Monograph |
Corresponding author: Elizabeth Anne Horvath ( horvath@westmont.edu ) Academic editor: James Reimer
© 2019 Elizabeth Anne Horvath.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Horvath EA (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, and Calcaxonia – Part I: Introduction, species of Scleraxonia and Holaxonia (Family Acanthogorgiidae). ZooKeys 860: 1-66. https://doi.org/10.3897/zookeys.860.19961
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Gorgonian specimens collected from the California Bight (northeastern Pacific Ocean) and adjacent areas held in the collection of the Santa Barbara Museum of Natural History (
Allan Hancock Foundation (AHF) – ‘Velero’ Expeditions, museum collection, new species, “red whips”, soft corals, Swiftia, “thread-like” forms
The trawl seemed to have gone over a regular field of a delicate, simple Gorgonid . . . . the stems, which were from eighteen inches to two feet in length, were coiled in great hanks around the beam-trawl and engaged in masses in the net; and as they showed a most vivid phosphorescence of a pale lilac colour, their immense number suggested a wonderful slate of things beneath – animated cornfields waving gently in a slow tidal current and glowing with a soft diffused light, scintillating and sparkling on the slightest touch, and now and again breaking into long avenues of vivid light indicating the paths of fishes or other wandering denizens of their enchanted region. Wyville Thomson (during the voyage of the ‘Challenger’)
With an extensive collection of California gorgonians housed at Scripps Institute of Oceanography (Univ. of California, San Diego) to the south and a large gorgonian collection housed to the north (California Academy of Sciences, San Francisco), the Santa Barbara Museum of Natural History (
Brief taxonomic reviews of previously described species (with representation in the collection), and additional information on species recorded from the Bight, regarding distribution, biology/ecology and noteworthy observations are presented in all three parts. Also included are more extensive descriptions for genera and species not previously reported for the California Bight, as are fuller descriptions for several potentially new species. It is apparent from the
For the purposes of understanding what is meant by a “gorgonian,” as discussed in all three parts of this work, the term “gorgonian” is used in reference to members of the Subclass Octocorallia, Order Alcyonacea, specifically the scleraxonians, holaxonians, and calcaxonians. Defining features (based on
The subclass is composed of 50 families (
While seemingly fragile and delicate in overall appearance, gorgonians are remarkably “plastic” (
The primary method used for identification of a gorgonian coral species is examination of the skeletal structures embedded in the gorgonian’s soft tissue; identification in this group has always required microscopic examination of the skeletal elements, the sclerites (
Other tools that are currently being explored and utilized, with varying degrees of success, for potential identification of organisms are genetic methods and molecular analysis (
As study and reorganization of the
A further area of study that needs to be more extensively undertaken is that regarding gorgonian coral ecology; studies that necessarily focus on the ways in which these organisms deal with the physical features of their environment, as well as the biological issues of finding food, defending themselves from predators and successfully mating.
Available literature on gorgonian species of the California Bight was, on one hand, somewhat abundant, but often not very accessible, being spread over so many different publications. The earliest works on California gorgonians date to those published by
The systematics of eastern Pacific coast octocorals, particularly with regards to the alcyonacean gorgonian corals from Baja, Mexico to the California, Oregon and Washington coasts, seemed sporadic and at times, unreliable. Part of the confusion generated was likely due to the cosmopolitan distribution of a number of species which, when first collected off California, were described as new species (but had actually already been described based on collection events undertaken elsewhere). As stated previously, taxonomists were often working with a very small (and not always stellar) sample from a larger colony, or did not take into account (perhaps because it was not known at the time) the environmental plasticity of these organisms; thus they would describe a species as always branching in one plane, when current flow, we now know (
A second problem related to the lack of types for some of the taxa described from the California coast. There are excellent reviews by
Nearly all of the specimens examined (housed currently as part of the Santa Barbara Museum of Natural History’s permanent research collection, Invertebrate Laboratory), were collected over a period of years dating from the 1930s to the present in either dry or wet condition. The collection of gorgonians housed at
Any new material that may have come in was supplied through standard collection procedures that are employed by others (
Approximately two-thirds of the
All colonies were examined for gross morphology; records of height and width of a colony, length of main and secondary branches, diameter of those branches, color of the axis (if visible), and color of the colony were made. No molecular taxonomic work was undertaken with specimens received from AHF. As no formalin should come in contact with the samples, and there was no certainty that any of the material in the
For this review, examination of the calcareous sclerites, present in different parts of the colony, was conducted for nearly all specimens. Two specimens lying side by side, and appearing similar in color and overall colony form could have very different sclerites, thus establishing them as potentially different species (however, as sclerites themselves can be environmentally “plastic,” which can complicate species identification). This was a common theme regarding certain genera present in the California Bight as reflected in the
This information regarding species and lots of specimens, examined for Part I, both for colony morphology and sclerites (either through light microscopy or SEM) is a summation of the more detailed information to be found in the Appendix
# of specimens analyzed with sclerite preparations | 32 |
# of specimens examined without sclerite preparation | 1 |
Breakdown of specimens examined: | |
# of specimens analyzed from |
13 |
# of specimens analyzed from |
13 |
# of specimens analyzed from |
2 |
# of specimens analyzed from other institutions | 4 |
Total # of species that received sclerite observations | 12 |
# of new species described | 1 |
Breakdown of species examined: | |
# of species from the |
6 |
# of species from |
11 |
# of species from |
1 |
# of species from other sources | 4 |
# of species shown in Figures (colony) | 5 |
# of species shown in Figures (either light microscopy and/or SEM of sclerites) | 4 |
|
Other institutions | Colony figure | Sclerite figure | |
Anthothela pacifica | Yes | Yes | Yes | Yes |
Paragorgia arborea | Yes | Yes | Yes | No |
Paragorgia regalis | No | Yes | No | No |
Paragorgia stephencairnsi | No | Yes | No | No |
Paragorgia yutlinux | No | Yes | No | No |
Sibogagorgia californica sp. nov. | Yes | Yes | Yes | Yes |
Hemicorallium ducale | Yes | Yes | No | No |
Hemicorallium imperiale | No | Yes | No | No |
Hemicorallium regale | No | Yes | No | No |
Acanthogorgia gracillima | Yes | ? | Yes | Yes |
Acanthogorgia sp. A | Yes | ? | Yes | Yes |
Muricella complanata | No | ? | No | No |
(Classification used throughout this paper conforms to that of
(Gorgonian corals, as defined previously)
Octocorals with uniformly short gastrovascular cavities; colonies typically arborescent, rarely lobate or incrusting, producing more or less specialized three-dimensional axial skeletal structures: either a distinct central axis of horny (gorgonin) or calcareous material (or both), or a central medullar zone of calcareous sclerites which are loosely or inseparably bound together by horny or calcareous matter.
Octocorals with central axis, medullar zone or inner layer containing sclerites bound together more or less solidly either by horny or calcareous material; outer layer of coenenchyme containing proximal portions of gastrovascular chambers of polyps, endodermal canals and solenia; cortical sclerites free, commonly appearing different from those in medullar region; axial cylinder/medulla may contain canals and solenia but polyp cavities do not penetrate; cross-chambered central chord absent; sclerites always present.
1 | Axis with spongy, horny nodes, filled with sclerites alternating with longer calcareous internodes composed of cemented sclerites | (not discussed in this publication) |
– | Axis not composed of alternating horny nodes and calcareous internodes | 2 |
2 | Axis a medullar region composed of completely fused calcareous sclerites forming a central cylinder of solid calcium carbonate | Family Coralliidae |
– | Axis a medullar region composed of multiple, separate, sclerites held together by a horny material | 3 |
3 | Axis with a cross-chambered central chord | (not discussed in this publication) |
– | Axis without a chambered central chord, but often with numerous gastrodermal canals | 4 |
4 | Cortex set off from medulla by a ring of boundary canals; polyps monomorphic and protruding | Family Anthothelidae |
– | Cortex not set off from medulla by a ring of boundary canals; medulla formed only by unfused sclerites, penetrated throughout by longitudinal canals; polyps retractile, not protruding | Family Paragorgiidae |
Class Anthozoa
Subclass Octocorallia Haeckel, 1866
Order Alcyonacea Lamouroux, 1816
Suborder Scleraxonia Studer, 1887
Family Anthothelidae Broch, 1916
Anthothela pacifica (Kükenthal, 1913)
Family Paragorgiidae Kükenthal, 1916
Paragorgia arborea var. pacifica (Verrill, 1922)
Paragorgia regalis Nutting, 1912
Paragorgia stephencairnsi Sánchez, 2005
Paragorgia yutlinux Sánchez, 2005
Sibogagorgia californica sp. nov.
Family Coralliidae Lamouroux, 1812
Hemicorallium ducale (Bayer, 1955)
Hemicorallium imperiale (Bayer, 1955)
Hemicorallium regale (Bayer, 1956)
Branches of colonies slender. Polyps monomorphic, with prominent calyces, anthocodiae usually exsert. Axis not jointed, without a cross-chambered central core. Medulla surrounded by longitudinal boundary canals (of roughly equal size) separating it from cortex; medulla only rarely perforated by gastrodermal solenia in smaller branches and even then, not as extensively as in lower parts of colony; in larger branches, medulla perforated by solenia. Generally, medulla with separable sclerites; medullar sclerites stout spindles (not needle-like), thorny, ornamented with warts, spines or branching processes, that may link sclerites together. Sclerites of coenenchyme longer fusiform spindles, sometimes clavate or bent, occasionally with radiate bodies and capstans (rarely).
Within the Subclass Octocorallia, taxonomic placement of this family reflects the changeable history the Order Alcyonacea has experienced since its inception. Currently, Alcyonacea is one of three orders in the subclass (
Any membranous octocoral colony currently held in the
Briareum Sars, 1856b: 63 [pars].
? Gymnosarca Kent, 1870a: 397. Stephens, 1909: 7.
Anthothela
Verrill, 1879a: 199; 1883: 40. Studer (and Wright) 1887: 28.
? Stereogorgia Kükenthal, 1916: 178.
Suberia Nutting, 1911: 15.
Briareum grandiflorum Sars, 1856b (by subsequent designation).
With slender, rounded, tortuous, commonly upright, abundant anastomosing branches producing tangled colonies. Branches always solid; no main stems developed, branches grading upwards from broadened membranous base. Polyps present on base as well as on branches, widely scattered on all sides, sometimes clustered into large masses. Polyps elongated in expansion arising from distinctly projecting, short yet elevated cylindrical calyces. Polyps partially retractile, seldom entirely retractile; large anthocodiae commonly preserved exsert, arising from either extended, rather thin, slightly sharp, spiculose, but spongy, basal membrane (encrusting) or from slender irregular stems (branched). Axis spiculose, well differentiated, not firm. Long, strongly warted, often irregular spindles and short, girdled rods in coenenchyme. Sclerites of axis more irregular; bear fewer, larger warts, knobs or lobes. Spongy base filled with thin spindles and rods, permeating tissue. Bright rosy-red or brownish in life, but other color forms likely.
Anthothela pacifica
(Kükenthal, 1913a): 237–239; text figs E–G.
(?) Clavularia pacifica Nutting, 1909: 686.
(?) Sympodium armatum Wright & Studer, 1889: 272.
(?) Anthothela argentea Studer, 1894: 60.
USA, California, China Point, (?) San Clemente Island, SW tip, 50 fm (91 m).
Location of type unknown.
One specimen in
Colony (Figure
Presumably named for type locality, northern Pacific Ocean.
Dwarf white gorgonian.
Not definitively known at this time for coastal western United States; potentially from southern California: USA, California, San Diego, Point Loma, 200 m (
According to Kozloff (1987), a shallow subtidal form.
From the World List of Octocorallia, the World Register of Marine Species (WoRMS), Anthothela pacifica is an accepted scientific name, while Anthothela argentea has been accepted as Victorgorgia argentea, and from that listing it is clear that these two are considered separate species (
Identification of specimen relied on notes made by Dr Hochberg, with a description given by Kükenthal; fragility of specimen did not permit an extensive examination, but as far as it could be done, one was done with the specimen to hand.
Location of type is at issue; someone (unknown) noted: “China Point, ‘San Clemente Island (SW tip).’ ” There was no way to confirm this statement, and there is the added problem of a China Point on Santa Catalina Island (SW side of island), as well. There was no means to identify which “China Point” was the correct collection location.
Robust, profusely branched colonies with dimorphic polyps (feeding autozooids, reproductive siphonozooids). Axial skeletal structure solely a continuous medulla, containing separable sclerites. Medulla perforated by gastrodermal canals all the way to branch tips, not separated from cortex by a ring of boundary canals.
Paragorgia
Milne Edwards & Haime, 1857: 190.
Alcyonium arboreum Linné, 1758; [= by subsequent designation, Paragorgia arborea (Linnaeus, 1758), by monotypy].
Massive, tree-like colonies with thick branches, measuring up to 7.0 meters tall, perhaps as much as 6.0–7.0 meters wide. Sclerites in axial medulla, long, ornate rods (spindles) with branching processes, derived from capstan type, up to 0.6–0.8 mm in length, colorless or pink; elsewhere (coenenchyme, tentacles, etc.) sclerites small (less than 0.15 mm in length), differing shapes, commonly pink or red. Surface sclerites six-, seven-, and eight-radiate capstans, always less than 0.1 mm long, with globular, smooth, grooved or lobulated ornamentation. Sclerites in subsurface/outer medulla of intermediate form, ranging between radiates and spindles. Autozooid polyp tentacles have distinctively blunt, stubby rods or ovals, less than 0.1 mm.
Paragorgia pacifica (Verrill, 1922): G16–G18; plate VIII, figs 3, 4b.
Alcyonium arboreum Linnaeus, 1758: 803. Pallas, 1787: 164.
Paragorgia arborea
(Linnaeus, 1758): 803. Milne Edwards 1857: 190.
Paragorgia nodosa
Koren & Danielsson, 1883: 19 [sensu
(?) Paragorgia nodosa Nutting, 1912: 99.
(?) Paragorgia regalis Nutting, 1912: 100.
Canada, British Columbia, Vancouver Island, Jervis Inlet, ~20 m.
Holotype YPM-5373 [dry].
~1–2 lots (see Appendix
Collection lot studied contains one branch fragment (Figure
The variety name pacifica was presumably proposed in reference to the location/distribution of the species in the Pacific Ocean.
Referred to frequently as “Bubblegum coral.” Cairns et al. (2003) referred to
Recorded from Alaskan waters, Bering Sea, ‘Albatross’, 54°02'40"N, 166°42'00"W, at a depth of 504 m;
Specimens collected or photographed in the Monterey Bay Marine Sanctuary have harbored polychaete worms, purple in color (species identification not determined), with the worms wound around the branches of the colony (
Sclerite examination of the sample shown in Figure
The California Academy of Sciences has approximately 30 lots of this, or other species, attributed to this genus; most specimens are from Alaska; as well, two are from the USSR, one is from Norway, and one is from Oregon. Nine of the remaining lots are specimens collected from California; most are from Monterey Bay, with one from the Davidson Seamount. Of these nine, only three have been identified as being this particular species. MBARI has extensive video records of this species from Monterey Canyon, as well as the Davidson Seamount. In the Moss Landing Marine Labs collection there is a small specimen of what may well be this species, collected in Monterey Bay, ~36°27'12"N, 122°04'02"W, ~450 meters; coll. G McDonald, 13 March 1974; C0067 [wet]. As well, there is an impressive, tree-sized dry specimen on display in the hallway near the museum door. (Collection data may be available for this specimen, but collection data could not be located.)
Based on multiple examinations of possible Paragorgia material in the
Paragorgia regalis
Nutting, 1912: 100.
Paragorgia dendroides
Bayer, 1956b: 69 [sensu
No material in
Inclusion of P. regalis is based on
Paragorgia stephencairnsi Sánchez, 2005: 57–60; figs 39–41.
No material in
A paratype specimen at NMNH (
Paragorgia yutlinux Sánchez, 2005: 53–57; figs 36–38.
No material in
A specimen at NMNH (
Sibogagorgia
Stiasny, 1937b: 80.
Sibogagorgia weberi Stiasny, 1937b: 80 [= Suberia köllikeri Nutting, 1911: 13, nec
A genus in the family Paragorgiidae with scleritic medulla showing no (or very few, one to two) large penetration canals; main solenia around subsurface/outer medulla as boundary canals, forming reticulate network; network of canals observable with light microscopy as a regularly reticulate and uniform mesh just beneath surface. Polyps without tentacular sclerites, outer surface with radiate sclerites; generally, medullary sclerites nearly bare of ornamentation. Autozooid polyps uniformly to randomly distributed along branches, throughout colony. Branching colonies often in one plane (but not always); main branches usually thicker than terminals; terminals clavate. Coloration either of uniform beige or bright orange-red to a beige coloring with slightly projected pinkish orange polyp apertures.
Holotype USA, California, Los Angeles County, West end, Santa Catalina Island, 300 meters. Paratype USA, California, Los Angeles County, NE × E of Long Point, Santa Catalina Island, 415–486 meters.
Holotype
~8 lots (see Appendix
Specimens rarely displayed growth in one plane. Sclerites of medulla with blunt tips, bearing minimal ornamentation, smooth in areas between widely spaced spiny processes; sclerites of colony surface and coenenchymal tissue intermediate between surface and medulla 7-radiates, but never 8-radiates. Thick, compact branches with color variation from pinkish orange to pale pink.
Colony (Figure
Sibogagorgia californica sp. nov.,
Sibogagorgia californica sp. nov.,
Nearly all specimens examined, with the exception of two, are from locations within the vicinity of the California Channel Islands, thus a reference to the state of California, where most of the specimens were collected.
Proposed, “California orange bubblegum” coral.
Based on the specimens in the
Preliminary examinations led to identification of this small group of specimens as Sibogagorgia cauliflora
From a taxonomic perspective, in their most recent molecular work Figueroa and Baco (2014) recommended that family Sibogagorgiidae be reinstated. According to WoRMS, the genus Sibogagorgia is still retained in the family Paragorgiidae (
The museum collection at Moss Landing Marine Labs held a specimen that might be this species, collected in Monterey Bay, 36°26'42"N, 122°01'56"W, ~684 m; coll. G McDonald, 14 August 1974; C0071 [wet]. Coloring of this specimen showed either a slightly bleached condition (storage artifact) or the beige coloring with orange pimples, a slightly brighter condition than that seen in
The
Axis an unjointed medullar region composed of completely fused calcareous sclerodermites (solid calcium carbonate) derived from sclerites;
Madrepora (pars) Linnaeus, 1758: 797.
Corallium [ächte rothe Steincoralle] Müller in Knorr, 1766: Delic. Nat. 1: 7, pl A I, figs 1, 2; 23, pl A VII, fig 1; 24, pl A VIII, figs 2–4; 127 (pars); nec. pp 9–13, 25, 128.
Isis (pars) Linnaeus, 1767: 1288.
Nec Isis Linnaeus, 1758: 799.
Nec Corallium Burman, 1769: [3] (= Isis Linnaeus, 1758).
Corallium
Cuvier, 1798[1797]: 673.
Pleurocorallium
Gray, 1867: 126.
Hemicorallium
Gray, 1867: 126.
Madrepora rubra Linnaeus, 1758 (by subsequent monotypy, the first species being assigned by
Sclerites of cortex numerous, regular capstans, often modified with six, seven or eight radii; or as double clubs (only some species), crosses and opera glasses; long spindles present in tentacles. Without axial pits bearing beaded rims beneath autozooids. Autozooids prominent, non-retractile (when contracted cannot fully retract into cortex) and ovate-cylindrical, usually distributed on one side of colony.
. An interesting genus; its connection to human enterprise and profit make it so. Collectively, “pink coral, red coral, noble coral, angel skin coral, Sardinia coral, midway coral” (CITES proposal, Convention of the Parties, CoP14 Prop. 21, June 2007) has had a long history, with the primary focus on harvesting of the coral for profit. At least one proposal, and multiple CITES conferences over the years (2007, 2010, 2013), have made this genus a focus of discussion. Numerous articles have been published concerning the impact of harvesting, management issues, etc. (
Because all species of coral in this genus tend to form tall, tree-like colonies, they likely increase three-dimensional complexity of the habitats they are found in and consequently, increase biodiversity where they occur. These colonies could easily provide valuable microhabitat for sessile, associated commensal invertebrates (
A number of species in the genus Corallium have recently been transferred to the genus Hemicorallium (encompassing species discussed here;
Corallium ducale
Bayer, 1955: 210–211, plate 1.
One lot, 2 specimens + fragments (recent addition) in
Distribution extends in Eastern Pacific from Mexico (Bayer, 1955), northern Baja Mexico (
The specimen in question does not easily identify to a species; in color it appeared more like that of Hemicorallium imperiale, but the polyps’ appearance and scleritic spindles were more like those seen in H. ducale. While all type material for this species (and the other two that follow) at NMNH were examined some years ago, this one specimen, recently received into the
WoRMS Data Base (
Corallium imperiale
Bayer, 1955: 209–210, plate 2, c-h.
No material in
Distribution in the Eastern Pacific, northern Baja, California (
Corallium regale
Bayer, 1956b: 70, 73–76; 77–78; figs 5c; 7e–g.
(see Remarks section below.)
No material in
Around the Pacific, from Hawaii (holotype,
Several species in the now updated genus Hemicorallium, including this one, found within the Hawaiian archipelago (and elsewhere in the western Pacific, including international waters) have been the focus of commercial exploitation (then recognized as species in the genus Corallium). Based on reports made public by CITES, regarding “consideration of proposals for amendment of Appendices I and II” (2007, 2010) and
In studying Bayer’s original 1956 description of C. laauense (correctly spelled) and C. regale, no determination could be made as to why these two were linked as synonymous. The now recognized species Hemicorallium regale has many double club sclerites, while H. laauense has none. As well, the entire colony of H. regale, as well as the axis, is pink, while the colony color of H. laauense is white or very faintly pink with a white axis. SEM images are on file (Bayer’s personal collection), SEM #2283,
The work by
With distinct central axis composed of horny material alone or of horny material more or less heavily permeated with calcareous substance, continuous or with alternating horny and calcareous joints. In center of axis is a relatively narrow, largely hollow, tubular space partitioned into series of small chambers, referred to as the cross-chambered central chord. Calcareous material of the peripheral zone of axis is in nonscleritic form (single exception in Keroeididae).
1 | Axis horny, with a chambered, hollow, soft central chord | 2 |
– | Axis not horny, but is a solid axis, with no soft, central, hollow core | Suborder Calcaxonia, Part III |
2 | Axis purely horny, composed of scleroprotein, without any calcareous deposits | 3 |
– | Axis horny, but some calcareous material may be present in some forms; hollow, horny, soft-chambered central chord is wide; there is a peripheral zone of hollow horny spaces containing calcareous material; cortex is thick, with an inner and outer layer, formed by systematic longitudinal canals; polyps retractile into prominent calyces | Family Plexauridae, Parts II & III |
3 | Axis perforated by a wide, cross-chambered central chord; cortex thin; polyps not retractile; sclerites spikey and conspicuous | Family Acanthogorgiidae, this part |
– | Distinct hollow, horny, soft-chambered central chord that perforates axis is narrow; axial cortex surrounding the core is very dense; polyps fully retractile, into calyces | Family Gorgoniidae, Part II |
Class Anthozoa
Subclass Octocorallia Haeckel, 1866
Order Alcyonacea Lamouroux, 1816
Suborder Holaxonia Studer, 1887
Family Acanthogorgiidae Gray, 1859
Acanthogorgia gracillima var. typica Kükenthal, 1909; (? A. gracillima var. lata Kükenthal, 1909)
Acanthogorgia species A
Muricella cf. complanata Wright & Studer, 1889
Axis purely horny (scleroprotein without calcareous deposits), dark-colored, predominantly black; very difficult to cut, with wide, hollow, cross-chambered central core. Coenenchyme very thin, polyps conspicuous, contractile, not retractile, completely covered with both straight and curved fusiform sclerites (forming, in appearance only, cylindrical calyces; no calyces actually present). Sclerites arranged in eight double rows, forming eight en chevron fields; no well-defined operculum; sclerites instead arranged as transverse ring and eight points of converging spindles on tentacle bases; thus, sclerites of polyps continuous with those of tentacular crown, latter being sharp spines arrayed conspicuously around top of polyp, with no intervening sclerite-free neck zone or transverse collaret. Consequently, no clear division between anthocodia and anthostele. Tentacles fold inward over oral disk. Predominant sclerites colorless, in form of prickly or warty spindles; sometimes, presence of three- and four-armed radiates.
Acanthogorgia
Gray, 1857a: 128, pl 3, fig 2 [1851]. (pars)
Blepharogorgia (pars) Duchassaing & Michelotti, 1864: 15.
Paracanthogorgia
Stiasny, 1943b: 130; 1947: 11, 53.
Acanthogorgia hirsuta Gray, by monotypy (? = A. aspera Pourtalès, 1867).
Colonies generally flattened (flabellate); commonly reticulate, or developed into dense bushy shrubs. Branches appear thin and delicate. Polyps tall, cylindrical, topped with thorny crown of strongly projecting spinous sclerites, embedded at tentacle bases. They lie, collectively, over infolded tentacles, protruding end of sclerites smooth. Polyps on all sides of branches, or roughly biserial; arise vertically at right angle to branch surface, acalycinous, not retractile. Coenenchyme between branches usually thin, axis visible through it. Sclerites in polyps slender spindles slightly bent, arranged en chevron in eight longitudinal double rows. Back of tentacles with only numerous small, flat, bent sclerites; stem coenenchyme with slender, generally bent or sinuous spindles sculptured by prickles or simple tubercles; in deeper layers of coenenchyme (some species), with radiates (tri-radiates and crosses, often with a projecting central spine). Axis dark, but coenenchyme usually colored; sclerites always colorless.
Only one collected specimen from the genus appeared in the ‘Velero’ material; it does not match any of the described species it has been compared it to, thus far. It was collected in Mexican waters, beyond the geographic range covered in this work; description of this specimen is in progress.
Acanthogorgia gracillima var. typica
Kükenthal, 1909: 73–76; pl 6, fig 33; (syn.) 1919: 763.
? A. gracillima var. lata Kükenthal, 1909: 71, 73, 75.
(?) Okinawa, Japan; 400 fathoms.
Zoological Museum of the University of Hamburg, Germany (formerly, Naturhistorisches Museum, Hamburg); Catalog Number 3298, with Catalog Number 3297. (Not labeled as type.)
1 lot (see Appendix
Colony (Figures
For designation gracillima, the Latin gracili- means slender; this may reference the conspicuous, very slender polyps and very slender points in the crown of this species.
None specifically designated; the Slender glass-shard gorgonian would be appropriate. Worth considering is whether or not this is the species that, predominantly, MBARI and
For genus in general, “species. . . inhabit moderate to considerable depths; various species of Acanthogorgia occur in all seas, some thriving in very cold waters” (Bayer, penciled personal annotations in
On this particular specimen, there were at least two scale worms wrapped around the base of polyps in two separate areas within the colony.
To handle the specimen during examination, although colony fairly hardy, gloves were necessary. Generally, all the long sclerites were very sharp, comparable to small shards of thin glass; those that got into the skin under the fingernail were rather painful. Given the delicate, and somewhat brittle/fragile nature of the longer boomerang-type sclerites, these sclerites are easily broken. It was often difficult to ascertain whether radiates and crosses really existed as such, or whether they were bits that had broken off from the longer spindle forms; this could account for some of the odder-shaped sclerites that previous description (
This description (and original description for the species) was based on the specimen used by
While the original specimen does indeed still exist, no opportunity to travel to Hamburg, Germany occurred and no request that the specimen be sent was made. The species is an accepted species in the WoRMS Data Base (
[Acanthogorgia (= ? Acalycigorgia) Grey, 1857)]; not accepted, WoRMS (
Acalycigorgia
Kükenthal, 1908b: 38; 1919: 298; 1924: 237–239.
? = Acanthogorgia Gray, 1857a: 128, pl 3, fig 2 [1851]. (pars)
? Paramuricea Moroff, 1902: 407.
A. grandiflora Kükenthal & Gorzawsky, 1908a; subsequent designation by
Polyps not functionally differentiated into anthocodia and anthostele; contractile but not retractile within common coenenchyme; tentacles fold over oral disk in contraction. Polyps similar to those of Acanthogorgia (without crown of strongly projecting spines, however), but polyps can be short and verruciform to prominent, tall and cylindrical, not clavate. Sclerites of polyp walls large spindles, very conspicuous; commonly arranged more or less distinctly en chevron in eight long, longitudinal double rows, but distal ones project little or not at all. Distal ends of sclerites around tentacle bases not specifically differentiated as spines, though the tips may project somewhat around polyps’ apex. Polyps are without suture separating tentacular/anthocodial from subtentacular sclerites. Sclerites of polyp body gradually merge with those of tentacle bases, which are not abruptly smaller; coenenchymal sclerites with tubercles of inner and outer sides similarly developed; inner layer of coenenchyme with more or less abundant radiates.
Cannot be indicated at this time.
Cannot be indicated at this time.
1 lot (see Appendix
Colonies (two) generally in one plane; one measures 7.0 cm × 5.5 cm (length to width); second (Figure
It is possible that these specimens, and several others in the collections either at
Lissogorgia (pars) Verrill, 1865: 187.
Muricea (pars) Verrill, 1868c: 411–416.
Muricella
Verrill, 1869a: 450 (= ? Anthogorgia Fabricius & Alderslade, 2001).
? Anthogorgia Verrill, 1868 (=?Acalycigorgia Kükenthal, 1908b).
? Acalycigorgia Kükenthal, 1908b (= ? Acanthogorgia or Astrogorgia [see
? Astrogorgia Verrill, 1868c: 413.
? Acanthogorgia Gray, 1857a: 128 [1851].
Lissogorgia flexuosa Verrill, 1865; [subsequent diagnosis by
Colonies fan-shaped, branching in one plane, some anastomosing. Larger branches with axis often tending to be flattened at right angles to plane of fan; in older colonies, smaller branches can bend and grow perpendicular from the fan. Polyps prominent, low, wart-like, non-retractile; coenenchyme thick between polyps. Sclerites of polyp tentacles small rods; below anthocodiae, sclerites large and bow-shaped, in angled double rows (chevrons), forming eight marginal points, forming strong collaret; tentacular operculum distinct. Sclerites of coenenchyme in two distinct layers, mostly spindles (coarse or densely warted), small capstans, with some clubs or discoidal forms.
Muricella complanata
Wright & Studer, 1889: 125–126.
No material identified as this species in collection at
Included here briefly, as there was a California collection location for a specimen that was identified as this species.
Based on
It is clear from this first part of the systematic review of alcyonacean gorgonian species in the
List of material examined – Part I
(Material examined = whole colony study plus multiple sclerite preparations; all with light microscopy, plus selected colonies under SEM, shown in figures associated with text)
Anthothela argentea Studer, 1894
Material examined. No specimens in
Other material, not examined. – ±1 colony; USA, off California coast, west of Channel Islands, Fieberling Guyot, 32°27'36"N, 127°47'36"W, 490 m; coll. R/V ‘Alvin’, 10 October 1990;
Anthothela pacifica (Kükenthal, 1913)
Material examined. 1 lot USA, California – 1 colony; San Diego County, edge of ridge running through Rodriquez Seamount, slightly northwest of West Cortes Basin, due west of southern tip, San Clemente Island, on sponge, 32°37'43"N, 120°05'04"W, 950–1150 m; coll. skipper of vessel ‘Calafia’ (California Department of Fish and Game – Long Beach, California), 9 August 1978;
Other material examined. – ±1 colony; USA, California, San Diego County, San Diego, Point Loma, ~32°40'38"N, 117°15'08"W, 201–262 m; coll. R/V ‘Albatross’, 4 March 1904;
Paragorgia arborea var. pacifica (Verrill, 1922)
Material examined. ~2 lots USA, California – 1 branch fragment; Orange County, San Clemente Island, between Pete Bay, 3–10 miles E X (N+S) of island, ~32°55'48"N, 118°24'19"W, 1000 m; coll. unknown, 19 July 1978;
Other material examined. – 1 colony/fragment; USA, California, Ventura County, off San Nicolas Island, (from fishermen’s net); coll. unknown, March 1985; donated by R Williams, of “The Sea;” CBA #85.14.2 [dry]. -- USA, California, Monterey County, Davidson Seamount, 1,313 m; coll. unknown, 2003;
Paragorgia regalis Nutting, 1912
Material examined. No apparent material in
Other material examined. – 1 colony fragment; USA, California, Santa Barbara County, west of San Miguel Passage on Rodriquez Seamount, 33°57'12"N, 121°08'41"W, 1,840 m; coll. unknown, 14 October 2003;
Paragorgia stephencairnsi Sánchez, 2005
Material examined. No material in
Other material examined. – 1 colony; USA, California, west of California Channel Islands, Fieberling Guyot, 36°26'00"N, 127°47'36"W, 490 m; coll. DS/V ‘Alvin’, 16 October 1990;
Paragorgia yutlinux Sánchez, 2005
Material examined. No material in
Other material examined. – 1 colony; USA, California, west of the California Channel Islands, Fieberling Guyot, from summit of seamount, 32°26'00"N, 127°47'42"W, 487 m; coll. DS/V ‘Alvin’, 21 October 1990;
Sibogagorgia californica sp. nov.
Material examined. ~6 lots USA, California – 1 branch fragment; Los Angeles County, Santa Catalina Island, 7 miles WSW of Church Rock, substrate loose rock, 33°14'40"N, 118°11'10"W, 276–282 m; coll. R/V ‘Velero III’, station 1323-41, 18 May 1941;
Hemicorallium ducale (Bayer, 1955)
Material examined. One lot (possibly this species). USA, California – 2 colonies + fragments; USA, California, Los Angeles County, Catalina Basin, N of San Clemente Island, 33°04'49"N, 118°29'52"W, 1,091 m; coll. P Gregory, California Fish and Game, Long Beach, 15 November 1977;
Other material examined. – 1 colony; USA, California, West of Channel Islands, Fieberling Guyot, “Sea Pen Rim,” 32°27'36"N, 127°49'30"W, 640 m; coll. DS/V ‘Alvin’, 14 October 1990;
Hemicorallium imperiale (Bayer, 1955)
Material examined. No material in
Other material examined. – multiple fragments; USA, California, Fieberling Guyot Seamount, 32°23'00"N, 127°47'00"W, ~600 m; coll. unknown, August 1989;
Hemicorallium regale (Bayer, 1956)
Material examined. No material in
Other material examined. –1 colony; USA, California, Davidson Seamount; 1,482 m; coll. unknown, 2003;
Acanthogorgia gracillima var. typica Kükenthal, 1909
(? A. gracillima var. lata Kükenthal, 1909)
Material examined. 1 lot USA, California – 1 colony; California, Ventura County, Channel Islands, 33°57'35"N, 119°28'34"W, 160 m; coll. M Love from submersible ‘Delta’, 8 October, 2005; with his number: A6662;
Other material examined. – 1 fragment (multiple colonies); USA, northern Pacific Ocean, Hawaii, Cross Seamount, 18°43'55"N, 158°15'41"W, 388.56 m; coll. Smith, 10 October 2004;
Acanthogorgia species A (= ? Acalycigorgia)
Material examined. 1 lot USA, California – 2 colonies; Monterey County, Monterey Bay, from Point Piños Light to Mid Point, ~36°41'36"N, 122°01'51"W, 439–814 m; coll. R/V ‘Velero IV’, stations 7462-61 or 7463-61, 10 October 1961;
Other material examined. – 1 colony; USA, California, off San Francisco, Mulberry Seamount, ~37°26'48"N, 123°23'22"W, 1,255–1,455 m.; coll. N/B ‘Scofield’, location 38, 13 February 1950;
Muricella complanata Wright & Studer, 1889
Material examined. No confirmed identification of material in
Other material examined (Identification not confirmed). – 1 colony; USA, California, Santa Barbara County, Santa Barbara Channel, ~12 south of Santa Barbara, midway between Santa Barbara city and Santa Cruz island, oil platform (settling plates), ~34°13'39"N, 119°41'12"W, ~182 m; coll. MMS III Voucher, Survey, settling plates (incidental), station: exploratory wellsite Lease Number 0522, 6 February 1994; #0001942 [wet]. --2 colonies (2 lots); USA, California, Monterey County, Monterey Bay, Point Piños Light House, bearing S 3 degrees: distant 9.3 miles, ~36°46'14"N, 121°55'23"W, 518–649 m; coll. R/V ‘Albatross’, California Coastal Cruise 1904, Station #4461, 12 May 1904;