Axis purely horny, composed of carbonate hydroxylapatite with narrow but distinct chambered central chord; cortex little loculated, if at all. Polyps fully retractile, some forming low calyces (polyp-mounds), scattered or biserially disposed. Axis/polyp coenenchyme moderately thick, packed with spindles and capstans with regular belts of tubercles; in certain genera modified into disc spindles, scaphoids, or unilaterally spinous forms. Anthocodial armature weak, in form of crown composed of flat rodlets with scalloped edges, or lacking entirely. Colonies of diverse form, from unbranched to pinnate, closely reticulate or foliate.

Adelogorgia phyllosclera Bayer, 1958.

Genus originally included in family Plexauridae (Bayer 1958). Presence of moderately thick coenenchyme; polyps communicate directly with system of longitudinal canals. Exterior coenenchyme contains derivatives of short, stout capstans called double wheels/discs; large, leaf-like expansions (on one side) up to 0.15 mm long; spindles with tubercles in transverse rows, to 0.2 mm, some developed as leaf clubs. Interior layers of coenenchyme contain only spindles. Anthocodiae weakly to moderately armed with flat rods, 0.15–0.3 mm long.

Adelo- is Greek for unknown. When Bayer described this genus in 1958 it was a new, unknown gorgonian genus; however, Bayer did not discuss the derivation.

USA, California, La Jolla, South of Scripps Institution, La Jolla Canyon, 30–33 m.

Holotype USNM 50186; [dry]. Paratypes listed under “other material” in “Appendix 1: List of material examined” for this species.

~26 lots (see Appendix 1: List of material examined). All specimens at USNM were examined for comparison purposes.

Colony (Figure 1) heavy; bushy to fan-shaped, branching strictly in single plane, particularly in young colonies (≤ 20 cm height), but with branches occasionally growing irregularly as colony gets older (up to 0.6 m in height, usually less than 0.3 m); branching dichotomous, irregular and lateral, not pinnate (Figure 2A), with knobby to smooth branches. Color of live colony red or orange-red, polyps yellow to yellow-orange; axis slender, orange. Dry specimens brilliant red, rusty red, maroon-rust to black. Branches 2.0–4.5 mm in diameter, ascending into a meandering, sinuous form; terminal branches short (3.0–4.0 cm), slightly swollen at distal ends (Figure 2B). Trunk diameter measures up to ~6.0 mm. Axis in dead/dry specimens, within all branches of older part of colony, black, smooth, without conspicuous striations; in branches of younger portions of colony, maroon. Outer layer of axis abundantly loculate, texture of axis weak, flexible in terminal portion of branches, rather brittle in base area. Polyps with weak operculum, composed of two to four curved spindles in every segment, arranged en chevron; sclerites in polyps not arranged transversely, not forming collaret. Polyps able to retract down to surface or upper marginal edge of low to moderate calyces, these moderately elevated as low bumps off branch surface (0.5–0.8 mm tall, 1.2 mm across), situated some 2.0–2.5 mm distance apart, distributed over entire surface of all branches; margin of calyces not dentate. Sclerites (Figure 3) of polyps straight or curved rods, sculptured with simple conical warts, arranged en chevron, two to four sclerites at base of each tentacle. Coenenchyme spiculation in two layers; layers determined by shape of sclerites seen in each. Exterior layers of coenenchyme with stout capstans (0.1 mm), and spindles (0.2 mm); (latter more common), some less commonly present as having leaves or scales over one surface; few appear as leaf clubs. Numerous sclerites with sculpturing on one side modified into leafy projections (double-discs) as seen in Figure 3B (appear occasionally as sclerites analogous to disc-spindles of Eugorgia); in many cases, sclerites (0.1–0.15 mm) strongly characteristic of this form; proportion of leafy sclerites to ordinary ones in outer layer of coenenchyme varies only slightly. Ordinary ones are most abundant, while leafy ones, though sometimes rare, are always present. Axial sheath (interior layer of coenenchyme) contains symmetrical spindles (0.16 mm) only; no capstans, clubs or leafy scales. Sclerites of outer layer red; of inner layer nearly colorless.

Figure 1. 

Adelogorgia phyllosclera, SBMNH 51252. Colony measures 16.5 cm × 15.0 cm.

Figure 2. 

Adelogorgia phyllosclera. A SBMNH 422403 [dry], slightly magnified image of branches B SBMNH 51252 [wet], rounded, slightly club-shaped branch tips.

Figure 3. 

Adelogorgia phyllosclera, SBMNH 51252, SEM image. Sclerites red-orange in color. A Short warted spindles (middle and right, potentially anthocodial sclerites) B Characteristic “leaf scale” sclerites C Warted coenenchymal spindles. Compare leaf scales shown here to those seen in Bayer 1979 (figs 5c, 6c).

The root phyllo- (Greek) = leaf; sclero- (Greek) = hard scale. Species is unusual in the leafy appearance of one sclerite type, a key characteristic in identifying the species. However, Bayer gave no explanation for either genus or species names.

Chuck’s gorgonian; Orange gorgonian; Shady-leafed gorgonian; Hard-leaved gorgonian; Hidden gorgon (these names appear in a variety of field/diving guides for the area such as Gotshall 2005, Kerstitch and Bertsch 2007).

Based on collection location data, from Upper Baja, California to southern California. Generally known from La Jolla area of southern California. One specimen from Catalina Island, Bird Rock, SBMNH 51252 (one of several from Santa Catalina Island), indicates this species does range a bit north of La Jolla, California.

Commonly encountered in southern California in kelp beds; depth range of 20–300 m (Gotshall 2005). An anecdotal note (J Ljubenkov, penciled notation on a species list) stated: “Adelogorgia phyllosclera is a deep water form; it is a major deep water gorgonian and replaces Muricea on sewage pipes” (verified by staff of LACSD and OCSD). Two specimens, USNM 50186 and SBMNH 422894 (Point Loma), support numerous epizootic anemones (perhaps Epizoanthus Gray, 1867). On others, a flat, grayish incrustation (perhaps bryozoan) can be seen. Some balanoid barnacles are present over the surface of some specimens examined, in the form of prominent cysts (“galls”) on the branches, which protrude out from the axis through a coenenchymal covering. One specimen (SBMNH 422893) harbors a small brittle star, wrapped around a portion of the main trunk. No zooxanthellae present in the tissues, particularly true of USNM 50186; specimen examined for their presence by Bayer.

Among the eight to ten specimens that Bayer examined in 1958, there appeared to be three main areas that showed variation: thickness of branches, development (size) of the calyces (if present), and proportions of different sclerite forms (those with leafy sculpturing as compared to common spindles/capstans) (Bayer unpublished ms 7, Cairns 2009). Specimens at SBMNH do illustrate variation in branch diameter. Terminal branches range in diameter from 2.0–4.5 mm. Branches with smallest diameter have very distinct, prominent calyces, arising conically from their base, but thicker branches have less conspicuously prominent calyces, actually coenenchymal mounds appearing as low bumps; polyps fold into simple openings that appear as pores. One specimen at NMNH, USNM 50187, is of an extreme form; some of the branches are quite slim, bearing very pronounced calyces. It became clear that similar variations were not of taxonomic significance. Other more slender specimens exhibit a wide range of variation in branch diameter.

Most field/diving guides imply that this species is fairly contained within, and to, the region of La Jolla, California. Several of the SBMNH specimens argue against this; it appears that this particular species ranges a bit further south (upper/lower Baja) than had been previously reported. Three lots examined and confirmed correct as to their genus identification (at least) implied either: 1) a range that extends further south and/or 2) the presence of several other species, including Adelogorgia telones Bayer, 1979, and one or more of the species recently described by Breedy and Guzmán (2018), in the collection. As to potential range of distribution, Bayer (1979) made the following comment in his description of A. telones: “Although it (A. phyllosclera) seem(ed) to be rather common in the vicinity of La Jolla, collections made farther to the south, in Baja California, by the same team of divers, (did) not include it (A. phyllosclera). Neither does it occur in other collections from Baja California and the Gulf of California taken by diving or dredging, nor in collections obtained by the US Fish Commission steamer ‘Albatross’ by dredging and trawling at many localities along the coast of Central America and South America”. There are, however, several specimens in the SBMNH collection taken from northern Baja California that clearly appeared to be this species (see Appendix 1: List of material examined). Further sightings/collections would help to confirm this species’ total range (where it may either transition to A. telones, or other recently discovered species, or has a definite southern limit, with A. telones and other species then appearing some distance further south). In the examination of specimens from California locations, and those from the Galápagos Islands, in the SBMNH collection, the distinctive differences that would separate species were not clear; all specimens, with one exception (completely bleached, SBMNH 422891 from Santa Cruz Island, Galápagos Islands), are the distinctive red color of this species. A. telones, by contrast, is typically either yellow or white (Bayer 1979, Breedy and Guzmán 2018). Bayer (1979) stated that, in a comparison of the two species, A. phyllosclera has: 1) branching that is more crooked and open, 2) polyps that form distinctly hemispherical or blunt-conical calyces (as opposed to calyces being inconspicuous or not really present at all), and 3) sclerites somewhat larger, with double wheels/discs somewhat different in shape along with being more elaborately sculptured. Based on the coloring of specimens (with one exception) in the SBMNH collection, along with Bayer’s (1958) discussion of variation in characters within this species, there are unanswered questions regarding distribution of this species and the potential for several other species, as represented in the SBMNH collection. Both A. phyllosclera and A. telones are accepted species in the WoRMS Data Base (Cordeiro et al. 2018a), along with three others.

Leptogorgia ampla Verrill, 1864; by subsequent designation Verrill 1868b: 386.

Breedy et al. (2009) did a thorough review of this genus (well represented in the SBMNH collection). Sclerites chiefly disc spindles, capstans or double discs (wheels); double discs up to 0.05 mm long, spindles 0.12–0.18 mm, not developed as clubs; ordinary spindles present in small numbers in some species. Anthocodiae unarmed; sclerites, if present, flat rods and platelets with lobed margins. Polyps fully retractile into coenenchyme, slightly raised to prominent mounds, forming polyp-mounds, in longitudinal rows; often evenly distributed on all sides of branches. Branching is lateral, dichotomous (partial) or pinnate-like, in one plane; if bushy, branches in multiple planes; no anastomoses. Axis contains network of frequently mineralized organic filaments. Colony colors quite variable, depending on species.

Central America: Pearl Islands; Costa Rica, Gulf of Nicoya.

Syntype series (Breedy et al. 2009): MCZ 723 [dry]; MCZ 7080 [dry]; YPM 1551a-d and 1629a, b [dry]; YPM 5146 [dry].

~10 lots (see Appendix 1: List of material examined). Was unable to examine the type specimens, but utilized descriptions and images noted in Breedy et al. (2009).

Collection lot examined, shown in Figures 4 (whole colony), 5 (branch magnified to show prominent polyps), 6 and 7 (sclerites, light microscopy and SEM, respectively), generally matches description given in Breedy et al. (2009: 17–20, 35). Color of branches tended generally to dark orangey-red, with coenenchyme base of polyp-mounds red, upper portion of polyp-mounds gold-orange; overall impression is that colony is basically red. Sclerites (Figure 6) either bright to deep red or yellow-green in color, mixed together. In some instances an individual sclerite can be bicolored (red at one end, other end yellow-green); sclerites as double discs, relatively large; inner wheels thin, with sharp edges, outer ones terminal, not half as large, also sharp-edged (Figure 7A) (see remarks below).

Figure 4. 

Eugorgia danianae, SBMNH 422897. Shows complex branching that creates wide, flat fan. Colony is 24 cm high × 25.5 cm broad at widest point.

Figure 5. 

Eugorgia danianae, SBMNH 422897. Close-up of pinnate branching pattern.

Figure 6. 

Eugorgia danianae, SBMNH 422897, light microscopy array, 10× magnification. Note jagged, sharp-edged, double discs and color variety. Sclerites will appear orangey-yellow collectively but individuals can be deep, bright red, yellow-green, or bicolored, with a maximum length up to ~100 µm.

Figure 7. 

Eugorgia danianae, SBMNH 422897, SEM image. Representative sclerite of each form. A Double-disc B Coenenchymal sclerite C Anthocodial sclerite. Images here compare well with those shown in Breedy et al. 2009 (fig. 8).

Red gorgonian (Gotshall 1998).

As recorded by Verrill (1868a and 1868b [1869]) found in Panama and the Pearl Islands, 11–15 m (this according to FH Bradley); also seen in the Gulf of Nicoya, Costa Rica (JA McNiel). Other notations indicated that it could extend down to Peru (see Breedy et al. 2009: table 4 for full, known distributional range). Based on one specimen, USNM, 57302, taken in Escondido Bay, near San Diego County, San Diego, California, the overall range would extend from San Diego, California (at least), through Central America, possibly down to South America. Assuming E. daniana is a distinct species, it is then present in Central and Lower Gulf of California, living in the same area with Eugorgia aurantiaca Horn, 1860. Thus, southern-most end of the California Bight may be the northern-most limit (and as a separate species, E. daniana extends just a bit further north and further south than E. aurantiaca).

Generally found on offshore reefs and islands; prefers clean, plankton-rich waters and generally found at depths to ~30 m (Gotshall 1998).

While description generally matches that given in Breedy et al. (2009), I would make the following caveats: first, the colony color as described by Verrill was as a bright yellow, streaked and blotched with dark red on both branches and polyps. The color seen in specimens here lies somewhat intermediate to that described in Olvera et al. (2018) for E. aurantiaca and E. daniana; overall, dark orange-red. Divers have anecdotally described the living colony as having slender red branches, with white (colorless) polyps.

For Verrill (1868b [1869]: 411), “In the mode of branching, the size and structure of the branchlets, and color (E. aurantiaca) closely resembles (E. daniana),” which he separated primarily on the basis of the very different size and form of the sclerites, especially of the double discs; for Verrill, these were the defining feature in designating this species as distinct from E. aurantiaca. In my examinations, the larger, “sharp” double discs were always present (distinctive of E. daniana). However, in a few specimens used for comparison, labeled as E. aurantiaca, the presence of sharper double discs was noted (misidentified specimens?); this condition was contrary to the generally slightly smaller size of the double discs that are commonly seen in E. aurantiaca, where discs are described as being generally rounded, inner and outer discs very close together. In subsequent comparisons, the polyps were not always so densely packed in E. daniana as was seen in E. aurantiaca (no overlapping of polyps). With regards to branching pattern, specimens of E. aurantiaca often had a far more distinctive (and decidedly symmetrical) pattern of pinnate branching, with majority of secondary branches (branchlets) of similar length, all generally lying in one plane as compared to that seen in E. daniana. SBMNH specimens of E. daniana nearly always had their terminal-most, thin, slender branchlets curving out of one plane. Without molecular investigation, there is no clear, definitive confirmation that Eugorgia aurantiaca and Eugorgia daniana are indeed separate species. There are many overlapping features, in terms of potential branch pattern, form, and size of sclerites, and general color. The differences could be accounted for as variation within one species. However, examination of numerous examples of both those identified as E. aurantiaca and E. daniana allowed for observation of the differences in the double discs that Verrill used to distinguish these two species (differences can be obvious). For now, it seems appropriate to recognize two separate species until further studies prove otherwise.

Of note is that WoRMS Data Base (Cordeiro et al. 2018b) does list E. aurantiaca and E. daniana as separate species, but that E. daniana has been accepted as Leptogorgia daniana. Very few species of Leptogorgia (L. ramulus Milne Edwards & Haime, 1857, is one of few) display the shorter branch lengths seen consistently in species of Eugorgia nor the irregular and pinnate branching of the Eugorgia, which are morphological characters; it is understood that this acceptance is based on the molecular work that was done by Soler-Hurtado et al. (2017). Of note is the rationale for this genus change, with Soler-Hurtado et al. (2017) noting the 1999 ICZN Principle of Priority (Article 23). In the work of Olvera et al. (2018), this species is listed as a species of Eugorgia and is not included in the list of Leptogorgia species that are discussed.

SE Pacific Ocean, South America, Peru, Piura Dept., Paita. The type locality is often incorrectly spelled as Payta.

Type–YPM 1779 [dry]; MCZ 36047 (slide of holotype).

~ 50 lots (see Appendix 1: List of material examined). Was unable to examine the type specimen, but examined many others at NMNH, CAS, etc.; this is easily recognizable and comparison to type was not necessary.

In general, all SBMNH material (along with supplemental lots) examined falls in line with the description and images provided for this species by Breedy et al. (2009: 5–7, 29, 31, 42). Whole colony (Figure 8), branch pattern (Figure 9), and basic sclerite forms (Figure 10) demonstrate the identifiable characteristics for the species.

Figure 8. 

Eugorgia rubens, SBMNH 422900. Colony 45 cm tall, 18 cm across (through broadest, middle section). Scale bar: 2 cm.

Figure 9. 

Eugorgia rubens, SBMNH 45562. Magnified, showing branching and extended polyps.

Figure 10. 

Eugorgia rubens, SBMNH 45562, SEM image. Color of sclerites purple. Representative sclerite of each form. A Anthocodial sclerites B Anthocodial sclerite or small sclerite of coenenchyme C Coenenchymal spindles D Double disc spindles. Images here compare well with those shown in Breedy, et al. 2009 (fig. 15).

Rube- is the Latin for red, or reddish, presumably in reference to this species’ color. However, there is no explanation for species name given by Verrill.

Purple gorgonian (Gotshall 1994); Purple sea fan; Reddish true gorgon. Common name not specified in Cairns et al. (1991; 2003).

From southeastern Pacific Ocean (type locality: Peru [Paita]), to southern and central California (Santa Barbara mainland and Channel Islands). Depth range from shallow subtidal to deeper than 100 meters. An extensive number of specimens were examined; collection location data shows geographic and depth ranges.

Found at depths usually greater than 10–30 m. Work by Lissner and Dorsey (1986) showed a range of depth for this species along Tanner and Cortes Banks and the Santa Rosa-Cortes Ridge off of southern California as follows: At depths < 49 meters, the species was sparse, at depths ranging from 49–79 meters, the species was very abundant to abundant, from 79–91 meters the species was commonly seen, and at depths below 91 meters was again sparse. In e-mail correspondence with C Bauder (and subsequently T Laidig at NOAA, December 2010), this species may actually occur at depths greater than originally thought, extending much further north than previously reported (the specimen that called attention to this greater depth was photographed by C Bauder at Point Lobos, Carmel Bay, at 66.5 m). A thorough examination of specimens taken from these greater depths at this, and other more northerly locations, should be done. From a list found in Museum records, depth ranges for this species from selected California sites (south to north) are as follows: Mainland: Tijuana River: 36 m; Point Loma: 21–42 m; La Jolla: 20–64 m; San Pedro: 17–33 m; Carpinteria: 36 m; Islands and Seamounts: Coronados Islands: 39 m; San Clemente Island: 9–20 m; also common around the San Benito Islands off Baja, California.

Several dry specimens examined showed the presence of distinct galls produced by a species of acorn barnacle, projecting out through the coenenchyme. One of the wet specimens examined had a pronounced mass of red algae, with sponge, hydroids, worm tubes, etc. On another wet specimen, white scaly-looking patches were present, one patch so dense it looked like a cushion. On both of these wet specimens, the masses of growth were generally only present on areas of the colony where the axis was fully exposed. There is also evidence of the presence of ovulid snails from the Genus Simnia (Neosimnia) Risso, 1826 (species S. barbarensis Dall, 1892 and S. loebbekiana Weinkauff, 1881) as well as Simnialena rufa (Sowerby III, 1832) in the branches of both California (Santa Barbara, East Beach, Slate Reefs, 24–27 m; 1 April 1967 and off Newport Beach, 6 m; 18 Dec. 1964) and Mexican-collected specimens (from Sonora, Guaymas, Miramar Cove, 0.9–2 m; October, 1965; however, latter of questionable species identification).

Type specimen donated to YPM by FH Bradley who originally received it from Mrs George Petrie. Not recorded in early monographs on the alcyonarian fauna of California (Nutting 1909; Kükenthal 1913). While I was unable to examine the type specimen, all specimens in the SBMNH research collection are readily identifiable as this species. In truth, many of the specimens in the SBMNH collection are excellent examples of this species; examination of the type was not necessary.

NMNH has several catalogued lots in their collection with location records from the Santa Barbara Channel and the California Channel Islands (many of these identified to genus only); also several lots identified to species, collected in the area of La Jolla: Scripps Canyon, La Jolla Canyon, as well as the southern part of California and Del Mar. In addition, NMNH has several lots collected by C Limbaugh. Several of these are from the La Jolla area, but as well, from a couple of locations not previously recorded, including the Richfield Oil Island, Redondo Beach, and Rocky Point, in close proximity to Point Vicente, at the south end of Santa Monica Bay. In addition, the Cabrillo Marine Aquarium, in San Pedro, California has a few dry specimens of this species in its museum, and as well, displayed live specimens in tanks on exhibit to the public; generally, all were collected from the local area. This is a very common species in Southern California waters and is an accepted species in the WoRMS Data Base (Cordeiro et al. 2018b).

Isla Cedros, Baja, Mexico.

Holotype Santa Barbara Museum of Natural History, SBMNH 422333.

~5 lots (see Appendix 1: List of material examined).

Colony an obvious whip-like form, no apparent holdfast, with minimal to no branching (not common to genus), branches fairly slender, with both branch ends pointed; sclerites double-disc spindles, with disc edges quite angular and sharp, characteristic of genus.

Colonies (Figure 11A) incomplete, with exception of one (total strand length of complete one, 58.5 cm; length of largest colony fragment, ~37 cm); diameter 1.0 mm (largest diameter up to 3.0 mm, when polyps included); few with tiny holdfast; long, thin, stiff, wiry strands, none or very few primary or secondary branches (unusual for species in this genus); what branches are present come off at right angles to main stem, then curve some five cm distant or more from branching point; branch pattern (one colony) more dichotomous or lateral. Tips of branches (both ends) terminate in small arrowhead configuration. Coenenchyme very thin; axis visible through it in some areas; color of colonies generally pure white, light creamy beige to very, very pale pink, both coenenchyme and polyps; axis red-gold, with greenish cast. Small polyp mounds (no more than 1.0 mm tall, 1.0–1.5 mm broad at base) moderately prominent, rounded, conically broad bumps arranged in nearly opposite (occasionally alternate) pattern, lateral, giving strands flattened appearance from front to back (closer examination revealed polyps on all sides); very thin ridge line (longitudinal ridge) runs down middle of both “front” and “back” of branch; appears as bare “thread,” slightly raised; overall, polyps give branches a distinct zig-zag edge on lateral sides. Polyps (Figure 11B) sit very close together, bases touching; greatest distance between polyps ~1.0 mm. Polyp aperture oval-shaped to thin slit, aligned with long axis of branch. Sclerites (Figures 12, 13) small; predominant type are double disc spindles (Figure 13B), with disc edges quite angular and sharp (common in species of Eugorgia); also, slightly longer, symmetrical spindles (more typical of species in genus Leptogorgia, Figure 13C) and occasionally, crosses (quadriradiates); all are colorless.

Figure 11. 

Eugorgia ljubenkovia sp. nov., SBMNH 472232. A View of long, coiled specimen strands; when fully extended, longest strand length ~37 cm. Strand diameter (excluding polyp mound) ~1.0 mm B Magnification of branch strand, showing small, rounded polyp mounds.

Figure 12. 

Eugorgia ljubenkovia sp. nov., SBMNH 472232, light microscopy sclerite array. A 10×, showing rounded spindles and double discs. Note apparent quadriradiate, upper right B Sclerite array at 10×, illustrating stout, sharp-toothed double discs (seen in some species of Eugorgia). Sclerite color is white, but may give colony a very, very pale pink cast when in the tissue. Largest sclerites up to 100 µm, those shorter, rounder up to 70–75 µm.

Figure 13. 

Eugorgia ljubenkovia sp. nov., SBMNH 472232, SEM image. Sclerite color is white, but may give colony a very, very pale pink cast when in tissue. A Anthocodial sclerites B Double-discs (teeth evident in most) C Coenenchymal spindles.

Proposing Eugorgia ljubenkovia, to honor John Ljubenkov, a southern California cnidarian biologist, colleague and friend of many Southern California Association of Marine Invertebrate Taxonomists (SCAMIT) members.

John’s wire gorgonian.

Known from collection events undertaken by staff of Orange County Sanitation District and one lot taken in South Bay, Isla Cedros (‘Velero IV’) in 1949; thus, at this time, known from southern California and northernmost Baja, Mexico.

Moderate occurrence, indicated by OCSD collection events; not occurring at great depth (~30–35 m). Hydroids (fuzzy mass) attached to bare axis on one colony (SBMNH 472233); elsewhere on same colony, barnacle galls, barnacles completely covered with gorgonian’s coenenchyme.

All colonies have shape of a thin, whip-like Leptogorgia species, and may exhibit the same presumed lifestyle as that of Leptogorgia filicrispa Horvath, 2011 or that of species in the genus Thesea Duchassaing & Michelotti, 1860. However, sclerites most distinctive in being largely double discs, characteristic of species in the genus Eugorgia. Originally, was tentatively identified as a possible Heterogorgia Verrill, 1868c by J Ljubenkov; these specimens did not show the characteristic collaret, point and thorn sclerites of that genus (and that genus does not display distinct double discs, as seen here). Originally, SBMNH 422333 was shelved with specimens of the genus Thesea; despite the long, thinner branch strands and possible lifestyle similarity, no large spheroidal bodies, characteristic of Thesea were found. Also, this species (in five lots), is not additional material of the species L. filicrispa (Horvath, 2011), as jagged double discs are not seen in that species, and sclerites in that species can be variably colored. This species is a unique mix of colony form seen in some Leptogorgia with the sclerites of a Eugorgia. The long, thin wiry condition of the stems may be the result of environmental circumstances, involving both substrate (sandy or soft bottom sediment) and water flow. From examination of all specimens, it seems possible that some strands have no attachment base, but instead have terminal tips at both ends of strand. Further in situ work would need to be undertaken to document the environmental conditions under which this species lives.

Gorgonia viminalis Pallas, 1766; subsequent designation by Verrill 1868a.

Sclerites primarily symmetrical spindles, most without unilateral fusion of warts to form discs; shorter ones may have warts on one side fused like those of disc spindles; long ones symmetrical or with warts on one side simple, conical, elsewhere complex tubercles in various arrangements (several whorls). Coenenchyme generally contains only spindles and radiate capstans with symmetrically developed tuberculation; warts/tubercles mostly in two whorls on capstans. Anthocodial armature flattened rods; sometimes, ovoid platelets. Colonies little-branched, long, slender, whip-like, or short with branching variable: pinnate, lateral or dichotomous, in one plane or bushy; color of colonies highly variable. Colonies either attached to substrate with holdfast or lying free on substrate. Axis consistent for family, containing network of organic filaments, frequently mineralized. Polyps fully retractile into coenenchyme; slightly raised, mound-like, around apertures.

Bayer (1951) stated that Leptogorgia contains many species in temperate and tropical waters; although represented practically around the world, the center of distribution seems to be the west coast of Central America (Breedy and Guzmán 2007, Breedy and Cortés 2011, Breedy and Guzmán 2012).

Apparently, originally collected from Chile, south of Valparaiso, and off Algarrobo. For neotype (designated here), northeastern Pacific Ocean, North America, USA, California, Santa Barbara County, Goleta, Sands Beach, ~6 m; coll. R/V ‘Vantuna’ Cruise #469, November 2001.

Location of original type specimen not known. Neotype (here designated) SBMNH 422953 [wet].

~25 lots (see Appendix 1: List of material examined).

Colony (Figure 14A) not reticulate; bushy, often lanky; branches spread out, in loosely subpinnate or dichotomous, irregular branching (Figure 14B) pattern; color of living colony orange-red to orangey salmon-pink. Limbaugh (unpublished key) described color as a rich salmon pink; polyps white; dry specimens pale orange to light salmon pink. Branches and branchlets very cylindrical, long, often greater than 30 mm in length, slender (2.0 mm), usually smooth and whip-like, with unbranched, pointed ends. Branches/branchlets lie roughly in one plane (not always); some branching in all directions. Colony height to 3 ft (~92 cm); usually 2 ft (~61 cm) or less. Polyps generally flush in complete retraction, forming oblong apertures, extending in all directions around the branches. Generally, several longitudinal grooves (in bare area between polyps) present. Tentacles on polyps taper at tips and bear two rows of lateral pinnules, slightly displaced to the oral side. Sclerites (Figure 15A–D) commonly spindles having acute or subacute warted ends extending beyond second ring of warts on either side of median girdle; also capstans (two whorls with end tufts), modified as disk-spindles. Anthocodial sclerites small rods, thin, sparsely ornamented; sclerites generally orange in color. What is shown here (Figure 15) comparable to that shown in Breedy and Guzmán (2007: Figure 14, page 24).

Figure 14. 

Leptogorgia chilensis, SBMNH 422953. A Colony measures 25 cm, maximum length, 13.5 cm, broadest width B Detail of branching pattern (branch diameter ≤ 2.0 mm), and gall formation created by species of barnacle, a common occurrence on this and other gorgonian species.

Figure 15. 

Leptogorgia chilensis, SBMNH 422953, SEM image. Color of sclerites is orange-red. A Anthocodial sclerites B–D Spindles of coenenchyme. Images closely match those shown in Breedy and Guzmán 2007 (fig. 14).

Lepto- is Greek for fine or slender; the root chilensis-, likely indicative of the original type locality. No discussion of the derivation of the species name was found.

Pink sea whip; Pink gorgonian; Red gorgonian; Common red sea whip; Chilean crested gorgon; Carmine sea spray; Violet sea spray (from a variety of field/diving guides, conversations with local divers, etc.).

Several general guidebooks, including that by Gotshall and Laurent (1979), state distribution as Monterey Bay to San Benitos Islands in Baja, California. Cairns et al. (1991, 2003) did not list this species. Specimens were collected locally (Santa Barbara area, 9–22 m) for studies done by Satterlie and Case (1978, 1979) on the neurobiology of gorgonian coelenterates. NMNH has numerous lots collected from La Jolla Canyon (USNM 50179), Scripps Canyon (USNM 50191), and southern California (USNM 52442). However, examination and comparison of sclerites taken from many “red whip” forms indicated that likely range of Leptogorgia chilensis is from Anacapa Island off the California coast (thus, from the middle of the California Bight), south, perhaps to the coast of Chile. Further discussion regarding distribution of this species can be found in the “Remarks” section of this description. That discussion may further clarify some of the confusion regarding this species of “red whip” amongst several others. Other “red whips” that extend from the middle of the California Bight northward, and overlap with L. chilensis in the extreme southern end of their range, may well be one or more different species (see “Discussion concerning diversity of “red whip” gorgonian forms,” following description of Chromoplexaura marki.)

Gotshall and Laurent (1979) mentioned that this species likes offshore pinnacles, depths of 50 to ~200 feet (15–61 m). Another guidebook (Snyderman 1987) stated that: “Reds are very common on the Channel Islands and on offshore pinnacles as far north as Monterey.” “Reds” would certainly include this species, but the term “reds” is not exclusively a reference to this species. Found at depths greater than 60 ft (18 m); at Catalina, 40 ft (12 m). Range given elsewhere as 15–60 m deep. Lissner and Dorsey (1986) recorded a maximum depth of 77 m for this species on Tanner and Cortes Banks, off southern California. From a list for California sites, both mainland and islands, with depth ranges indicated, we see: Mainland: Tijuana River: 36 m; Point Loma: 18–42 m; La Jolla: 17–64 m; San Pedro: 12–33 m; Redondo Beach: 12 m; Santa Barbara: 9 m; Islands: Rock Pile (Seamount 8 miles S. Coronados Islands): 30 m; Coronados Islands: 15–39 m; San Clemente Island: 5–21 m; Santa Catalina Island: 8–26 m; Anacapa Island: 6–9 m; Santa Cruz Island: 2–6 m; Santa Rosa Island: 5–8 m.

This species has been studied both electrophysiologically and morphologically by Satterlie and Case (1978, 1979), and has been the subject of several studies regarding its (and other gorgonian species) relationships with other organisms, such as the obligate commensal barnacle Conopea galeata (Linnaeus, 1771), formerly Balanus galeatus Linnaeus, 1771(Gomez 1973, Lewis 1978, Standing et al. 1984, Crisp 1990, Langstroth and Langstroth 2000). I have seen in multiple instances that these barnacles cluster as galls, attached to the axial skeleton of this species and are overgrown by the gorgonian’s soft outer tissue. Balanus nubilis Darwin, 1854 is recorded as having been seen on the axial skeleton of dead “Lophogorgia” (Leptogorgia chilensis) in Monterey Bay (Langstroth and Langstroth 2000) (questionable gorgonian species identification); this may be opportunistic as it populates widely different sites in addition to this species. As well, several mollusk species have been recorded in association with this species, such as Tritonia festiva Stearns, 1873, as reported by Gomez (1973) and several snails of the genus Neosimnia (now Simnia), such as Neosimnia barbarensis Dall, 1892 (Theodor 1967, as referenced in Langstroth and Langstroth 2000). An unidentified field guide indicated that the ovulid snail Delonovolva Sowerby III, 1881 lives and feeds on the branches of this gorgonian. Still other organisms may be seen associated with this species, such as other species of cnidarian; “red” gorgonian is often colonized by the zoanthid anemone, Parazoanthus lucificum, now Savalia lucifica (Cutress & Pequenat, 1960), and likely other species, ultimately resulting in the death of all or most of the red gorgonian polyps (Patton 1972, as referenced in Langstroth and Langstroth 2000). In the SBMNH collection, data for several wet specimens on the Zoanthinaria shelves indicated this gorgonian as the substrate. A specimen, SBMNH 45570, collected from Avalon area of Santa Catalina Island, has Epizoanthus induratum Cutress & Pequenat, 1960 attached, while SBMNH 45549M, collected from the NE end of Anacapa Island, has Epizoanthus leptoderma Cutress & Pequenat, 1960 attached to it and SBMNH 45550, collected from the Pinnacle off the quarry, near Avalon on Catalina Island, has Savalia lucifica (Cutress & Pequenat, 1960) attached to it. As well, a specimen of L. chilensis (SBMNH 265962), recently collected by Scott Clark in 2010, on Platform A as part of a survey for Milton Love, has approximately one third of its branches festooned with a creamy yellow zoanthid. There is some specific substratum choice indicated here, and is apparently common among colonial zoanthids. On SBMNH 422944 (see Appendix 1: List of material examined–Part II), there are large clumps of hydroid, but only on bare axis portions of the branches in the colony.

According to Langstroth and Langstroth (2000), other organisms may be found associated with this species (although there is a question as to species identification of the gorgonian they discussed, as the examples are all from Carmel and Monterey Bays, in northern California; I suspect they may actually be looking at organisms on Chromoplexaura marki). They mention the bryozoan Celleporina robertsonae, now recognized as Costazia robertsonae (Canu & Bassler, 1923), the Broken-back shrimp Heptacarpus flexus (Rathbun, 1902), which may scavenge on sclerites, mucus and even toxic tissues from the surface of the gorgonian, a caprellid amphipod, specifically a skeleton shrimp, Metacaprella anomala (Mayer, 1903), whose color may derive from their acquiring the pigment ingested while scavenging the gorgonian’s sloughed off debris, and the very small hermit crab Parapagurodes hartae McLaughlin & Jensen, 1996 (now recognized as Pagurus hartae (McLaughlin & Jensen, 1966), as noted in McLaughlin and Asakura 2004), recorded as being found only at depths of several hundred meters (presumably on this species; identification of host gorgonian may be incorrect), in southern California.

As described by Fenical et al. (1981), work was undertaken to extract what has been described as a neuromuscular toxin, lophotoxin, from several species of Lophogorgia (Leptogorgia); L. chilensis has subsequently been found to produce this chemical, as well (Fenical et al. 1981). “Therefore, the distribution of toxin-producing gorgonians extends from Panama Bay northward to Point Conception, California” (Fenical et al. 1981). Also, it appears that gorgonians are able to distinguish (chemically?) between self colonies and not-self colonies (Langstroth and Langstroth 2000, citing Theodor 1970).

Harden (1979?) stated this as being one of the most common sea whips from southern California; my examinations confirm this. However, the fact that this is such a common species in southern California has led many to assume that all “red whip” forms (or those red and moderately branched forms), are this species, to the exclusion of others. The reality is that there are other red whip species which can easily be mistaken for L. chilensis; a cursory look by eye alone can (and has led) to misidentification.

Characteristics ordinarily used for separating Lophogorgia from Leptogorgia, the flattened branches and arrangement of zooids all around the branches and branchlets, are so variable as to be useless for generic distinctions (Bayer 1951). Round as well as flattened branches may occur in the same colony, and biserial zooid distribution can be found with little difficulty. Furthermore, specimens of Leptogorgia (typical in all other respects) may have zooids distributed all around the branchlets. Bayer first placed the genus Lophogorgia in synonymy with Leptogorgia in 1951; based on the work done by Grasshoff (1988), Bayer (2000) was then able to support that synonymy. Bayer used a specimen from Santa Catalina Island, collected at approximately 15 m to conclude that: “Leptogorgia chilensis (= Lophogorgia, as labeled).”

As this is one of the most common sea whips from southern California, it is not surprising that it has often appeared in live aquarium displays. The Cabrillo Marine Aquarium, in San Pedro, California, had a number of live colonies of this species on display in the public area; all were collected in the local area. As well, the Aquarium of the Pacific, in Long Beach, had a live display of this species in one of its tanks; these also were collected in local southern California areas. However, it is not likely the only species of “whip” that appears in southern California. In any event, it is likely not a common form in northern California. The transitional areas around Point Conception (and waters northward beyond the Bight) offer some intriguing distributional scenarios that will require further exploration. While of similar appearance in general colony form and color, material collected by staff of Olympic Coast National Marine Sanctuary that I examined in Washington State the summer of 2006 (tentatively identified as Swiftia spauldingi) indicated the possibility of several other species of “red whip” along the western United States’ coast. Based on examinations of several other “red whip” forms, it appeared that the upper geographic limit for this species would be Point Conception, California. “Red whips” further north were determined to be one or more different species. This is covered further in the “red whip” discussion included in the remarks made regarding the quintessential “red whip” of the northwestern California coast, Chromoplexaura marki.

Looking at location records for specimens collected and identified as L. chilensis (with confirmed identification), I noted that if type locality is correct, L. chilensis should range from the colder waters off the coast of Chile up through the warmer waters of Central America and Mexico before again encountering the cooler waters of the southern California Bight. How is this possible? What would the depth parameters, substrate features, distance from shore and specific distributional pattern (continuous or fragmented) look like for this species? The missing type material confounds the issue. The material used for Philippi’s (1866) description of L. rosea could not be located and no recently collected material from Chile resembling this species is apparently available. Breedy and Guzmán (2007) used California specimens for their description of this species. They “do not exclude the possibility (that) the material from California actually represents another species, but (to date) it most resembles Philippi’s description.” What is needed is a new specimen collected from the Valparaiso area of Chile, so that a definitive neotype could be established, and then used for comparisons. Until that occurs, I have designated a neotype from among the specimens in the SBMNH collection. Perhaps what we now call L. chilensis in southern California waters is actually a southern California endemic in need of its own species name, with a far more restricted range than has been implied previously. This is a prime example of a situation where the presumption that any “red whip” found in the southeastern or northeastern Pacific Ocean is likely L. chilensis (considered to be quite common), is faulty. The only way to resolve questions surrounding this species is to intentionally examine all “red whip” specimens (in any collection) that may have been collected from California and to intentionally undertake the collecting of material from Chile to southern California, at discrete intervals noting not only latitude/longitude but depth.

In the multiple examinations made of “red whip” forms, a specimen of what had been identified as Leptogorgia caryi Verrill, 1868 was examined. This was a dry specimen from NMNH (USNM 5988), collected at Catalina Island, California. In examining it, along with several specimens of L. chilensis, there appeared no marked differences between these specimens, either in overall colony form or in the appearance of sclerites. My initial impression was that L. caryi Verrill, 1868 might not stand as a valid species. Interestingly, Breedy and Guzmán (2007) examined the same specimen from NMNH. Independently, they came to the same conclusion regarding L. caryi that I did; the specimen was L. chilensis, with L. caryi a dubious species designation. Cordeiro et al. (2018c) shows L. chilensis as an accepted species, while L. caryi is designated as nomen dubium. Additionally, L. caryi was linked to another “red whip,” Euplexaura (now the genus Chromoplexaura, Williams 2013a) marki, with the species E. (Chromoplexaura) marki being a junior synonym of L. caryi (Cairns et al. 2003). The sclerites in this NMNH specimen definitively put it in the genus Leptogorgia; sclerites of any specimen identified as E. (Chromoplexaura) marki certainly did not fit the sclerite description of any in the genus Leptogorgia. Thus, E. (Chromoplexaura) marki is not a junior synonym of L. caryi. This was subsequently verified (ITIS Report, accessed online June 2011). Further, a specimen identified by Nutting (1909) as L. caryi was collected near San Francisco (and well above what I believe is the upper geographic limit for L. chilensis). This specimen was supposedly deposited in the Museum of Comparative Zoology at Harvard University, but Breedy and Guzmán could not locate it for examination. Based on its collection locality, it would seem that the original identification of this missing specimen is in error. I have concluded that the range of distribution that was stated in the discussions above, and confirmed generally by Breedy and Guzmán, seems to accurately pinpoint where this particular species is found. That it may overlap several other red whip forms in the northern end of its range only emphasizes the need for thorough sclerite examination of specimens collected, particularly in the transitional area from southern to northern California. Future molecular work on “red whips” in the transitional area of southern to northern California may answer the question more definitively.

(Lectotype) Gulf of Panama, Panama, Pearl Islands; additionally (Paralectotypes) Gulf of Nicoya, Costa Rica.

Holotype, as Litigorgia diffusa Verrill, 1868); YPM 1659a [dry]. Lectotype Breedy and Guzmán 2007: YPM 1659 [dry]. Paralectotypes Breedy and Guzmán 2007: MCZ 7081 [dry]; YPM 5151 [wet]. Breedy and Guzmán (2007) believe it unlikely that YPM 1659a or 1659b are this species.

4 lots (see Appendix 1: List of material examined). I was unable to examine the designated type specimens, but again, distinctive characters of the species made this unnecessary for ID of SBMNH specimens.

Colony form (Figure 16A), appearance of branch and polyp placement (with overall effect of polyp placement that of serrated or zig-zag appearance; Figure 16B) and sclerites (Figures 17, 18A–D) correspond with that given in description of the species in Breedy and Guzmán (2007: 32–36), although in the SBMNH material examined, the anthocodial sclerites (Figure 18A) seem exceptionally long.

Figure 16. 

Leptogorgia diffusa, SBMNH 423090. A Colony measures 19–20 cm tall, 13.5 cm wide, at widest point B Close up, branch tips and placement of polyps on branch surface.

Figure 17. 

Leptogorgia diffusa, SBMNH 423090, light microscopy, 10×, showing sclerites typical of genus and this species. Note pale yellow anthocodial sclerite, upper middle portion of field, amongst deep red to mauve-pink spindles. Longest sclerites may be part of the anthocodial armature.

Figure 18. 

Leptogorgia diffusa, SBMNH 423090, SEM image. A Anthocodial sclerites; some were surprisingly and unexpectedly long B–D Coenenchymal spindles. Images, for the most part, match those shown in Breedy and Guzmán 2007 (figs 23, 24).

Latin diffuses- means spreading, perhaps in reference to open shrub-like appearance that the branches create. No discussion of the species name is given by Verrill.

From Panama and Costa Rica to southern California, at least.

The lax, flattened branches, large polyps that produce zig-zag appearance, large (and in this case, long) anthocodial rods and the dull brick-red coloring are clear diagnostic features for this species. The species is listed as an accepted form in Cordeiro et al. (2018c).

Mexico, Baja, California, off Boca Flor de Malva, SE of Punta Tosca, ~24°11'07.04"N, 111°21'03.08"W, 69–87 m.

Holotype SBMNH 423057; [dry]; Paratypes SBMNH 423079 [dry]; USNM 1106683 [dry]; USNM 1106684 [dry]; USNM 1106685 [dry].

~9 lots (see Appendix 1: List of material examined).

Colony primarily unbranched; if branched, loosely and little branched in one plane, lateral or pinnate to subpinnate, occasionally dichotomous, not usually bushy; many long (~20–30 cm), slender (0.5–1.0 mm, excluding polyps), whip-like branches (many collected and, presumably, found together as shown in Figure 19), somewhat flattened but never greatly expanded to form lamellar ridges, with free ends more than 50 mm long. Branches very slender, somewhat sinuous from end to end, seeming to curve loosely back on themselves, like fine wire, yet stiff and brittle (Figure 20); tapering towards tips, also very slender. Branches likely grow from these tips; some strands with growth tip at both ends. With specimens available, base seen on several colonies (each colony usually a single strand with none, one or two branches) quite small, and usually affixed to a small rock or pebble; majority of colonies without a base. Not definitively known whether lack of a base (no attachment) is an artifact of collection, or common condition; those with attachment are a more rare situation (no attachment far more commonly seen in collected specimens). Axis very slender; ranging in color from black/dark brown to a translucent brown or reddish brown. Color of living colony, in situ, unknown. In all specimens examined, several uniform color phases were seen. Dry specimen strands exhibited color range from mauve to salmon pink to a much lighter cotton-candy pink to cream to pure white. Polyp-mounds small, conical projections (roughly 1.0 mm in height) on each side of branch (Figure 21); polyps not crowded (1.0–3.0 mm apart), arranged alternately in one or two lateral rows along sides of the branchlets. On some branches, a thin, medial line can be seen running down middle of the flattened branch, between alternate-situated polyps. Sclerites (Figure 22) are spindles, as described by Bayer (1956) for genus, typically with an absence of other specialized forms of sclerite. In this species, spindles thick, tapered; with warts low, rounded; with acute or subacute warted ends, extending beyond a second ring of warts on either side of median girdle. Long spindles generally symmetrical; some with warts on one side simple and conical, elsewhere more complicated. Very few shorter ones with warts of one side fused like those of disc spindles. Flat, tentacular sclerites large; on average sclerites can measure 0.1 mm long by 0.05 mm wide. Generally, sclerites without much color; if colored, typically light pink.

Figure 19. 

Leptogorgia filicrispa, USNM 1106683-Paratype. Full view of an amazingly large collection of branch strands, showing the multi-colored strands; entire mass measuring 20 cm × 12–13 cm.

Figure 20. 

Leptogorgia filicrispa, SBMNH 423057-Holotype. Two single strands, each showing a different pink color variant. Image originally published as figure 1B in Horvath (2011, Proceedings of the Biological Society of WA 124: 1, 47).

Figure 21. 

Leptogorgia filicrispa, SBMNH 423057-Holotype. Close-up of branch, showing spacing of polyps; Scale bar: 1.0 mm. Image originally published as figure 1C in Horvath (2011, Proceedings of the Biological Society of WA, 124: 1, 47).

Figure 22. 

Leptogorgia filicrispa, SBMNH 423057-Holotype. SEM image, originally published as figure 2B–N in Horvath (2011, Proceedings of the Biological Society of WA, 124: 1, page 48). Image prepared by D Geiger, SBMNH. Color of sclerites variable, from white, cream, to pale, bright or deep pink, generally measuring in the range of 55–80 µm; coenenchymal spindles, scale bar 20 µm (B–F, H–J), coenencymal sclerite, scale bar 100 µm (G); anthocodial sclerites, scale bar 20 µm (K–N).

The species designation is derived from the Latin root fili- for thread, and the Latin root crispa- for curled or twisted; designation reflects overall strand appearance, which is thread-like, stiff and wiry; strands of this species reminiscent of the stiff, wiry, curled body of an adult horsehair worm.

Multi-colored wire gorgonian.

Based on collection locations for specimens in SBMNH collection, LACoMNH collection and those examined in collection at NMNH, from at least Ventura, California south to coast of Baja, into Gulf of California. Perhaps southern end of the California Bight is the northern limit for this species.

A comment was made (Grigg 1972) about gorgonian colonies having “a loosely branched and whip-like shape when located in circular basins where water flow is turbulent.” Apparently this condition can be seen on shallow reefs to depths of 25 m (Grigg 1972). In this instance, it is suspected that regardless of water motion, the species consistently displays this distinctly thread-like form. Noteworthy point: an Atlantic form, Leptogorgia stheno Bayer, 1952, is normally unattached to any substrate; generally, many of the strands in the SBMNH material appear to exhibit that condition, as well. Having never seen this species in situ, and with no confirmed reports from other observers, thus far, nothing more can be stated about this species’ biology.

The species as described (Horvath 2011), shown as an accepted species in Cordeiro et al. (2018c), had strong similarities to Leptogorgia setacea Pallas, 1766 and shared many with L. stheno Bayer, 1952, both of which are species found in the western Atlantic. Differences were apparent in geographic location, coloring of sclerites, and subtly, in branch diameter; may be a twin species of an Atlantic form. Based on numerous strands in the NMNH material as well as material from Boca Flor De Malva at SBMNH and LACoMNH (and two other examples, both dry that were examined [see Appendix 1: List of material examined, Other material examined], one specimen from SW Punta San Juanico, outer coast, Baja, California and the other from deep water off Redondo Beach, California, US), it is possible that this species is actually very common. While perhaps not very obvious, looking nothing like the standard of a sea fan (or even a sea whip), it is suspected that this species is routinely overlooked or regarded as nothing more than a batch of dead coralline algae. Further collection is necessary, and when found, depending on environmental conditions, the question of colonies attached or not should be addressed.

Archipelago Las Perlas, Panama, 11–15 m.

Syntypes Breedy and Guzmán 2007: YPM 1553a, b [dry]; MCZ 4123 (722) [dry].

5 lots (see Appendix 1: List of material examined). Designated types not examined.

An examination of SBMNH material revealed that colony form (Figure 23A), branch and polyp appearance (Figure 23B) and sclerites (shown here, Figure 24), are comparable with images shown in Breedy and Guzmán (2007: 40–44).

Figure 23. 

Leptogorgia flexilis, SBMNH 422941. A shows colony color, and interesting disposition of branches, with tendency to droop. Colony, gently extended, ~30–40 cm tall B Close up of several branches. Note marked point of branch tip on end of lowest branch.

Figure 24. 

Leptogorgia flexilis, SBMNH 422941, SEM image. Sclerite color deep orange. A Anthocodial sclerite B Sclerites of coenenchyme (last of which may actually be an anthodocial sclerite) C Quadriradiate from coenenchyme. Images match those shown in Breedy and Guzmán 2007 (fig. 30).

The root flexi- is Latin for pliant, bendable, referring to the apparently flexible, droopy, slender branchlets of the live colony. However, Verrill does not give any rationale for the species name.

Panama, north into lower third of California Bight (off Santa Catalina Island and adjacent California mainland sites).

Initially, the drooping branches were considered to be more an artifact of preservation and the containers initially used when collected (branches bent downward so specimen would fit in the jar). However, descriptions by others (Breedy and Guzmán 2007), indicated that this is a normal branch configuration. Initial preservation in harsh chemicals caused drooping, pliant branchlets to become anything but; now quite brittle and easily broken. SBMNH 422942 is a nice, large colony but badly fragmented due to those early preservation efforts. As well, much of its color has leached out, with any particular branch colored from almost white to tan to pinkish red.

Leptogorgia flexilis is an accepted species in the WoRMS Data Base (Cordeiro et al. 2018c).

There is a need for further confirmation of species identification regarding SBMNH specimens, through examination of other definitively identified specimens, as well as the type specimens for L. tricorata (Holotype UCR 1833; Paratypes UCR 1834, 1835, 1836 and 1837). The holotype was collected in Cocos Island National Park, Isla Manuelita NW, taken on 8 September 2006 at a depth of 14 m. The paratypes were collected from Cocos Island National Park as well, either at Isla Manuelita or Roca Sucia.

~11 lots (see Appendix 1: List of material examined).

Colonies (Figure 25A) non-reticulate; main stem ~14 cm long, arising from thin, flat attachment structure; latter gives off generally dichotomous (or irregular), mostly lateral, few to moderate, elongated, sometimes slightly crooked branches; these may divide again, often not; upright growth pattern in most, overall giving colony the appearance of a candelabra. Stem and branches rounded, nearly uniform, 1.0–2.0 mm diameter, not including polyps. Branches bend outwards in broad curve at axils; terminal branches from 2.5–7.5 cm long, without division, blunt at end. Few branchlets, rounded and slightly crooked. (One lot, SBMNH 422334, a simple, single whip-like, unbranched to minimally branched fragment, where diameter tends to smallest measurements of range, length ~37 cm, but not complete; other fragments much shorter, as above). Stem, branches and branchlets covered on all sides with prominent conical polyps, when extended (Figure 25B); when contracted, nearly flush with branch surface; apertures circular. Polyps measure 0.2 mm tall (extended), 1.7 mm wide; spacing between them 2.0–2.5 mm apart. Arrangement of polyps does not delineate median groove. Color of all colonies, regardless of colony shape, bright lemon yellow or gold; most sclerites bright lemon-yellow or gold; the few straight, less warted sclerites, pale or colorless. Sclerite shapes (Figures 26A–C, 27A–D) not diverse; mostly spindles, heavily warted; warts form regular belts; belts either evenly spaced (six to seven belted rings) or belts much closer together, largest at middle of spindle and outwards toward spindle tips progressively smaller, creating in silhouette sclerites that appear in elongated diamond shape (Figures 26C, 27C); some few (Figure 27D) of these with dense triangular collection of warts at each end with very narrow, median waist; very few straight, not as heavily warted, spindles. In a comparison with images from Breedy and Cortés (2011, Figure 2), similarities between the sclerites shown in their image and the one included here in Figures 26C and 27 are strong, with exception of tentacular sclerites (rods); SBMNH specimens may be L. tricorata Breedy & Cortés, 2011.

Figure 25. 

Leptogorgia species A, SBMNH 423080. Two specimens from same lot. A shows branching, the tortuous condition seen in some branches, and bright yellow color, with prominent calyces/polyps. Larger colony measures 6.0 cm × 3.0 cm B Close up of branches, showing bright yellow coenenchyme and prominent, conical mounds with white polyps.

Figure 26. 

Leptogorgia sp. A, SBMNH 423080, Light microscopy image. All sclerites in SBMNH material are deep yellow in color. A–C Increasing magnifications of coenenchymal sclerites. In some specimens examined, the longest sclerites measured ~280–360 µm, and the smaller, more slender sclerites measured somewhere between 180–260 µm.

Figure 27. 

Leptogorgia sp. A, SBMNH 423080, SEM image. A Anthocodial sclerites B Coenencyhmal sclerites of odd shape C–D Coenenchymal spindles. Compare this SEM with SEM images shown in Breedy and Cortés (2011).

From specimens examined within the California Bight, limited range from Cortes Bank up to California Channel Islands, but see also Breedy and Cortés (2011) and “Remarks” below.

Barnacle galls present on a number of specimens (SBMNH 423084 and SBMNH 422903).

This assemblage of specimens still not identified with certainty; despite the apparent similarity with Leptogorgia tricorata Breedy & Cortés, 2011, it seemed unlikely that a species from the shallow waters of Cocos Island would be seen in the California Bight. Yet, its species name, using an adjective derived from the Latin root tricoratus-, meaning to make tricks, is applicable, as no one I spoke to who regularly collects within the Bight (LACSD, OCSD) recalled ever seeing this species. Nearly all specimens in the SBMNH collection were collected in southern California in 1940 and 1941. A few specimens more recently collected (recent being late 1970s) are also included in the collection. Since then, however, no specimens that might be this species have been encountered or reported in any collecting events to the present. All specimens examined have slightly thicker branch diameter than that seen in Thesea Duchassaing & Michelotti, 1860 (which they can resemble on a superficial level; this especially true of fragments of SBMNH 422334; one other specimen in collection, SBMNH 13304, from a station off Point Loma, is identical), and color that is generally bright lemon yellow-gold, with very markedly colored sclerites, which display a very angular, elongated diamond-shape (refer to Figures 26C, 27C here and Figure 2 in Breedy and Cortés 2011). None of the large, spheroidal bodies common to Thesea were seen in these specimens; Thesea was eliminated as a possibility. What prevents a positive identification (as L. tricorata) was a difference in aperture shape (circular vs. oblong) when polyps are contracted and complete absence of the tentacular rods, characteristic of L. tricorata. Multiple sclerite arrays were prepared, none of which displayed even a hint of the tentacular rods. While the material is older, it is in very good shape, having always been kept as wet specimens, with no evidence of formalin contact. Until further specimens can be found and collected from waters in southern California, within the Bight, and thoroughly examined, it seemed best to place these in the genus Leptogorgia without species designation (L. tricorata does seem a strong possibility; however, lack of tentacular scleritic rods is problematic).

An additional piece of information regarding L. tricorata can be found in the work of Soler-Hurtado et al. (2017). Based on molecular analysis, they have proposed that L. tricorata should now be considered a derived form in the genus Pacifigorgia Bayer, 1951 (see page 226, Soler-Hurtado et al. 2017). Cordeiro et al. (2018d) does not show this proposed emendation of the genus Pacifigorgia in the WoRMS Data Base; L. tricorata, however, is shown as an accepted species in the genus Leptogorgia. I would add that the branching morphology in the SBMNH specimens is not reflective of branching patterns usually seen species of Pacifigorgia.

Colonies of very diverse form, generally with thick branches arising laterally, dichotomously (in some, pinnately). Polyps completely retractile or forming distinct calyces into which anthocodiae can be withdrawn. Axis with wide, chambered central chord; peripheral zone of loculated horny material, usually containing nonscleritic calcareous matter (common tendency toward heavy calcification of base in old colonies). Coenenchyme thick, perforated by system of longitudinal canals surrounding axis, delimiting outer coenenchymal layer from inner one (axial sheath), which differ in spiculation. Sclerites usually include some form of club; some with spindles only, oval bodies, rods or large quadriradiates.

Due to the highly variable nature of genera and species placed in this family, this is a complex, often confusing group of organisms. Ultimately, the best means to understanding this family was to study, in total, each of the several genera placed in it that are seen in California waters. In this part (Part II), emphasis has been placed on Chromoplexaura (formerly Euplexaura) marki Williams, 2013a (part of a collective group referred to as the “red whip” species) and genera Muricea Lamouroux, 1821 and Placogorgia Studer, 1887 (Wright and Studer 1889). While the genera Swiftia Duchassaing & Michelotti, 1864 and Thesea Duchassaing & Michelotti, 1860 are also included in this family, it was necessary to cover those with a more extensive study, discussed in Part III. The genus Thesea, as represented in California waters, additionally requires still further examination; ongoing study of that genus is in progress, and will require a separate discussion, to be presented at a later date.