Gorgonia exserta Ellis & Solander, 1786: 87 (non Thesea exserta Duchassaing & Michelotti, 1860); [= Stenogorgia Verrill, 1883].

Colonies chiefly in one plane, with lax branching (dichotomous or pinnate-like); branches/branchlets tend to curve upwards; in some species, anastomoses possible (fan-like); in others, minimal branching or none. Polyps widely scattered, or crowded; often lateral or biserial, forming prominent conical or cylindrical mounds; on tips of branchlets, two polyps always opposed; conical anthostele seldom retracted; generally, polyps retractile. Anthocodiae commonly tall, exsert. Coenenchyme thin to moderate, somewhat rough/granular, outer layer filled not only with spinous rods or spindles, but with capstans having warts more or less conspicuously modified as double disks; some capstans quite foliate; inner layer mostly restricted to areas between longitudinal canals, containing only small capstans. Mound margins, base of tentacles, with numerous rows of conspicuous, stout spindles as bar-like rods, characteristic for species in the genus (fingerbiscuit shaped; see Bayer et al. 1983, pp 72–73, pl 19, figs 184–185). Axis is horny, flexible, somewhat flattened. Colony colors generally red, red-orange, pink or white.

Deichmann (1936) stated that the definition of the genus Swiftia corresponded exactly with Stenogorgia Verrill, 1883; the problem discussed there stemmed from a misinterpretation of G. exserta Ellis & Solander, 1786 by Verrill (also by Kükenthal 1924). See remarks, following.

The stout, anthocodial rods (seen at mound margins and bases of tentacles), are definitive for this genus. Examinations of multiple specimens (several different species) within this genus usually revealed the appearance of these rods; when present, looking much like the fingerbiscuit sclerite form shown in Bayer et al. (1983) for the genera Clavularia Blainville, 1830 and Ptilosarcus Verrill, 1865 (neither of these gorgonian genera), where sclerites are described as minute, flattened rods (rods here have a bit of depth). Further examinations (multiple species) revealed that some individual colonies of species in the genus did not have these conspicuous rod forms (having only spindles and capstans). Other colonies displayed spinous spindles and/or capstans and anthocodial rods; some few species had only the fingerbiscuit rods, numerous throughout all tissue structures. The rod form is not always easy to obtain in a sclerite array; some specimens without rods may actually have them, but they may be quite small, not very numerous and very widely scattered. A trend observed is that colonies further north in the Pacific (Alaska) have very obvious rods, while specimens of some of the same species collected in California (specifically central and southern California) may have rods, but infrequently. By way of comparison, in several species from the genus examined from waters in/near New Zealand, some had only rods, and no other form of sclerite. It appeared that colder, temperate to subpolar species had the rods (to the exclusion of all others) but species from warmer, albeit temperate water, tended to display a minimal number or complete absence of rods. Examination of many more specimens, collected in both hemispheres from poles to equator, could reveal further insight into the appearance of this key sclerite form. To further clarify questions surrounding location ranges for each of the Swiftia species discussed, Appendix 2: Map A1 shows the distributional range of each and Appendix 1: Table A1 shows key features used to distinguish one species from another.

Regarding use of the generic name Swiftia, Muzik (1979) stated: “(t)o preserve the generic name Swiftia a petition to” the International Commission on Zoological Nomenclature “(ICZN) must be made;” uncertain as to whether this was ever done. “For a full explanation, see Challenger Reports 31: 146 and Deichmann 1936: 185” (Muzik 1979). The complete explanation can be found in Deichmann (1936) and Muzik (1979: 168); they serve to confirm the confusion that had developed, through the work of previous investigators, regarding generic status for the species discussed below. In Madsen (1970: 5), “A total of about a dozen gorgonarian species referred or referable to Swiftia (syn. Stenogorgia) from widely scattered localities in all three oceans have been recorded, but only a few of them are sufficiently described.” In the WoRMS Database (Cordeiro et al. 2018), status of this genus has been accepted, but Breedy et al. (2015: 329) stated that a “thorough review is needed in order to clarify taxonomic problems related to Swiftia.” This paper attempts to clarify some of the issues related only to those species that are found in, near, or extending geographically slightly south or north of the California Bight.

USA, California, Monterey Bay, bearing S 67°E, 3.7 mi off Point Piños light-house, ~36°38'00"N, 121°55'00"W, 119–135 m.

Holotype USNM 25432 [wet]; specimen was examined.

~20–25 lots (see Appendix 3: List of material examined).

Colony (Figure 1A) flabellate, loosely branched, sparsely reticulate; often large areas of open space between branches; on rare occasions, branches anastomose; irregular, pinnate branching chiefly in one plane. Appears moderately delicate; some distance from base, main stem generally divides into several main branches, center one ascending nearly unbranched, lateral ones at first widely divaricating, then ascending, giving forth pinnate branches which tend to be opposite, often irregular. Branchlets ~6.0 mm apart (where regular), somewhat flattened (0.5–1.0 mm); terminal branchlets usually end with two oppositely disposed polyps (Figure 1B). Polyps form prominent, conical, volcano-shaped mounds, with broad base (not boxy), with distinct exsert anthocodiae; polyps scattered or biserial (Figure 1B), in two irregular, lateral rows, creating a very narrow front and back; laterally, opposite or (often) alternate on branchlets; colony and branch profile appearing in form of zig-zag pattern. Polyps more numerous on front than behind; mound summits 2.0–4.0 mm apart on one lateral side, 0.6–1.0 mm high, surmounted by 0.4–1.0 mm tall anthocodiae, ~1.5 mm across. Margins with eight-lobed edge; outer sides of tentacles crowded with sclerites, tentacles retractile, bent inward at rest. Coenenchyme thin; outer layer filled with spinous rods, spindles, and capstans. Color of living colony typically true salmon, also bright orange to deep coral red; polyps may be same color (paler), cream or white; axis dark greenish-brown, lightening distally. Sclerites (Figure 2A–G) can be always exclusively small to somewhat long, warty (thorny), slender spindles, larger ones often curved. On shorter, thorny ones, some have very prominent, jagged teeth, projecting off one side (reminiscent of sclerites seen in some Muricea); also few small, elongated ones with median waists (“some as granules, some as foliate capstans,” Nutting 1909). In current examinations, those that almost appeared as radiate nuggets could be the granular-appearing or foliate capstans mentioned by Nutting (1909). As well, some can resemble torches, and can be quite evident; these may be the shorter, thorny ones mentioned above. Mound surface sclerites lie transversely. Sclerites in tentacles described as stout, blunt rods; in all specimens examined (identified as this species), rods were sometimes rarely seen in very small numbers, but never obvious.

Figure 1. 

Swiftia kofoidi, SBMNH 422965 (delicate orange “morph”). A Colony 7.5 cm at tallest point, 7.5 cm broad at widest point B Closer view of several branch tips showing distinctive placement of the calyces, creating zig-zag profile of each branch. Scale bars: 2 cm (A); 1 cm (B).

Figure 2. 

Swiftia kofoidi, SBMNH 422965, SEM image. A Potential anthocodial sclerites B Very elongated spindles, prevalent in species in CA and Mexican waters C Irregular spindles D Truncated, disk-spindle/capstan forms E Unusual quadri-radiate F Jagged torch-like form G Shorter, jagged spindles

Named in honor of Professor CA Kofoid from the University of California.

None specifically indicated. Could appropriately be called the Orange ‘rick-rack’ gorgonian.

Found off California and Oregon coasts (refer to Appendix 3: List of material examined). Off California in locations between 175 and 218 m, off Oregon in 101–106 m, at 126 m and at 138 m. Speculative range from Upper Baja, California, to NE of Tanner Bank in the south (USNM 59817), along southern California coast through Monterey Bay (Holotype, USNM 25432), to south of the Farallon Islands, California (USNM 57235). May continue range northward, off the coast (NW) of Portland, Oregon (USNM 53972), at least (as well, possibly USNM 49538 and USNM 56989, both from Heceta Bank, Oregon). Extreme range makes identification of some specimens questionable; possible range overlap with similar-looking species, S. pacifica (Nutting, 1912). If following a slightly deeper, moderately cooler, water gradient in southerly locations, then a possibility. Appendix 2: Map A1 shows apparent range of this species in relation to those such as S. pacifica and S. torreyi (Nutting, 1909) that it might be confused with when viewed in situ.

Appears to be a moderate depth to deep-water species, generally deeper off California coast, slightly shallower in more northerly locales. Several lots of this species in SBMNH collection displayed associated organisms: on several, the attached organism looked like a clump of small bubbles, reminiscent of that which “spittle bugs” produce on branches of some land plants; at least some of these clumps were small anemones. Some specimens had species of hydroid attached; few colonies harbored tiny Ophiuroidea.

Multiple labels associated with some specimens examined (some with as many as three different labels, each with a different genus/species name, as determined by three separate investigators), complicated identification. This called into question whether all of them were indeed S. kofoidi, a different species in the genus Swiftia or some other gorgonian (belonging to an entirely different genus); some specimens had been identified previously as belonging to the genus Psammogorgia (sclerites did not support this).

Bayer made personal notations in his copy of Nutting (1909), indicating that this species was “stouter than torreyi, calyces closer, bars in anthocodiae large and stout. Similar to Callistephanus pacificus Nutting, 1912: Pg. 96.” This species is stouter than S. torreyi (in the form of the polyps, and in total colony height), but polyps rarely as close together as those in S. torreyi, nor were the large, stout bars that Bayer mentioned obvious; additionally, the color is completely different for these two. In other respects, specimens strongly match images for this species given by Nutting (1909). Bayer’s notation regarding similarity between it and Callistephanus pacificus Nutting, 1912 is generally affirmed. Overall, most similar to Swiftia pacifica, but S. kofoidi is often more open and delicate; many colonies of S. pacifica can be thicker-branched and bulky in overall appearance (Figures 15, 21). The bright orange color (salmon), overall delicate appearance, widely-spaced polyp mounds and very distinct, jagged/sharp zig-zag profile (due to predominantly alternate and lateral placement of prominent polyps) of this species set it apart from the other two species (S. pacifica or S. torreyi) that it is often compared with. As indicated in the WoRMS Database (Cordeiro et al. 2019), this species has accepted status.

The California Academy of Sciences (CAS) has some thirty cataloged records (only some identified to species) from this genus in its collection. Eleven of them are from Alaska and are likely Swiftia pacifica. Of the remaining, sixteen or so were identified as this species. Most of these however, were collected from northern California (Monterey Bay, Humboldt County, Sonoma County, Marin County; many likely the more northerly-dwelling S. pacifica). The Monterey Bay Aquarium Research Institute (MBARI) collection records indicated specimens of this species, collected and/or surveyed/photographed throughout their study areas (of those identified as this species, many may actually be S. pacifica). One colony examined via video/photo, was T1101-A21; based on colony shape and color, it appeared to be this species. Sclerite examination of the actual specimen could have confirmed it as such but specimen was not collected. Two other data collection events should be considered. However, neither of the specimens in question were located; they apparently are not housed at the National Museum of Natural History, Smithsonian (NMNH). Both from: North Pacific Ocean, USA, California, Monterey County: [Monterey Bay], bearing S 46°E, 8.4 miles off Point Piños light-house and S 76°E, 3 miles off Point Piños light-house, at 1,544 m and 109 m, respectively. Both collected by USBCF ‘Albatross’, at stations 4546 and 4554, respectively.

For holotype, unknown (erroneously labeled); for type, ‘Albatross’ Station 4781, 52°14'30"N, 174°13'00”W, south and east of the Bering Sea. (See Remarks below.)

Holotype USNM 49513 (colony portion only); Type USNM 30024; both specimens were examined.

~23 lots (see Appendix 3: List of material examined).

Colony moderately sized (up to 18–19 cm tall), planar, flabellate, flexible, rubbery in appearance (Figure 3A); branches nearly forming net-like reticulations, but usually not anastomosing; generally, moderate open appearance to branches. Main stem extends upwards some few cm (above base), 1.0–2.0 mm wide; branches from main stem opposite or alternate, coming off at 45° to 90° angles, then tending upwards; distance between branches 0.5–2.0 cm; terminal branches to 2.0 cm long, l.0 mm in diameter. Polyps lateral (mostly alternate, sometimes opposite), very few on front, with back generally free of polyps, thus flat; polyps conical (sometimes tubular), three, four or five per cm, arising from small mounds (Figure 3B); anthostele 0.5–2.0 mm H, 1.0–2.0 mm W (seeming rather broad and boxy), anthocodiae preserved exert up to 2.0 mm long, but often appearing as dense tuft with 1.0 mm or less showing above polyp mound. At distal end of terminal branches, two (or two pair) oppositely disposed polyps. Color of colony bright to deep crimson or muddy red (brick-red) in life, but both darker and lighter red (dull pink) colonies occur; polyps sometimes dark greenish-grey; sclerites bright red or orange (rods) to pale pinkish-red (most common color, usually true of spindles and capstans). Sclerites (Figures 4A–D, 5A, B, 6A–F) symmetrical; unilaterally developed superficial capstans and spindles, 0.08 to 0.17 mm long in coenenchyme; flatter warty spindles to 0.35 mm in layer below; also eight radiates. Axial sheath sclerites short (to 0.12 mm), narrow-waisted, blunt-ended. Anthocodial bases can have numerous prominent blunt bars (fingerbiscuit rods, in shape comparable to a bacterial rod), curved or straight (Figures 5B, 6D); arranged transversely at bases, more longitudinal at distal end. Examination of multiple specimens did not always reveal presence of rods, but when present, very obvious. Pinnular scales 0.06–0.1 mm long.

Figure 3. 

Swiftia kofoidi [? Swiftia pacifica] (thicker dark red “morph”), SBMNH 232036. A Colony measures 6.0 cm tall, 5.5 cm broad at widest point, demonstrating zig-zag appearance of branches due to calycular placement B Branch close-up, showing placement of prominent conical calyces on branches; calyces measure ~1.0 mm tall.

Figure 4. 

Swiftia kofoidi [? Swiftia pacifica], SBMNH 232036, SEM image. A Truncated, jagged disk-spindle/capstan-like forms B Elongated spindles C Might be a typical “fingerbiscuit-rod” form typical of the genus; seen very rarely, if at all in this species D An irregular spindle.

Figure 5. 

Swiftia pacifica, specimen 41-39-1 (Alaska Fisheries Service, Gulf of Alaska); looking in same form of that seen in Figure 3, light microscopy arrays. A (4×) showing variety of sclerites, particularly the characteristic “fingerbiscuit-rod” seen in the genus Swiftia. Sclerites from specimen examined for Bob Stone, Alaska Fisheries Service B Higher magnification, 10×, showing all sclerite forms, including obvious anthocodial fingerbiscuit-rods. The larger spindles measure ~300 µm long, smaller spindles of ~200 µm, and the rods range from 308–370 µm in length.

Figure 6. 

Swiftia pacifica, specimen 41-100A-2 (Alaska Fisheries Service, Gulf of Alaska, via Bob Stone), SEM image. A Long spindles B Jagged disk-spindle/capstan-like sclerites C Thick, elongated rod-like spindles D Anthocodial “fingerbiscuit-rod” forms characteristic of the genus E Irregular spindles F Odd irregular spindles.

Species name likely refers to locality where type specimen was collected, outer Aleutian Islands, Alaska in the North Pacific.

North Pacific Ocean, Aleutian Islands, Alaska down to California (not a common occurrence), and from Alaska down through western Pacific to Hawaii. Range determined from collection location information provided with specimens examined (see Appendix 3: List of material examined).

Usually bathyal, but depth range extends from ~18 to ≥ 2,000 m, based on depth information provided with specimens examined.

Bayer made personal notations in his copy of Nutting (1909); species is quite similar to Swiftia kofoidi. In overall shape, this species ranges from a rather open-spaced and delicately appearing colony (rarely) to one that seems bulkier. The zig-zag profile is evident but is much more rounded (less jagged-looking) than that of S. kofoidi. Polyp mounds in S. pacifica are somewhat lower, more rounded than those seen in S. kofoidi. In Nutting’s 1912 description, he noted this species’ “very close resemblance to Calllistephanus koreni Wright and Studer” but also added that “(g)eographical considerations render it unlikely that the two are identical.” Madsen (1970) noted that the species described here so closely agreed “with the Scandinavian rosea (it too, has radiate capstans among the spicules) that it is reasonable to consider it its amphiboreal representative, and to regard it as a subspecies of rosea, stat. nov. [Swiftia rosea (Grieg 1887)];” an example of a discontinuous circumboreal octocoral.

Examination of specimens collected in the Gulf of Mexico, 2009 (provided by P Etnoyer, NOAA’s National Ocean Service Office, South Carolina), indicated Madsen may have been correct. Three specimens were sent (without collection data). Sclerite preparations were performed, and specimens were tentatively identified as S. pacifica; when informed (pers. comm. from A Quattrini, then a doctoral candidate, Temple University) that these three were actually collected from the Gulf of Mexico, further investigation was warranted. Specimens of S. pallida Madsen, 1970 had been examined and sequenced (via barcoding of those specimens), and a close match was found between S. pallida specimens and other lots of the same specimens examined. Referring back to Madsen’s (1970) discussion, S. pallida was considered by Madsen to be, at most, a subspecies of Swiftia rosea (Swiftia rosea pallida Madsen, 1970), of the north Atlantic, based on the color of its colony form (pale gray) and sclerites (colorless). S. pacifica from the Pacific Ocean, examined here (considering Madsen) looked very similar to the species S. rosea (which could certainly be the correct species identification for the three specimens from the Gulf of Mexico sent by Etnoyer) in its colony color, branch detail, arrangement of polyps and shape of its sclerites. It appeared that S. pacifica (eastern North Pacific), S. rosea of the Atlantic and its subspecies, S. pallida (a northern Atlantic bathyal form) were strongly related. S. rosea is the nominate form, found not only in the bathyal North Atlantic, but also in the Scandinavian sublittoral (Madsen 1970). It would appear that the species described here could be the Pacific Ocean extension (of the Atlantic species S. rosea) in its distribution, having moved into the Pacific Ocean via waters circumscribing the North Pole. It can be inferred that as S. pacifica appeared in the Pacific, it dispersed down the western coast of the North American continent (at least as far as, generally, central California), just as S. rosea has apparently spread across the north Atlantic and down the eastern side of the North American continent (and presumably, into the Gulf of Mexico, perhaps as a new subspecies). The WoRMS Database (Cordeiro et al. 2019) shows the accepted status of this species.

Muzik (1979) noted discrepancies regarding locality for the holotype. The specimen is housed in the Bishop Museum, Hawaii (as Allogorgia exserta, #101), with a portion of it housed at NMNH (USNM 49513). This specimen “agrees in details of color, sclerites, and polyp size and shape with the type from Alaska of Swiftia pacifica (USNM 30024) collected from the Aleutians in 1906 and described by Nutting in 1912. One can conclude that there was an error in the locality of the so-called Hawaiian specimen. It is entered in the Bishop Museum catalog as ‘Albatross’ 2742 without locality. Entry 2741 is from Station 3353 off Panama. Prior to that station, the ‘Albatross’ had been collecting in the Pacific Northwest, so it is conceivable that this S. pacifica was collected there and later confused.” It appeared that normal range for Swiftia pacifica is from central/northern California northward, but occasionally may appear south of that range. CAS has thirty cataloged records from this genus; of these, eleven lots are from Alaska, and are likely Swiftia pacifica.

There appeared to be a distinct morphological trend, from southern to northern waters, along the California coast up through the coastal areas of Oregon, Washington and Alaska that required discussion; a proposed explanation for the range distribution of this species follows the descriptions of all species (with red color) found in or near the Bight covered in this paper. Briefly, an extensive examination of colonies collected from Baja California to the remote northern aspect of the Bering Sea (see Appendix 3: List of material examined) revealed a very distinctive trend in the appearance of colonies and sclerites from south to north. The sequential trends seen within the two species, S. kofoidi and S. pacifica (or morphs of one) are discussed in the Further Remarks section (along with variation in sclerite morphology of other eastern Pacific Swiftia species). The observation of this phenomenon has never been discerned or noted previously.

[USA], California, Santa Cruz Island, bearing N 35°E, 7 miles off Point San Pedro, ~34°02'02.76"N, 119°31'11.77"W, 447–510 fm [813–927 m].

Syntype USNM 25431 and USNM 43130 [both wet]; both specimens were examined.

~24 lots (see Appendix 3: List of material examined).

Colony (Figures 7, 8A) straggling, whip-like, not always erect; branched slightly, mostly unbranched; largest specimens ~13+ cm tall. Stem round, slender, of uniform diameter throughout. On stem/branches, polyps uniformly distributed on all sides, not crowded (Figure 8B), up to 2.0 mm apart; tubular, small, ~1.0 mm high, usually higher than broad; when polyps contracted, nearly flush with branch surface. Coenenchyme moderately thick; color of living colony, including polyps, salmon, brick reddish-pink (commonly) to coral-red throughout; sclerites reddish-pink; rods reddish-orange; sometimes long, warty spindles colored and colorless. Sclerites (Figures 9A, B, 10A–H) of three main kinds: 1) small double-spindles, rosettes, stars and/or small clubs, found mostly in superficial layer of coenenchyme (all much less numerous than second kind; clubs much less numerous than other kinds); 2) larger spindles, slender, pointed, some slightly curved, covered with regularly distributed small warts (Figure 10A–C); 3) not always numerous, but when present, very conspicuous, colored anthocodial rods (fingerbiscuit rods; Figures 9A, B, 10H); moderately to heavily warted, much shorter than long spindles, longer than first type. Polyps, generally, with spindle-shaped sclerites in walls and near/on tentacular bases, arranged more or less in chevrons. Otherwise, longitudinally arranged.

Figure 7. 

In situ shot, identified as Swiftia simplex. This is what one would expect to see of live colony. Notice strong similarity to that seen in Figure 28, Part II. As specimen was not collected, identification cannot be confirmed. Image (02_57_27_20) courtesy of Lonny Lundsten and Kim Fulton-Bennet, “(c) 1992 MBARI.”

Figure 8. 

Swiftia simplex, SBMNH 422979. A Entire colony, ~21 cm long, diameter ≤3 cm, including calyx B Branch close-up showing calyx placement on branch. Scale bar: 3 cm (A).

Figure 9. 

Swiftia simplex, specimen #81-99B-1 (Alaska Fisheries Service), light microscopy arrays. A 4× magnification, showing long, slender spindles and anthocodial fingerbiscuit-rods B 10× magnification of sclerites, emphasizing fingerbiscuit-rods of anthocodium. These rods measure between 340–350 µm, while long, thin spindles measure between 430–500 µm. Specimen provided by Bob Stone, Alaska Fisheries Service.

Figure 10. 

Swiftia simplex, specimen #81-99B-1, SEM image. A Elongated spindles B Moderate-length spindles C Shorter spindles D Thick, elongated rod-like spindles E Less jagged double-disk/capstan-like forms F Tiny, odd spindle G Irregular spindles H Anthocodial “fingerbiscuit-rod” forms typical of genus. Alaska Fisheries Service.

Earlier genus designation (Nutting, 1909), psammo- = sand); simplex- = simple, perhaps referencing the very simple, usually unbranched colony, found generally on soft-bottomed sites. However, no derivation for species name given in Nutting’s (1909) description.

Whip coral (suggested: Brick-red whip coral).

Kükenthal (1919), in Chun: California, coast to abyssal. Total distributional range (surmised from collection location data reported by various institutions) extends from southern California Channel Islands (and further west--San Juan and Rodriquez Seamounts), up the coast of California (Monterey Bay, Carmel Canyon), sparsely along Oregon coast (Tillamook Head, Columbia River), to Washington coast (Grays Harbor, Quinault Canyon, Queets; general site locations off Oregon and Washington gleaned from NMNH material), up to Gulf of Alaska, found on seamounts and elsewhere (NMNH material, Alaska Fisheries Service). Fairly recent collection event (2008, Olympic Coast National Marine Sanctuary) produced at least one sample that may be this species, collected at ~48°07'53"N, 125°05'20"W at 335 m.

Appears to prefer at least subtidal depths, generally deeper, according to collection location data; frequently encountered on Seamounts (~190–900 m). MBARI T630-A13 had attached to it what appeared to be a cluster of white eggs (cluster identity not determined); these flexible branch strands, projecting up into localized water currents, would make good attachment sites for eggs needing oxygenation and/or flow to keep them clean, being suspended above muddy bottoms found at depth.

CAS has three specimens (likely this species), all from northern California (Cordell Bank, W of Point Reyes, CAS-IZ-96739; off Pigeon Point in San Mateo County, CAS-IZ-96744 and Eel River Canyon in Humboldt County, CAS-IZ-96758), labeled as Euplexaura simplex. This is a hitherto unknown application of a genus name, done by D Harden in the early 1970s (likely, an attempt to be comparable with the then named Euplexaura marki). This genus designation is incorrect; the specimen is the species Swiftia simplex. While Nutting (1909) placed it in the genus Psammogorgia Verrill, 1868a, geographic location of specimen(s) he described, geographic locations of specimens examined here, appearance of sclerites, along with molecular work conducted by M Everett et al. (2016), do not warrant that species designation either. While the WoRMS Database (Cordeiro et al. 2019) did indicate an accepted status for this species designation under the genus Swiftia, it also showed accepted status for the species as Psammogorgia simplex (Cordeiro et al. 2018). Based on the genus description for Psammogorgia by Verrill (1868a), Bayer (1958) on the morphological characters mentioned above and discussion provided by Bayer and Deichmann (1960), which also discussed the probability of appearance in the Panamanian province, material examined in this work warrants placement in the genus Swiftia, not in the genus Psammogorgia.

Sclerite examinations revealed a few individual colonies (several species) in the genus Swiftia (such as that shown in Figure 8), with minimal/no fingerbiscuit rods. Nothing examined and identified as Chromoplexaura marki (Kükenthal, 1913) (species closest in superficial colony appearance) ever displayed these rods, as expected for this genus. It was easy to understand how identification done in the field, on in situ colonies (with water depth distorting color), could label colonies from the two species (this and C. marki) as the same organism. Current examinations discussed here shed some light on the confusion. The explanation provided regarding S. pacifica in Further Remarks section, is an attempt to clarify (and explain) why some colonies of Swiftia have scleritic anthocodial rods and others do not.

California specimens identified from the genus Euplexaura (now the genus Chromoplexaura Williams, 2013) on several MBARI video clips that were viewed could actually be this species. C. marki (which this species can so closely resemble), is usually bright deep red, with white or pale yellow anthocodiae/polyps (Kükenthal 1913, 1924; also Johnson and Snook 1927) but not always (see discussion, Part II, this work) while this species is a dull pinkish to brick red color, with colony coenenchyme and anthocodiae/polyps the same color; sclerites are very different for the two genera. It is likely that MBARI is seeing both this species and C. marki, but not able to clearly distinguish between the two due to color distortion at depth under field conditions, if not collecting.

A specimen (R1159_EPI_164_0015) collected by Olympic Coast National Marine Sanctuary in 2008 superficially appeared to be this species; polyps were mostly contracted into very round, prominent mounds, although these had larger dimension than that given in the above description (tentacles were more or less the same salmon color as the coenenchyme, but polyp bodies, closely proximal to branch, were white when dissected out). Based on further examinations, specimen was tentatively identified as Swiftia spauldingi (Nutting, 1909); however, lack of fingerbiscuit rods points in the direction of Chromoplexaura marki. Recent DNA sequencing (communications with M Everett, NOAA affiliate, 2013–2014) indicated that some Swiftia species might need subdividing (three different species or variants a possibility).

[USA], North Pacific, California, Monterey Bay, Pacific Grove.

Holotype; transferred from Hopkins Marine Laboratory Collection; [USNM 91854, wet]; specimen was examined.

~9 lots (see Appendix 3: List of material examined). [See also discussion regarding “red whip” forms, in Remarks section for species Chromoplexaura marki discussed in Part II of this work.]

Colony (Figure 11) low, moderately bushy (tending to one plane); flabellate. Sparsely branched; irregularly dichotomous, subdividing some distance above base; branches round in cross section. Terminal branches somewhat stout, 5.0–10.0 cm long, as large as main stem, nearly as round (2.0–3.0 mm thick); slender, whip-like. Polyps scattered closely, uniformly, over surface on all sides, as very low, fairly large, somewhat rounded warts; in some specimens, scarcely raised above general surface of colony, almost entirely included, hardly evident (flush), seeming to be nearly absent, yet in others readily visible; less than 1.0 mm across, 1.0 mm apart. Color of living colony bright coral or salmon-orange; sclerites range from yellowish to very pale red (most commonly, moderate to pale salmon pink), with orange rods; polyps, in preserved specimens, appear to be pure white, while in some colonies (preserved) can appear light salmon-pink. Sclerites (Figures 12A, B, 13A, B) of several kinds, but generally small, short, exceedingly warty spindles and double-spindles. Sclerites in body wall of polyps somewhat longer, more slender spindles (double-spindles), with more delicate warts and points, tapered with wide, median space, but stout, scarcely acute (but never as long as those seen in other eastern Pacific species of Swiftia). Bayer (1956) noted these longer sclerites as symmetrical or with warts on one side simple and conical, elsewhere compound. With appearance of two or three whorls of large, compound, rough warts on each end, those nearest middle usually the largest. These longer sclerites tend to longitudinal arrangement in body wall in eight rows; rows sometime extending part way up outer sides of tentacles. Stouter sclerites (double-spindles) also tapered with wide median space, but shorter, blunt, each end with two or three crowded, usually somewhat confused whorls of large rough warts, forming large terminal cluster. (Bayer, 1956 noted these sclerites as having warts of one side fused like those of disc spindles). In some colonies identified as this species, presence of sclerite form approaching that of double heads (Figure 13A, top row), with narrow median space and large cluster of closely crowded warts on each end, resembling dark, dense triangular tip; these sclerites are of particular interest in comparison with Chromoplexaura marki and the double dunce-cap sclerite. Other heads shorter, lacking median space, entirely covered with crowded warts. Crosses, with four short, roughly-warted branches said to occur frequently; not evident in examinations undertaken. Fingerbiscuit rods (Figure 13B) more heavily warted than those seen in other species from genus (but may not be abundantly present).

Figure 11. 

Swiftia spauldingi, CB#34806-455. Full colony height approximately 12 cm. Image taken by Carla Stehr, courtesy of Ewann Berntson, National Fisheries Science Center, Port Orchard, WA.

Figure 12. 

Swiftia spauldingi, CB#34806-455, light microscopy arrays. A 4× magnification B10× magnification; anthocodial fingerbiscuit-rods very obvious, measuring from 128–171 µm. The dense “ovals” measure between 100–115 µm; smallest sclerites are ~86 µm.

Figure 13. 

Swiftia spauldingi, CB#34806-455, SEM image. A An array of coenenchymal sclerites (spindles) B An array of, primarily, “fingerbiscuit-rods,” the characteristic sclerites for the genus. Images prepared by Carla Stehr, courtesy of Ewann Berntson, National Fisheries Science Center, Port Orchard, WA.

Named in honor of Mr MH Spaulding from Stanford University.

Rarely, southern California (Los Angeles County); may extend from central California, northern California Channel Islands, north to coast of Washington State (Strait of Juan de Fuca). Distribution based on specimens examined with collection location data, from several sources (NMNH, NOAA offices, MBARI). The specimen collected by Olympic Coast National Marine Sanctuary in 2008, collected at 335 m, ~48°07'53"N, 125°05'20"W, confirmed WA coastal waters as a location for this species.

Conspicuous Ophiuroidea found intertwined on branches, as those seen on specimen from “Oregon State, R/V ‘Yaquina’ NH15” (SBMNH 423073) and that collected by Olympic Coast National Marine Sanctuary in 2008.

Multiple labels (NMNH) were associated with some specimens examined, along with differences in literature usage of the genus name Swiftia and at least one specimen at NMNH had been given the name Psammogorgia spauldingi (while elsewhere, Bayer’s SEM files, the folder of SEM images for this same specimen, was labeled “Leptogorgia caryi = Swiftia spauldingi,” with the numbers “57157, SEM 2787 & 2790” [note use of the genus name Swiftia]; however, this synonymy designation is in error).

The discussion of Chromoplexaura marki in Part II (along with remarks given for S. simplex) is pertinent. At least one specimen of C. marki examined had sclerites very similar to those seen in S. spauldingi, but there were no anthocodial fingerbiscuit rods. Could S. spauldingi sometimes be seen as a less-branched colony, resembling C. marki (or be very unbranched, and look more like S. simplex)? S. simplex, S. spauldingi and C. marki can have similar colony appearance; but the first two will have the fingerbiscuit rods, and only the latter two will have sclerites showing other similarities of form (but not entirely). A key difference (and justification for keeping the two, C. marki and S. spauldingi, as separate species) is the consistent lack of fingerbiscuit rods in C. marki, but which does have the unusual double dunce-cap spindles that are only uncommonly seen in S. spauldingi (and in this latter species, usually smaller-sized; refer to Figure 13A). Bayer (1979: 1034) offered support for a synonymy between the two, but this synonymy seems questionable. More specimens will need to be collected and examined. Despite confusion regarding the status of this species, Cordeiro et al. (2019), shows S. spauldingi as the accepted name, with Psammogorgia spauldingi the only synonymized name.

[USA], California, Monterey Bay, 36°38'00"N, 121°55'00"W (bearing S 78°E, 6.8 miles) off Point Piños light-house, 755–958 fm [1373–1742 m].

Holotype USNM 25433, [wet]; specimen was examined.

None of the material examined (~16 lots) came from the SBMNH collection (see Appendix 3: List of material examined).

Colonies strictly flabellate (usually), ~15–30 cm tall, ~16–17 cm in breadth. Branches commonly anastomosed; branches dense, closely spaced. Main stem bears branches on opposite sides separated by distance of 4.5 mm to +7.0 mm; branches generally thin (no more than 1.0 mm wide) in appearance. The whole forms a loose reticulation, somewhat comparable to that seen in a few species of the genus Pacifigorgia, such as P. gracilis (Kükenthal, 1924). Polyp mounds slightly truncated to (commonly) tubular cones, 1.0 mm high or less, can be as wide as high; extended polyp can add ~1.0 mm to height; distributed primarily on sides of branches, ~2.0 mm or less apart on one lateral side. In front view, there appeared to be two opposite rows, but can be alternate; body and tentacles of polyps tend to bend (curl) toward front of colony somewhat, giving appearance of numerous polyps on the colony’s front, when just a very few, scattered, are present; often back of colony without polyps or very few. Curling of polyp body and polyp tentacles gives colony a somewhat lacy look. Color of living colony dark, purplish-red (maroon), deep brick red to nearly black throughout; when placed in alcohol, tends to nearly black. Sclerites warty spindles, generally; those on stem, branches smaller than those on polyps. Largest appear to be those in polyp walls and basal parts of tentacles; large, warty, fusiform, sometimes curved, arranged longitudinally, extending downward in meridional bands to near base of polyps. Smaller spindle-types almost with appearance of a radiate (capstan) shape; some few almost appear as disk spindles. Some few club-shaped sclerites, nearly all of which are the warty, fusiform type. Rods (fingerbiscuit shape) very conspicuous, when present, though not always numerous; generally not heavily warted; most sclerites rich reddish-purple; conspicuous rods vibrant pumpkin orange. The color combination of purple-red and orange is unmistakable.

Named in honor of Dr Harry B Torrey from the University of California.

Dwarf red gorgonian.

From MBARI, CAS and Moss Landing Marine Lab (MLML) collection data, found in Monterey Bay (‘Albatross’ stations 4514, 4537, 4546). Range may extend from northern-most end of California Bight to areas off mid-Washington coast (Quinault Canyon, 47°28'59"N, 125°11'45"W; some specimens listed as this species may be S. pacifica; further examination required). If specimens, identified as this species off Oregon and Washington coasts, are actually S. pacifica, then range for this species from at least Rodriquez Seamount, ~33°59'16"N, 120°59'52"W (west of San Miguel Passage) to Pioneer Seamount (~37°17'44"N, 123°29'58"W) off California; there are ample records at MBARI for sightings of this species in the region between these two seamounts. NMNH records extend the range up through Oregon and Washington, as far north as the mouth of the Strait of Juan de Fuca (48°32'42"N, 124°52'44"W). A recent collection by Olympic Coast National Marine Sanctuary (2008), with specimen collected at ~47°55'16"N, 125°30'00"W, 429 m, was examined, confirming species does extend up into waters of Washington State. As well, one specimen (apparently this species, not examined) was collected from San Diego, Point Loma, at 201–262 m [USNM 49522]. This would put the species in the Bight, but identification may be incorrect. Species most commonly found in the region of Monterey Bay.

MBARI records would indicate a moderately deep-water form (1,029–2,200 m). It also seems to prefer steep walls of seamounts based on collection details.

A brief description is included here as this species is often confused with others in the genus by field investigators, when simply viewing colony morphology in situ (it has been found just north of the California Bight’s upper geographic limit), and completes, to date, descriptions for all colored species within the genus Swiftia currently known to exist in the waters along the western North American continent.

In minor ways, previously published descriptions roughly matched that published for Muricella complanata Wright & Studer, 1889; Harden (1979) listed M. complanata as synonymous with this species, and an unpublished Bayer annotation noted: “Muricella complanata = a Swiftia?” Overall, however, descriptions did not match. A brief study of CAS specimens identified by Harden was undertaken, but did not sufficiently clear up his proposed synonymy. For two specimens identified by Harden as this species, one had no locality data; the other was from Monterey Bay. Identifications made by Harden often proved problematic. While definitive specimens with correct identification were needed, was not able to locate specimens with confirmed identification as Muricella complanata in any of the research collections examined so as to compare known specimens of Swiftia torreyi against it; study of new material, which needs to be collected, is required. As well, Harden (1979: 171, unpublished PhD dissertation) did designate Psammogorgia torreyi Nutting, 1909 (= Swiftia torreyi Nutting, 1909). Cordeiro et al. (2018) does show P. torreyi with accepted status, but that as well, Swiftia torreyi is also given accepted status (Cordeiro et al. 2019). Based on a number of descriptions given for members of the genus Psammogorgia and Bayer and Deichmann’s (1960) statement regarding the marine province where Psammogorgia is likely to be found, specimens examined and studied, identified as Swiftia torreyi, cannot be synonymously identified with any Psammogorgia species. At the time that Bayer and Deichmann were working (1960), they suggested that Psammogorgia would not/does not occur anywhere outside of the Panamanian province, which then encompassed the area from Cape Blanco, Peru to Lower Baja California, including the Gulf of California (Verrill 1868a, b, c, 1870). As discussed in current literature (Briggs and Bowen 2011), there now exists a California Transition Zone (CTZ), within the Oregon province, extending from Monterey, California to Los Angeles; the California province then extends from Los Angeles to Magdalena Bay, Mexico. Running south of Magdalena Bay around the tip of the Baja California Peninsula, and including all of the Gulf of California, is the Cortez province, with the Panamanian province now extending from the mouth of the Gulf of California to the Gulf of Guayaquil on the border between Ecuador and Peru. Specimens of Swiftia described in this work barely make an appearance in upper portions of the California province, but appear no further south, based on a review of all the collection location data for all specimens examined. What is of interest is their appearance in the upper California province, the CTZ and the northeastern Pacific province. In any event, that still definitively puts all specimen/species of the genus Psammogorgia outside of the California Bight, either in the Cortez or Panamanian provinces.

Bayer (unpublished annotations) contemplated differences between this species, S. kofoidi and S. pacifica; his comments do not entirely fit with what has been determined for the species here, and in Nutting’s 1909 description. However, he stated that the species is a “slender form, . . .” whereas S. kofoidi is “stouter than torreyi. . .” (this can be confirmed). “Similar to Callistephanus pacificus Nutting, 1912, pg. 96.” “P. pacificus is Swiftia pacifica, a brighter red species, with more robust branches, found commonly in waters of Washington and Alaska (bathyal North Pacific). S. pacifica is, generally, comparatively more sparsely-branched, with distinctive bar-like sclerites on the anthocodiae and eight-radiates,” than is this species.

Several portions of statements in the discussion section of Breedy et al. (2015: 332) were of interest. Those sections read much the same as several statements this author made regarding the above five species of Swiftia in an earlier, pre-revision draft of this volume submitted for review in the spring of 2014. It was interesting to see those comments used as contrast for the new Chilean species that was described.

In the MLML collection, one specimen (C0072) of this species was found; the orange rod sclerite form, generally seen below the tentacles, and anastomosing branches were present (a note furnished with the specimen made a point of the distinctive rods). Two others were labeled as such, but either color was markedly off or, more significantly, branching pattern did not match (no branch anastomoses). Of the MBARI specimens examined, at least six appeared to be this species. Some were originally identified as S. kofoidi, but it is fairly certain they are this species; the deep purple-red color is a consistent characteristic, along with many anastomosed branches. As well, presence (or absence) of the vibrant orange rods was a telling feature; if other colonies seen in videos were collected, they should be examined for their sclerites. Overall, colony C0072 has a very distinct deep red-wine color, numerous, dense, thin, anastomosing branches, with polyps having a tendency to curl. In most colonies, a definite front and back is present; the sclerite form that is most evident and obvious in this species is the vibrant orange bacillus-shaped, fingerbiscuit rod, easily seen in a light microscopy array.

No material in the SBMNH collection (see Appendix 3: List of material examined).

Colonies likely small; branching presumed irregular; with material available, not possible to confirm plane configuration; may present only a few branches or is unbranched. A main stem could give rise to roughly alternate branches, at irregular intervals. Main stem and branches may tend to curve upwards, almost running parallel to one another; stem and branches with nearly same diameter; branches can be slightly swollen. Polyps on all sides of branches, fairly dense, roughly arising off branch surface at right angles; occasionally slanting, bending upwards; may give appearance of biserial rows, but often not distinct, usually sitting on opposite sides of branch. Polyps vertically placed, conical and prominent, perhaps slightly raised; distal-most end somewhat widened, showing eight-rayed figure in retraction. Anthocodiae appear to retract vertically into truncated tips, with polyps completely able to retract tentacles. Very few sclerites that could be extracted were generally sharp, acute needles (spindles). Present in coenenchyme (relatively thin) of polyps, coarse spindles; many unsymmetrical spindles bearing crenulated warts, jagged edges and processes. Marginal sclerites tending to converge as eight calycular processes, tips projecting more or less distinctly. In polyp body walls, spindles may be partially overlapping, transverse in orientation; not arranged in convergent double-rows. No presence of any fingerbiscuit rods could be detected. Colonies (when live?) colored in shades of brown; faded to gray or white with time (preservatives).

Holotype USA, California, San Diego County, San Diego, Point Loma, 176 meters.

Holotype NMNH 25430 [wet/dry]; all material was examined (as well as could be done), several times.

Examination of preserved material at NMNH, both wet and dry, was not at all enlightening. Specimens very small; wet material in very bad shape, due to protracted storage in formalin (while now water washed and placed in 70% ETOH, the damage had already been done, long ago). The dry fragment was very small, thin and whitish, with zig-zag appearance. This correlated with photographs shown in Nutting’s (1909) work. No other institution, where collections were examined, had any material with this species designation. Nutting’s (1909) description of the colony, being more or less flabellate and in one plane, does not negate the possibility of his specimen being in the genus Swiftia (appearance of polyps on branches is similar), but there is doubt as to whether this is a separate species; specimens in question may be badly preserved or bleached examples of something else. In general, appearance of fragments most closely resembled a species of Thesea seen in southern California waters; coloring, however, does not match most Thesea (fragments bleached?) and any species in the genus Thesea should have the distinctly large, spheroidal sclerite form (not seen in this specimen, but minimal material available to work with, highly degraded). Or it may be a species belonging to genera that can display long, thread-like colonies, such as Leptogorgia, Eugorgia (new species described in this work, Part II) or even an aberrant, bleached Swiftia. The notion that this colony form, described by Nutting, is not an accurately named species, or even a member of the genus Eumuricea, has support in final comments made by Kükenthal (1924), translated here: “(i)n no case does this form belong to Eumuricea, arguing against it in comparison is the overall construction (shape), the arrangement of the polyps in two lateral rows, their wide distance from one another, as well as the form of the coenenchymal sclerites.” This means that Nutting’s material at NMNH does not belong in the genus Eumuricea; unfortunately, with the material in such poor condition, it may never be possible to clearly confirm what genus and species the specimens do belong to. As Breedy and Guzmán (2015) have elected to place it in the genus Swiftia, Kükenthal’s comments are supported. Notably, no mention of this species is made in the WoRMS Database listing of accepted species in the genus Swiftia.

SBMNH has several lots (provided by both OCSD and LACSD) in its possession that closely resemble the fragments held by NMNH. They are without color (white) or very, very pale yellow, and show the polyp pattern seen in Breedy and Guzmán (2015: fig 11). However, they also very closely resemble, in branch form (diameter, polyp placement), some sclerites (in the predominance of longish spindles, and the unsymmetrical sclerites with jagged edges), and the dull coloring, the specimens that many field investigators in southern California are calling a paler, less common species of Thesea. The fragments shown in Nutting (1909: figs 3, 4), and those shown in Breedy and Guzmán (2015: fig. 11A), closely resemble what is seen with local, somewhat less abundant, specimens of a species of Thesea. In sclerite examinations of these paler, less common, “Thesea-like” specimens, they do not clearly match the sclerite forms that are seen in species of Swiftia, and while they come closer in matching the sclerites seen in the commonly encountered Thesea species, they do not exactly match those sclerites either. Based on that seen in numerous examinations of both locally collected specimens of both Swiftia and Thesea, encompassing a number of species, the suggestion would be that Nutting’s Eumuricea pusilla might be a species of Thesea rather than a Swiftia. Nothing can be certain until more specimens that fit his original description can be located, collected and studied. With Eumuricea rigida having been recently assigned to the genus Thesea (Ofwegen, 2014), there is a likely possibility that S. (E.) pusilla might need to likewise be assigned to the genus Thesea.

Thesea exserta Duchassaing & Michelotti, 1860 (non Gorgonia exserta Ellis & Solander, 1786) = Thesea guadalupensis Duchassaing & Michelotti, 1864.

Colonies moderately threadlike, some (rare) sparsely branched in one plane; slightly flexible branches slender, long, each ascending to slightly expanded, stout, possibly truncated, distal branch tip; terminates with flattened arrow-head-like tip; proximal end, when free, also drawn into arrow-head (looking as distal end), or with small attachment disk; axis horny; coenenchyme thin. Calyces distinct, roughly placed alternately; low-domed with eight marginal teeth formed by simple converging spindles. Sclerites of coenenchyme in two layers: outer one containing large, spheroidal/oval or plate-like bodies, outer faces of which are commonly undulated, generating a wash-board appearance (key sclerite form for genus); inner layer including warted spindles of smaller diameter.

Kinoshita (1909) and Aurivillius (1931) considered the genus Filigella synonymous with Elasmogorgia. This synonymy was called into question by Matsumoto and Ofwegen (2016) in statements made regarding two distinct species. They stated that Filigella mitsukurii is actually Euplexaura mitsukurii and that there is only one species in the genus Elasmogorgia, that being Elasmogorgia filiformis Wright & Studer, 1889 (closely resembling a species in the genus Astrogorgia Verrill, 1868); in actuality, there now are three accepted species in this genus, listed accordingly by Cordeiro et al. (2019) in the WoRMS Database. Considering the characteristics of the two genera (Euplexaura Verrill, 1869 and Elasmogorgia), neither E. mitsukurii or E. filiformis belong in the genus Thesea; thus Elasmogorgia is not synonymous with Thesea. However, Matsumoto and Ofwegen (2016) did state that the genus Filigella is a synonym of Thesea. The basis for this might be the fact that the genus descriptions given by Bayer (1956a) for Thesea and Filigella overlap, in part; this would explain the suggestion of synonymy between the two genera made by Bayer in 1958. Bayer (1958) synonymized the two genera Filigella Gray and Elasmogorgia Wright and Studer with the West Atlantic genus Thesea Duchassaing & Michellotti, and transferred the genus to the family Plexauridae from the family Paramuriceidae (latter no longer a currently recognized taxon). Bayer (1981) then stated that Filigella was a synonym of Thesea; based on the recent work of Matsumoto and Ofwegen (2016), Elasmogorgia must be removed from Bayer’s 1958 synonymy, while Filigella’s synonymy might be retained. According to Utinomi (1961) however, the coenenchyme of Filigella is thinner and less distinctly displays the two layers seen in most plexaurids; as well, anthocodial armature is more powerful so as to form an operculum, typical of the group formerly known as the paramuriceids. Based on this, he considered it better to retain the genus name Filigella than to unite it with the plexaurid genus Thesea. Muzik (1979) also did not synonymize Thesea with Filigella; her rationale was that Filigella had a distinct collaret (having something more like a true operculum), along with numerous scales forming the tentacle backs. While Filigella was considered to be similar to Thesea, Thesea was stated to have “bulky boots” (Muzik, 1979: 143) forming its (collaret’s) points. She went on to surmise that, “depending on the importance of anthocodial armature, these three genera (Thesea, Filigella, along with Muricella) may remain distinct or one day be merged into one genus.” The NMNH did not use/recognize Filigella during times when author visited and worked in the collection; Thesea was the genus designation used. From examinations of specimens at NMNH, etc., noting specimen identification while also considering the synonymy discussion given here, Thesea and Filigella may or may not be synonymous; the genus Thesea is used here for colonies from California (and Mexican) waters. Elasmogorgia (and its species, including E. filiformis) is not considered, based on the recent work of Matsumoto and Ofwegen (2016). Cordeiro et al. (2019), in the WoRMS Database, indicated the genus Thesea as having accepted status, and they list some twenty-eight species within the genus; however, neither Thesea [non Elasmogorgia] filiformis (Nutting, 1909) or Thesea variabilis (Studer, 1894) are included in that listing.

As a determination of species encompassed within this assortment of specimens from the eastern Pacific has not yet been established/confirmed, information regarding type locality and identification of any type specimens must await further study.

~65 lots (see Appendix 3: List of material examined).

Colony (Figure 28A) simple; long, thin, single, whip-like (wiry), unbranched stems or sparingly branched (branch can arise near small attachment disk/base, if present, or anywhere along length of long strand, at sharp angle or nearly perpendicular to primary strand; not usually longer than main strand); many specimens show neither end of branch/stem as having a base, each terminating with a bluntly-pointed arrowhead (usually three small polyps arranged in one plane) as do all branches coming off of main stem; all strands slender and flexible. Branchlets somewhat flattened, 1.0–3.0 mm wide, 1.0 mm thick; stem/branch length generally no more than 0.3 m (≤1 foot). Calyces (Figure 28B) low (~ ≤ 0.5–0.7 mm high), conical, broad domes with basal diameter ~1.8 mm (hard to determine; calyx walls slope very gradually into general surface of thin, transparent [can be, but not usually] coenenchyme); each rising slightly above general surface, nearly right angled, on all sides of stem and branches (in some colonies, appear to stand taller). Placement slightly alternate; although some calyces tend to be lateral, and alternate, actually present on all sides of stem and any branches, somewhat distant, irregularly separated by a space ~0.0–3.0 mm. Calyces ovate (sometimes round) in cross-section; longer diameter parallel with stem. Polyps usually completely retracted, almost entirely concealed by indrawn margins. Sometimes, polyps fully retracted, with large, visible opening above tentacles; margin edge easily seen, usually displaying eight triangular teeth-like projections; individual sclerites on tentacles not easily seen. Collaret not easily seen on most specimens; may not be present at all. Color of living colonies range from yellowish-beige or tannish-beige, dirty white to bright white (also perhaps bright golden yellow?); with white or cream polyps; axis pale yellow to yellowish brown to dark brown or black. Sclerites (Figures 29A, B, 30A–C, 31A–E, 32A–C) generally medium-sized spindles (average ~0.2 mm L X 0.07 mm W); largest-sized heavy, conspicuous, densely warted (dense, elongate footballs), often one-sided, covered with very jagged projections. Also many smaller sclerites: some slender spindles with surfaces covered. Largest sclerites found on stem between calyces; although often invading walls of the latter, usually of a slightly more slender type, appearing as small, short spindles arranged transversely on lower parts; a few (almost as blunt-ended scales) with several closely layered (two or three deep), vertically placed around margin, their ends forming moderately conspicuous circlet of points, annulations or oval markings around margins when viewed from above. Collaret consists of two or more circular rows of spindles; difficult to see in many colonies (see Remarks section, below). Coenenchyme filled with compact layer of short stout spindles lying lengthwise of stem; in gross examination of coenenchyme surface, stout spindles and/or spheroidal bodies very evident, dense in number. However, largest spheroidal or plate-like bodies common to genus not always abundant in sclerite arrays, but always present, exceedingly evident, very densely warted with jagged, bumpy edges.

Figure 28. 

Thesea, spp., SBMNH 422414. A Colony 23 cm from base attachment point on rock to tip (attachment was very tenuous; rock now separated from colony) B Closer view of branches, branch tip and pattern of calyces on branch surface.

Figure 29. 

Thesea spp., SBMNH 422414. A Light microscopy sclerite array, 4× magnification, showing different sclerite forms seen in specimens of Thesea; large, dense sclerites characteristic for members of the genus B 10× magnification, illustrating not only dense sclerite form, but also common spindle form. Very densely warted sclerites range from 317–450 µm, slightly shorter, dense spindles ~220 µm, and those sclerites that are thinner, less warted measure 190–200 µm in length.

Figure 30. 

Thesea spp., SBMNH 265941, light microscopy array. A 4× magnification, showing dense sclerites as typical “football (arrow),” a key characteristic of the genus B Same specimen, SBMNH 265941, 10× magnification C One additional array, 10× magnification; shows distinctly dense warting of “typical” sclerite. Sclerites in C extracted from specimen T0-61, provided by research/survey staff, Los Angeles County Sanitation District.

Figure 31. 

Thesea spp., SBMNH 422352, SEM image. A Possible, small developing inner coenenchymal sclerites B, E Characteristic, large, spheroidal sclerites of outer coenenchyme C Tentacular sclerites D Inner coenenchymal sclerites

Figure 32. 

Thesea spp., SBMNH 422344, SEM image. A Possible tentacular sclerites B Inner coenenchymal sclerites C Large spheroidal sclerites of outer coenenchyme, characteristic of genus.

The Latin fili- = “thread;” and form-, the Latin for “form or shape;” could refer to the slender thread- or wire-like nature of the branches; this branch pattern was seen consistently in all specimens examined (regardless of what species might be represented). Thus, specimens from at least one species group could potentially be named Thesea filiformis, but could not be synonymous with Elasmogorgia filiformis Wright & Studer, 1889. For T. variabilis, variability is implied, but whether that is in regards to such characters as colony form, branch thickness or sclerite appearance, the description given by Bayer (1958) is not clear; he does, however, indicate that the colonies he examined displayed variety of color.

For this assemblage of specimens, from the northern Channel Islands, California, south to Baja, California, based on location data (see Appendix 3: List of material examined). NMNH has T. filiformis in their collection; those specimens were examined and they indeed belong in the genus Thesea (for example, USNM 30295, from San Diego, Point Loma, resembles those in SBMNH collection). It has been determined that many specimens collected by both LASD and OCSD are similar to those in SBMNH collection. NMNH has several other specimens (USNM 57172, Baja, CA; 57173 Baja, CA and 57525, from California, Point Loma) that also resembled those in SBMNH collection, or those collected by both LASD and OCSD. As to T. variabilis, the two specimens examined at NMNH (USNM 50633 and 50634) were both collected from La Jolla Canyon, San Diego County, CA.

Found on sand/soft mud bottoms to depths of at least 30 m, based on collection data for many specimens in SBMNH collection. Branches of these colonies can be found with any number of other organisms growing on, or associated with, them. On specimens examined, both wet and dry, were found: 1) round, gall-like growths formed by a species of acorn barnacle (protruding out through the coenenchyme, 2) presence of a Lepas-like barnacle (found primarily on bare, exposed axis, rather than on strands with coenenchyme intact; one specimen with a note: “Scalpellum,” where barnacle was conspicuously affixed to strand), 3) presence of ovulid snails (genus Neosimnia), 4) intertwined with arms of Ophiuroidea, 5) sometimes with other kinds of cnidarian attached at base of colony; often mixed in with species of sea pen, having the same body form (thin strand-like colonies) and 6) some with what appeared to be a kind of worm (? ribbon worm) curled around the branch strands.

Muzik (1979) made the comment that “colony shape is largely environmentally, not genetically determined” in the Thesea-Filigella-Muricella genera cluster. Colonies will be “large, planar, even branched . . . when on a large firm substrate, but filiform on rubble.” The species group discussed here consistently grew on something other than large, firm substrates (based on consistent slender, filiform appearance of branch strands), confirmed through images provided by staff of LACSD and OCSD. Their growth and presence on a softer or rubble bottom would dictate aspects of their biology, such as materials fed upon, means of anchorage (or lack thereof), etc.

Of interest are specimens belonging to the genus Thesea that are consistently, and regularly collected by Los Angeles and Orange County Sanitation District (LACSD and OCSD) staff; these look very comparable to many lots of specimens housed in SBMNH collection. Both LACSD and OCSD regularly label the majority of their collected samples as “Thesea species B;” it appeared that this is the species described here, so plentiful in the SBMNH collection. These will have a tan, light beige to dull cream coloring. Based on the number of lots housed in SBMNH collection (see Appendix 3: List of material examined), and the consistent collection records that both LACSD and OCSD report, Thesea is very common in California waters. Also of interest are specimens (determined as belonging to this genus), present in both SBMNH collection (in fewer numbers) and collected by LACSD and OCSD, that have a noticeably yellower color than the typical Thesea described above. It was the range of color displayed in the SBMNH collection lots (and of those collected regularly by LACSD and OCSD) that was of more than passing interest. Nutting (1909) stated color for Elasmogorgia filiformis from California (incorrect genus) as clear, light gray, with axis dark brown, and Nutting (1912) stated color (for same species, again incorrect genus) as rather dark brown; axis almost black (was unable to determine if these two statements by Nutting, regarding color, were for dry specimens, but likely). Multiple wet specimens examined revealed variable color in both coenenchyme and axis, without consistent color pairing, despite no real differences in appearance of sclerites (aside from color). With no apparent, clear-cut differentiation in variation of the sclerites (aside from color) in most of the arrays examined from colonies colored beige, white, or yellow, initial consideration was that all were several color morphs of the same species. At present, colonies are being treated as such; more extensive examinations currently being conducted may reveal differences that would warrant colonies of varying color being given separate taxonomic designations. Additionally, a small number of lots within the collection have far smaller, stouter (almost equal in width and length) dense sclerites; these are very similar in length to the less heavily warted spindles that are seen. The possibility that these specimens need to be separated out has not been excluded, but different environmental circumstances may account for these variations.

Determination as to whether or not an actual operculum was present on calyces was (and is as yet) not clarified. Kinoshita (1909), Kükenthal (1924) and Deichmann (1936) all specifically made mention, and described arrangements of, sclerites that form the operculum in this genus. In one known species the operculum was described as being strong, each flap consisting of three spindles forming an acute-angled triangle, reinforced by others lying parallel to these, or disposed longitudinally on distal parts of tentacles. In another species, the operculum was described as being composed of three to six pairs of spindles in converging double rows. Neither pattern was clearly seen in specimens examined. Considering the accepted definition of an operculum, and those families where one is quite evident, it did not seem that in this genus there is such a structure in the correct sense of that term; it might be more accurate to speak of a collaret, or a crown and points arrangement. Neither Kinoshita (1909), Kükenthal (1924) or Deichmann (1936) provided any clear illustrations of the situations they discussed regarding an operculum. Fabricius and Alderslade (2001) speak of the family as having crown and points, with no mention of an operculum. Despite an issue of semantics, or incorrect use of the term operculum, the simple fact was that very few colonies examined provided even one or a few calyces where a clear view could be had of what was covering over the tentacles. The eight tooth-like projections of the calyx margin were evident (on many), but any pattern of sclerites that might have been overlying the indrawn tentacles was another matter. The dry, often brittle nature of many of the specimens compounded the problem. With fresh material, more malleable to work with, this question could definitively be answered.

Regarding the species T. [non Elasmogorgia] filiformis (Nutting, 1909), Nutting reported a specimen taken from off San Pedro, California coast, in the University of California collection, as well as one described in Nutting (1909), from ‘Albatross’ station 4349, Point Loma light-house, NE 6.5 miles, 136–244 m (unable to locate them, thus these specimens could not be examined). He also reported specimens taken in the Dutch East Indies at 112 meters, and the specimen described in Nutting (1912) was taken from ‘Albatross’ station 4837, Tateisha Zaki Light, S 53 E 8 miles, 104 m. These latter two may not be the same species as the other two mentioned.

Regarding Thesea variabilis, there are no apparent specimens of the species in the SBMNH collection (see Appendix 3: List of material examined). Based on collection data for the specimens at NMNH (USNM 50633 & USNM 50634, from La Jolla Canyon, San Diego), this species required inclusion here but was unable to determine if these specimens are this species or actually a morph of T. [non Elasmogorgia) filiformis. Commonly called the White gorgonian, “Thesea variabilis resembles very narrowly by its exterior aspect some of the Thesea’s of the ‘American Indies’ ” (Bayer 1958). Drawings of sclerites from Thesea guadalupensis that Bayer (1958) provided for comparison with T. variabilis showed that the resemblance is more than superficial. The only real difference seen was that the sclerites of T. variabilis were smaller than those of T. guadalupensis (and of roughly the same size as those seen in T. [non E.] filiformis). “The sclerites are in limited accord, as are the external characters. Thus, there is no doubt that the eastern Pacific species belongs to the genus Thesea, a genus that was formerly thought to be restricted to the ‘American Indies.’ ” While noted on a list found in SBMNH files, indicating California sites and depth ranges (40–46 m, in the La Jolla area, only), there is no certainty that other coastal areas of California (even just southern California), would be an actual locality for this species. In Bayer’s personal SEM files, images were found for Thesea variabilis Studer = Psammogorgia variabilis. These did not fully resolve identification issues but does lead to further consideration of this species belonging to the genus Psammogorgia rather than the genus Thesea. Found at depths greater than 100 ft [~33 m], on the two specimens Bayer examined (1958), barnacles that were present formed prominent cysts on the branches.

Overall branching pattern described puts T. variabilis at odds with the colony form commonly seen and named as Thesea spp. in southern California, where branches simply appear as long thin strands, often with no apparent base, both ends of each strand/branch frequently terminating in a flattened, arrowhead shape. From images sent by LACSD and OCSD, many have an attachment to the substrate simply by being partially buried in the soft bottom sand or mud. USNM 50633 and USNM 50634 appeared more as a flattened bush, with indication of an actual base structure, and had branches of a thicker diameter, with a slightly more yellow coloration. They did not appear as the many colonies in the SBMNH collection, but their sclerites were very comparable. Thus, there might be the possibility that T. variabilis and T. [non E.] filiformis are the same, with variable colony form (attached forms, with actual base structure, more extensively branched, even almost as a fan, while those with no attachment base more thread-like); perhaps this is a case of different living conditions dictated by the surrounding environment resulting in different colony morphologies.

Generally, this multiple-lot assemblage is composed of specimens that are best described as a conglomerate of what could be called Thesea filiformis (with few possible morphs) or is composed of a few different species. Based on location data for all, T. filiformis for all may be the better choice; further studies will need to be done, but there is no doubt that the genus Thesea is commonly encountered in southern California waters.

Axis of strongly calcified material embedded in gorgonin, unjointed, arranged in undulated concentric layers; core not a soft, chambered central chord. Attachment base a calcareous disc; rarely, a branched, rhizoidal structure. Colonies usually profusely branched, rarely flagelliform. Polyps single, in pairs, or in regular whorls, heavily armored with calcareous scales (sclerites primarily scales in all species), permanently exsert; in contraction, tentacles in-folded. Polyps protected by eight triangular scales making up distinct operculum, below which scales of polyp body aligned in eight rows, some of which may be reduced or missing on adaxial side; rarely (single species) scales not regularly arranged, operculum undifferentiated. In coenenchyme, a layer of plates or scales, commonly elongate, some with inner layer of stellate sclerites. Scales always distinguished by cruciform extinction pattern seen in polarized light.

A rationale for the distinction between the use of the words calyx and polyp required in reference to the family. S Cairns (pers. comm.), in a conversation with P Alderslade some years ago, determined that the term calyx should be reserved for those polyps that can contract to a small mound (such as those seen in the plexaurids), and that the primnoid morphology is a polyp. Thus, there is no calyx to be seen in this family; projections and living animals are called polyps; that usage has been incorporated here.

Gorgonia verticillata Pallas, 1766 (by monotypy).

Colonies usually branched pinnately, some rarely dichotomously, mostly in one plane; axis longitudinally striated, commonly iridescent. Polyps in regular whorls, strongly bent inward toward axis. Adaxial rows of body scales reduced; opercular scales distinctly differentiated from body scales, not overreached by marginals (which do not bend inward over them); sclerites usually elaborately sculptured externally, with ridges, crests or small granules; cortical sclerites thick, pebble-like or more elongate.

WoRMS Database (Cordeiro et al. 2019) gives this genus accepted status, with this spelling.

USA, California, 218–2472 m. Possible collection location for type La Jolla, San Diego, based on work of Kükenthal (1913; 1924).

Repository of type(s) unknown.

6 lots (see Appendix 3: List of material examined).

Colony (Figure 33A) flabellate, usually branched in regularly alternate, pinnate pattern; some colonies (often main branches) rarely dichotomous; most branches in one plane. Maximum colony height over 30 cm (base excluded); average height of colonies in SBMNH collection ±15 cm. Central stem slightly bent in geniculate (jointed, zig-zag) pattern, giving off branches at angles or joints; few branches give off branchlets in similar manner. Distance between branches/branchlets on same side of central stem roughly one cm (slightly larger than one cm closer to base and less than one cm toward tips of branchlets). All branchlets unbranched, parallel to each other. Distal ends of branchlets extremely thin, more flexible, with branchlets often recurved back on themselves. Axis stiff, longitudinally striated; creamy yellow to tan, covered with fairly thin coenenchyme. Color of living colony (?)white to creamy-white; perhaps very light pinkish-beige; color in alcohol creamy whitish-beige to light tan. Five or six polyps (rarely four) regularly arranged in each whorl (most common number being five); whorl diameter 2.1–2.2 mm; generally, four to five whorls per centimeter of branch length. Minimal distance between whorls no more than 1.0 mm (often less), but evident. Polyps 2.0 mm tall; slightly clavate, covered in four to eight rows of nearly spindle-shaped (rods) sclerites; polyps strongly curved from base outward, upward and inward toward axis (Figure 33B), thus apertures directed toward stem or branch. Sclerites (Figures 34A, B, 35A–E) predominantly scales, flattened (sometimes oblong, fusiform; some appearing as flattened caveman clubs) on stem and branches, with long tooth-like spines, and radiating ribs. Outer sclerite surface may also have many small to medium-sized warts, bumps and granules. Scales imbricating (like roof tiles), fan-shaped on polyp walls. Aperture edge of polyp with ctenate marginal scales, inside of which are bases of eight opercular scales; these form tall, pyramidal opercula, with height ~0.5 mm. Opercular scales (Figures 34Bd, 35D) distinctly differentiated from body scales, not overreached by marginals, not bending inward over them. Individual opercular scales elongate triangles, especially on abaxial side, forming a conspicuous, elongated spine when polyp is fully retracted; these scales bear thickened, longitudinal ridges on their inner surface, ending with truncated points. Opercular scales up to ~0.65 mm long by 0.2 mm wide at broader end; adaxial opercular scales much smaller. Upper layer (ring) of marginal scales (Figure 35E) large, with radiating ribs, furrowed at their edge; others (proceeding proximally) show these markings feebly, if at all. Longitudinal rows of scales on polyp body commonly numbering seven (rarely eight), best seen on abaxial and lateral sides (inner lateral scales number four on each side of polyp); only abaxial rows of body scales complete. Adaxial rows reduced or absent; if present, generally two scales placed distally, two proximally, revealing large area of naked adaxial wall; thus, total number of scales within a row varies, but typically eight (with six to nine possible) scales in row; most numerous on exposed, abaxial side. Largest body wall scales, abaxial (Figures 34Ba, b, 35E); abaxial scales near tip of polyp smaller, those of adaxial side up to ~0.1 mm across by 0.1 mm tall. Lateral scales slightly smaller (Figure 34Bc). Coenenchymal sclerites (Figure 35A–C) dense, as elongated, nearly spindle-shaped rods often covered with numerous thorns or prickles.

Figure 33. 

Callogorgia kinoshitai, SBMNH 422990. A Colony; height (base missing) ~30 cm B SBMNH 422982, branch close-up, illustrating arrangement of polyps in whorls, each polyp curving strongly upward and inward toward branch.

Figure 34. 

Callogorgia kinoshitai, SBMNH 422982. A Light microscopy array, 4× magnification, showing variety of large scales (body wall, coenenchymal, opercular, etc.) as well as very small coenenchymal rod-like sclerites B Array of representative scales, 10× magnification, SBMNH 422982; a abaxial body wall scale, indicated by arrow b possible inner lateral or abaxial from polyp base c outer lateral scale d opercular scale. Opercular scales average 239 µm in length, body wall scales range in breadth from 117–217 µm and very small spindles ~28 µm in length.

Figure 35. 

Callogorgia kinoshitai, SBMNH 422982, SEM image. A Small, developing coenenchymal rods B Very flattened, “odd” coenenchymal rods C Coenenchymal rods D Opercular scales E Highly ridged abaxials (polyp tip). Sclerite forms shown here correspond to figures shown in Cairn and Bayer 2002 for species in the genus.

Named in honor of Kumao Kinoshita of Japan (Cairns 2018).

Callogorgia kinoshitai appears to extend from Monterey Bay to as far south as Chile, based on collection location data found recorded at several institutions (see Appendix 3: List of material examined). Based on specimens housed in SBMNH collection, it would appear that the species does extend further north, into waters off Oregon and Washington (USA).

Generally found in deep water (averaging 800–1,000 meters). Intertwined amongst branches may be found moderate to large Ophiuroidea, along with either what appear to be anemones (quite large, very fleshy and wrinkled) and/or possibly a type of acorn barnacle, attached to stems and branches.

Kükenthal (1919, 1924) speculated that the species C. kinoshitae(i) might be a junior synonym of Caligorgia sertosa Wright & Studer, 1889 (note error in spelling of genus name), as described by Nutting (1909). Nutting (1909) indicated five localities for what he called C. sertosa, all in the vicinity of USA, California, San Diego, Point Loma light-house. Nutting also established the type for C. sertosa, that being collected at Station 192, off Kei Island, South Pacific, 255 m, by R/V ‘Challenger’. Perhaps Nutting’s specimens from the San Diego area should be ascribed to this species rather than to C. sertosa. In any event, the two are indeed separate species. Researchers with greater exposure to, and expertise on, this species (SD Cairns) should be consulted regarding legitimacy of C. sertosa as senior synonym. Cordeiro et al. (2019) does not show this synonymy in the WoRMS Database. Earlier descriptions for both C. kinoshitae(i) and C. sertosa found in Kükenthal (1919) clearly indicated the distinct differences used to distinguish between these two species. Based on locations of collection events, with C. sertosa having its type collected from Kei Island in the South Pacific, the two appear to be separate, distinct species.

Stenella doederleini Wright & Studer, 1889; subsequent designation Bayer 1956a.

Colonies primarily branched, planar dichotomous; occasionally slightly bushy. Polyps arranged in either whorls of up to four, in pairs, or isolated, generally standing perpendicular to branch. Operculum well developed, opercular scales decidedly keeled on inner surface. Marginal scales eight, in alternate position with respect to opercular scales. All polyps, generally, completely covered with five to eight longitudinal rows of body wall scales; outer surfaces covered with small granules. Coenenchymal scales arranged in one layer. Tentacular rods sometimes present.

Genus holds accepted status, shown in WoRMS Database (Cordeiro et al. 2019).

USA, Oregon, west of Cape Meares, 45°25'18"N, 125°11'01"W, 1498–1527 m.

Holotype USNM 1071799 [dry]; type was not examined.

1 lot (see Appendix 3: List of material examined).

Colony (Figure 36A) with dichotomous branching, somewhat irregular, generally in one plane; some SBMNH specimens slightly bushy, flabellate, up to +30 cm in height (next largest, 14 cm tall). In largest colonies branchlet tips tend to droop down, curling slightly back on themselves (Figure 36A); branching intervals vary from ~3.5 cm distance at lower end of main stem (near base) to less than 1.0 cm near branch/branchlet tips. Polyps (Figure 36B) with opercula well differentiated; usually spaced 0.5–1.5 mm apart in/as singles, pairs or whorls of up to three, erect (perpendicular to axis), or slightly bent downward toward stem. Polyp height 2.0–3.5 cm, flared distally with slender, delicate stalk, heavily armored with calcareous scales. Polyps found on numerous branches, tending to favor one side of colony. Axis as described for family; visible through single layer of white, translucent coenenchymal scales; dark to light brown in color. Color of living colony (?)cream or white; in alcohol, cream to light tan. Sclerites are scales (Figures 37A–C, 38A–E); marginal scales (standard number eight) alternating in position from opercular scales (latter forming distinct operculum, creating obvious projection out from polyp). Marginal scales (Figures 37Aa, 37B, 38C) all of similar shape and size, most showing broad, shallow apical flute; these generate symmetrical rosette when viewed from above. Submarginal body wall scales (Figures 37Ab, 38A) roughly arranged in eight longitudinal rows, each row with three to four scales that appear to overlap those in adjacent rows; distal end obviously rounded, no fluting apparent; flutes absent on submarginal abaxial body wall scales. Otherwise, polyp completely covered with body wall scales, including adaxial region. Opercular scales (Figures 37Ac, 37C, 38D) alternate with marginal scales (as opposed to overlapping them) around polyp; triangular shape, prominently keeled on inner surface; most all of similar size (0.5 mm in length, on average). Coenenchymal scales (Figure 38B, possibly) generally elliptical, very evident on branches (resembling sea pansy rachis or water lily pad), in one thin layer; few with irregular shape. Pinnular sclerites (Figure 38E) small rods, with granular surface.

Figure 36. 

Parastenella pacifica, SBMNH 422983. A Colony (attachment base missing), ~15 cm × 13 cm, at widest point B Branch tip close-up.

Figure 37. 

Parastenella pacifica, SBMNH 422983. Light microscopy arrays, scleritic scales. A An assortment a possible marginal scale b abaxial body wall scale c opercular scale B 10×-magnified image; marginal scale C 10×-magnified image; opercular scale.

Figure 38. 

Parastenella pacifica, SBMNH 422983, SEM image. A Abaxial body wall scales B Coenenchymal scale C Marginal scales D Opercular scales E Pinnular sclerites. Compare to those shown in Cairns 2007 (Figure 9).

The species name pacifica- in reference to its general location; stated to be closely similar to Parastenella atlantica (Cairns (2007b). Cairns suggested that these could form a geminate (twin species) pair, differing largely in having ranges in different oceans. The species designation is listed as accepted in WoRMS Database (Cordeiro et al. 2019)

Deep-water species (~1,500–2,086 m, currently known to live on the continental slope off Oregon up to British Columbia (Queen Charlotte Islands); see Appendix 3: List of material examined. Material in the SBMNH collection came from an area north of the California Bight’s northern limit; whether it will be found further south (into the California Bight) remains to be seen.

Of the many fragments/partial colonies present in the one lot from the SBMNH collection, one of the fragments has bits of a distinguishable, pale orange ophiuroid (brittle/basket star) intertwined/tangled within it. This could either be an artifact of collection or a true living condition. The specimen from Moss Landing Marine Labs (see Appendix 3: List of material examined) also showed presence of Ophiuroidea; based on the nature of their location, etc. within the colony, likely a living situation, not an artifact.

Sclerites in specimens from SBMNH were consistently a bit smaller than those from holotype shown in Cairns (2007b). Furthermore, the SBMNH material not generally in good condition; was often difficult to get good microscopic arrays showing enough of the different forms of body wall scales (abaxials, laterals and adaxials) so as to see clear differences. The coenenchymal sclerites on branches were very evident, however, and examination of sclerites showed clearly the broad, shallow fluting. While tentacular rods are considered common in this species, sometimes very few would be found in the fragments examined; the condition of many of the polyps may partly explain their absence. Further examination of undamaged colonies, collected from the same area, may better reveal their presence.

No specimens in SBMNH collection (see Appendix 3: List of material examined).

Species included here as collection records examined (see Appendix 3: List of material examined) show a distributional range that includes the California Bight. Based on those collection records, this is a deep-water species that has been collected off the west coast of Central America, and from areas of the California coast, to just north of the Santa Barbara Channel Islands (Rodriquez Seamount). It has also been collected from Monterey Bay (Davidson Seamount), north to Oregon, Washington, Vancouver Island to Gulf of Alaska; 665–1750 m. Cairns (2011) stated that the known distributional range of this species now extends west to Adak Canyon in the Aleutian Islands and the Commander Islands, Russia. This represents a substantial range, encompassing the California Bight region.

Cairns (2007b) stated that given the similarity of Parastenella doederleini (Wright & Studer, 1889) and Parastenella ramosa, “it is likely that Studer’s (1894) identification of S. doederleini from off Panama at 1,429 m (specimen missing from MCZ), taken quite close to the type locality of P. ramosa, is probably also P. ramosa.” This species is generally most similar to the type for the genus, P. doederleini (Wright & Studer, 1889). That specimen was collected from off Sagami Bay, Japan at 3,427 m. Cairns (2007b) differentiated between the two; P. doederleini has more elongated and slender marginal flutes, more delicate polyps and coenenchymal scales with one or more small rounded knobs either at their center and/or on their perimeter. Specimens of P. ramosa examined (indicated in the Appendix 3: List of material examined), exhibited polyps more distinctly directed downwards and the marginal flutes were slightly broader than those seen in P. doederleini. A check of the WoRMS Database (Cordeiro et al. 2019) show both P. ramosa and P. doederleini as separate, accepted species. As well, P. ramosa can easily be distinguished from P. pacifica by the latter having obviously broad, shallow marginal flutes, eight rows of submarginal body wall scales and the absence of flutes on submarginal abaxial body wall scales; P. ramosa exhibits narrow, tubular marginal flutes, five rows of submarginal body wall scales and abaxial body wall scales with flutes. Additionally, Cairns (2011) stated that the confirmed presence of nematocyst pads on the inner surface of the marginal scales in this species might be the case for all species in the genus, as suggested in Cairns, 2010.

Gorgonia penna Lamarck, 1815; subsequent designation by Wright & Studer, 1889: 73.

Branching in one plane, pinnate, with branches close together in many colonies. Polyps biserial, alternate (rarely, opposite) or irregularly scattered; never in whorls or pairs. All eight rows of body scales present; adaxial surface usually has fewer scales; inner face of opercular scales with inconspicuous apical keel, or none; opercular scales aligned with marginals. Sclerites of coenenchyme (some species) as scales or warty radiates in lower parts of colony and inner cortex.

Genus bears accepted status in WoRMS Database (Cordeiro et al. 2019).

N. Pacific Ocean, Japan, Honshu Island, Sagami Bay, Okinose Bank, 600 m.

Holotype USNM 50117 [dry]; branch (from holotype), donated by Tokyo Imperial Museum; this material was examined. Main colony presumably still housed in collection at Tokyo Imperial Museum (all scientific and “natural materials” collections housed separately at what is now called the National Museum of Nature and Science); was unable to verify or confirm catalog number.

~33 lots (wet/dry) (see Appendix 3: List of material examined).

Colony (Figure 39A) exhibits dense, alternate, pinnate branching in one plane, leading to flabellate form. Main stem somewhat flattened, giving rise to alternate main branches at irregular distances; both main stem and branches may subdivide. Each main branch gives forth regularly alternate, slightly smaller branches that do not subdivide. Branchlets flattened, 1.5 mm thick (Figure 39B). Polyps small, short, cylindrical projections, 0.5 mm tall (to summit of operculum), 0.5 mm across, 1.5 mm apart; arranged laterally in two opposite rows on flattened stems, branches and branchlets; some polyps placed such that they project toward a front side of colony, with back of colony smooth; strictly alternate to strictly opposite in different parts of colony, with upper edge of one polyp ordinarily reaching to base of next one above. Polyp aperture pointed upward, slightly outward. Walls of polyps armed with sclerites; these conspicuous, flattened scales, vary greatly in size and form in different polyps. Color of colony (? alive) generally white; dry or in alcohol, dull creamy-white; some preserved colonies light grayish-brown, with surfaces of stem and branches being more distinctly gray. Sclerites (Figure 40A–H) quite varied in form, generally more or less flattened into scales; thin, cycloid. Key characteristic sclerite a flattened basal portion bearing on its distal edge long thorn-like processes (spines) projecting above margin of polyp (Figure 40E). Many scales ornamented with convex, ctenate margin. Surfaces of scales ornamented with evenly, closely distributed granules, irregularly placed nodular warts and occasional spines. Typical arrangement of scales on polyp wall is eight longitudinal rows, each row having roughly four scales in a ring; two proximal rings composed of broad curved scales with their distal convex edges ctenate, distal-most marginal ring composed of scales (with no keel), bearing prominent thorn-like, unwarted spines extending beyond end of operculum. Marginal spines usually number from two to six, two of which (abaxial) are often distinctly longer than the others. Operculum composed of eight irregularly shaped scales, not keeled, points of which often joined into spine-like processes (Figure 40F–H). Adcauline opercular scales reduced to narrow band, the antero-lateral processes from proximal rings of sclerites being the only ones that meet to complete the ring on abcauline side.

Figure 39. 

Plumarella longispina, SBMNH 422394. A Colony, 14 cm tall × 15 cm wide B Branch tips. Scale bar: 2 cm (A).

Figure 40. 

Plumarella longispina, SBMNH 422394, SEM image. A, B Body wall scales C Flatter coenenchymal scales D Marginal scales E Marginal spinous sclerites F–H Opercular scales (worn). Compare/contrast these images with those shown in Cairns 2011 (Figure 5, P. spicata).

From the Latin, longi- = long and spina- = spine; long-spined, referencing the spinose marginal sclerites that extend beyond end of operculum on polyps.

Found off California coast between ~55–735 m. Of specimens examined, could not confirm that this species is found off the Oregon coast (thus far, all specimens examined were collected either from Baja California [Mexico] and California [USA] or Washington [USA]; it seems odd that it would skip an entire area between CA and WA). Based on material collected by staff of Olympic Coast National Marine Sanctuary (May, 2006 and July, 2008) that was examined, seen off northwest Washington coast at depths of at least ~208–309 m. Specimens from the genus have been taken in Alaskan waters (Bering Sea, etc.) in depths from 85–2514 m; collection data for these specimens can be found by doing a search of the online data base for the NMNH, Smithsonian, Invertebrate Collection. Listings of this particular species (by Wing and Barnard 2004; Heifetz et. al. 2005; and Stone and Shotwell 2007) mentioned in Cairns, 2011, could not be confirmed.

Work by Lissner and Dorsey (1986) along Tanner and Cortes Banks and the Santa Rosa-Cortes Ridge area off southern California showed a depth range as follows: at depths <67 m the species is sparse, at depths ranging from 67–122 m the species is common to abundant, and at depths below 122 m, again becoming sparse. Deep-water video images taken by MBARI indicated the possibility of the genus (perhaps this species) being more common at greater depth (at least in some areas) than once thought.

In all specimens examined, only one had any other organism associated with it; on this specimen there appeared two anemones, both on branches near the tip. One, the larger of the two, is on the exposed axis. On this same specimen, on the area of branches just above the base, there appeared to be the anchor tendrils from the egg case of a shark. These tendrils are quite thin, but with the stiff curl they usually display. Egg cases were noticeable on specimens collected by OCNMS in May 2006. Colonies of this species are quite rigid, so it is likely that they provide good anchorage.

A key data point in the distribution of this species was Nutting’s specimen locality (1909): ‘Albatross’ station 4359, Point Loma light-house, 32°42'00"N, 117°14'00"W (N 85, E 9 miles), 191 fathoms (347 m). This specimen currently housed at NMNH (USNM 25429); specimen was examined.

In a comparison with a different species (from the Aleutian Islands, Plumarella spicata Nutting, 1912), it presented marginal scales that were similar in shape to those seen in this species, but the spinous process of the marginal scales in P. longispina are much less ornamented. As well, all of the operculars in the species described here display areas of surface that appear very smooth and undecorated; in P. spicata, surface ornamentation is more prevalent, although perhaps not continuous along entire surface. Colony form of P. spicata (delicate and flimsy, more or less dichotomously branched), does not match what is seen for this species.

Unless there are very subtle differences, e.g., characteristics that might specify several subspecies, this species seemed to be one of the most abundant deep-water primnoids occurring in the California Bight (and elsewhere). Its overall colony form is quite distinctive, and easily recognizable. While appearing to be quite delicate, closer examination and handling indicated that it is actually fairly hardy. In the near future, an examination of all specimens in the SBMNH collection will have to be undertaken, with special attention paid to any feature(s) that could be assessed as a key characteristic that might show some degree of variability. The question arose as to whether there are transitional variations over the entire range of this species, and if so, whether those variations might subdivide the specimens, such that they point in the direction of distinct subspecies (or for that matter, species). Molecular studies on any of those groupings could add further clarity. However, it may be that this is simply an enormously successful species, thus very common, with adequate and successful dispersal abilities. Cordeiro et al. (2019) shows P. longispina with accepted species status.

Gorgonia lepadifera Linnaeus, 1767 (= Gorgonia resedaeformis Gunnerus, 1763), by monotypy.

Dichotomously branched, arborescent form with polyps not arranged in whorls but closely crowded on all sides of branches and branchlets; polyps distinctly curved downward toward axis. Marginal scales eight, operculum strongly developed. Tentacles bear small, thorny rods.

This genus, in the form of P. resedaeformis (Gunnerus, 1763), has been known since the earliest days of science (1605), and P. resedaeformis (Atlantic species to which the species discussed here is most closely related) is one of the most often reported deep-water octocorals. It is likely that the Atlantic Primnoa were some of the very first deep-water octocorals to be seen and acknowledged (Cairns and Bayer 2005).

Genus, with accepted status in WoRMS Database (Cordeiro et al. 2019), mentioned here due to evidence provided by collection records examined (see Appendix 3: List of material examined), which showed a distributional range that includes the California Bight. Based on those collection records (NMNH), this is a deep-water form that has been collected off the California coast, from a southern location of La Jolla, San Diego to a northern location of Monterey Bay. As well, NMNH, OCNMS and MBARI, through collection and video records, also indicated sightings or collections off Oregon, Washington, British Columbia to the Gulf of Alaska, at 64-≥1000 m. This represents a substantial range but does include the entire California Bight region. Additionally, Cairns and Bayer (2005, 2009), along with Heikoop et al. (2002) and Risk et al. (2002), reported species of Primnoa from the subantarctic Pacific sector. Sánchez (in Gordon 2009), reported species of Primnoa off New Zealand coasts.

No specimens in SBMNH collection (see Appendix 3: List of material examined).

Members of this genus display, in texture and color (when preserved in alcohol) that reminiscent of large-curd cottage cheese, arranged into branches. P. resedaeformis from the Atlantic is known to Canadian fishermen commonly as Seacorn or Popcorn coral. (On a first examination of preserved specimens, which were creamy yellow-white in alcohol, the appearance of popcorn immediately came to mind.) Information given here primarily focuses on Primnoa pacifica typical; known distribution ranges from Honshu, Japan; California, north to at least the Aleutian Islands and Gulf of Alaska (Cairns and Bayer 2005). P. pacifica is known by some (anecdotal, via fishermen working in the Pacific) as Red tree coral (when living, the colony’s pink color is quite beautiful), as well as Seacorn or Popcorn coral.

This species has now been synonymized with P. willeyi Hickson, 1915, following work done by Cairns and Bayer (2005, 2009); this is shown in the WoRMS Database (Cordeiro et al. 2019), where it is considered in the Database as Primnoa pacifica var. willeyi (Hickson, 1915).

Research staff at OCNMS originally believed that Primnoa occurred only on hard substrates (such as large boulders, and exposed bedrock) in areas of low turbidity, at a minimum yearly temperature of 3.7 °C, at depths of at least 9–800 m (Brancato et al. 2007). However, the OCNMS expedition in May of 2006 noted its location at several sites having muddy or sandy bottoms.

Verrill noted, in his original unpublished notes for the ‘Blake’ Expedition manuscript (transcribed by Bayer in personal notes but not published with the plates in Bayer and Cairns 2004) that in the deep sea, because of the “absolute stillness of the water,” many deep-dwelling forms exhibited extreme delicacy and fragility. He noted that numerous examples of the more delicate features occurred in the family Primnoidae. At depth, this species has been found with crinoids intertwined amongst its branches; it may further provide shelter within its branches for species of deep-water rockfish (Brancato et al. 2007). While Verrill noted (unpublished personal note transcriptions made by Bayer) that many of the deep-sea Alcyonaria are “phosphorescent” (bioluminescent), no recent information was found that could confirm/deny that characteristic for this species, or any other in the family.

The specimen mentioned in the Appendix 3: Other material, may be the southern-most report in eastern Pacific (USA) waters for a specimen of this genus and species. Occurring as far south as La Jolla, California, it may also be found further north into the California Bight, perhaps off the Channel Islands, in deep water. Of the thirteen genus records noted at CAS, the majority of specimens are from Alaska; their only record of this species is from the Sea of Japan. MBARI has records (provided by L Lundsten) for colonies known to belong in the Family Primnoidae, but most are not identified to genus or species. It would not be surprising if some of those specimens represent species within this genus, if not this species. As this manuscript was in preparation, a colony fragment (this genus and likely this species) was located (by myself and my research student, C Schaefer, in 2015) in material sampled from LACoMNH; fragment was found in fishing nets in 1981, set in SW Alaskan waters.

Primnoa regularis Duchassaing & Michelotti, 1860.

North Atlantic Ocean, Caribbean Sea, St. Lucia, south of 13°36'27"N, 61°03'36"W, 514 m.

Neotype of type housed at NMNH (USNM 49385, wet); not examined.

None housed at SBMNH.

Colonies of moderate size (to 50 cm height), branched dichotomously or pinnately (some few trichotomously) in single plane, or unbranched. Polyps conspicuous, facing downward, in discrete whorls or pairs, non-retractile. Axis continuous; strongly calcified, especially in lower branches; generally grey to black, sometimes with metallic sheen; down center of axis (longitudinally grooved) is solid core of calcareous material. Base a discoidal holdfast, for attachment to solid substrates. Sclerites are scales, on polyps, usually numbering sixteen to eighteen on each polyp (not counting tentacular sclerites), arranged in three or four pairs of large unfused abaxial body wall scales that partially encircle polyp, but rarely meet adaxially; arranged so as to have definite pattern and number. With adaxial buccal scales commonly present, one well developed buccal in each row. Operculum consists of eight (four pairs) generally triangular scales, each with distinct longitudinal medial keel on inner surface, with corresponding trough on outer surface. Tentacles can contain few to numerous, minute, flat rodlets; coenenchymal sclerites elongate or elliptical scales, often with tall longitudinal keels.

Exclusively deep water (55–4,594 m), found worldwide (Cairns and Bayer 2008; 2009). The genus is noted (Cairns and Baco 2007) as having the second deepest location record of all primnoid genera (4,594 m in the Gulf of Alaska). In addition to species from the Atlantic, there are some 23 species recorded (Cairns and Baco 2007) from regions in the Pacific (Alaska, Japan, Hawaiian Islands, Indonesia and eastern Pacific). Also, a few species are recorded from either the SW Indian Ocean, the Galápagos Islands or off Antarctica (Cairns and Baco 2007). Cairns stated (2007b) that species of Narella have been found along the southern California coast, from both San Marcos Seamount (2,193 m), and Rodriquez Seamount (664 m). These specimens were very fragmented and could not be definitively identified, but each one may represent an undescribed species. Some 54 named species are considered valid within this genus currently, as seen in the WoRMS Database, listed by Cordeiro et al. (2019).

In Studer’s 1894 description of N. ambigua, he discussed the presence of an annelid worm from family Eunicidae Berthold, 1827 that had established itself on the coenenchyme. It apparently sought shelter under the wing-shaped extensions, in a space (a tunnel of sorts) produced by the greatly enlarged basal scales of each of the neighboring polyps. In personal note transcriptions (unpublished) made by Bayer, Verrill had outlined thoughts he had concerning the deeper water gorgonians. Regarding the annelid worm found in Studer’s specimen, Verrill (unpublished personal note transcriptions made by Bayer) discussed a comparable situation and referred to Studer’s 1894 examination.

The genus is presented here; based on collection records examined (NMNH), there is indication of a distributional range that includes the California Bight. Based on those collection records (NMNH), this is a deep-water genus that has been collected (if only as fragments) several times off the southern California coast (Cairns 2007b).

According to Cairns and Baco (2007) and Cairns (2007b), there were some 38 recognized species (that number has increased, according to Cordeiro et al. 2019), making this a species-rich genus; in fact, it is said to be the most prolific of the primnoid genera (Cairns 2007b). Of those, there may be at least a few species that could potentially be found in or near the California Bight; Narella ambigua Studer, 1894 is one species that might yet be found in the Bight. CAS has seven records of this genus (none identified to species), coming from Hawaii and Alaska. MBARI has posted on-line images of those in this genus found on Davidson Seamount, photographed at depths of 2,669 and 3,079 m. Only one specimen identified to this genus has been recorded as having been collected by MBARI staff, but there are a few additional video observations. This one collected specimen was taken in the general area slightly north and west of San Miguel Island, California Channel Islands. Of interest is a specimen housed at NMNH; from California, Rodriquez Seamount, W of San Miguel Passage, 34°02'17"N, 121°02'49"W, 662 m; coll. unknown, date unknown; USNM 1027059 [wet]. The MBARI specimen and the one at NMNH appeared to be from the same collection event. It is the shape and sculpturing of the abaxial body wall scales that are the best means to identifying a species in the genus; however, finer details regarding sculpturing of scales can only be seen with SEM. Further work with unidentified species housed at CAS and NMNH should be undertaken.

Axis distinctly segmented, composed of alternating purely horny (gorgonin) nodes and nonscleritic calcareous, mostly solid, internodes (in some, hollow); internodes may be colored, quite smooth or with small projections or ridges. Base may be either a root-like calcareous structure for anchoring colony in soft substrate or a basal disc for attachment to hard object. Colonies whip-like, profusely branched, bushy or fan-like, with polyps retractile (or not). Majority of species in family found in deeper waters; all members of family commonly called Bamboo coral.

Mopsea arbusculum Johnson, 1862.

Atlantic Ocean, Canada, Nova Scotia, Sable Island, ~43°56'10"N, 59°56'10"W, 503 m.

Type (status not researched); YPM 4744 [dry]; as Acanella normani Verrill, 1878a, now considered synonymous with Acanella (Mopsea) arbuscula (Johnson, 1862).

No specimens of this genus in collection at SBMNH.

Colonies densely or openly bushy, moderate-sized (no more than 20 cm); usually anchored in soft substrates (ooze or fine sand) by lobate, root-like holdfast, in deep water. Colonies generally larger and compressed (to one meter in height) when attached to hard substrates. Internodes white; nodes generally some shade of brown. Branched in whorls (three to six, at least in upper parts) from horny nodes; internodes solid, shorter (up to 2.0 cm). Polyps generally non-retractile, often prominent, columnar; coenenchyme thin. Sclerites of polyps mostly spindles; some flattened blunt rods, with fine prickles or low warts. Larger spindles and/or rods in body wall; sometimes rods conspicuously projecting between bases of tentacles. Small, slightly flattened, sometimes thorny, rods and/or double stars in pharyngeal walls.

While members of this genus are commonly referred to as a type of Bamboo coral, no discussion of genus name derivation was found. Genus is listed with accepted status by Cordeiro et al. (2019).

Deep water, throughout all oceans, based on an examination of collection records for specimens housed at various institutions (MBARI, NMNH, CAS).

Verrill (unpublished personal note transcriptions made by Bayer) stated that most of the deep water Alcyonaria are bioluminescent; “among the ‘phosphorescent’ gorgonians, the abundant deep-sea species, Acanella normani Verrill, 1878 was very ‘phosphorescent.’ It is also very well protected by sclerites and has a highly developed root-like branching base for anchorage in the deep-sea ooze. This has allowed it to become one of the commonest and most widely diffused of all deep-sea genera.”

From examinations of recent deep-water video and digital stills (MBARI), species in this genus are usually seen on a muddy/sandy soft bottom. Acanella dispar Bayer, 1990 (a species that was described from material taken in Hawaii, and thus, found in the Pacific Ocean) is the only species noted (thus far) that inhabits a hard bottom and has a stout trunk.

Discussion of this genus included as there are reports of unidentified species (noted by MBARI in collection/video records undertaken by them) found north of the California Bight. It is not certain what, if any, species from this genus occur within the Bight, geographically lying some distance south of MBARI’s usual study locations. However, the California Bight has not been fully explored specifically for deeper water gorgonian forms; there is the possibility of species from this genus being found within it.

Andrews et al. (2005) discussed a specimen of this genus collected off San Francisco, California that was used in an age determination study of a gorgonian colony, and MBARI (posting on-line) displayed an image of a specimen, identified to this genus, sighted on Davidson Seamount, at a depth of 1,682 m (photograph taken 28 January 2006). From the MBARI data lists, roughly four specimens collected have been identified to this genus. Several other observations, without collection, have also been recorded in the area extending from southwest of Morro Bay to off the coast of Oregon (lat./long range of 35/36–45°N, 122–130°W). As for the total number of species within this genus, most are from the Atlantic; Cordeiro et al. (2019) in the WoRMS Database list 13 species. CAS has five specimens recorded, three from Japan and two from USA, Massachusetts, off Martha’s Vineyard, while the NMNH has quite a few specimens (~305), from either Hawaii, Japan, the Philippines, or Indonesia; however, the vast majority are from the North Atlantic. Pacific Ocean species include the previously mentioned A. dispar Bayer, 1990 as well as A. sibogae Nutting, 1910b and A. weberi Nutting, 1910b. Further expeditions, with collection and study, need to be done to determine if species from this genus occur within the California Bight.

Isis elongata Esper, 1788.

Generally, eastern North Atlantic; likely, Mediterranean Sea.

Location of type specimen unknown.

2 lots (see Appendix 3: List of material examined.)

Colonies sparsely branched from horny nodes, dichotomously (at ~30–35° angle; also trichotomously or lateral), generally in one plane, thus colony usually open, flat and spreading; a candelabrum shape possible. Internodes long, with axis in preserved colonies white; axis of nodes orangey-gold/brown; coenenchyme colorless. Branching not in whorls; branches moderately slender; distance from one branch to next (thus from node to node, establishing internode length) long, 3.5–4.0 cm; calcareous internodes hollow (distal tips; solid at proximal ends), longitudinally grooved, straight (or nearly so); horny nodes three-pronged, 3.0–5.0 mm tall at joints of older branches, but a simple cylinder (~1.0 mm tall) at joints of younger branches. Base of main stem a calcareous root, lobate, for anchoring in soft substrate or discoid, calcareous holdfast for anchorage on hard substrate. Polyps non-retractile and cylindrical. Sclerites of polyps mostly long rodlets that do not project between bases of tentacles; or stout, slightly prickly needles. Verrill (unpublished personal note transcriptions made by Bayer) made reference to “girdled ellipses,” which are elongated scales with rounded ends having a notable median constriction or emargination on each edge; these sclerites are normal in all Isididae.

All members of this genus are commonly called Bamboo coral, but no discussion of exact derivation could be found; genus has accepted status in the WoRMS Database (Cordeiro et al. 2019).

Deep water, likely worldwide; at depths averaging ~1,000 m (determined from collection records of various institutions, such as MBARI and NMNH).

Can grow to very large size, perhaps able to attain great age (Andrews et al. 2005). Despite the calcareous nature of the internodes, as is true of many deep-sea gorgonians, species in this genus can be somewhat delicate and fragile. Quieter waters of the deep sea likely allow for the larger size.

Inclusion of the genus reflects locality data for the few collection and video records made by MBARI and NMNH off southern California. Of particular interest is USNM 1082174; specimen collection by D Clague (MBARI) on ‘Tiburon’ dive #630, 16 October 2003 (see Further remarks, below). No sclerite preparations could be done for specimens in SBMNH collection, as no coenenchyme tissue is present.

There are some six species recognized in this genus, according to Cordeiro et al. (2019); at least two are from the Atlantic. The species described by Bayer (1990), Isidella trichotoma, Etnoyer (2008), Isidella tentaculum, and Cairns (2018a), Isidella tenuis, are confirmed from the Pacific. As access to deeper areas becomes more common, it is certainly possible that new species will be found.

Of specimens examined at CAS, none were identified to species. One specimen came from California, Humboldt County, two were from Oregon and two were from Alaska. None (as able to determine) are recorded in web-posted MBARI images, but some two dozen-plus specimens have been sampled (with even more video observations made) by MBARI in a region encompassing an area just west of San Miguel Island in the northern California Channel Island group extending northward to an area SW of San Francisco (lat./long range = 34–37°N, 121–123°W). From the collection at NMNH, the one specimen (identified as Isidella tentaculum Etnoyer, 2008) is of interest; taken off California at Rodriquez Seamount, 34°01'26"N, 121°05'59"W, 846.9 m; USNM 1082174. The holotype for this species, is USNM 1076658, collected by P Etnoyer in the Gulf of Alaska, on Dickins Seamount, 7 August 2004 (see Etnoyer 2008). A paratype of this species is found in the SBMNH collection, SBMNH 369349 (Gulf of Alaska, Welker Seamount). Only further study with collection can determine how much further south members of this genus can range, and whether or not they are present in the California Bight.

Keratoisis grayi (Wright, 1869). Some few years ago, UNESCO-IOC Register of Marine Organisms proposed the possibility of Keratoisis ornata Verrill, 1878 being a synonym of the type. Information provided on World Register of Marine Species (WoRMS) indicated that that synonymy is now accepted (Cordeiro et al. 2019).

Specific locality of type unknown; generally, bathyal, from NE to NW Atlantic Ocean; also Mediterranean Sea.

Location of the type species could not be determined.

~3 lots (see Appendix 3: List of material examined.)

Colonies (Figure 41A) branched (few and distant, ~5.0 cm from one branch to next), with branches arising at nearly 90° angle, on same side, or opposite(?); near a node or from middle to end of long (4.0–5.0 cm), calcareous internode, then slightly curving; no secondary branching; some unbranched; generally uniplanar. Base can be either root-like calcareous structure for anchoring into soft substrate or a basal disc for attachment to a hard substrate. Axis as seen in the family; internodes calcareous, white, not composed of fused sclerites, hollow (often) or solid; and purely proteinaceous, horny, shorter (2.0 mm tall), reddish-brown to dark brown nodes, alternating with internodes. Overall color of colony (preserved) creamy yellowish; coenenchyme translucent yellow. Polyps (Figure 41B) cylindrical, height between 4.0–8.0 mm; not retractile. Polyps irregularly arranged, but with tendency toward biserial arrangement; in general, somewhat curved, distal part of polyp body with eight longitudinal rows of spindles and needles, some projecting beyond tentacles. Tentacles of polyp form a rounded top, like a mushroom, with individual tentacles usually visible. Distance between polyps no more than 1.0 cm but usually less. Coenenchyme very thin, transparent; straw-yellow in specimens examined. Sclerites (Figure 42A–C, 43A–E) generally long, fusiform spindles; some (Figure 43B) very long (needles) and others (Figure 43A, C–E) more numerous, of moderate length, in coenenchyme and polyp bodies; those in coenenchyme not always obvious; polyps armed with eight-plus, needle-like sclerites (largest), often (not always) projecting beyond tentacles as sharp marginal spines between bases of tentacles, coming from eight longitudinal rows of spindles and needles. Sclerite surfaces seemingly smooth, or (if present) with dense low warts, in parallel. Stellate forms seen in pharynx. Sclerites colorless to light tan, depending on species.

Figure 41. 

Genus Keratoisis, SBMNH 422980. A Branch fragment; coenenchyme thin, translucent yellow, easily coming off underlying axis; the fragment measures ~15 cm long B Close up of polyps and very thin coenenchyme on branch fragment.

Figure 42. 

Genus Keratoisis, SBMNH 422980, light microscopy arrays. A 4× magnification, showing sclerites, most notably very long needle form B Array of shorter needles, 10× magnification C Image specifically highlights very long needles characteristic of species in genus Keratoisis. Long needle-like sclerites range from 620–775 µm in length, while very small spindles average ~80 µm.

Figure 43. 

Genus Keratoisis, SBMNH 422980, SEM image. A, C–E Moderate-length spindles B Very long, fusiform spindles (needles) characteristic of genus.

No clear derivation for this genus name was found. All members of this genus are referred to as species of Bamboo coral. Genus Keratoisis is accepted; WoRMS Database (Cordeiro et al. 2019) shows the spelling variation, Ceratoisis Verrill, 1883, as synonymized with Keratosis Wright, 1869.

It had been noted (Verrill, 1922) that this genus included some of the largest known species of the family; specimens of K. ornata (now K. grayi), from considerable depth, on the banks off Newfoundland and Nova Scotia, can be ~ four feet high. As well, this genus is composed of species that may live to considerable age. Andrews et al. (2005) stated that age for some of these coral colonies may exceed 200 years.

Discussion of the genus included here, as several fragmented specimens in the collection of SIO and NMNH were collected off California (see Appendix 3: List of material examined and discussion in this section). The speculation is that this genus may be seen throughout much of the Pacific Ocean, at depth, based on collection location data found via on-line databases, etc. Several species that might be of interest with regards to the California Bight could include K. paucispinosa (Wright & Studer, 1889) ranging from Alaska to Hawaii, K. philippinensis (Wright & Studer, 1889), which is generally a western Pacific form, ranging from Russia to Indonesia and K. flabellum (Nutting, 1908), which apparently has only been recorded from Hawaii. K. profunda (Wright, 1885), recorded from Alaska and Japan (as noted by B Wing and G Williams in Andrews et al. 2005), was at first thought to be the only species in this genus actually listed for the northeastern Pacific Ocean. However, K. profunda is no longer recognized as belonging to this genus, having been accepted as Bathygorgia profunda Wright, 1885 (Cordeiro et al. 2019).

Of the approximately fourteen specimens identified as belonging to this genus, housed in the collection at CAS, roughly half are from California; the other half from either Alaska or Hawaii. Of these fourteen, most are not identified to species, but of those that are, three of the four species mentioned above are listed, with the two records from California that have species identification listed as K. flabellum and K. philippinensis; no opportunity to verify those identifications. The southernmost California records seen previously were from Monterey Bay and from the now extensively studied Davidson Seamount; one other specimen from this genus collected/photographed by MBARI (shown on a public website) at Davidson Seamount noted it as being collected at 1,455 m. Searching Excel data sheets from MBARI (provided by L Lundsten in 2008), three dozen or so specimens have been collected over the last few years, but there are many more video observations (many of those will be multiple observations of the same organism) that have been identified as belonging to this genus. NMNH has records of specimens belonging to this genus from both Oregon and Alaska, and of note are several other specimens: those from Fieberling Guyot, 32°26'00"N, 127°47'36"W, 490 m, 16 October 1990; USNM 94443, and those from Rodriquez Seamount, 33°57'12"N, 121°08'41"W, 1840 m, 14 October 2003; USNM 1027077, both of these areas off the California coast.

Lepidisis caryophyllia Verrill, 1883; subsequent designation Kukenthal, 1915a (L. caryophyllia accepted species; proposed synonymy for Lepidisis vitrea Verrill, 1883 has been accepted as shown in WoRMS Database, Cordeiro et al. 2019).

Generally, northern and western Atlantic Ocean; bathyal.

Location of type unknown.

No specimens of this genus in collection of SBMNH.

Colonies simple, unbranched, or (rarely) sparsely branched from horny nodes; internodes hollow. In overall shape, whip-like, often exhibiting spiral growth form. Base root-like, for anchorage in deep-water bottoms of soft ooze or fine sand. Polyps non-retractile. Sclerites of polyps projecting needles and elongate scales.

No explanation was found for the rationale behind the naming of this genus; they are however, commonly called Sea whips.

A deep-water genus, likely found worldwide.

The apparent fragile and delicate nature of many deep-sea species of gorgonian in this suborder, including this genus, may demonstrate the relaxation of certain selection pressures in the deep sea, as proposed by Childress (1995) for deep-water forms. As well, many deep-water forms of Alcyonaria can be bioluminescent. This was certainly true for species described by Muzik (1978) seen in Hawaiian waters; further studies should reveal whether that feature is true of other members in this genus.

To date, there are approximately a dozen species recognized and accepted within the genus (Cordeiro et al. 2019); brief discussion is included based on location data for specimens collected (or at least noted) by both MBARI and NMNH. Both institutions have specimens that were either collected or note locations that put them in close proximity of, if not actually in, the California Bight, but only a very few described species have potential for being located within the region (although new species are certainly possible as deep-water sites are further explored). There are two specimens of interest housed at NMNH: one from California Channel Islands, San Nicolas Island, ~40 miles SW of the island, 32°31'08"N, 119°42'10"W, 950 m; coll. J Ljubenkov, no date given; USNM 59821 [wet], the other from California, Fieberling Guyot, W of Channel Islands, 32°27'36"N, 127°49'30"W, 640 m; coll. un-known, via submersible ‘Alvin’, 14 October 1990; USNM 94447 [wet]. A posted MBARI image showed a pink specimen from Davidson Seamount, at 2,683 m. MBARI data records indicated that approximately a dozen different samples have been taken, classified as belonging to this genus, with many more video-recorded sightings, in the vicinity of 32–35–37°N, 121–122–123°W. CAS has only three specimens, none of them from California waters; none of these specimens have been identified to species. This is a genus that requires further study; only with collection events south of Monterey Bay, in and near central California or some distance west of the northern California Channel Islands, will we know the extent to which members of this genus are present within the California Bight.

The description given by Studer, 1894 for Lepidisis inermis originally did not seem to fit with general characteristics ascribed to members of the genus. He did, however, in his description, mention similarities with Ceratoisis (Keratoisis) nuda Wright & Studer, 1889; this was later recognized as synonymous with Lepidisis nuda (Wright & Studer, 1889); the species L. inermis has branching from the internodes. It would appear that in some instances, sparse branching does occur in some species within the genus Lepidisis.