Leptogorgia chilensis (Verrill, 1868)

Horvath, Elizabeth Anne, 2019, A review of gorgonian coral species (Cnidaria, Octocorallia, Alcyonacea) held in the Santa Barbara Museum of Natural History research collection: focus on species from Scleraxonia, Holaxonia, Calcaxonia - Part II: Species of Holaxonia, families Gorgoniidae and Plexauridae, ZooKeys 860, pp. 67-182 : 67

Leptogorgia chilensis (Verrill, 1868) Figures 14A, B, 15 A–D

Plexaura rosea Philippi, 1866: 118 (junior homonym, Breedy and Guzmán 2007).

Leptogorgia rosea Phillipi, 1892: 7 (as: Verrill 1868b: 406 (nec Leptogorgia rosea Milne Edwards & Haime, 1857: 134).

(?) Litigorgia (?) rosea : Verrill 1868a; 1868b: 406. Philippi 1892: 7.

Nec Litigorgia flexilis Verrill, 1868a.

Leptogorgia (?) chilensis Verrill, 1868b: 406.

Leptogorgia chilensis Kükenthal, 1919: 772; 1924: 355. Bielschowsky 1929: 132. Breedy and Guzmán 2007: 22-25.

Type locality.

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.

Type specimens.

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

Material examined.

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

Description.

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 Limbaugh (unpublished key) described color as a rich salmon pink; polyps white; dry specimens pale orange to light salmon pink.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 15 A–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).

Etymology.

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.

Common names.

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.).

Distribution.

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 .)

Biology.

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).

Remarks.

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.