancato et al. 2007) that these large, aged colonies provide critical habitat for such organisms as Northern Rockfish, Pacific Ocean Perch, species of King Crab, and Pacific Cod. An expedition undertaken by Olympic Coast NMS (May 2006) lent credence to this speculation.

Sclerite examination of the sample shown in Figure 3 agreed with those seen in Sánchez (2005: fig 9); the six-radiates distinctive of this species were confirmed. Verrill (1922), reporting on specimens of Paragorgia arborea, discussed the possibility of the existence of this variant. Bayer stated in his unpublished manuscript (ms 2, Cairns 2009)) that “P. arborea seems limited to latitudes of 40° or higher . . . . P. arborea seems to be truly bipolar, since no reliable finds have been made south of British Columbia in the Pacific. It is impossible to be sure that the species does not show equatorial submergence.” Thus, “it appears that the genus Paragorgia, an inhabitant of cold waters, . . . whose various species occur at moderate depths in boreal and anti-boreal regions, follow the cool water to greater depths in low latitudes.” Bayer additionally stated that a specimen obtained in British Columbia, as P. pacifica could be a variety of P. arborea. Kozloff et al. (1987) stated that Paragorgia pacifica is “the most commonly encountered gorgonian of” the Pacific Northwest region; “it has been called Paragorgia arborea Linnaeus, 1758.” The final word comes from Sánchez (2005); he stated that a comprehensive review of North Pacific populations of P. arborea, “including type material and genetics is needed before reaching conclusions on P. pacifica and the differentiation of south vs. north P. arborea populations.”

The California Academy of Sciences has approximately 30 lots of this, or other species, attributed to this genus; most specimens are from Alaska; as well, two are from the USSR, one is from Norway, and one is from Oregon. Nine of the remaining lots are specimens collected from California; most are from Monterey Bay, with one from the Davidson Seamount. Of these nine, only three have been identified as being this particular species. MBARI has extensive video records of this species from Monterey Canyon, as well as the Davidson Seamount. In the Moss Landing Marine Labs collection there is a small specimen of what may well be this species, collected in Monterey Bay, ~36°27'12"N, 122°04'02"W, ~450 meters; coll. G McDonald, 13 March 1974; C0067 [wet]. As well, there is an impressive, tree-sized dry specimen on display in the hallway near the museum door. (Collection data may be available for this specimen, but collection data could not be located.)

Based on multiple examinations of possible Paragorgia material in the SBMNH research collection, none (one exception) were examples of P. arborea var. pacifica; numerous sclerite preparations never revealed the six-radiate sclerite form that is characteristic of this species; only a very few displayed eight-radiates. As well, no polyp sclerites were ever obtained. As most of the SBMNH specimens examined clearly lacked the “key” identifying sclerites, no request was made to obtain the holotype specimen from Yale Peabody Museum. P. arborea (though not the variety pacifica suggested by Verrill) is an accepted species in the WoRMS listings (Cordeiro et al. 2018b). As well, the three following species are also accepted species in the WoRMS registry. These are included as they have been collected in very close proximity to the California Bight region. None however, are represented in the SBMNH collection and research indicated that they were never collected on any of the ‘Velero’ expeditions. In point of fact, there were no records of any specimens in the genus Paragorgia having been collected.

No material in SBMNH collection (see Appendix 1: List of material examined).

Inclusion of P. regalis is based on USNM 1027063, which was collected just west of the California Channel Islands, in proximity to western boundary edge of Bight. Moss Landing Marine Labs may have a representative sample of this species in their collection, from Monterey Bay Canyon, 36°25'54"N, 122°00'03"W, ~900 m; coll. D Rold and H King, December 1978; C0074 [wet]. It is possible that some of the Paragorgia seen in MBARI video segments could be this species; this species may have been seen when examination of samples of gorgonian collected by J Barry (MBARI) was done, made available for examination by Lonny Lundsten. Distribution ranges across the Pacific Ocean (based on collection records for specimens in NMNH). Collection depths in Hawaii and Japan range from 452–1840 m (Sánchez 2005), and as noted from NMNH collection. For a common name, this species might be called the “Regal bubblegum tree.”

No material in SBMNH collection (see Appendix 1: List of material examined).

A paratype specimen at NMNH (USNM 94437), identified as this species, was collected in proximity of the southern California Channel Islands, in an area lying close to the western boundary of the California Bight; range seems to extend northward to an area off British Columbia, Canada (location data for USNM 57982, the holotype; discussed in Sánchez 2005). As this species was named in honor of Dr Stephen Cairns of the National Museum of Natural History, Smithsonian, it would be fitting to call this “Stephen’s bubblegum tree.” Any similarities between this and other species in the genus were thoroughly discussed in Sánchez (2005).

No material in SBMNH collection (see Appendix 1: List of material examined).

A specimen at NMNH (USNM 90345), identified as this species, was collected west of the southern California Channel Islands in an area that lies near the western boundary of the California Bight. A MBARI sample taken from Monterey Canyon, near Point Sur, on 7 April 2008 could also be this species. A number of other samples, examined cursorily, housed at MBARI, could be this species.

Sibogagorgia weberi Stiasny, 1937b: 80 [= Suberia köllikeri Nutting, 1911: 13, nec Studer 1879].

A genus in the family Paragorgiidae with scleritic medulla showing no (or very few, one to two) large penetration canals; main solenia around subsurface/outer medulla as boundary canals, forming reticulate network; network of canals observable with light microscopy as a regularly reticulate and uniform mesh just beneath surface. Polyps without tentacular sclerites, outer surface with radiate sclerites; generally, medullary sclerites nearly bare of ornamentation. Autozooid polyps uniformly to randomly distributed along branches, throughout colony. Branching colonies often in one plane (but not always); main branches usually thicker than terminals; terminals clavate. Coloration either of uniform beige or bright orange-red to a beige coloring with slightly projected pinkish orange polyp apertures.

Holotype USA, California, Los Angeles County, West end, Santa Catalina Island, 300 meters. Paratype USA, California, Los Angeles County, NE × E of Long Point, Santa Catalina Island, 415–486 meters.

Holotype SBMNH 422974; Paratype SBMNH 422973.

~8 lots (see Appendix 1: List of material examined).

Specimens rarely displayed growth in one plane. Sclerites of medulla with blunt tips, bearing minimal ornamentation, smooth in areas between widely spaced spiny processes; sclerites of colony surface and coenenchymal tissue intermediate between surface and medulla 7-radiates, but never 8-radiates. Thick, compact branches with color variation from pinkish orange to pale pink.

Colony (Figure 4, 6) fragments robust, tree-like, with thick, conspicuous branches. Specimen of Figure 4 approximately 18 cm tall, that of Figure 6 roughly 36 cm long (when gently stretched out). Coenenchyme is thick and tough (like cutting through raw carrot). Branches moderately smooth in appearance, although lumpy in many spots, with small calyces evident; appear somewhat moderately spaced, scattered irregularly, on all sides of branches; distal or lateral branch tips each end with round, swollen knob. Color of branch coenenchyme (Figure 4) bright reddish orange; specimen shown in Figure 6 creamy beige with orange polyp apertures (this could be normal color or could have bleached out due to earlier storage solutions); in both specimens, polyps of same orange hue, with tentacles white (more visible in specimen of Figure 4). Cross section samples of both colonies revealed obvious boundary canals, and both colonies have very few, but rather large penetration canals in the medulla. No blunt, stubby, ornate polyp tentacular sclerites (rods) were ever found in any of numerous tissue samples examined; outer surface sclerites are radiate (Figures 5C, 7C), most closely matching a 7-radiate configuration, with ornamentation often jagged and extensive; color of these sclerites a pale pinkish orange; medullary sclerites (Figures 5A, 7A) are spindles, with moderate ornamentation, not as bare as seen in most other species of the genus; these spindles are more or less white, but may often have a very pale yellow color.

Figure 4. 

Sibogagorgia californica sp. nov., SBMNH 422974 (Holotype). Large fragment, ~18 cm long, showing branching pattern; intertwined tendrils from shark egg case are visible.

Figure 5. 

Sibogagorgia californica sp. nov., SBMNH 422974, SEM image. Sclerites either bright salmon in color, or pale yellow to white. A Sclerites from colony medulla B sclerites from colony surface and possible intermediates toward the medulla C 7-radiate sclerites from colony surface.

Figure 6. 

Sibogagorgia californica sp. nov., SBMNH 422978. A Large, ~36 cm long colony fragment B close-up view of polyps, illustrating color and position on branch; note contrast color between polyps and surrounding coenenchyme, characteristic of species in the genus. This may however, be a partially bleached specimen, rather than one exhibiting true coloring.

Figure 7. 

Sibogagorgia californica sp. nov., SBMNH 422978, SEM image. Color of sclerites from this colony very pale pinkish orange or white. A Sclerites from colony medulla B intermediate form between surface and internal medulla C 7-radiates from the surface. Compare this plate with that of Figure 5; these two appear extremely similar, indicating that SBMNH 422978 is likely a color variant (or very bleached) colony of that species represented by SBMNH 422974.

Nearly all specimens examined, with the exception of two, are from locations within the vicinity of the California Channel Islands, thus a reference to the state of California, where most of the specimens were collected.

Proposed, “California orange bubblegum” coral.

Based on the specimens in the SBMNH collection, ranges from at least Lincoln County, Oregon through southern California waters.

Preliminary examinations led to identification of this small group of specimens as Sibogagorgia cauliflora Herrera et al., 2010. The two colony morphs shown in their description matched well with that seen in the SBMNH specimens: either a vibrant orange-pink (Figure 4) or a more dull cream color with orange polyp apertures (Figure 6). Unfortunately, the paler-colored specimens at SBMNH had been stored in less than desirable conditions for a lengthy period of time; still, the resemblance was strong. However, the SBMNH specimens were decidedly different in the appearance of their radiate sclerites, taken from the surface cortex. In the species described here, surface radiates were not oval, with 8-radiate origination; this form of sclerite was never seen in examined specimens, despite numerous tissue/sclerite preparations, and is the key distinguishing feature of S. cauliflora. Several possibilities emerged: 1) that the SBMNH specimens were an endemic subspecies of S. cauliflora, or (because of their strictly southern California location), 2) a very similar, but different species, or 3) the specimens collected were an isolated group of S. cauliflora that just happened to be found in a location where, for some environmental reason, the distinctive 8-radiates that normally would form in development and growth, did not. Because of their collection location, and differences in sclerite forms, a new species designation is proposed for what may be a very closely-related, species.

From a taxonomic perspective, in their most recent molecular work Figueroa and Baco (2014) recommended that family Sibogagorgiidae be reinstated. According to WoRMS, the genus Sibogagorgia is still retained in the family Paragorgiidae (Cordeiro et al. 2018c).

The museum collection at Moss Landing Marine Labs held a specimen that might be this species, collected in Monterey Bay, 36°26'42"N, 122°01'56"W, ~684 m; coll. G McDonald, 14 August 1974; C0071 [wet]. Coloring of this specimen showed either a slightly bleached condition (storage artifact) or the beige coloring with orange pimples, a slightly brighter condition than that seen in SBMNH 422978.

The SBMNH holotype specimen has, tightly wound around/across its branches, the attachment tendrils from the ends of a swell shark’s mermaid’s purse. The tree-like nature of species not only in this genus, but also in the genus Paragorgia, likely provides anchorage and hiding places for a number of organisms, ranging from brittle stars to fish.

Axis an unjointed medullar region composed of completely fused calcareous sclerodermites (solid calcium carbonate) derived from sclerites; Grillo et al. (1993) indicated calcium carbonate axis of family is not derived from the fusion of sclerites, but rather that there are two different origins for the sclerites and the axial skeleton (concentric addition of CaCO₃). Sclerites present as numerous regular capstans, or as rods, plates, and irregular forms without capstans.

Madrepora rubra Linnaeus, 1758 (by subsequent monotypy, the first species being assigned by Lamarck 1801).

Sclerites of cortex numerous, regular capstans, often modified with six, seven or eight radii; or as double clubs (only some species), crosses and opera glasses; long spindles present in tentacles. Without axial pits bearing beaded rims beneath autozooids. Autozooids prominent, non-retractile (when contracted cannot fully retract into cortex) and ovate-cylindrical, usually distributed on one side of colony.

Bayer (1956b) stated that the name Corallium “is an old name of dubious origin, going back to the ancient Greeks, classically applied to the red coral of commerce, the ‘true red stony coral’.”

. An interesting genus; its connection to human enterprise and profit make it so. Collectively, “pink coral, red coral, noble coral, angel skin coral, Sardinia coral, midway coral” (CITES proposal, Convention of the Parties, CoP14 Prop. 21, June 2007) has had a long history, with the primary focus on harvesting of the coral for profit. At least one proposal, and multiple CITES conferences over the years (2007, 2010, 2013), have made this genus a focus of discussion. Numerous articles have been published concerning the impact of harvesting, management issues, etc. (Tsounis et al. 2010, 2013, to name but a couple).

Because all species of coral in this genus tend to form tall, tree-like colonies, they likely increase three-dimensional complexity of the habitats they are found in and consequently, increase biodiversity where they occur. These colonies could easily provide valuable microhabitat for sessile, associated commensal invertebrates (Baco and Shirley 2005, Baco 2007), protecting them from strong currents and predators. With regards to species occurring in the Pacific Ocean, “one of the more notable commensal relationships is the association of polynoid polychaetes with species in (this genus). Each Corallium species appears to have its own species of polynoid polychaete, which can reach high densities within individual colonies” (Baco 2007). They would also provide structural relief that fishes and mobile invertebrates could use as feeding, spawning and resting grounds (CITES proposal, Convention of the Parties, June 2007). Thus, they contribute far more to their natural living situation than perhaps had been considered when commercial harvesting for species in this genus was first instituted.

A number of species in the genus Corallium have recently been transferred to the genus Hemicorallium (encompassing species discussed here; Cordeiro et al. 2019); this is based on the work of Tu et al. (2015a, b, 2016). Species in the genus Hemicorallium are represented in the SBMNH collection by a single specimen, SBMNH 471940 (likely Hemicorallium ducale; see Appendix 1: List of material examined), but three species have been recorded (collected west of the California Channel Islands), in proximity to the western boundary edge of the Bight (location data for specimens housed at NMNH). The California Academy of Sciences has seven separate lots of specimens in this genus. Generally, all are from the Hawaiian Islands, with the exception of one. That one, identified only to genus, is from California, Davidson Seamount, 128 km SW of Monterey, taken at a depth of 1481 m, 21 May 2002. Further documentation of the presence of Hemicorallium in or near the California Bight is needed; in consideration of their rarity and commercial value, their presence would need to be very carefully monitored.

One lot, 2 specimens + fragments (recent addition) in SBMNH collection, most likely this species (see Appendix 1: List of material examined).

Distribution extends in Eastern Pacific from Mexico (Bayer, 1955), northern Baja Mexico (USNM 50111 (type locality) to at least California Channel Islands (USNM 94459). SEM images on file (Bayer’s personal collection): SEM #2284, for USNM 94459 and SEM #s 2483 and 2484 for USNM 50111. Specimen USNM 50111 represented one of the first finds of the genus Hemicorallium in North American waters.

The specimen in question does not easily identify to a species; in color it appeared more like that of Hemicorallium imperiale, but the polyps’ appearance and scleritic spindles were more like those seen in H. ducale. While all type material for this species (and the other two that follow) at NMNH were examined some years ago, this one specimen, recently received into the SBMNH collection, requires further study.

WoRMS Data Base (Cordeiro et al. 2019) verifies that H. ducale, with the other two listed below, are accepted species. They are included, with brief comments, due to their collection locations and proximity to the region of the California Bight. It should be noted that there are no specimens that came into the SBMNH collection from the ‘Velero’ expeditions.

No material in SBMNH collection (see Appendix 1: List of material examined).

Distribution in the Eastern Pacific, northern Baja, California (Bayer 1955), as seen for the holotype (USNM 50110) to California Channel Islands, in proximity of western boundary edge of Bight (USNM 85082). No SEM images for this species were found in Bayer’s personal collection at NMNH. Could be called “Imperial red coral.”

(see Remarks section below.)

No material in SBMNH collection (see Appendix 1: List of material examined).

Around the Pacific, from Hawaii (holotype, USNM 49520) to offshore seamounts some miles west of California coast (outer edge of Bight western boundary, USNM 94460), certainly at substantial depth (based on specimens housed in NMNH collection). Not enough specimens examined (or collected with attention to specific collection locations) to determine extent of north-south range. Bayer commented (1956b), “of all the Hawaiian precious corals, C. regale has the best color and might be of commercial value if it could be fished in quantity.” Thin, calcareous extensions of axis extending outward to thick coenenchyme can support expansion of coenenchyme near sides of branches as recurved flaps, a distinctive feature. These can form tunnels inhabited by polychaete commensals (Baco and Shirley 2005, Baco 2007). Could be given the common name “Regal red coral.”

Several species in the now updated genus Hemicorallium, including this one, found within the Hawaiian archipelago (and elsewhere in the western Pacific, including international waters) have been the focus of commercial exploitation (then recognized as species in the genus Corallium). Based on reports made public by CITES, regarding “consideration of proposals for amendment of Appendices I and II” (2007, 2010) and Bruckner (2009), there was much discussion, and confusion, as to whether this species (using older genus designation) was valid and/or whether it could be synonymous with Corallium laauense (misspelled as C. lauuense). Grigg and Bayer (1976: 169) and Grigg (2002: 17–19) specifically mentioned C. regale and/or C. laauense as separate species. CoP14 Prop. 21 (CITES, 2007) listed this species as a potential synonym for C. laauense; this based on Baco and Shank (2005: 664). However, Baco and Shank (2005) did not treat this species as a synonym of C. lauuense (note the misspelling), but a comment they made regarding the work of Grigg (2002) may have inadvertently lead some to assume that was the case. Cairns (CITES, 2007) did not consider these two taxa to be synonymous and Bruckner (2009: 321) also discussed these taxa as two separate species. Bruckner indicated that although he discussed them as two separate species, “these 2 species may be synonymous.” In the document, Proposal, CoP15 Prop. 21 (CITES 2010), C. regale was still shown as a synonym of C. lauuense (sic); note the following statement: “C. lauuense and C. regale are listed as separate species in the US Precious Coral Fishery Management plan, but these species are usually considered synonymous (Parrish 2007)”. Again, C. laauense and C. regale were treated as synonymous; however, regarding C. regale, Baco and Shank (2005) stated: “C. lauuense was previously misidentified and referred to as C. regale which is not an indication of synonymy. There may still be some unresolved taxonomic problems concerning these two species.” Additionally, “Bayer and Cairns (2003) differs from the SS species list in a number of ways: C. boshuense, C. niveum, C. porcellanum, C. pusillum, C. vanderbilti, and C. variabile are not mentioned; C. regale is treated as valid.” (CITES 2010); of these species, C. boshuense and C. variabile have been moved to the genus Hemicorallium (Cordeiro et al. 2019) in the WoRMS Database. The final implication was that Corallium sp. was found throughout “the Hawaiian archipelago and into the Emperor Seamount Chain” (Baco 2007), but that the certainty of species identification was still in question.

In studying Bayer’s original 1956 description of C. laauense (correctly spelled) and C. regale, no determination could be made as to why these two were linked as synonymous. The now recognized species Hemicorallium regale has many double club sclerites, while H. laauense has none. As well, the entire colony of H. regale, as well as the axis, is pink, while the colony color of H. laauense is white or very faintly pink with a white axis. SEM images are on file (Bayer’s personal collection), SEM #2283, USNM 94460.

The work by Tu et al. (2015a, b; 2016) resulted in most species of Corallium being placed in the genus Hemicorallium. WoRMS (Cordeiro et al. 2019) confirms the placement of the species discussed here: H. ducale, H. imperiale, H. regale (as well as H. laauense); all are listed as accepted, separate species.

Axis purely horny (scleroprotein without calcareous deposits), dark-colored, predominantly black; very difficult to cut, with wide, hollow, cross-chambered central core. Coenenchyme very thin, polyps conspicuous, contractile, not retractile, completely covered with both straight and curved fusiform sclerites (forming, in appearance only, cylindrical calyces; no calyces actually present). Sclerites arranged in eight double rows, forming eight en chevron fields; no well-defined operculum; sclerites instead arranged as transverse ring and eight points of converging spindles on tentacle bases; thus, sclerites of polyps continuous with those of tentacular crown, latter being sharp spines arrayed conspicuously around top of polyp, with no intervening sclerite-free neck zone or transverse collaret. Consequently, no clear division between anthocodia and anthostele. Tentacles fold inward over oral disk. Predominant sclerites colorless, in form of prickly or warty spindles; sometimes, presence of three- and four-armed radiates.

Acanthogorgia hirsuta Gray, by monotypy (? = A. aspera Pourtalès, 1867).

Colonies generally flattened (flabellate); commonly reticulate, or developed into dense bushy shrubs. Branches appear thin and delicate. Polyps tall, cylindrical, topped with thorny crown of strongly projecting spinous sclerites, embedded at tentacle bases. They lie, collectively, over infolded tentacles, protruding end of sclerites smooth. Polyps on all sides of branches, or roughly biserial; arise vertically at right angle to branch surface, acalycinous, not retractile. Coenenchyme between branches usually thin, axis visible through it. Sclerites in polyps slender spindles slightly bent, arranged en chevron in eight longitudinal double rows. Back of tentacles with only numerous small, flat, bent sclerites; stem coenenchyme with slender, generally bent or sinuous spindles sculptured by prickles or simple tubercles; in deeper layers of coenenchyme (some species), with radiates (tri-radiates and crosses, often with a projecting central spine). Axis dark, but coenenchyme usually colored; sclerites always colorless.

Only one collected specimen from the genus appeared in the ‘Velero’ material; it does not match any of the described species it has been compared it to, thus far. It was collected in Mexican waters, beyond the geographic range covered in this work; description of this specimen is in progress.

(?) Okinawa, Japan; 400 fathoms.

Zoological Museum of the University of Hamburg, Germany (formerly, Naturhistorisches Museum, Hamburg); Catalog Number 3298, with Catalog Number 3297. (Not labeled as type.)

1 lot (see Appendix 1: List of material examined).

Colony (Figures 8, 9A) richly branched, not entirely in one plane, forming bushy fan or tree no more than 10 cm at widest point. Colony appeared fragile and delicate, but actually a tough, spiky bush/fan, branches reminiscent of those in a bottlebrush (Figure 9B); not greatly flexible. Colony height (dictated by main, central, generally straight stem), base to tip, 15–16 cm; 7.0 cm broad; holdfast remnant present. If any regularity to branching pattern, slightly dichotomous to pinnate, usually in one plane; all lateral branches, of differing lengths, project at nearly right angles, extending/curving quickly upward, even those with more lateral placement. Branch diameter averages 2.0–3.0 mm. Polyps distributed over entire surface (not so much on lower portion of main stem, just above base), nearly in rings around branches, closely placed but not crowded (Figure 10); sometimes two with bases contiguous, generally separated by 1.0 mm, perhaps more; terminal twigs rounded, almost clavate in appearance; numerous polyps at apex, completely covered with straight and curved spindle-shaped sclerites. Polyps not retractile; very conspicuous, decidedly slender, columnar in shape, height between ~4.0–7.0 mm (most average 5.0–6.0 mm tall); diameter generally 1.0 mm for most of polyp length, narrowing slightly, then increasing to ~1.5 mm wide at the crown. Easily recognized by crown of sharp spines encircling top of polyp. Coenenchyme very thin and translucent, with axis color showing through. Color of freshly collected specimen bright lemon yellow (M Love, pers. comm.; Figure 8); on being placed in alcohol quickly turned, generally, light olive-green towards base, becoming slightly darker grey-green at uppermost branch tips. (While species in family are described as having a predominantly black, purely horny axis, color of axis showing through extremely thin coenenchyme appeared to account for overall olive-green color. Having now sat in 70% ETOH for some time, the colony has turned more yellowish brown.) Sclerites (Figures 11, 12A, B) mainly spindle-shaped; straight or curved, showing arrangement of eight double rows, forming longitudinally-placed chevrons (obliquely angled double-rows) characteristic of genus. A very few oddly branched; some, more a tripod shape. Sclerites appear mostly tuberculate, with distinct boomerang bend (Figure 12A), easily removed from surface of colony. The longest sclerites, with distinct bend roughly a third of the way along their length, range from 1.0–2.0 mm in length (average 1.6 mm L × 0.17 mm W); one third of surface of sclerite bears tubercles, while other two-thirds is generally smooth; this smooth section, thin, rounded, somewhat beveled, is the distal, prominent spike that projects from the thorny basal portion embedded in the mesoglea of the body wall, in nearly longitudinal direction; lower, embedded portion, ~0.5 mm long, appears to cross over into the neighboring angled rows, these basal portions not much different one from another. These sclerites form the crown of thorns seen around top of polyp. Two very long spines project upwards to form the points of the crown at distal end of each of the eight double-rows. Smooth portion of these sclerites sit with approximately 1.0 mm of their length free of polyp. Numerous, slightly smaller, flatter sclerites have bend more centrally located, their entire surface covered with tubercles (average 1.0 mm L × ~0.08 mm W). These primarily cover outside surface of polyp (Figure 11), illustrating chevron pattern (eight longitudinal double rows) in placement. Sclerites of coenenchyme similar to those seen on polyps’ surface (~0.8 mm long), perhaps slightly thicker in width and slightly more fully covered with tubercles; tentacular sclerites smaller still, bent, flattened (up to ~0.18 mm long), completely covered with tubercles; more prominent, dense, on dorsal side of tentacles. If present, a few smaller (0.1 mm long) radiate or cross-shaped sclerites may be seen. In the coenenchyme covering the base, exclusively, are 0.25 mm long, bent, strongly spined spindles. All sclerites completely colorless, reminiscent of thin, bent shards of glass.

Figure 8. 

Acanthogorgia gracillima var. typica. In situ, living colonies, illustrating vibrant yellow color, as seen in Santa Barbara Channel, photograph by D Schroeder; image courtesy of M Love, UCSB.

Figure 9. 

Acanthogorgia gracillima var. typica, SBMNH 423074. A Whole colony preserved in 70% ETOH. Maximum height of colony 15–16 cm, ≤ 10 cm at widest point B magnified view of several branches; pronounced, rather spikey appearance of polyps is evident.

Figure 10. 

Acanthogorgia gracillima var. typica, greater magnification of SBMNH 423074. Shows position of polyps on branches. Note prominent crown of sclerites at distal end of each polyp, polyp height averages 5.0–6.0 mm.

For designation gracillima, the Latin gracili- means slender; this may reference the conspicuous, very slender polyps and very slender points in the crown of this species. Kükenthal (1909) gave no explanation for the derivation of the species name.

None specifically designated; the Slender glass-shard gorgonian would be appropriate. Worth considering is whether or not this is the species that, predominantly, MBARI and NOAA researchers are seeing and calling the “Gold gorgonian.“

For genus in general, “species. . . inhabit moderate to considerable depths; various species of Acanthogorgia occur in all seas, some thriving in very cold waters” (Bayer, penciled personal annotations in Kükenthal 1924). Examinations of the literature revealed numerous species in the Indo-Pacific region. Recent MBARI on-line postings indicated this genus is found in northern California waters, often at great depth. Milton Love (description based on specimen he collected) indicated (pers. comm.) that the color of this gorgonian is very vibrant, thus easily seen; it is quite abundant in certain areas, such as the Footprint, a feature outside the Anacapa/Santa Cruz Island Passage. Thus, for this species, range seems to extend around the Pacific Ocean from Japan (Kükenthal’s specimen) to eastern Pacific waters of California; further study will be required to determine whether, and how far, it extends north and south of California.

On this particular specimen, there were at least two scale worms wrapped around the base of polyps in two separate areas within the colony.

To handle the specimen during examination, although colony fairly hardy, gloves were necessary. Generally, all the long sclerites were very sharp, comparable to small shards of thin glass; those that got into the skin under the fingernail were rather painful. Given the delicate, and somewhat brittle/fragile nature of the longer boomerang-type sclerites, these sclerites are easily broken. It was often difficult to ascertain whether radiates and crosses really existed as such, or whether they were bits that had broken off from the longer spindle forms; this could account for some of the odder-shaped sclerites that previous description (Kükenthal 1909) made mention of. Microscopic examinations were always done without a cover glass, when possible, so that sclerites were not crushed.

This description (and original description for the species) was based on the specimen used by Kükenthal (1909). In initial comparisons of known, and described species, using characteristics of the colony (lying more or less in one plane, with polyps generally more than 4.0 mm in length), the key that Kükenthal (1924) provided in his overview of the genus was relied upon, but it took some time to rule possibilities out. Bayer (1996b) stated that all the sclerites are so similar in form within all members of the genus, that distinguishing species is difficult; this was confirmed. As well, there are very few recent descriptions of any Acanthogorgia species that are truly adequate to use for comparative purposes. Few specimens had been collected from California waters; thus, not a lot of material to compare against. In examinations of collections at other institutions, only two/three specimens were found at NMNH that approximated the appearance of this colony (USNM 1071429, dry and USNM 1072361, wet, in two lots). The dry specimen is from Cross Seamount, Hawaii, USA and the wet lots are from a seamount east of Necker Island, Hawaii, USA. Both are identified to genus, but not to species; in color, branch pattern, pattern of sclerite positioning on polyps and sclerite morphology, specimen USNM1071429 appeared most similar. If NMNH specimens can be accurately identified to species, they could confirm (or negate) the notion of trans-Pacific distribution. In any event, this discussion/description represents one of the first published reports for this particular species in southern to central California waters; overall the genus needs further work, not only on those specimens being frequently noted in underwater explorations (cruises and surveys being done by MBARI and NOAA), but also on those that have already been collected, perhaps described or need description, currently housed in other museum collections. With greater access to locations of depth, and the use of ROV, AUV, digital imaging and improved ability to collect specimens at depth, many currently described species can be confirmed and many new species are likely to come to light. As well, with careful attention to locations where specimens are found, it should be possible to develop a far more accurate picture of the distribution of these deep-water forms.

While the original specimen does indeed still exist, no opportunity to travel to Hamburg, Germany occurred and no request that the specimen be sent was made. The species is an accepted species in the WoRMS Data Base (Cordeiro et al. 2018e).

[Acanthogorgia (= ? Acalycigorgia) Grey, 1857)]; not accepted, WoRMS (Cordeiro et al. 2018e).

Figure 11. 

Acanthogorgia gracillima var. typica (light microscopy image), SBMNH 423074. 4X magnification, demonstrating position of long, boomerang-shaped sclerites, arranged longitudinally on surface of polyp, mid-section to distal end.

Figure 12. 

Acanthogorgia gracillima var. typica, SBMNH 423074, SEM image. These sclerites are colorless. A Sclerites from crown B sclerites from polyp surface C small sclerites from coenenchyme D odd coronal or polyp surface sclerite.

A. grandiflora Kükenthal & Gorzawsky, 1908a; subsequent designation by Kükenthal and Gorzawsky (1908b).

Polyps not functionally differentiated into anthocodia and anthostele; contractile but not retractile within common coenenchyme; tentacles fold over oral disk in contraction. Polyps similar to those of Acanthogorgia (without crown of strongly projecting spines, however), but polyps can be short and verruciform to prominent, tall and cylindrical, not clavate. Sclerites of polyp walls large spindles, very conspicuous; commonly arranged more or less distinctly en chevron in eight long, longitudinal double rows, but distal ones project little or not at all. Distal ends of sclerites around tentacle bases not specifically differentiated as spines, though the tips may project somewhat around polyps’ apex. Polyps are without suture separating tentacular/anthocodial from subtentacular sclerites. Sclerites of polyp body gradually merge with those of tentacle bases, which are not abruptly smaller; coenenchymal sclerites with tubercles of inner and outer sides similarly developed; inner layer of coenenchyme with more or less abundant radiates.

Cannot be indicated at this time.

Cannot be indicated at this time.

1 lot (see Appendix 1: List of material examined).

Colonies (two) generally in one plane; one measures 7.0 cm × 5.5 cm (length to width); second (Figure 13) measures 9.0 cm × 4.5 cm, at widest, halfway up colony. Thin, delicate-looking branches (round to slightly square in shape); branching more or less dichotomous; closely monopodial. No flattening at branch origins. Base 2.0 mm wide, main branch 1.0 mm wide; branchlets vary between 0.5–0.75 mm wide and tips of branchlets very thin, thread-like; all branches quite stiff. Coenenchyme very thin (very little still present in these specimens); axis predominantly exposed, yellow-gold to rusty-brown. Of the few polyps present on a few branches, most located near branch tips (Figure 14A); coenenchyme and polyps creamy-white. Polyps primarily sit lateral to branch, at distance of ~1.0 mm or less from each other; closer to branch tip sitting literally side by side; some few branches indicate that polyps can be found on all sides. There are marked, longitudinal grooves/ridges at distal ends of polyps; there is barely apparent a very short little spiny crown at their very tip (Figure 14B). The ridges, eight in number, are each formed by a parallel collection of two or three bent spindle-type sclerites. Polyp surface densely covered with sclerites; no calyx apparent. Polyp height 3.0 mm, 2.0 mm from base to area of longitudinal grooves with another 1.0 mm of height when area of grooves/ridges included. All approximately 1.0 mm wide, distal end slightly wider, somewhat obvious, ~1.5 mm wide. No expanded tentacles readily visible (contracted over mouth); all heavily covered or encased by sclerites. Sclerites (Figures 15, A particularly) predominantly bent spindles; all tuberculated across entire surface, averaging 0.5 mm long by 0.08 mm wide. The largest (~0.7 mm × ~0.1 mm), decidedly bent spindles; these form the eight ridges mentioned above; others appear to lie in longitudinal direction up to and beyond upper edge of polyp, barely showing as short points of a crown. Bent spindles, somewhat smaller, almost tend to the formation of the en chevron, double-row pattern at the proximal end of ridges and down on to lower end of polyp. Also, less bent ones, seemingly very narrow spindles (0.6 mm × 0.05 mm); few appear slightly club-shaped (average 0.4 mm × 0.06 mm), primarily from lower polyp wall and coenenchyme. Sclerites with boomerang shape scarce or not present. From initial light microscopy examination, apparent that many of these spindles can be broken; many odd-shaped bits seen in arrays, with some of the spindles having oddly truncated ends, where some aspect of the sclerites likely had broken off. All forms quite densely arrayed on specimen’s surface, giving polyp and branch coenenchyme a distinct white to glassy appearance; all sclerites colorless. Inner coenenchyme radiates not found.

Figure 13. 

Acanthogorgia species A, SBMNH 423075. Of tissue present, polyps readily visible, colony measuring ~ 9.0 cm in height.

Figure 14. 

Acanthogorgia species A, SBMNH 423075. A Closer view of polyps present at tips of several branches; polyp height 3.0 mm tall, 1.0 mm broad B further magnification of polyps. Barely visible at distal end of several polyps, the weakly developed crown of sclerites can be seen.

Figure 15. 

Acanthogorgia species A, SBMNH 423075, SEM image. Sclerites from these colonies are white, rather than clear. A Coronal sclerites B, C smaller coenenchymal sclerites.

It is possible that these specimens, and several others in the collections either at CAS or NMNH, Smithsonian (not yet identified to species), could be first records of appearance for this species in, or near boundaries of, the California Bight. Description here is based on two specimens, which in overall colony appearance superficially look very similar to Nutting’s photographs (1912: pl 11, fig 1, 1a), shown for Muriceides cylindrica, which Nutting established as a new species. However, the sclerites illustrated in Nutting (1912: pl 20, fig 3), while of comparable size to that indicated for the species described here, do not match. In considering overall appearance of polyps on the branches, and the manner in which sclerites covered the surface of the polyps from distal to proximal ends, a generic best fit occurred working from an illustration that was given in Bayer (1956a) for Acalycigorgia. However, Fabricius and Alderslade (2001) have stated that “(m)any species of Acanthogorgia, in which the polyp sclerites are so short that there is little or no projecting crown, erroneously appear in the literature under the name Acalycigorgia Kükenthal, 1919. Acalycigorgia is actually a synonym of Acanthogorgia and most species referred to the former should be called Acanthogorgia.” This is supported in the WoRMS Data Base (Cordeiro et al. 2018e) for taxon information regarding status of the genus Acalycigorgia. It now remains to be seen if the species described here is indeed one of the 61 species listed as accepted by the Data Base.

Lissogorgia flexuosa Verrill, 1865; [subsequent diagnosis by Nutting 1910].

Colonies fan-shaped, branching in one plane, some anastomosing. Larger branches with axis often tending to be flattened at right angles to plane of fan; in older colonies, smaller branches can bend and grow perpendicular from the fan. Polyps prominent, low, wart-like, non-retractile; coenenchyme thick between polyps. Sclerites of polyp tentacles small rods; below anthocodiae, sclerites large and bow-shaped, in angled double rows (chevrons), forming eight marginal points, forming strong collaret; tentacular operculum distinct. Sclerites of coenenchyme in two distinct layers, mostly spindles (coarse or densely warted), small capstans, with some clubs or discoidal forms.

Kükenthal (1924) articulated the characteristics of this genus, which include: condition of the branch ends, position, orientation, height, and form of polyps, presence of an operculum, and arrangement of sclerites in the coenencyme. Fabricius and Alderslade (2001) discussed aspects of comparison/contrast between this genus and others; it was stated, “not much separates this genus from Anthogorgia, and a study of a large suite of specimens could see the two synonymized.” Additionally, a characteristic feature pertinent to the genus diagnosis came from the Octocoral Taxonomy Laboratory Manual (2007): There are no calyces in the genus Muricella; “almost 90% of the species attributed to this genus do not belong there. A major revision is needed.” In reviewing older literature there were references to calyces in this genus; Bayer (1956a) comments on the “truncated cones” or “rounded, low verrucae.” In descriptions given by Verrill (1869a), reference is made to calyces, as well. The concept of how to define a calyx may need revisiting, notably with regards to this genus.

No material identified as this species in collection at SBMNH (see Appendix 1: List of material examined).

Included here briefly, as there was a California collection location for a specimen that was identified as this species.

Based on Kükenthal (1924), the species ranges from coastal to abyssal depths; likely can be found in a wide and diverse number of locations around the world. The type was collected off Japan, but present location of the type specimen could not be determined. The specimen upon which Nutting’s 1909 description was based, taken by the Fisheries Steamer ‘Albatross’, Station 4461, Point Loma Light-house, S 3, E 9.3 miles, 285 fathoms, was said to have differed slightly from the type, particularly in having a well-marked collaret and in the arrangement of the calycular sclerites. Kükenthal (1924) indicated that the California location (at an approximate depth of 250 m) is questionable. There is no certainty that any specimens found in the California Bight will be specimens of Muricella complanata, or whether the species actually exists. Harden (unpublished dissertation, 1979) suggested that Muricella complanata was synonymous with Swiftia torreyi. Both were stated to display anastomosing (S. torreyi is always anastomosed; was unable to determine if that is always case with Muricella complanata, as thus far no confirmed specimen of this species has been located). As well, I would disagree with a comment found in an unpublished Bayer annotation: “Muricella complanata = a Swiftia?” in that S. torreyi is a plexaurid, while Muricella is a recognized genus within the family Acanthogorgiidae (Bayer 1981c, Fabricius and Alderslade 2001, Cordeiro et al. 2018f) with no mention of any species of Muricella being synonymized with any Swiftia species.