![Adelogorgia phyllosclera. ASBMNH 422403 [dry], slightly magnified image of branches BSBMNH 51252 [wet], rounded, slightly club-shaped branch tips.](https://www.researchgate.net/publication/334848645/figure/fig2/AS:11431281210378332@1702041319522/Adelogorgia-phyllosclera-ASBMNH-422403-dry-slightly-magnified-image-of-branches_Q320.jpg)
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A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 67
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
Elizabeth Anne Horvath1,2
1 Westmont College, 955 La Paz Road, Santa Barbara, California 93108, USA 2 Invertebrate Laboratory,
Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road, Santa Barbara, California 93105, USA
Corresponding author: Elizabeth Anne Horvath (horvath@westmont.edu)
Academic editor: James Reimer|Received 1 February 2019|Accepted 27 March 2019|Published 4 June 2019
http://zoobank.org/128BC183-0A6A-4234-8893-1CBD2D2AF962
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,
Calcaxonia – Part II: Species of Holaxonia, families Gorgoniidae and Plexauridae. ZooKeys 860: 67–182. https://doi.
org/10.3897/zookeys.860.33597
Abstract
Gorgonian coral specimens from the Holaxonia, families Gorgoniidae and Plexauridae held in the col-
lection of the Santa Barbara Museum of Natural History (SBMNH) were reviewed and evaluated for
species identication. e specimens were collected from within, and adjacent areas of, the California
Bight. e SBMNH collection has encompassed within it a large percentage of specimens collected by
the Allan Hancock Foundation (AHF) ‘Velero’ Expeditions of 1931–1941 and 1948–1985. is historic
collection displays an emphasis on species belonging to the Holaxonia, particularly the gorgoniids and
plexaurids; thus, this second part presents a thorough discussion of well-known genera from within the
California Bight, with more extensive discussions of several genera that have historically, and currently, led
to confusion (and thus, misidentication). A brief discussion of a California Bight grouping, referred to
within as the “red whips,” is presented; this grouping encompasses several species with very similar colony
appearance across a number of genera. Two species, the gorgoniid Leptogorgia chilensis (Verrill, 1868) and
the plexaurid Chromoplexaura marki (Kükenthal, 1913) each required the designation of a neotype from
within the collection. A new species in the genus Eugorgia Verrill, 1868, a whip or thread-like form be-
ZooKeys 860: 67–182 (2019)
doi: 10.3897/zookeys.860.33597
http://zookeys.pensoft.net
Copyright 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.
RESEARCH ARTICLE
Launched to accelerate biodiversity research
A peer-reviewed open-access journal
Elizabeth Anne Horvath / ZooKeys 860: 67–182 (2019)
68
longing to the family Gorgoniidae, is described. One additional plexaurid genus (Placogorgia) is discussed,
a genus not commonly reported for the California Bight region. is is the rst comprehensive work, in
three parts, focusing on all species of gorgonian coral known to inhabit the California Bight. is paper,
Part II of the full work, continues the systematic review of all species represented in the Santa Barbara
Museum of Natural History research collection begun in Part I.
Keywords
Allan Hancock Foundation (AHF) – ‘Velero’ Expeditions, cryptic species, local endemics, museum collec-
tion, new Eugorgia species, Placogorgia, “red whips”, soft corals, “thread-like” forms
Table of contents
Class Anthozoa, Subclass Octocorallia, Order Alcyonacea (Gorgonian corals) Part II
Introduction ........................................................................................................... 69
Materials and methods ........................................................................................... 69
Systematic accounts ................................................................................................ 71
Diagnosis ......................................................................................................... 71
Holaxonia Studer, 1887 ................................................................................... 71
Key to Families represented in SBMNH collection (Holaxonia) ....................... 72
List of species ................................................................................................... 72
Descriptions of species ..................................................................................... 73
Family Gorgoniidae Lamouroux, 1812 ...................................................... 73
Adelogorgia phyllosclera Bayer, 1958 ............................................................ 73
Eugorgia daniana Verrill, 1868 ................................................................... 79
Eugorgia rubens Verrill, 1868 ...................................................................... 83
Eugorgia ljubenkovia sp. nov. ...................................................................... 87
Leptogorgia chilensis (Verrill, 1868) ............................................................. 92
Leptogorgia diusa (Verrill, 1868) ............................................................... 98
Leptogorgia licrispa Horvath, 2011 ......................................................... 101
Leptogorgia exilis (Verrill, 1868) .............................................................. 106
Leptogorgia species A ................................................................................ 106
Family Plexauridae Gray, 1859 ................................................................. 112
Chromoplexaura marki (Kükenthal, 1913) ................................................ 113
Discussion concerning diversity of “red whip” forms ............................................ 125
Muricea californica Aurivillius, 1931 ............................................................... 129
Muricea plantaginea (Valenciennes, 1846) ...................................................... 136
Muricea fruticosa Verrill, 1868 ........................................................................ 138
Placogorgia species A ....................................................................................... 148
Conclusions ......................................................................................................... 153
References ............................................................................................................ 153
Appendix 1 ........................................................................................................... 164
Appendix 2 ........................................................................................................... 181
Appendix 3 ........................................................................................................... 182
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 69
Introduction
As dened in Part I of this work, the term gorgonian used in this paper refers to those
alcyonacean octocorals belonging to the groups Scleraxonia, Holaxonia and Calcaxo-
nia. ese organisms are modular colonies, usually extensively branched, displaying a
sti central, internal axis (composed of calcite and gorgonin), in both main stem and
all branches, composed of either fused sclerites, or sclerites composed of scleropro-
teinous gorgonin. e entire axial skeleton is covered with soft tissue coenenchyme
lled with numerous calcareous sclerites, either embedded in it or lying on its surface.
e supporting axial skeleton allows for colonies to achieve large size (some species)
and allows for the display of both highly branched colonies, known as sea fans, as well
as long, slender forms known as sea whips.
e gorgonian Holaxonia are the most numerous of the gorgonian corals found in
the Santa Barbara Museum of Natural History’s (SBMNH) research collection. While
a fair number of specimens, representing the families Gorgoniidae and Plexauridae
were already present, with the acquisition of gorgonian materials from the Allan Han-
cock Foundation ‘Velero’ Expeditions, the holdings within the collection were greatly
enhanced. Many of the specimens not only needed rebottling, but extensive work had
to be done to identify (or to correct identication of), not only the specimens that
were already in the collection, but the many specimens collected during the ‘Velero’
years of operation. As many of the genera in these two families have been thoroughly
reviewed elsewhere (Breedy and Guzmán 2007, 2016, 2018, Breedy et al. 2009), I am
providing only brief descriptions for most. More problematic genera, or genera and
species that have not been well studied, are given more extensive coverage and discus-
sion. Not all of the Holaxonia holdings in the SBMNH collection are covered here.
Part III of this review will cover two of the genera that were most in need of work and
required more thorough discussion.
Materials and methods
Nearly all of the specimens examined in this work (housed currently as part of the
Santa Barbara Museum of Natural History’s permanent research collection, Inverte-
brate Laboratory), were collected over a period of years dating from the 1930s to the
present, in either dry or wet condition. A large percentage of these specimens came to
the SBMNH through a diverse 10,000-lot cnidarian collection, a portion of the Allan
Hancock Foundation (AHF) collection built upon the historic ‘Velero’ expeditions of
1931–1941 and 1948–1985. Not only are gorgonian specimens housed in the cnidar-
ian section of the entire invertebrate collection, but there are gorgonians housed else-
where within the collection; for instance, gorgonian coral fragments are housed in the
museum’s mollusk collection (the mollusks in question were found on, and collected
with, a species of gorgonian), or in other sections of the museum’s cnidarian collection
(such as zoanthid anemones collected on gorgonian corals). Scattered throughout other
Elizabeth Anne Horvath / ZooKeys 860: 67–182 (2019)
70
portions of the museum’s invertebrate collection are bryozoans, barnacles, or brittle
stars that were collected from within or on gorgonian coral colonies and were preserved
with the gorgonian they were living with. To assist with the identication of the SBM-
NH specimens, examinations of specimens of known species from or collected in the
Bight were performed on material found in the collections of the National Museum
of Natural History, Smithsonian (USNM = NMNH), the California Academy of Sci-
ences, San Francisco (CAS), the Los Angeles County Museum of Natural History (LA-
CoMNH), Scripps Institute of Oceanography (SIO), the Monterey Bay Aquarium Re-
search Institute (MBARI), Moss Landing Marine Laboratories (MLML) and the small
museum which is a part of the Cabrillo Marine Aquarium in San Pedro, California
(CMA) (see Appendix 1: List of material examined). ese were compared to SBMNH
specimens, informing the identication of species represented in the SBMNH collec-
tion. Additionally, several National Oceanographic and Atmospheric Administration
(NOAA) oces throughout the country provided further material for study.
All specimens were examined for gross colony morphology; more importantly,
examination of the calcareous sclerites, present in dierent parts of the colony, was
conducted for nearly all specimens. e standard method for sclerite extraction (tis-
sue sample in common household bleach) was performed, and light microscopy via
a compound Olympus (CH) microscope, was used initially to determine the genus
to which a specimen belonged. Scanning Electron Microscopy (SEM) of the sclerites
was then undertaken. All samples were coated with gold, using a Cressington Sput-
ter Coater Unit, 108auto. Samples were examined and digital images taken, using a
Zeiss Scanning Electron Microscope EVO 40, at 10 kV. is second part covers some
fourteen species, classied as holaxonians belonging to the families Gorgoniidae and
Plexauridae. A summative overview of species housed in the SBMNH research collec-
tion, from these specic groups, is included below.
is information regarding species and lots of specimens examined for Part II for
colony morphology and sclerites (either through light microscopy or SEM) is a sum-
Part II: Collective specimen and species data.
# of specimens analyzed with sclerite preparations ~260
# of specimens examined without sclerite preparation 0
Breakdown of specimens examined:
# of specimens analyzed from SBMNH collection ~184
# of specimens analyzed from USNM-Smithsonian 19
# of specimens analyzed from CAS 13
# of specimens analyzed from other institutions 54
Total # of species that received sclerite observations 14
# of new species described 1
Breakdown of species examined:
# of species from the SBMNH collection 14
# of species from USNM-Smithsonian 7
# of species from CAS 3
# of species from other sources 10
# of species shown in Figures (colony) 13
# of species shown in Figures (either light microscopy and/or SEM of sclerites) 14
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 71
mation of the more detailed information to be found in the Appendix 1: List of mate-
rial examined – Part II. It is evident from this summative overview that the SBMNH
research collection illustrates diversity and abundance of species from the holoaxonian
group found within or near the California Bight.
Systematic accounts
(Classication used throughout this paper conforms to that of Bayer 1981c)
Diagnosis of the Order
Alcyonacea Lamouroux, 1816
(Gorgonian corals, as dened 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 ma-
terial (or both), or a central medullar zone of calcareous sclerites which are loosely or
inseparably bound together by horny or calcareous matter.
Holaxonia Studer, 1887
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-cham-
Species covered in this part.
SBMNH Other institutions Colony gure Sclerite gure
Adelogorgia phyllosclera Yes Yes Yes Yes
Eugorgia daniana Yes Yes Yes Yes
Eugorgia rubens Yes Yes Yes Yes
Eugorgia ljubenkovia sp. nov. Yes Yes Yes Yes
Leptogorgia chilensis Yes Yes Yes Yes
Leptogorgia diusa Yes Yes Yes Yes
Leptogorgia licrispa Yes Yes Yes Yes
Leptogorgia exilis Yes Yes Yes Yes
Leptogorgia sp. A Yes Yes Yes Yes
Chromoplexaura marki Yes Yes Yes Yes
Muricea californica Yes Yes Yes Yes
Muricea plantaginea No Yes No Yes
Muricea fruticosa Yes Yes Yes Yes
Placogorgia sp. A No ? Yes Yes
Elizabeth Anne Horvath / ZooKeys 860: 67–182 (2019)
72
bered central chord. Calcareous material of the peripheral zone of axis is in nonscleritic
form (single exception in Keroeididae).
Key to Families represented in SBMNH collection (Holaxonia)
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 .............
.............................................................................. See 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; hol-
low, 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
3 Axis perforated by a wide, cross-chambered central chord; cortex thin; polyps
not retractile; sclerites spikey and conspicuous ..............................................
............................................... Family Acanthogorgiidae (covered in Part I)
– Distinct hollow, horny, soft-chambered central chord that perforates axis is
narrow; axial cortex surrounding the core is very dense; polyps fully retractile,
into low calyces ............................................................ Family Gorgoniidae
List of species
Class Anthozoa
Subclass Octocorallia Haeckel, 1866
Order Alcyonacea Lamouroux, 1816
Holaxonia Studer, 1887
Family Gorgoniidae Lamouroux, 1812
Adelogorgia phyllosclera Bayer, 1958
Eugorgia daniana Verrill, 1868
Eugorgia rubens Verrill, 1868
Eugorgia ljubenkovia sp. nov.
Leptogorgia chilensis (Verrill, 1868)
Leptogorgia diusa (Verrill, 1868)
Leptogorgia licrispa Horvath, 2011
Leptogorgia exilis (Verrill, 1868)
Leptogorgia species A [? = Leptogorgia tricorata Breedy & Cortés, 2011]
Family Plexauridae Gray, 1859 [= Muricidae]
Chromoplexaura marki (Kükenthal, 1913)
Discussion concerning diversity of “red whip” forms
Muricea californica Aurivillius, 1931
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 73
Muricea plantaginea (Valenciennes, 1846) = M. appressa Verrill, 1864
Muricea fruticosa Verrill, 1868
(following genus formerly part of: [Stenogorgiinae = old Paramuriceidae]
Placogorgia species A
Descriptions of species
Family Gorgoniidae Lamouroux, 1812
Diagnosis. Axis purely horny, composed of carbonate hydroxylapatite with narrow
but distinct chambered central chord; cortex little loculated, if at all. Polyps fully re-
tractile, 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 modied into disc spindles, scaphoids, or
unilaterally spinous forms. Anthocodial armature weak, in form of crown composed
of at rodlets with scalloped edges, or lacking entirely. Colonies of diverse form, from
unbranched to pinnate, closely reticulate or foliate.
Genus Adelogorgia Bayer, 1958
Adelogorgia Bayer, 1958: 46; 1979: 1026–1027. Breedy and Guzmán 2018: 329.
Type species. Adelogorgia phyllosclera Bayer, 1958.
Diagnosis. Genus originally included in family Plexauridae (Bayer 1958). Pres-
ence 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 at rods, 0.15–0.3 mm long.
Etymology. 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.
Adelogorgia phyllosclera Bayer, 1958
Figures 1, 2A, B, 3A–C
Adelogorgia phyllosclera Bayer, 1958: 46–47; gs 3a–f, 4a, b, 9b, c. Breedy and Guzmán
2018: 330–333.
Type locality. USA, California, La Jolla, South of Scripps Institution, La Jolla
Canyon, 30–33 m.
Elizabeth Anne Horvath / ZooKeys 860: 67–182 (2019)
74
Type specimens. Holotype USNM 50186; [dry]. Paratypes listed under “other
material” in “Appendix 1: List of material examined” for this species.
Material examined. ~26 lots (see Appendix 1: List of material examined). All
specimens at USNM were examined for comparison purposes.
Description. Colony (Figure 1) heavy; bushy to fan-shaped, branching strictly in
single plane, particularly in young colonies (≤ 20 cm height), but with branches occa-
sionally 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 (Figure2B). 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, exible 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 o 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 modied 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 or-
dinary 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 cap-
stans, clubs or leafy scales. Sclerites of outer layer red; of inner layer nearly colorless.
Etymology. e 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.
Common name. Chuck’s gorgonian; Orange gorgonian; Shady-leafed gorgonian;
Hard-leaved gorgonian; Hidden gorgon (these names appear in a variety of eld/diving
guides for the area such as Gotshall 2005, Kerstitch and Bertsch 2007).
Distribution. 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.
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 75
Figure 1. Adelogorgia phyllosclera, SBMNH 51252. Colony measures 16.5 cm × 15.0 cm.
Biology. 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” (veried by sta of LACSD
and OCSD). Two specimens, USNM 50186 and SBMNH 422894 (Point Loma),
Elizabeth Anne Horvath / ZooKeys 860: 67–182 (2019)
76
support numerous epizootic anemones (perhaps Epizoanthus Gray, 1867). On others,
a at, 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 coenenchy-
mal covering. One specimen (SBMNH 422893) harbors a small brittle star, wrapped
around a portion of the main trunk. No zooxanthellae present in the tissues, particu-
larly true of USNM 50186; specimen examined for their presence by Bayer.
Remarks. Among the eight to ten specimens that Bayer examined in 1958, there ap-
peared to be three main areas that showed variation: thickness of branches, development
(size) of the calyces (if present), and proportions of dierent 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 signi-
cance. Other more slender specimens exhibit a wide range of variation in branch diameter.
Most eld/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. ree lots examined and conrmed correct as to their genus
identication (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”.
ere 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 conrm this species’ total range
(where it may either transition to A. telones, or other recently discovered species, or has
a denite 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 dierences that
would separate species were not clear; all specimens, with one exception (completely
bleached, SBMNH 422891 from Santa Cruz Island, Galápagos Islands), are the distinc-
tive 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
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 77
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 caly-
ces being inconspicuous or not really present at all), and 3) sclerites somewhat larger,
with double wheels/discs somewhat dierent in shape along with being more elaborately
sculptured. Based on the coloring of specimens (with one exception) in the SBMNH
Figure 2. Adelogorgia phyllosclera. A SBMNH 422403 [dry], slightly magnied image of branches
BSBMNH 51252 [wet], rounded, slightly club-shaped branch tips.
Elizabeth Anne Horvath / ZooKeys 860: 67–182 (2019)
78
Figure 3. Adelogorgia phyllosclera, SBMNH 51252, SEM image. Sclerites red-orange in color. A Short wart-
ed 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 (gs 5c, 6c).
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.
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 79
Genus Eugorgia Verrill, 1868
Lophogorgia (pars) Horn, 1860: 233.
Gorgonia (pars) Verrill, 1864: 33; 1866: 327.
Leptogorgia Verrill, 1864: 32.
Eugorgia (pars) Verrill, 1868c: 414.
Eugorgia Verrill, 1868b: 406–407. Studer 1887: 64–65. Bielschowsky 1918: 39. Kü-
kenthal 1924: 343. Bielschowsky 1929: 170. Stiasny 1951: 63. Bayer 1951: 99;
1981: 921. Breedy et al. 2009: 8.
Type species. Leptogorgia ampla Verrill, 1864; by subsequent designation Verrill
1868b: 386.
Diagnosis. Breedy et al. (2009) did a thorough review of this genus (well repre-
sented in the SBMNH collection). Sclerites chiey disc spindles, capstans or double
discs (wheels); double discs up to 0.05 mm long, spindles 0.12–0.18 mm, not devel-
oped as clubs; ordinary spindles present in small numbers in some species. Anthoco-
diae unarmed; sclerites, if present, at rods and platelets with lobed margins. Pol-
yps 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 min-
eralized organic laments. Colony colors quite variable, depending on species.
Eugorgia daniana Verrill, 1868
Figures 4, 5, 6, 7
Eugorgia daniana Verrill, 1868a; 1868b [1869]: 409–410; pl V, g. 14; pl VI, g. 7.
Bielschowsky 1918: 45. Kükenthal 1924: 346. Bielschowsky 1929: 181. Stiasny
1951: 65. Prahl et al. 1986: 17. Breedy et al. 2009: 17–20.
Type locality. Central America: Pearl Islands; Costa Rica, Gulf of Nicoya.
Type specimens. Syntype series (Breedy et al. 2009): MCZ 723 [dry]; MCZ
7080 [dry]; YPM 1551a-d and 1629a, b [dry]; YPM 5146 [dry].
Material examined. ~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).
Description. Collection lot examined, shown in Figures 4 (whole colony), 5
(branch magnied 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-
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Figure 4. Eugorgia danianae, SBMNH 422897. Shows complex branching that creates wide, at fan.
Colony is 24 cm high × 25.5 cm broad at widest point.
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).
Common name. Red gorgonian (Gotshall 1998).
Distribution. As recorded by Verrill (1868a and 1868b [1869]) found in Pana-
ma 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 Die-
go, 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.
us, 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).
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 81
Biology. Generally found on oshore reefs and islands; prefers clean, plankton-
rich waters and generally found at depths to ~30 m (Gotshall 1998).
Remarks. While description generally matches that given in Breedy et al. (2009),
I would make the following caveats: rst, the colony color as described by Verrill was
as a bright yellow, streaked and blotched with dark red on both branches and polyps.
e color seen in specimens here lies somewhat intermediate to that described in Ol-
vera 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 dierent size and form of the sclerites,
especially of the double discs; for Verrill, these were the dening feature in designating
this species as distinct from E. aurantiaca. In my examinations, the larger, “sharp” dou-
ble 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 (misidentied 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 pat-
tern, specimens of E. aurantiaca often had a far more distinctive (and decidedly sym-
Figure 5. Eugorgia danianae, SBMNH 422897. Close-up of pinnate branching pattern.
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Figure 6. Eugorgia danianae, SBMNH 422897, light microscopy array, 10× magnication. Note jagged,
sharp-edged, double discs and color variety. Sclerites will appear orangey-yellow collectively but individu-
als can be deep, bright red, yellow-green, or bicolored, with a maximum length up to ~100 µm.
metrical) pattern of pinnate branching, with majority of secondary branches (branch-
lets) 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, denitive conrmation that Eugorgia aurantiaca and Eugorgia daniana
are indeed separate species. ere are many overlapping features, in terms of potential
branch pattern, form, and size of sclerites, and general color. e dierences could be
accounted for as variation within one species. However, examination of numerous ex-
amples of both those identied as E. aurantiaca and E. daniana allowed for observation
of the dierences in the double discs that Verrill used to distinguish these two species
(dierences 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 Eugor-
gia 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
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 83
Figure 7. Eugorgia danianae, SBMNH 422897, SEM image. Representative sclerite of each form.
ADouble-disc B Coenenchymal sclerite C Anthocodial sclerite. Images here compare well with those
shown in Breedy et al. 2009 (g. 8).
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.
Eugorgia rubens Verrill, 1868
Figures 8, 9, 10A–D
Eugorgia rubens Verrill, 1868b [1869]: 411 [not gured]. Studer 1894: 69. Biels-
chowsky 1918: 45.
Kükenthal 1924: 346. Bielschowsky 1929: 183. Breedy et al. 2009: 29–31.
Type locality. SE Pacic Ocean, South America, Peru, Piura Dept., Paita. e type
locality is often incorrectly spelled as Payta.
Type specimens. Type–YPM 1779 [dry]; MCZ 36047 (slide of holotype).
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Figure 8. Eugorgia rubens, SBMNH 422900. Colony 45 cm tall, 18 cm across (through broadest, middle
section). Scale bar: 2 cm.
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 85
Material examined. ~ 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.
Description. In general, all SBMNH material (along with supplemental lots) exam-
ined 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 identiable characteristics for the species.
Etymology. 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.
Common name. Purple gorgonian (Gotshall 1994); Purple sea fan; Reddish true
gorgon. Common name not specied in Cairns et al. (1991; 2003).
Distribution. From southeastern Pacic 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 speci-
mens were examined; collection location data shows geographic and depth ranges.
Biology. 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 o 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
Figure 9. Eugorgia rubens, SBMNH 45562. Magnied, showing branching and extended polyps.
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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
CCoenenchymal spindles D Double disc spindles. Images here compare well with those shown in Breedy,
et al. 2009 (g. 15).
occur at depths greater than originally thought, extending much further north than
previously reported (the specimen that called attention to this greater depth was pho-
tographed by C Bauder at Point Lobos, Carmel Bay, at 66.5 m). A thorough examina-
tion 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
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 87
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 o 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. ere is also evidence of the presence of ovulid snails from the Genus Simnia
(Neosimnia) Risso, 1826 (species S. barbarensis Dall, 1892 and S. loebbekiana Weinkau,
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 o 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 identication).
Remarks. 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 identiable
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 identied to
genus only); also several lots identied 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 ad-
dition, 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 Richeld 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 Aquar-
ium, 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 col-
lected from the local area. is is a very common species in Southern California waters
and is an accepted species in the WoRMS Data Base (Cordeiro et al. 2018b).
Eugorgia ljubenkovia sp. nov.
http://zoobank.org/48F9BA66-D012-44A5-8C8F-1EEAE2279548
Figures 11A, B, 12A, B, 13A–C
Type locality. Isla Cedros, Baja, Mexico.
Type specimens. Holotype Santa Barbara Museum of Natural History, SBM-
NH 422333.
Material examined. ~5 lots (see Appendix 1: List of material examined).
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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 Magnication of branch strand, showing small, rounded polyp mounds.
Diagnosis. Colony an obvious whip-like form, no apparent holdfast, with mini-
mal 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.
Description. 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, sti, wiry strands, none or very few primary or secondary
branches (unusual for species in this genus); what branches are present come o at
right angles to main stem, then curve some ve cm distant or more from branching
point; branch pattern (one colony) more dichotomous or lateral. Tips of branches
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 89
(both ends) terminate in small arrowhead conguration. 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 green-
ish 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 attened appearance from front
to back (closer examination revealed polyps on all sides); very thin ridge line (longitu-
dinal 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 be-
tween 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.
Etymology. Proposing Eugorgia ljubenkovia, to honor John Ljubenkov, a southern
California cnidarian biologist, colleague and friend of many Southern California As-
sociation of Marine Invertebrate Taxonomists (SCAMIT) members.
Common name. John’s wire gorgonian.
Distribution. Known from collection events undertaken by sta of Orange Coun-
ty 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.
Figure 12. Eugorgia ljubenkovia sp. nov., SBMNH 472232, light microscopy sclerite array. A 10×, show-
ing 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.
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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.
Biology. Moderate occurrence, indicated by OCSD collection events; not occur-
ring 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 com-
pletely covered with gorgonian’s coenenchyme.
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 91
Remarks. All colonies have shape of a thin, whip-like Leptogorgia species, and may
exhibit the same presumed lifestyle as that of Leptogorgia licrispa Horvath, 2011 or
that of species in the genus esea Duchassaing & Michelotti, 1860. However, sclerites
most distinctive in being largely double discs, characteristic of species in the genus Eu-
gorgia. Originally, was tentatively identied 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 esea;
despite the long, thinner branch strands and possible lifestyle similarity, no large sphe-
roidal bodies, characteristic of esea were found. Also, this species (in ve lots), is not
additional material of the species L. licrispa (Horvath, 2011), as jagged double discs
are not seen in that species, and sclerites in that species can be variably colored. is
species is a unique mix of colony form seen in some Leptogorgia with the sclerites of a
Eugorgia. e long, thin wiry condition of the stems may be the result of environmen-
tal circumstances, involving both substrate (sandy or soft bottom sediment) and water
ow. 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.
Genus Leptogorgia Milne Edwards & Haime, 1857
Gorgonia (pars) Pallas, 1766: 160. Milne Edwards and Haime 1857 (pars): 157.
Leptogorgia Milne Edwards & Haime, 1857: 163. Verrill 1864 (pars): 31, 33; 1868b:
387; 1869b: 420. Studer 1887 (in Wright): 64. Bielschowsky 1918: 18. Kükenthal
1919: 851; 1924: 324–325. Bielschowsky 1929: 81. Stiasny 1943a: 87. Bayer
1951: 98–99; 1956a: F212; 1961: 214. Grassho 1988: 97; 1992: 54. Williams
1992: 231. Williams and Lindo 1997: 500. Bayer 2000: 609. Breedy and Guzmán
2005: 2; 2007: 6. Horvath 2011: 46. Breedy and Cortés 2011: 63.
Lophogorgia Milne Edwards & Haime, 1857: 167. Nutting 1910c: 3, 4. Kükenthal 1924:
322. Bielschowsky 1929: 73. Stiasny 1943: 87. Bayer 1951: 99; 1956a: F212; 1961:
194. (Type species by monotypy: Gorgonia palma Pallas, 1766 [South Africa]).
Type species. Gorgonia viminalis Pallas, 1766; subsequent designation by Verrill 1868a.
Diagnosis. Sclerites primarily symmetrical spindles, most without unilateral fu-
sion 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 arma-
ture attened rods; sometimes, ovoid platelets. Colonies little-branched, long, slender,
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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 laments, frequently mineralized. Polyps fully retractile into coenenchyme;
slightly raised, mound-like, around apertures.
Remarks. 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).
Leptogorgia chilensis (Verrill, 1868)
Figures 14A, B, 15A–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 exilis 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 o
Algarrobo. For neotype (designated here), northeastern Pacic Ocean, North Amer-
ica, 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 (un-
published 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 ush 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
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 93
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 cre-
ated by species of barnacle, a common occurrence on this and other gorgonian species.
ring of warts on either side of median girdle; also capstans (two whorls with end tufts),
modied 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 ne 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 eld/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 south-
ern California (USNM 52442). However, examination and comparison of sclerites
taken from many “red whip” forms indicated that likely range of Leptogorgia chilensis is
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Figure 15. Leptogorgia chilensis, SBMNH 422953, SEM image. Color of sclerites is orange-red. A An-
thocodial sclerites B–D Spindles of coenenchyme. Images closely match those shown in Breedy and
Guzmán 2007 (g. 14).
from Anacapa Island o 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. at discus-
sion 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 Califor-
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 95
nia Bight northward, and overlap with L. chilensis in the extreme southern end of their
range, may well be one or more dierent 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 oshore
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 oshore 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, o 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.
is 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 Lin-
naeus, 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 identication); this may be op-
portunistic as it populates widely dierent sites in addition to this species. As well, several
mollusk species have been recorded in association with this species, such as Tritonia festi-
va Stearns, 1873, as reported by Gomez (1973) and several snails of the genus Neosimnia
(now Simnia), such as Neosimnia barbarensis Dall, 1892 (eodor 1967, as referenced in
Langstroth and Langstroth 2000). An unidentied eld 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
lucicum, now Savalia lucica (Cutress & Pequenat, 1960), and likely other species, ulti-
mately 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 sev-
eral 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 o the
quarry, near Avalon on Catalina Island, has Savalia lucica (Cutress & Pequenat, 1960)
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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 approxi-
mately one third of its branches festooned with a creamy yellow zoanthid. ere is some
specic substratum choice indicated here, and is apparently common among colonial zo-
anthids. 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 identication 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 Chro-
moplexaura marki). ey mention the bryozoan Celleporina robertsonae, now recognized
as Costazia robertsonae (Canu & Bassler, 1923), the Broken-back shrimp Heptacarpus
exus (Rathbun, 1902), which may scavenge on sclerites, mucus and even toxic tissues
from the surface of the gorgonian, a caprellid amphipod, specically a skeleton shrimp,
Metacaprella anomala (Mayer, 1903), whose color may derive from their acquiring the
pigment ingested while scavenging the gorgonian’s sloughed o 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; identication 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). “erefore, 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 eodor 1970).
Remarks. Harden (1979?) stated this as being one of the most common sea whips
from southern California; my examinations conrm 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 exclu-
sion of others. e reality is that there are other red whip species which can easily be mis-
taken for L. chilensis; a cursory look by eye alone can (and has led) to misidentication.
Characteristics ordinarily used for separating Lophogorgia from Leptogorgia, the
attened 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
attened branches may occur in the same colony, and biserial zooid distribution can be
found with little diculty. Furthermore, specimens of Leptogorgia (typical in all other
respects) may have zooids distributed all around the branchlets. Bayer rst placed the
genus Lophogorgia in synonymy with Leptogorgia in 1951; based on the work done by
Grassho (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).”
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 97
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. e Cabrillo Marine
Aquarium, in San Pedro, California, had a number of live colonies of this species on dis-
play in the public area; all were collected in the local area. As well, the Aquarium of the
Pacic, 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. e transitional areas around Point Conception (and waters
northward beyond the Bight) oer some intriguing distributional scenarios that will re-
quire further exploration. While of similar appearance in general colony form and color,
material collected by sta of Olympic Coast National Marine Sanctuary that I examined
in Washington State the summer of 2006 (tentatively identied 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 dierent species. is is covered
further in the “red whip” discussion included in the remarks made regarding the quintes-
sential “red whip” of the northwestern California coast, Chromoplexaura marki.
Looking at location records for specimens collected and identied as L. chilensis (with
conrmed identication), I noted that if type locality is correct, L. chilensis should range
from the colder waters o the coast of Chile up through the warmer waters of Central
America and Mexico before again encountering the cooler waters of the southern Cali-
fornia Bight. How is this possible? What would the depth parameters, substrate features,
distance from shore and specic distributional pattern (continuous or fragmented) look
like for this species? e missing type material confounds the issue. e material used
for Philippi’s (1866) description of L. rosea could not be located and no recently col-
lected material from Chile resembling this species is apparently available. Breedy and
Guzmán (2007) used California specimens for their description of this species. ey “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 denitive 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. chil-
ensis 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. is
is a prime example of a situation where the presumption that any “red whip” found in
the southeastern or northeastern Pacic Ocean is likely L. chilensis (considered to be quite
common), is faulty. e only way to resolve questions surrounding this species is to inten-
tionally 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 identied as Leptogorgia caryi Verrill, 1868 was examined. is was a dry specimen
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from NMNH (USNM 5988), collected at Catalina Island, California. In examining
it, along with several specimens of L. chilensis, there appeared no marked dierences
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 speci-
men 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). e sclerites in this NMNH
specimen denitively put it in the genus Leptogorgia; sclerites of any specimen identied
as E. (Chromoplexaura) marki certainly did not t the sclerite description of any in the
genus Leptogorgia. us, E. (Chromoplexaura) marki is not a junior synonym of L. caryi.
is was subsequently veried (ITIS Report, accessed online June 2011). Further, a
specimen identied 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). is specimen
was supposedly deposited in the Museum of Comparative Zoology at Harvard Univer-
sity, but Breedy and Guzmán could not locate it for examination. Based on its collec-
tion locality, it would seem that the original identication of this missing specimen is in
error. I have concluded that the range of distribution that was stated in the discussions
above, and conrmed generally by Breedy and Guzmán, seems to accurately pinpoint
where this particular species is found. at it may overlap several other red whip forms
in the northern end of its range only emphasizes the need for thorough sclerite ex-
amination 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 denitively.
Leptogorgia diusa (Verrill, 1868)
Figures 16A, B, 17, 18A–D
Litigorgia diusa Verrill, 1868a; 1868b: 397–398.
Gorgonia (Litigorgia) diusa Verrill, 1868c: 415.
Leptogorgia diusa Verrill, 1868a; 1868b: 397–398; pl V, g. 6; pl VI, g. 3; 1869b:
421. Nutting 1910d: 5. Bielschowsky 1918: 30; 1929: 112. Kükenthal 1919: 771;
1924: 329–330. Hickson 1928: 413–414. Stiasny 1935: 29. Breedy and Guzmán
2007: 32–37.
Nec Leptogorgia diusa: Stiasny 1951: 71 [Guyane Française, Ile Royale] = (Leptogorgia
punicea (Milne Edwards & Haime, 1857) [see Bayer 1961]).
Leptogorgia rubra Bielschowsky, 1918: 29 [nomen nudum]; 1929: 92–94. Kükenthal
1919: 911–912; 1924: 325.
Lophogorgia diusa: Prahl et al. 1986: 21.
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 99
Type locality. (Lectotype) Gulf of Panama, Panama, Pearl Islands; additionally (Para-
lectotypes) Gulf of Nicoya, Costa Rica.
Type specimens. Holotype, as Litigorgia diusa Verrill, 1868); YPM 1659a [dry].
Lectotype Breedy and Guzmán 2007: YPM 1659 [dry]. Paralectotypes Breedy and
Figure 16. Leptogorgia diusa, 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.
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Figure 17. Leptogorgia diusa, SBMNH 423090, light microscopy, 10×, showing sclerites typical of
genus and this species. Note pale yellow anthocodial sclerite, upper middle portion of eld, amongst deep
red to mauve-pink spindles. Longest sclerites may be part of the anthocodial armature.
Guzmán 2007: MCZ 7081 [dry]; YPM 5151 [wet]. Breedy and Guzmán (2007) be-
lieve it unlikely that YPM 1659a or 1659b are this species.
Material examined. 4 lots (see Appendix 1: List of material examined). I was un-
able to examine the designated type specimens, but again, distinctive characters of the
species made this unnecessary for ID of SBMNH specimens.
Description. Colony form (Figure 16A), appearance of branch and polyp place-
ment (with overall eect of polyp placement that of serrated or zig-zag appearance;
Figure 16B) and sclerites (Figures 17, 18A–D) correspond with that given in descrip-
tion of the species in Breedy and Guzmán (2007: 32–36), although in the SBMNH
material examined, the anthocodial sclerites (Figure 18A) seem exceptionally long.
Etymology. Latin diuses- means spreading, perhaps in reference to open shrub-like
appearance that the branches create. No discussion of the species name is given by Verrill.
Distribution. From Panama and Costa Rica to southern California, at least.
Remarks. e lax, attened branches, large polyps that produce zig-zag appear-
ance, large (and in this case, long) anthocodial rods and the dull brick-red coloring are
clear diagnostic features for this species. e species is listed as an accepted form in
Cordeiro et al. (2018c).
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 101
Leptogorgia licrispa Horvath, 2011
Figures 19, 20, 21, 22B–N
Leptogorgia licrispa Horvath, 2011: 45–52.
Type locality. Mexico, Baja, California, o Boca Flor de Malva, SE of Punta Tosca,
~24°11'07.04"N, 111°21'03.08"W, 69–87 m.
Figure 18. Leptogorgia diusa, SBMNH 423090, SEM image. A Anthocodial sclerites; some were sur-
prisingly and unexpectedly long B–D Coenenchymal spindles. Images, for the most part, match those
shown in Breedy and Guzmán 2007 (gs 23, 24).
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Figure 19. Leptogorgia licrispa, 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.
Type specimens. Holotype SBMNH 423057; [dry]; Paratypes SBMNH 423079
[dry]; USNM 1106683 [dry]; USNM 1106684 [dry]; USNM 1106685 [dry].
Material examined. ~9 lots (see Appendix 1: List of material examined).
Description. 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 attened 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 ne wire, yet sti and brittle (Fig-
ure20); 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 axed to a small rock or pebble; majority of colonies without
a base. Not denitively 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; rang-
ing 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
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 103
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 (Fig-
ure21); 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 attened 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 sec-
ond 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. Gener-
ally, sclerites without much color; if colored, typically light pink.
Etymology. e species designation is derived from the Latin root li- for thread,
and the Latin root crispa- for curled or twisted; designation reects overall strand ap-
Figure 20. Leptogorgia licrispa, SBMNH 423057-Holotype. Two single strands, each showing a dif-
ferent pink color variant. Image originally published as gure 1B in Horvath (2011, Proceedings of the
Biological Society of WA 124: 1, 47).
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Figure 21. Leptogorgia licrispa, SBMNH 423057-Holotype. Close-up of branch, showing spacing of
polyps; Scale bar: 1.0 mm. Image originally published as gure 1C in Horvath (2011, Proceedings of the
Biological Society of WA, 124: 1, 47).
pearance, which is thread-like, sti and wiry; strands of this species reminiscent of the
sti, wiry, curled body of an adult horsehair worm.
Common name. Multi-colored wire gorgonian.
Distribution. 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.
Biology. A comment was made (Grigg 1972) about gorgonian colonies having
“a loosely branched and whip-like shape when located in circular basins where water
ow 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 mo-
tion, 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
conrmed reports from other observers, thus far, nothing more can be stated about
this species’ biology.
Remarks. e 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. Dierences were apparent in geographic location, coloring of scler-
ites, 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
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 105
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 o Redondo Beach, California, US), it is possible that this species is actu-
ally 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.
Figure 22. Leptogorgia licrispa, SBMNH 423057-Holotype. SEM image, originally published as gure
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, gener-
ally measuring in the range of 55–80 µm; coenenchymal spindles, scale bar 20 µm (B–F, H–J), coenency-
mal sclerite, scale bar 100 µm (G); anthocodial sclerites, scale bar 20 µm (K–N).
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Leptogorgia exilis (Verrill, 1868)
Figures 23A, B, 24A–C
Gorgonia (Eugorgia) exilis Verrill, 1868c: 415.
Litigorgia exilis Verrill, 1868a; 1868b: 400–401.
Leptogorgia exilis Verrill, 1868b: 400–401; pl V; g. 11; 1869b: 421. Nutting 1910d:
5. Bielschowsky 1918: 29. Kükenthal 1919: 771; 1924: 326. Hickson 1928: 414–
416. Bielschowsky 1929: 96. Stiasny 1943: 82. Breedy and Guzmán 2007: 40–44.
Type locality. Archipelago Las Perlas, Panama, 11–15 m.
Type specimens. Syntypes Breedy and Guzmán 2007: YPM 1553a, b [dry];
MCZ 4123 (722) [dry].
Material examined. 5 lots (see Appendix 1: List of material examined). Desig-
nated types not examined.
Description. 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).
Etymology. e root exi- is Latin for pliant, bendable, referring to the apparently
exible, droopy, slender branchlets of the live colony. However, Verrill does not give
any rationale for the species name.
Distribution. Panama, north into lower third of California Bight (o Santa Cat-
alina Island and adjacent California mainland sites).
Remarks. Initially, the drooping branches were considered to be more an arti-
fact of preservation and the containers initially used when collected (branches bent
downward so specimen would t in the jar). However, descriptions by others (Breedy
and Guzmán 2007), indicated that this is a normal branch conguration. Initial pres-
ervation 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 eorts. As well, much of its
color has leached out, with any particular branch colored from almost white to tan
to pinkish red.
Leptogorgia exilis is an accepted species in the WoRMS Data Base (Cordeiro
etal.2018c).
Leptogorgia species A
Figures 25A, B, 26A–C, 27A–D
[? = Leptogorgia tricorata Breedy and Cortés 2011]
Type locality and type specimens. ere is a need for further conrmation of species
identication regarding SBMNH specimens, through examination of other denitive-
ly identied specimens, as well as the type specimens for L. tricorata (Holotype UCR
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 107
Figure 23. Leptogorgia exilis, 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.
1833; Paratypes UCR 1834, 1835, 1836 and 1837). e holotype was collected
in Cocos Island National Park, Isla Manuelita NW, taken on 8 September 2006 at a
depth of 14 m. e paratypes were collected from Cocos Island National Park as well,
either at Isla Manuelita or Roca Sucia.
Material examined. ~11 lots (see Appendix 1: List of material examined).
Description. Colonies (Figure 25A) non-reticulate; main stem ~14 cm long, aris-
ing from thin, at attachment structure; latter gives o 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
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Figure 24. Leptogorgia exilis, 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 Quadriradi-
ate from coenenchyme. Images match those shown in Breedy and Guzmán 2007 (g. 30).
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 diam-
eter 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
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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.
with prominent conical polyps, when extended (Figure 25B); when contracted, nearly
ush 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 silhou-
ette 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 compari-
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Figure 26. Leptogorgia sp. A, SBMNH 423080, Light microscopy image. All sclerites in SBMNH mate-
rial are deep yellow in color. A–C Increasing magnications 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.
son 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.
Distribution. 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.
Biology. Barnacle galls present on a number of specimens (SBMNH 423084 and
SBMNH 422903).
Remarks. is assemblage of specimens still not identied with certainty; de-
spite 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 be-
ing late 1970s) are also included in the collection. Since then, however, no speci-
mens 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 esea Duchassaing & Michelotti, 1860 (which they can resemble
on a supercial level; this especially true of fragments of SBMNH 422334; one
other specimen in collection, SBMNH 13304, from a station o Point Loma, is
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 111
identical), and color that is generally bright lemon yellow-gold, with very mark-
edly 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 esea were seen in these specimens; esea was
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).
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eliminated as a possibility. What prevents a positive identication (as L. tricorata)
was a dierence 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 ten-
tacular 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 Pacigorgia Bayer, 1951
(see page 226, Soler-Hurtado et al. 2017). Cordeiro et al. (2018d) does not show this
proposed emendation of the genus Pacigorgia 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 reective of branching
patterns usually seen species of Pacigorgia.
Family Plexauridae Gray, 1859
Diagnosis. Colonies of very diverse form, generally with thick branches arising lat-
erally, 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 nonscle-
ritic calcareous matter (common tendency toward heavy calcication of base in old
colonies). Coenenchyme thick, perforated by system of longitudinal canals surround-
ing axis, delimiting outer coenenchymal layer from inner one (axial sheath), which
dier in spiculation. Sclerites usually include some form of club; some with spindles
only, oval bodies, rods or large quadriradiates.
Remarks. 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), em-
phasis 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 esea Duchassaing
& Michelotti, 1860 are also included in this family, it was necessary to cover those
with a more extensive study, discussed in Part III. e genus esea, 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.
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Plexaurinae
Family Plexauridae [= Muricidae]
Genus Chromoplexaura Williams, 2013
Chromoplexaura Williams, 2013a (part): 31, 34–35.
Type species. Euplexaura marki Kükenthal, 1913.
Diagnosis. Tall, erect, generally planar colonies, bright red; if branched, lateral
(not extensively branched, if present, at all), from single, basal stem. Upper branches
slender, elongate, most slightly curved, distally less dense; denser proximally and low-
er in colony. Polyps fully retractile; on all sides of branches and stem, as numerous
slightly rounded, low to at protuberances. Sclerites also red; robust spindles, and
radiates, some ellipsoidal to sub-spherical in shape; prominent sclerite a long spindle
with prominent, cone-shaped caps at each end and obvious median “waist” (herein
referenced with new terminology: the double-dunce cap or double-dunce). Contains a
single species from the temperate eastern Pacic (generally, California to Washington;
slight possibility of presence in Canadian (even Alaskan) waters).
Etymology. Derived from the Greek chroma- referring to color, and the gorgonian
generic name plexaura- in reference to the bright color of the colonies.
Remarks. Diagnosis for the genus Euplexaura (Kükenthal, 1913a) was examined for
comparison with that of the recently proposed genus Chromoplexaura Williams, 2013a.
e species placed in this new genus is well represented in the SBMNH collection, t-
ting the description given by Williams (2013a, 32–39). e original placement of the
temperate Eastern Pacic species in the genus Euplexaura was adhered to for an entire
century, based on the original description of Kükenthal (1913a). at original descrip-
tion was little referenced, and specimens of the species in this genus collected along the
California coast post-1913 were often misidentied. Both the genus (as represented in
CA) and the locally collected species received virtually no further attention until my work
began on the SBMNH collection in 2002; a subsequent inquiry of Dr Williams (e-mail
conversation, March 2011) was made, regarding what his perspective on the species was.
Williams’ (2013a) establishment of a new genus for this temperate gorgonian is justied.
Chromoplexaura marki (Kükenthal, 1913)
Figures 28, 29A, B, 30A, B, 31A–C, 32A, B, 33A, B, 34A–C, 35A–E, 36, 37A1–4,
B1–3, 38A–E
Euplexaura marki Kükenthal, 1913: 266–269; text gs G, H, J, K, pl 8 g. 11; 1924:
93–94.
Chromoplexaura marki (Kükenthal, 1913): Williams 2013a: 36–39; gs 12–17.
Type locality. For the original specimen, USA, Southern California, 64–616 m. (Iden-
tication cannot be conrmed.) For proposed Neotype, collected in Northeastern
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Figure 28. In situ image of what appears to be Chromoplexaura marki, DSCN5297. Colony seen o
Oregon coast. Without initial examination of sclerites, originally thought to be Swiftia simplex; examina-
tion of sclerites aligned it with C. marki. ere can be remarkable similarity in external gross appearance
between the two species. Image courtesy of Peter Etnoyer, NOAA, 2010.
Figure 29. Chromoplexaura marki, SBMNH 423060 (Neotype). A is specimen, measuring 24–26 cm
at greatest length, bears two dierent types of barnacle, a species that appears to be that of an acorn barnacle
(forming the galls on this specimen), and clusters of a barnacle species that appears to be a “Lepas-type.” e
sclerites from this colony aligned with known specimens of C. marki B Branch tips at greater magnication.
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Figure 30. Chromoplexaura marki, SBMNH 423062. A Specimen (whole colony). Sclerites did not
always consistently match sclerites seen in the species, illustrating variable nature of sclerite forms. Colony
measures ~16 cm × 10 cm B Close up, branches and branch tips, clearly showing bright red-orange color
and distinctly white polyps of the species C. marki.
Pacic, USA, California, Monterey County, Monterey, BLM Reference Station 360
(Burch #40128), ~22 m; coll. T Burch, 18 August 1940.
Type specimens. Repository for the original type specimen unknown. Proposed
Neotype (designated here), SBMNH 423060 [dry].
Material examined. ~60 lots (see Appendix 1: List of material examined).
Description. Colony (Figures 28, 29A, 30A, 31A, 32A, 33A) shape can be a wide,
broad, moderate to sparsely branched fan, typically in one plane or simple, unbranched;
initial branching lateral, progressing to branches that tend to project more up than out;
sometimes projecting/winding more in a “front” or “back” direction; with broader
membranous base; main branches can divide repeatedly, all secondary branches o
larger ones having same diameter; round in cross-section; distal ends often slightly
swollen; see Figures 29B, 30B, 31B, C (See Remarks below for further discussion of
overall colony shape). Axis proteinaceous, generally well calcied, not very exible,
with hollow core. Color of axis variable between white, yellowish, and light brown.
Color of living colony base, stem and branches bright coral red (orange-red), bright
red, or dark red; red color enhanced by color of the sclerites, which are a bright, pale,
transparent red. Polyps are (pale) yellow-colored when living; (Johnson and Snook
(1927) stated the color of E. marki to be coral-red, with the living polyps yellowish
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Figure 31. Chromoplexaura marki, SBMNH 423072. A Colony with coenenchyme of a bright orange-
red color with conspicuous white polyps. Sclerites did not always consistently match sclerites seen in the
species, illustrating variable nature of sclerite forms. Specimen measures ~15.5 cm × ≤2 cm B Branch
close-up, showing conspicuous white polyps C Close up of branch tip.
white). When polyps visible in preserved specimens, they appeared white/cream. Coe-
nenchyme moderately thick, rising in wall of polyp as eight very short folds. Polyps
sit ~2.0 mm distant; polyps ~2.0 mm high, 1.0 mm broad, when extended; polyps
fully retractile into coenenchyme surface, forming very low, rounded bumps (“polyp-
mound”); aperture suggestive of a goblet/chalice shape; fortication of polyps weak.
Kükenthal (1913a) stated that there are no calyces in this species, or at least are so neg-
ligible as to be virtually nonexistent (in describing Euplexaura marki, he indicated that
there should be two transverse rings of large sclerites, one over folds in the polyp head
and a second just below insertion of the tentacles). Sclerites of polyp body red, while
sclerites of tentacles smaller, colorless, transparent, but of similar form. Occasionally,
tentacle sclerites can be a bent spindle (Figures 34A–C, 35A–E, 36, 37A1–4, B1–3,
38A–E). Found initially in a specimen identied as this species (USNM 51500) was
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 117
a sclerite form that was not clearly featured in sclerite descriptions for the genus, or
other known species (Figures 34A–C, 35C, 36, 37A1, B1, 38B): in outer surface of
branch coenenchyme lie sclerites whose basic form is a thick spindle, but on these sit
two or more belts of very big, jagged warts. rough the development of these large
warts, the spindle can have a contour that is nearly oval (these might be the tuberculate
spheroids mentioned in original description; apparently a key sclerite form, for the
genus Euplexaura, at least) to a distinct diamond shape. Occasionally, one can nd
Figure 32. Chromoplexaura marki, SBMNH 265948. A Very small colony, 4.0 cm in length, excluding
mass at base, which appears to be a sponge. Sclerites did not always consistently match sclerites seen in
the species, illustrating variable nature of sclerite forms B Close up of extended polyps on single branch,
each measuring ≤ 1.0 mm in height.
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Figure 33. Chromoplexaura marki, SBMNH 265935. A Two colonies, larger of the two (only 7.0 cm
tall) with conspicuous brittle star attached. Sclerites did not always consistently match sclerites seen in
the species, illustrating variable nature of sclerite forms B Close up of branch tip, showing placement of
calyces, color of polyp body and tentacles.
on both ends of these spindles, dense triangular caps (Figure 34A–C, 35C, 36, 37A1,
B1, 38B) separated by a smooth, usually thin, median waist, so that in this case, the
term double spindle (double-head) could apply; these are characteristic and conspicu-
ous of multiple specimens examined, and henceforth referred to as the double-dunce
cap. Size of these outer spindles uctuated considerably, with smallest only 0.05 mm
in length, but often bigger (~0.2 mm). ose deeper into coenenchyme of branches
had similar form, but warts were more rounded (Figures 35B, 37A3, B3, 38C). All of
the more supercial sclerites are red, making the strong bright red color of the colony
fairly pronounced.
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Figure 34. Chromoplexaura marki, SBMNH 423060, light microscopy arrays. A 4× magnication. Note
in particular the “triangular-capped” spindle designated with arrow. is sclerite form is never seen in speci-
mens from the genus Swiftia (some species in genus Swiftia can look supercially like this species in overall
colony form) B Additional image at 4× magnication of sclerites, illustrating further examples of “trian-
gular-capped” spindles appearing commonly in Chromoplexaura marki C 10× magnication of sclerites,
showing variety, with particularly clear example of the “triangular-capped” spindle, indicated with arrow.
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Figure 35. Chromoplexaura marki, SBMNH 423060, SEM image. A Anthocodial sclerites B Small
coenenchymal forms C Larger, distinctive “double dunce-cap” spindles D Odd spindle E Still smaller
coenenchymal sclerites. Images match variety shown in Williams 2013a (gs 14–17).
Etymology. Species named in honor of EL Mark of Harvard University.
Common names. MBARI (seen in a hall display, Summer 2008) referred to this
species (and to any species from northern California appearing as a “red whip”) as “Red
licorice gorgonian”.
Distribution. Southern California; littoral and coast-abyssal (Kükenthal, 1924, as
Euplexaura marki). Johnson and Snook (1927) noted the species living in deep water,
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taken with a dredge; specimens were collected o the Oregon coast, and are either in
the Oceanography Department of Oregon State University, or in the personal collec-
tion of FP Belcik. Nutting (1909) reported numerous collection points, at stations
near San Nicolas Island, and for stations near Point Piños Lighthouse, Monterey Bay.
Likely, range extends from southern California to waters o Washington coast. ere
is the possibility that the species extends further north, to Alaska; further examinations
of specimens from that area are in progress.
Biology. An unidentied, anecdotal comment indicated that this form is seen in
the assemblage of organisms found at the head of Carmel Submarine Canyon, located
oshore at San Jose Creek Beach, near Carmel, California; considered part of a deep-
water assemblage that begins to appear at depths between 21–30 m, where turbulence
is minimal and ne sediments accumulate on surface irregularities of rock walls. Be-
tween 30–61 m, the fauna appears to change very little, suggesting that many of these
deep-water forms extend to greater depths.
e neotype designated here (SBMNH 423060, Figure 29) bears on several
branches, enlargements that are in actuality gall-like growths, containing epizoic bar-
nacles of the genus Conopea (likely Conopea galeata). is is a consistent, common
obligate commensal barnacle of gorgonians (Langstroth and Langstroth 2000). On
SBMNH 423069 (previously SBMNH 40612), a large cluster of commensal acorn
barnacles was seen, on bare-axis portions of the branches. Another specimen, SBMNH
423078, had attached to its bare axis something having, in general appearance, the
wooly, cotton-like spittle-bug mass that insects are known to produce on plant stems.
Figure 36. Chromoplexaura marki, SBMNH 423062, light microscopy array of sclerites, 10× magnica-
tion. Two sclerites, marked with arrows, are those that dene the species (compare with those shown in
Figure 34). e large central sclerite, just to left of image center, is > 100 µm in length, while non-capped
spindles measure 100–130 µm.
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Figure 37. Chromoplexaura marki, SEM images. A SBMNH 423061 A1 Double dunce-cap forms A2
Slightly smaller double dunce-cap forms A3 Coenenchymal sclerites A4 Anthocodial form B SBMNH
423072 B1 Double dunce-cap sclerites B2 Elongated spindles from coenenchyme B3 Coenenchymal
sclerites. Images match variety shown in Williams 2013a (gs 14–17).
Conspicuous brittle stars are intertwined on branches on the specimen from o New-
port, Oregon, SBMNH 423073.
Remarks. Cordeiro et al. (2018f) lists Chromoplexaura marki as the only species in
the genus Chromoplexaura in the WoRMS Database.
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Figure 38. Chromoplexaura marki, SEM images, representative forms. SBMNH 265937. A Anthocodial
form B Typical double-dunce-cap C Smaller (developing?) double dunce-cap D–E Coenenchymal scler-
ites. Images match variety shown in Williams 2013a (gs 14–17).
Kükenthal (1913a) had initially suggested that this species may equal Psammogorgia
arbuscula (Nutting, 1909) but later stated that characteristics of this species were com-
pletely dierent. e separation of these two species is reected in Kükenthal (1924),
with separate descriptions for E. marki and P. arbuscula (syn. Echinogorgia arbuscula).
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Bayer (1956a), in his description of the two genera in question, Euplexaura Verrill, 1869
(colony in one plane) and Psammogorgia Verrill, 1868 (colony bushy), indicated some
slight overlap.
Kükenthal (1913a; 1924) noted that prior to the discovery of this species, other
species in the genus (Euplexaura) had only been found in the area of East Asia, from
Japan to Singapore and West Australia. It appeared that this was the rst red-colored
member seen in Euplexaura and was the rst of the genus from the west coast of
North America.
In overall colony shape, some branching occurs; however, more often colony is a
single, or rarely branched, stem; any branching from base results in a single or very
scarcely branched “whip.” is would have been unique to this eastern Pacic spe-
cies, along with its obvious, predominantly red sclerites, if it were truly in the genus
Euplexaura (in most species of the genus, the sclerites are colorless); hence the need for
the establishment of the new genus by Williams (2013a). In general colony color and
shape, it would be quite easy to simply assume that this organism is Leptogorgia chilen-
sis, but an examination of sclerites reveals the distinct dierences.
Cairns et al. (2003) had this species listed as a junior synonym of Leptogorgia caryi
Verrill, 1868; as noted previously, this is not the case. According to unpublished notes
by Bayer, C. (E.) marki might have been synonymous with Psammogorgia spauldingi
(now referred to as Swiftia spauldingi). A possible synonymy was considered, with
both Swiftia spauldingi (Nutting, 1909), and/or Swiftia simplex (Nutting, 1909). After
examinations of multiple samples of what has been labeled as this species and those
labeled as S. spauldingi or S. simplex, if any synonymy were to exist, it would be that
between C. (E.) marki and S. spauldingi. With the very obvious large, broad spindles,
the double-dunce sclerite, I consider this a separate species, but it does exhibit a strong
supercial similarity to S. spauldingi and there are some shared sclerite forms. An initial
conclusion arrived at some years ago (regarding synonymy with S. spaulding), seemed
to have support with the discovery of a comment made by Bayer (1979). While the
statement was an unexpected one to nd in this particular article, nding it was note-
worthy. It read “e colonies of A(delogorgia) telones are similar in general aspect to
those of Euplexaura marki Kükenkthal (= Psammogorgia abuscula sensu Nutting, not
Verrill) and the closely related (if not identical) Psammogorgia spauldingi Nutting, both
of which have longer and less sinuous branches.” However, no anthocodial rods in the
form of a ngerbiscuit, a key characteristic sclerite of species in the genus Swiftia Du-
chassaing & Michelotti, 1864, have been found in C. (E.) marki specimens, and the
initial conclusion was dismissed. A further discussion of California “red whip” diversity
follows below and correlations are discussed in Part III of this collection review, on
Swiftia cf. spauldingi, but also in the description for Swiftia simplex.
An examination of several specimens (collected by P Etnoyer on a West Coast
Survey for NOAA, in the Fall of 2010) was done at Etnoyer’s request. Made available
were actual specimens, along with several in situ shots. In digital images, the little-
branched colonies were a dirty brick-red or pink (Figure 28); coloring was seen in
both extended polyps and throughout branch coenenchyme. An initial diagnosis of the
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specimens via still images was Swiftia simplex. However, upon physical examination,
the polyps themselves were actually white (indicating that only the tentacles were the
pinkish color of the coenenchyme) and an examination of the sclerites revealed the
large, broad double-dunce spindles so characteristic of C. (E.) marki. If one were to see
a colony with little branching, having an overall dirty brick-red to pink color, and did
not dissect out a polyp to reveal their white color, or examine the sclerites, an errone-
ous identication could be made. ese specimens presented something of a quandary.
Supercially, they looked very much like conrmed Swiftia simplex, yet the sclerites
revealed something dierent. ere is a possibility that C. (E.) marki has color variants,
with one looking very much like Swiftia simplex. us, specimens previously identied
in various museum collections as C. (E.) marki may not belong to the genus Chromopl-
exaura at all if double-dunce sclerites are not found, but nger-biscuit rods are.
MBARI has encountered many single or few-branched whips in their investiga-
tions. Many of these specimens have been recorded in video and in still photography; a
few have actually been collected. A number of principal investigators identify many of
these distinct whips as being this species (in genus Euplexaura, now Chromoplexaura).
However, some of those identied as this species may actually be Swiftia simplex; in
overall shape very comparable, but in S. simplex, the color leans to a dull brick red
rather than the usual bright red hue, and polyps of S. simplex are always the same color
as the coenenchyme, not the “. . . .white, cream or yellow” polyps described for this
species by Kükenthal (1913a, 1924), Johnson and Snook (1927) or Williams (2013).
Collection of an array of these “whips” when encountered, along with examination of
their sclerites and molecular testing of tissue could help to clear up any confusion sur-
rounding these red whip species; in collaboration with E Berntson, M Everett and their
colleagues at Northwest Fisheries Science Center (Port Orchard and Seattle, Washing-
ton), those needed examinations are currently being conducted.
Discussion concerning diversity of “red whip” forms
From a morphological perspective, with numerous sclerite preparations having been
conducted by myself and my research students, it is reasonable to discuss the “red
whip” Chromoplaxaura marki, and its high degree of variability that could mistakenly
lead investigators to name it something else (especially true if colony samples are not
taken and/or no sclerite examinations are done). “Red whip” gorgonian species within
the California Bight (and in geographic areas immediately adjacent, north- and south-
wards) present challenges. In the SBMNH collection were found a number of “red
whip” specimens, identied by earlier investigators, as the widely common Leptogorgia
chilensis, but whose sclerites and/or colony form were not that of L. chilensis. L. chilensis
appears as a predominantly thin-branched, but none-the-less, highly branched spe-
cies, usually seen as a broad fan. Based on location records found for specimens at the
NMNH, CAS, etc., it appears that L. chilensis is the major “red” species from approxi-
mately Central California southward, primarily inhabiting the subtidal zone, often
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sharing space with Eugorgia rubens and both Muricea californica Aurivillius, 1931 and
Muricea fruticosa Verrill, 1868a. (See distribution map for “red whips,” Appendix2:
Map A1 and Appendix 3: Table of characteristics, Table A1. For comparison, distribu-
tion of another species of Leptogorgia, L. exilis, which could be mistaken for L. chilen-
sis, is shown along with distribution range for several of the whip-like species from the
genus Swiftia.) e other specimens of “red whip” in the SBMNH collection, usually
(but not always) appeared as less branched or completely unbranched, slender, but not
thin, colonies with very dierent arrays of sclerites. Locality data indicated their ap-
pearance as being far more common in waters of the northern Central California coast,
extending northwards, slightly overlapping the range of L. chilensis, but usually from
waters of greater depth. Potential species would thus be: Chromoplexaura (Euplexaura)
marki, Swiftia simplex, or Swiftia spauldingi.
Within this group of predominantly California-collected “red whip” specimens,
further subgroupings could be made (based on the appearance of the sclerites): Sub-
group 1, “red whip” forms that closely resembled examples of colony variation in Chro-
moplexaura (E.) marki, but which might be specimens of Swiftia spauldingi and a few
indeterminate specimens, looking far more like S. spauldingi than anything else. ese
few specimens are colonies with shorter, chunkier, more heavily warted spindles, along
the lines of those from the specimen at NMNH (USNM 78385), labeled as Swiftia
spauldingi, collected in Monterey Bay. Further collection eorts are required, particu-
larly in the transitional area of California’s Point Conception, so as to document ex-
tent of occurrence in and near the California Bight. e notion of cryptic species or
transitional endemics may apply here, not an unusual occurrence in a region like the
California Bight; an equally likely possibility may be that of hybridization between
certain species.
Other remaining small, odd whip-like colonies (in ve separate lots), SBMNH
265946, 265947, 265948, and 265949, comprising Subgroup 2, bore a stronger su-
percial resemblance to a specimen collected by J Ljubenkov (which he misidentied
as Muricella complanata Wright & Studer, 1889, and as yet, are still not conrmed as
to their identication). ese are the most dicult to link with a known genus or spe-
cies. Most of these very small fragments are a bright, vivid red (or reddish pink), and
have noticeably white polyps. At this point, they most closely t very small specimen
examples of the morphological variety seen in C. marki. But, the sclerites generally did
not closely match those seen in known “red whip” species. ere are few to none of the
double dunce-caps of C. marki, the simple spindles of L. chilensis, or the spindles and
rods that can typically be seen in any species from the genus Swiftia (however, Swiftia
kofoidi (Nutting, 1909) does not always display the denitive rod-shaped sclerite).
Unfortunately, compounding the problem is the overall condition of the specimens;
they are very small, without much coenenchymal tissue. ere is now so little mate-
rial to work with that without collecting more material, continued work with these
specimens would destroy what is left; none of these are identied to my satisfaction.
Without more material, collected in the same areas as these lots, I cannot proceed
further with a conclusive identication (see Appendix 1: List of material examined.)
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Two additional lots (SBMNH 423066 and 423067) presented with sclerites some-
what intermediate between that seen in those labeled as L. chilensis and the above men-
tioned subgroups. Location data indicated the possibility of them being examples of L.
chilensis, but sclerites did not t as cleanly as would have been expected. I considered
the possibility of these specimens being part of an array of cryptic species (within one
or more of these genera and species) or that these gorgonians are either transitional
endemic or hybridized forms. Currently, these are listed as material examined labeled
L. chilensis, but further examinations may alter that placement. Based on location re-
cords, each of these several specimens may demonstrate a need for establishing dif-
ferent colony types as dierent subspecies (cryptic species?), but could equally show
degree of overlap (and possible morphological transitions) of various known species.
An attempt has been made to show key characteristics of each (refer to Appendix 3:
Table A1). Overlap of these gorgonians conrmed my understanding of the California
Bight as being an area of tremendous diversity (and as a result, an area where confusion
regarding very similar-looking forms can occur), an area understood ecologically as an
environmental disjunction. is overlap supports the admonition to anyone working
in this geographical area, encountering these gorgonians, to collect and examine mul-
tiple specimens and to do extensive sclerite preparations in those comparisons. Further
collection eorts will be helpful in documenting the extent of occurrence of each of
these species, not only in the California Bight, but in areas adjacent to it, both north
and south, with nal determination likely coming from molecular comparison studies,
which could conrm such concepts as cryptic species or regional (Point Conception)
transitional endemics. Somewhat fortuitously, an interesting and pertinent statement
was discovered in the introduction to the rst volume of “New Zealand Inventory of
Biodiversity: Volume I: Kingdom Animalia,” edited by Dennis P Gordon (2009). To
quote: “Often, what we think is a single species turns out to be a complex of closely
related cryptic (hidden) species that resemble each other so closely their existence had
been overlooked.” is may well be the situation for “odd red whips” and several spe-
cies of Swiftia (along with other genera) in the eastern Pacic. Appendix 2: Map A1
shows the distributional ranges and geographical overlap of known species and possible
endemic intermediates.
Pertinent specimens at CAS were examined. For a substantial gorgonian collec-
tion of eastern North Pacic Ocean species, CAS has taken the correct stance on “red
whips” from California and parts north; twenty four out of twenty ve lots (identied
as Euplexaura marki, now Chromoplexaura marki) are from Morro Bay and areas north,
with most from Monterey Bay and Oregon. According to the database, they do not
have specimens of either Swiftia simplex or Swiftia spauldingi; three specimens are listed
as Euplexaura simplex (genus name usage not seen elsewhere, erroneously proposed by
Harden). Material from several other sources (NOAA, NMFS, MBARI, etc.) were
examined to broaden the scope of my understanding of eastern Pacic “red whips”.
ere was little doubt that “red whips” seen by MBARI would likely not be L. chilensis.
at would be the case for two reasons: depth at which colonies are seen, and location
where MBARI researchers are doing their work (well north of the California Bight). A
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128
dry specimen of Chromoplexaura (Euplexaura) marki collected in Monterey, California,
housed in the Smithsonian collection (USNM 51500), was of interest. It displayed
sclerites very dierent from what would be seen in L. chilensis, but also did not clearly
display the key sclerite form for the genus Chromoplexaura, the large double dunce-
cap sclerite. Of further interest was the fact that NMNH listed only three specimens
of Leptogorgia chilensis, all from La Jolla, San Diego or southern California, but had
two lots of Lophogorgia chilensis, taken o the Washington coast! ese were collected
in 2006 at depths of 84 and 232 meters, on that year’s NOAA “Deep Sea Coral and
Sponge Habitat Expedition.” ese latter two are more likely either Swiftia spauldingi
or one of the other species mentioned above.
Genus Muricea Lamouroux, 1821
Muricea (pars) Lamouroux, 1821: 36. (pars) Blainville 1834: 509. (pars) Ehrenberg
1834: 134. Dana 1846: 673. (pars) Milne Edwards 1857: 142. Duchassaing and
Michelotti 1864: 14. (pars) Kölliker 1865: 135. (pars) Verrill 1868b: 418–419;
1868c: 411; 1869a: 449–450. Kent 1870: 84, pl 41, gs 13–17, (?) 36–37. (pars)
Studer 1878–1879: 649. Studer 1887: 58. Wright and Studer 1889: 93, 133 + pl.
Gorzawsky 1908: 8. Nutting 1910a: 9. Kükenthal 1919: 752, 835; 1924: 141.
(pars) Riess 1929: 383–384. Aurivillius 1931: 102–103. Deichmann 1936: 99.
Bayer 1956a: F210; 1959a: 12; 1961: 179–180. Tixier-Durivault 1969; 1970a, b,
c: 154. Bayer 1981c: 930 [in key only]; 1994: 23–24. Marques and Castro 1995:
162. Hardee and Wicksten 1996: 127–128. Castro et al. 2010: 779. Breedy and
Guzmán 2015: 6–7; 2016b: 7–8.
Emuricea (pars) Verrill, 1869a: 449. Studer 1887: 58. Wright and Studer 1889: pl LVI.
Nutting 1909: 718. omson and Simpson 1909: 258. Kükenthal 1919: 836.
Reiss 1919: 397–398. Kükenthal 1924: 149–150. omson 1927: 48–49. Reiss
1929: 397. Aurivillius 1931: 50 (emended). Deichmann 1936: 104.
Eumuricea (Muricea) Bayer, 1981: 930 (key). Breedy and Guzman 2015: abstract, 28.
Type species. Muricea spicifera Lamouroux, 1821, by subsequent designation Milne
Edwards and Haime 1850: lxxx. Kükenthal (1924) listed Muricea muricata (Pallas,
1766) as the type. [M. specifera was later synonymized with Muricea muricata Pallas,
1766 apud Bayer, 1961: 179–180]
Diagnosis. Arborescent colonies richly branched (dichotomously or laterally), of-
ten in one plane. Branch diameter moderate to very thick, tendency to curve upwards,
most nearly parallel to one another, tips tending to slightly swollen. Calyces shelf-
like, on all sides, close-set, prickly, tubular or distinctly projecting (at right angle or
upwards); stiened by large, fusiform sclerites; aperture wide and eight-rayed; polyps
fully retractile. Axis purely horny; weakly loculated (if at all). Sclerites usually fusiform,
long, often massive, spindles (up to 3.0 mm in length), obviously sculptured, with
strong outer or terminal spines, or both, arranged in calycular wall longitudinally;
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 129
rarely some irregular forms. Anthocodial sclerites numerous, small spindles, forming
at most weak, slightly dierentiated transverse crown or collaret below tentacles, con-
verging on bases of tentacles. Sclerites in genus stated as generally, markedly stockier,
denser and thicker; a bit larger overall, than those seen in many other genera.
Remarks. e genus Muricea may contain at least a dozen species specically
found in the eastern Pacic; however, species descriptions, and their potential syn-
onymies, needed review. e work of Breedy and Guzmán (2015, 2016a, 2016b) has
been of help. But, the California Bight is a complex area in the eastern Pacic, and may
hold some surprises with regards to this genus. To date two, perhaps three, species are
commonly recorded from the California Bight; others, however, occasionally may ap-
pear. A more extensive and thorough discussion of this genus as it appears specically
in the California Bight may be required. A further investigation is also necessitated by
the fact that the number of species of Mexican Muricea, in both the SBMNH collec-
tion and other collections, is far greater than the number of species currently known
to occur within the California Bight. While the review of the genus Muricea and its
species by Breedy and Guzmán (2015, 2016a, 2106b) is helpful, further investigation
of Muricea found in the eastern Pacic waters of California and upper Baja, Mexico is
still required; specimens from the SBMNH and LACoMNH will be helpful in such
investigations.
Muricea californica Aurivillius, 1931
Figures 39A, B, 40A–D, 41A–C
? Gorgonia plantaginea Valenciennes, 1846: pl 15 (non Lamarck).
? Muricea appressa Verrill, 1864: 37; 1866b: 329; 1869a: 44. Grigg 1970: xiv, 20, 25,
207; 1977: 280.)
? Muricea appressa avescens Verrill, 1868a; 1869a: 446 (? nec Verrill, 1864: 37).
Muricea californica Aurivillius, 1931: 111–114, g, p 113. Hardee and Wicksten 1996:
130–132. Breedy and Guzmán 2016b: 32–34.
Type locality. North Pacic Ocean, California Channel Islands, Santa Catalina Island,
Gulf of Santa Catalina, 2–27 m.
Type specimens. Syntype USNM 44188 [wet]; Lectotype USA: Swedish Mu-
seum of Natural History 1122 [wet]. Syntype specimen was examined; a common
form in California waters, often easily identied.
Material examined. ~32 lots (see Appendix 1: List of material examined).
Description. Colony (Figure 39A) non-reticulate; up to 100 cm wide, 120 cm high,
usually 60 cm or less. Loose, dichotomous, irregular branching primarily in one plane,
forming heavy fan-shaped colony; some primary and secondary branches extend out of
plane. Branches thick, averaging 2.0–5.0 mm in width; curve to lie parallel with main
branch. Branching lateral, terminals of even thickness or tapering slightly. Outer coe-
nenchyme mostly occluded by calyces (Figure 39B). Calyces distally open cups, erect,
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Figure 39. Muricea californica, SBMNH 422921. A Whole colony, 25.5 cm × 15 cm B View of branch
tip magnied to show prominent, somewhat rounded calyces.
very elongated, prominent, conical in shape, 0.8–1.1 mm tall, 1.0–1.5 mm across, 1.0–
2.0 mm apart (close together, but not overlapping), protruding 45 to 90 degrees away
from branch when polyp extended (extendable to 3.0 mm). When polyp not extended,
calyx lying close to and curving into stem, broad and smooth (like bracts in a partially
closed pine cone). Upper lip varies, from those without sclerites to having denite lip.
Calyces extend in all directions around branches. Tentacles taper at tip; bear two rows of
lateral pinnules that are slightly displaced to the oral side. Color of living colony gener-
ally rusty brown; ranges from golden-brown to dark reddish orange to reddish brown
to brown to dark brown. Axis reddish brown at base; becomes light yellow-brown at
tips. Polyps most commonly yellow; golden orange, bright yellow, pale yellow, creamy
yellow, even white; all polyps of a branch the same color. Possibility that more than one
color of polyp per colony occurs (demonstrated in digital images sent to me by Mary
Wicksten, and examination of specimens). Dry colonies dark rusty brown. Sclerites
(Figure 40A–D) rust red to golden-brown. Sclerites predominantly club-shaped with
large, rounded spines or pointed tubercles projecting from broader, club-shaped end;
other end tapering, covered with tubercles (Figure41C). Outer coenenchyme consist-
ing of small sclerites, to 0.5 mm long, torch-like, with processes often continuing ir-
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 131
Figure 40. Muricea californica. A–C SBMNH 422911, SBMNH 422914, and SBMNH 422361, re-
spectively; light microscopy arrays, 4× magnication, showing diverse mix of sclerites seen in the species
D Light microscopy array, the “torch-like,” multi-toothed sclerites so typical of M. californica, shown at
10× magnication. In D the middle, upside-down torch measures some 580 µm, the middle upright torch
measures ~400 µm, while the torch lower down measures 480 µm.
regularly down one side, other end tapering, covered with tubercles. Some spindles
fusiform, bent very slightly in middle, or having large processes in middle projecting
outward. Inner coenenchyme spindles small, fusiform, set with distinct tubercles.
Sclerites examined compared to those shown by Hardee and Wicksten (1996);
there can be dense coverings of warts, but condition not seen on all sclerites; some
sclerites have dense covering of warts at one end, but not at other end. On largest scle-
rites, warts are large, very few a bit bigger than those shown by Hardee and Wicksten
(1996) for M. fruticosa. Many more of largest sclerites have ame-like teeth at one end,
not down entire length of one side. In drawings and photographs examined of those
believed to be, or labeled, this species, much larger tubercle bumps are seen. Regarding
ame-like teeth, on some sclerites there are scattered, randomly-placed spines running
down the side, but many more have ame-like teeth only coming o one end (reminis-
cent of a aming torch; Figure 41C). Surface bumps can be dense, but not always, not
over whole surface (true of some). For this species, stocky, dense clubs and torches very
evident; overall, sclerites give impression of being a bit larger and more densely warted
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132
than those that may be from Muricea appressa. Refer to Figure 42, shown as Muricea
plantaginea (Valenciennes, 1846); largest sclerites, however, not nearly as large as those
seen in M. fruticosa.
Etymology. Surmise that the name californica refers to the species type locality; no
explanation for the species derivation was found.
Figure 41. Muricea californica, SBMNH 265938, SEM image. A, B Axial sheath sclerites A Very jag-
ged spindles B Moderately “toothed” spindles C Calycular/coenenchymal torch-type spindles. Compare/
contrast with those shown in Breedy and Guzmán 2016 (g. 21).
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 133
Common name. California golden gorgonian; California’s purple one; California
rust gorgonian; Brown sea fan (so called in a variety of eld/diving guides).
Distribution. Stated as ranging from Point Conception, through Baja (Santa Ma-
ria) to the Gulf of California; range extending further south is possible. Grigg (1977)
remarked that if M. californica is identical to M. appressa (See Harden 1969; Grigg
1970 for rationale), southern limit for this species is then Zorritos, Peru. From the
following list of California sites, running from south to north, note depth ranges indi-
cated. On the Mainland: Point Loma: 5–18 m; La Jolla: 3–27 m (USNM 50190, was
collected at 11 m, in La Jolla, at the Torrey Pines kelp bed, 5 miles north of Scripps
Institution); USNM 77286 was collected at Corona Del Mar, Newport Bay; Newport
Beach: 2–12 m; San Pedro: 3–18 m; yet another at NMNH (USNM 52485) was col-
lected along the southern coast. Islands: Coronados Islands: 5–24 m; Santa Catalina
Island: 5–30 m.
Biology. Common in kelp beds (Ricketts, 4th Ed. 1968); found at depths greater
than 3 m, perhaps being one of the most common gorgonians from southern Califor-
nia (Hardee and Wicksten 1996). Grigg (1977) noted it as a species seen in the rocky
sublittoral zone o California at depths between one and 30 meters.
As colonies grow, they form annual growth rings in the skeleton (Grigg 1974;
Bertsch 1984). Colonies grow separately sexed, requiring 5–10 years to reach sexual
maturity with a maximum longevity of ca. 50 years (Grigg 1975). Grigg (1977) indi-
cated that mortality could be caused by abrasion from suspended particulate matter
when there are high waves, smothering by sand and by encrustation due to presence of
zoanthid cnidarians Savalia lucica (Cutress & Pequenat, 1960) and Epizoanthus indu-
ratum Cutress & Pequenat, 1960. On specimens of Muricea (potentially this species)
located on Shale Reef, between Corona del Mar and Laguna Beach, Savalia lucica
was found (a wet SBMNH specimen, as yet uncataloged). As well, SBMNH 422359
has a heavy growth of some form of epizoanthid. Presence of the colonial anemone
Corynactis californica (Carlgren, 1936) may also be an important source of mortality.
Satterlie and Case (1978) used this species extensively in studies on the neurobi-
ology of gorgonian coelenterates, which they obtained locally (Santa Barbara area, at
depths of 4–11 m). In a study done by Lissner and Dorsey (1986), it was noted that
while this species was common around the Channel Islands and rocky areas of the
mainland, it was conspicuously absent on the Tanner and Cortes Banks, and on the
Santa Rosa-Cortes Ridge. In an anecdotal notation made by R Grigg, the reason could
be that populations of Muricea species may be limited by cold water and/or poor dis-
persal abilities of the larvae. Grigg (1977) stated that Muricea species rarely cover more
than 1% of the sea bed, where space is fully occupied.
Considering associations this species has with other organisms, Humes and Lewbel
(1977) reported two species of Acanthomolgus Humes & Stock, 1972 (cyclopoid cope-
pods) for the rst time from the eastern Pacic in association with this gorgonian, from
an area near La Jolla, California (Quast Reef). ey indicated that these copepods are
consistent members of the epifaunal community on the gorgonian, found with the gor-
gonian throughout the year. In association with SIO/BIC #CO 1600, there appeared
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134
to be the exoskeletal remains of either skeleton shrimp, or some other small, (and now
pale) crustacean. Notes from H Bertsch (1984) indicated that sometimes the ovulid
snail Simnia (Neosimnia) vidleri (GB Sowerby III, 1881) could be found eating this
gorgonian; Grigg (1974) reported that the only sh known to feed on Muricea species
is the Garibaldi Hypsypops rubicunda Girard, 1854 (from Clarke 1970). A specimen,
from Baja, South Bay, Isla Cedros (SBMNH 422363), had very well developed acorn
barnacle galls, completely overgrown and covered by healthy-looking coenenchyme;
only the barnacles’ uppermost valves were visible. ey appeared to have been well
protected and secured to their gorgonian host. One additional specimen (SBMNH
422362) displays a bare axis at the tip of several branches; obvious barnacles are at-
tached to these bare-tip sites. Overall, extensive organismal growth is uncommon on
Muricea specimens, both wet and dry, housed in the SBMNH collection.
Remarks. Conicting comments about this species (comparing it with Muricea
fruticosa Verrill, 1868a) have been made, particularly with regards to polyp color (Rick-
etts 3rd Ed, 1962; Ricketts et al. 5th Ed, 1985; Harden MS thesis 1969; unpublished
pencil notation, Harden; Harden PhD dissertation 1979). In reading these it was clear
to me that some identied the species by presence of yellow polyps, while others did
so by white ones. ere is no doubt that discrepancies regarding polyp color have
carried into current identication of the species. It would be well to remember that
color is hard to determine in underwater situations, in ambient light conditions, or
with articial light sources. In some of the oldest descriptions, polyp color was not
always clearly stated, most likely due to collection procedures and gaps of time that
ensued between collection and actual examination. ere is evidence, based on my
own examinations, that M. californica can have yellow or white polyps (even dier-
ent colored polyps in dierent locations within the same colony). Conrmation came
via e-mail correspondence with M Wicksten, and is stated in Hardee and Wicksten
(1996: 129, 138). Divers comment that often colonies closely situated side-by-side,
with basically the same colony form (thickly branched, usually in one plane), with the
distinctively common brown coenenchyme, can display markedly yellow polyps in one
colony and in the other (closely adjacent) one, obvious white ones. Because the colony
form is so similar (perhaps due solely to current ow in the immediate area, thus an
environmental condition), but polyp color so dierent, the two are considered (and
identied) as being dierent species, that with yellow polyps, M. californica and those
with white polyps, M. fruticosa. Yet, there is the question as to whether polyp color
can be an accurate means of identication in situ, especially under low light condi-
tions, or if color variation is genetically inherent. Questions that still require further
study: 1) are colonies (yellow polyps vs. white polyps) two distinct species? 2) If so, are
the two species so similar that they can hybridize? Or, 3) do two color morphs of this
species exist (in southern California, at least, each with the characteristic large, planar,
generally dichotomously branched colony shape)? Only an examination of sclerites,
with notation made of polyp color at the time of collection, could clearly answer these
questions. Channel Islands National Park, in annual sh surveys, identies M. fruticosa
as that with white polyps, while M. californica has yellow polyps. e Aquarium of
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 135
the Pacic in Long Beach (via phone communication with P Clarkson, March 2003)
identies them in the same way. Unfortunately, most specimens originally identied
as this species in the SBMNH collection had either sat in formalin since initial collec-
tion, had been in alcohol quite a long time, or are extremely dry; there is no color to
be seen in the polyps (even if at one time they had color) and data labels usually do
not indicate polyp color at time of collection. Fresh material has been requested from
local sources, as they incidentally collect, to help clarify identication of this, and very
similar looking species.
Allen (1976) stated that M. californica had been common at Corona del Mar in
the past, and D Kushner from Channel Islands National Park (phone comm., March
2003) stated that there has been an enhanced abundance of M. californica observed
in recent years at San Onofre, almost to the point of taking over, becoming quite
abundant in the area. is is an area where I would expect to nd this species, as it
appears to be common in southern California, extending into Baja, but this growth
spurt seemed a somewhat unusual event. e small collection of dry specimens (and of
living specimens on public display) at the San Pedro Cabrillo Marine Aquarium holds
many colonies identied as this species, collected in the surrounding local area and o
the southern California Channel Islands.
Coloring of actual sclerites was often of little help, but observation of sclerite size
and shape was crucial. Some of the specimens identied originally as M. californica
(and for that matter, some identied as M. appressa) were actually the typical vari-
ant of M. fruticosa, but because of condition of colony, method of preservation and
transitional sclerite forms, it was initially dicult to clearly separate the species; calyx
appearance was used as well, but again, groupings into distinct clusters was not always
easy/clear. ere was inclination to think that some specimens, listed as M. californica
(likely because of overall colony shape), but from far more southern (Mexican or even
Central American) collection locations, might actually be M. appressa or M. fruticosa.
Without fresh material (having specic data location, observation of polyp color re-
corded, along with more extensive and careful extraction and preparation of the large
and jagged sclerites, so as not to break them or break o the teeth), can these Muricea
specimens be condently assigned a species name. As Muricea californica seems the
most common form, those listed in the Appendix 1 (List of material examined) are
listed as M. californica, unless it was very clear, based on calyx appearance and scler-
ite form/size, that they were another species. Examination of recent, locally collected
colonies (SBMNH Sea Center, Aquarium of the Pacic, Long Beach and OCSD),
while not numerous in quantity or large in size, was invaluable. Sclerite size and shape
were noted, along with calyx shape and conrmed polyp color. It would appear that
M. californica has a wide range of polyp coloring (rarely white to commonly yellow
or rich gold), while its sclerites are distinct in not having the large, densely warted,
rounded spindles common to M. fruticosa (which only displays white polyps). In situ
identication, therefore, can be challenging if the colony in question has white polyps;
both M. californica (occasionally) and M. fruticosa (typically) have white polyps, while
M. appressa (as M. plantaginea, included briey below, for comparison) seems to always
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136
display yellow polyps, (which is normal for M. californica). Despite the challenges re-
garding in situ identication, M. californica (as well as M. fruticosa and M. appressa (=
M. plantaginea) are all recognized as separate species in the WoRMS Database (Cord-
eiro et al. 2018g).
Muricea plantaginea (Valenciennes, 1846)
Figure 42A–K
Gorgonia plantaginea Valenciennes, 1846: pl 15.
nec Gorgonia plantaginea Lamarck, 1815: 163.
nec Eunicea plantaginea Valenciennes, 1855: 13; Milne Edwards and Haime 1857:
146, 151.
Eunicea tabogenesis Duchassaing & Michelotti, 1860: 17; 1864 [1866]: 111. Kü-
kenthal 1924: 145.
Eunicea ransoni Stiasny, 1937: 331, 334–336, gs 5, 6, 7.
? Muricea appressa Verrill, 1864: 37 [January]; 1866: 329; 1868a: 412; 1869a: 444–
446; pl VIII, g. 13. Kükenthal 1919: 752; 1924: 145. Reiss 1929: 390–391.
Hardee and Wicksten 1996: 132–136 (syn. n).
Muricea appressa var. avescens Verrill, 1869a: 446. Kükenthal 1919: 752; Kükenthal
1924: 145 (syn. nov.). Hickson 1928: 371–372. Reiss 1929: 389–390. Stiasny
1943: 72–74.
Muricea plantaginea Lamarck, 1836: 333. Breedy and Guzmán 2016b: 25–32.
Muricea californica Aurivillius, 1931: 111–114 [according to Grigg, 1977: 280, after
Grigg, 1970: xiv, 20, 25, 207].
Muricea tenella Verrill, 1869a: 446–448. Kükenthal 1919: 752; 1924: 145. Hickson
1928: 371–372. Reiss 1929: 389–390. Stiasny 1943: 72–74.
Type locality. Holotype Mazatlán, Mexico, Voyage sur la Frégate La Vénus, MA Du
Petit ouars, 1836–1839. Also, Peru, Tumbes Department, Zorritos, 3–5 fm [6–9
m]. Specimen from NMNH (USNM 33585, and many others) collected in the North
Pacic, Panama.
Type specimens. Syntype YPM 1616A of Muricea appressa var. avescens. As well,
housed at NMNH, USNM 33585, listed as a Syntype, with SEM image #2517; [dry].
Syntype specimen at NMNH was examined.
Material examined. A number of lots housed in SBMNH collection (see Appen-
dix 1: List of material examined).
Remarks. Included here is a brief commentary on this species, and an SEM plate
(Figure 42A–K) is provided as a means of comparison, because as Grigg (1977) stated
“this could be synonymous with M. californica.” However, the work of Hardee and
Wicksten (1996) led them to conclude that M appressa (= M. plantaginea) is not syn-
onymous with M. californica. Based on my own observations and research, M. ap-
pressa is more likely to be found south of the California Bight (Baja, CA Sur, Ecuador,
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 137
Figure 42. Muricea plantaginea (= Muricea appressa), SBMNH 422909 (Ecuador) and SBMNH 422417
(Baja CA Sur). Image included for comparison between sclerites found in M. californica and those seen
in M. plataginea [= M. appressa Breedy & Guzmán, 2016]. SEM image. A–F SBMNH 422909 (A–C);
E–F Calycular/coenenchymal sclerites A Elongated, jagged spindles B Small torch-type C Rounded,
stout torches D Shorter, jagged spindles (axial sheath) E Unilateral spinous spindle F A second unilat-
eral spinous form tending to torch shape G–K SBMNH 422417 Calycular and coenenchymal sclerites
GFan-shaped sclerite H Large, truncated unilateral spinous spindles I Stout prickly spindle J Rounded,
unilateral spinous type K Torch-types typical of M. californica. Compare sclerites A–F with those shown
in Breedy and Guzmán 2016 (g. 15, for M. plantaginea) and sclerites G–K with those shown in Breedy
and Guzmán 2016 (g. 21, for M. californica).
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138
Galápagos, etc.). Johnson and Snook (1927) made mention of storm-washed living
specimens of Eunicea Lamouroux, 1816 (but no reference to a species) with the liv-
ing polyps yellowish white (the black and white photograph of a specimen shown in
that volume looked most like a somewhat worn specimen of either M. californica, or
perhaps M. fruticosa typical). While those identied as M. appressa in the SBMNH col-
lection generally seemed more prickly in overall appearance (as compared to M. cali-
fornica), along with slightly smaller-diameter branches, any cursory visual inspection
of gorgonian specimens from this genus could misidentify species. A more intentional
study of calyx shape along with further comparisons of sclerites from freshly collected
specimens over the total range of the Bight to clarify the possible synonymy of this spe-
cies with Muricea californica is underway. I am inclined to keep Muricea appressa (= M.
plantaginea) a separate species while this further study is being conducted.
Several additional locations were noted for this species in Verrill’s (1864; 1869a)
description: Panamá and the Pearl Islands, in pools at extreme low water; ex. FH Brad-
ley; also, JH Sternbergh and FH Bradley. Also, records from Nicaragua, Corinto; ex.
JA McNiel and from Mexico, La Paz; ex. J Pedersen. Note that all of these locations lie
well south of the California Bight’s southern boundary.
Compounding the confusion surrounding Muricea species, particularly in the
southern portion of California’s geographical range, is that in the following description
of Muricea fruticosa, two very distinct colony forms must be mentioned: that which
looks very much like the typical Muricea californica (the typical colony shape, albeit
with white polyps, according to most encountering it in the eld) and that with a far
smaller, stier, shorter-branched cespitose or fruticose bushy shape, a distinctly dier-
ent variant of M. fruticosa according to Verrill (1868a; 1869a). As this latter variant is
not encountered in the southern California Bight it is not discussed here.
Muricea fruticosa Verrill, 1868
Figures 43A, B, 44A, B, 45A–D, 46, 47A, B, 48A–E
Muricea fruticosa Verrill, 1868a; 1869a: 428–430; pl 7, g. 2. Kükenthal 1919: 752;
1924: 142–143. Hardee and Wicksten 1996: 129–130. Breedy and Guzmán
2016b: 9–14.
Muricea fruticosa var. typica Verrill, 1868a; 1869a: 428–430. Kükenthal 1924: 142.
Muricea fruticosa var. miser Verrill, 1869a: 430. Kükenthal 1919: 752; 1924: 143.
esea crosslandi Hickson, 1928: 354–356 (syn. nov.).
Pseudothesea crosslandi (Hickson, 1928). Stiasny 1943: 64–66 (syn. nov.)
Type locality. (Lectotype) Panama, Pearl Islands, 11–14 m.
Type specimens. Lectotype YPM 1574C [dry]; additionally, YPM 1660 [wet], YPM
1792 (fragment from Lectotype) [dry], and YPM 3067C [dry], are all listed as Syntypes.
ere are several specimens listed as Syntypes at NMNH: for instance, USNM 52292
[dry], with SEM images from stub #239. Syntype material at NMNH was examined.
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 139
Figure 43. Muricea fruticosa, SBMNH 422430 (as seen in California waters). Colony shows pale base
area tending to darker, colored branch tips. A Whole colony, 15 cm tall × 11.0–11.5 cm broad B Closer
view of branch tip showing conical calyces forming shelf-like projections o branch surface.
Material examined. ~14 lots (see Appendix 1: List of material examined).
Description. Colony (Figures 43A, 46) fairly large; dense, abundant branching; not
reticulate. Branching irregularly dichotomous (also seen as cespitose/fruticose, tightly
bush-shaped variant, with rather small, somewhat clavate branchlets outside California
Bight; variant description not discussed here). Prominent, spreading, spinose calyces
create rough texture to branches of colony. Colony (of Figure 46) very bushy; not in
one plane. Colony stands up to three feet (90 cm) tall (Figure 43A), but usually shorter
(30 cm). Main stem stout, short, arising from large, irregular base, usually dividing at
once into several large, thick, unequal main branches, rapidly dividing and subdividing
in irregular manner; branching extensive, such that main branches soon lost among
crowded, crooked secondary branches. Branches and branchlets usually not more than
7.0–12 mm apart; branching can be in one plane, but not always. Small branches near
ends often divide in irregular dichotomous manner, sometimes coalesce; very numer-
ous, nearly equal in size, usually distinctly curved and crooked, spreading out at origin
with a broad curve. Terminal branchlets short, 7.0–40 mm long, 2.0–3.0 mm thick,
often curved, not tapering, either ending evenly or clavately, with obtusely rounded
ends. In overall appearance, can look much like the colony shape of M. californica (per-
haps reective of similar environmental conditions). Color of living colony generally
brown (darker) with white polyps; mostly deep reddish orange, rusty-brown; branch-
let tips and calyces deep reddish brown, color generally fading to yellowish brown in
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Figure 44. Muricea fruticosa, SBMNH 265940, light microscopy arrays. A, B 4× magnication, with
diverse sclerites shown for the species. Of particular interest are prominent sclerites, indicated by arrows,
in B (the tardigrade-like form mentioned in text description). In B the lower, middle sclerite measures
some 1300 µm in length, the one up and to the right of that measures 1133 µm; thinner spindles measure
between 200–300 µm in length.
proximal portion of branchlets, fading into light yellow, tinged with brown in main
branches and trunk. Dry specimens orange-rusty brown, while polyps are pure white,
situated on upper side of calyces; near distal end, aperture lled with yellow polyp
sclerites, arising from bases of tentacles. Horn-like axis yellowish wood-brown at base
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 141
Figure 45. Muricea fruticosa, SBMNH 265940, SEM image. A, D Calycular and coenenchymal uni-
lateral spinous spindles B, C Axial sheath A Large nudibranch-type sclerites B Typical spindle C Smaller
spindles D tardigrade-type spindles.
and in larger branches (Hardee and Wicksten 1996 stated axis dark brown at base);
darker reddish brown, translucent in smaller branches; light amber-yellow, translucent,
slender up into branchlet tips. Calyces close together, but not overlapping, spread-
ing outward and upward, 45 degrees from branch when closed, nearly 90 degrees
with polyp extended; prominent, with conspicuous shelves opening distally, conical
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to columnar, larger and closer (1.0 mm) toward tips (Figure 43B), approximately as
high (1.0–2.0 mm) as they are broad (1.0–1.5 mm); on larger branches low, rounded,
without prolonged lower lip, better developed than at base where they are atter, small
and spread apart (2.0 mm). ose better developed have an obvious lower lip, sharp
and long with very large, long, stout sclerites as spindles (some of which approximately
as long as the calyces) which lie parallel to each other, projecting past upper margin of
lower lip, giving colony a prickly feel when touched; upper lip small or barely notice-
able. As calyces do not overlap, outer, thin coenenchyme easily seen lying between
calyces, characterized by extremely large, stout, sclerites, visible to naked eye, and curv-
ing around them, often larger than calyces; these sclerites may be missing near base of
colony or in poorly preserved specimens. Sclerites (Figure 44A, B) vary in color from
brownish yellow and yellowish white to deep reddish brown. Largest sclerites (of outer
coenenchyme), reddish orange-brown in color, up to 3.0 mm long; several shapes vis-
ible (Figures 44A, B, 45, 47A, B, 48A–E). One shape, very distinct, mostly stout, blunt
and truncate, almost rectangular; longer, large, massive ones rather thick in middle,
tapering somewhat abruptly at ends, densely, evenly covered by small tubercles (Ver-
rill 1868 stated longer ones covered by small, sharp spinules on one side; other parts
covered with crowded rough warts; these I refer to as tardigrade-like sclerites; Fig-
ures44A, B, 45D, 47B, 48E). Second largest form irregularly fusiform, covered with
tubercles; third form (Figure 48D) hook-shaped either on one end, both ends curving
inward, or one end forked, the other tapering to a point; the latter often covered on
ends with small, sparse, occasionally spiny tubercles, becoming more densely covered
with tubercles toward the center, usually found around base of calyces. Medium-sized
sclerites (~1.0 mm) more regular, fat (stout) in middle usually tapering to acute points;
one side or one end covered with quite large, very sharp, simple projections, other side
with densely crowded, rough microtuberculate warts. Sclerites of inner coenenchyme
distinctly smaller, color ranging from yellow to white, fusiform, slender, often tapering
to sharp point, covered with distinctly raised tubercles or warts (Figure 48B). Calyx
sclerites long, up to 1.6 mm, very irregular and oddly shaped, mostly fusiform, one end
often forking slightly or very noticeably, evenly covered with distinct tubercles, with
some unilaterally spinose projections.
Generally, sclerites shown in Hardee and Wicksten (1996) appear to have extreme-
ly dense coverings of warts and bumps, the latter smallish in size. As well, they seem
to cover a greater portion of the surface area of the sclerite as compared to those seen
in drawings for sclerites of M. californica. Examinations of sclerites, for specimens
identied as this species, reveal that sclerites can have ame-like teeth, these running
almost the full length of the sclerite, on a side (what are termed nudibranch sclerites
(Figure45A), as those of Matamoros-Rosales (1984), rather than at an end such as seen
in a torch. Very largest sclerites (tardigrade-like) rounded, densely warted, with projec-
tions coming o one of the longer sides, creating the appearance of legs and claws (like
those of a tardigrade; Figures 44, 45D, some in Fig. 48E). Generally, sclerites have very
dense tubercle coverings. e largest sclerites in this species are decidedly larger than
those of M. californica; this will be the case despite what might have been concluded
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 143
Figure 46. Muricea fruticosa, SBMNH 265945. Colony measures ~7.5 cm × 4.0 cm. Visible in image is
the pale cream/yellow base/lower trunk; trunk intensies and darkens to brownish orange towards branch
tips. Notice swollen appearance of branch tips.
from having only looked at polyp color. is agrees with data provided in both Harden
(1969) and Grigg (1970).
e colony shown in Figure 46 displays a few odd features, warranting further
mention. Branching primarily pinnate to dichotomous. Colony measures ~7.5 cm
tall, 4.0 cm wide. Slightly central main stem runs entire height of colony, with some
slight curving laterally in random sections of stem; multiple branches all begin directly
above or from base, many coming o of main stem. Branches often angle out a very
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short distance then curve upwards. Primary branches average 3.0–4.0 mm long, di-
ameter ~1.0–2.0 mm (excluding calyces); diameter appears consistent from axillary
branch points to branch tips. Calyces very columnar (1.0–2.0 mm wide, ~3.0 mm
tall), heavily covered with longitudinally-oriented sclerites; polyps, many partially to
fully extended from calyces, cream or light yellow; appear smooth. Calyces on all sides
of branches, in some areas of colony very dense, in other areas calyces with some little
distance between themselves, where sclerites can be seen on coenenchyme surface, ly-
ing sometimes longitudinally with the branches, sometimes not; latter more transverse
(or oriented slightly in a triangular pattern) at base of calyces; calyces cover entire
colony, right down onto coenenchymal surface of colony’s base. Calyces generally ap-
pear distinct reddish brown due to large, conspicuous sclerites. Color of colony gener-
ally darker distally, reddish brown, grading from free branch tips of colony to much
lighter yellow or cream proximally, in stem and base (due to light-colored sclerites,
usually small in size); polyps appear very light yellow, cream to white. Majority of
calyx-bearing polyps appear such that they give colony an overall swollen appearance
(as though branches of colony are covered with small, round grapes). is is the most
unique feature of this colony; this may reect an active reproductive state at the time of
collection; specimen’s reproductive condition is still a question. Some calyces, scattered
on all branches, tend to curve upwards slightly. e sclerites shown in both light mi-
croscopy (Figure 47A, B) and SEM (Figure 48A–E) in some ways match those for the
colony of Figures 43–45, and yet in other ways do not clearly point to it being the same
species. Based on this single specimen in the SBMNH collection (SBMNH 265945),
from Long Beach, California, it requires further study. e biology of this colony, and
its swollen appearance, has not been further explored or explained to date, other than
to indicate that specimen was collected from very far into the back channels of Long
Beach Harbor, quite a distance from open water (D Cadien, LACSD, pers. comm.).
Etymology. Latin, fruticosu- meaning shrubby (bush-like); Verrill gave no specics
as to the derivation of the species name.
Common name. Brown gorgonian; Fruitful purple one; Bushy rust gorgonian;
Robust gorgonian (as indicated in various eld/diving guides).
Distribution. Potentially from Panama, up along California coast, perhaps as far
north as Los Angeles County, California, with maximum, though infrequent, north-
ern limit Point Conception, California. In a list of California sites, showing depth
ranges, the following are indicated: Mainland: Point Loma: 5–14 m; La Jolla: 3–12
m; (USNM 50192 was collected at 11 m at La Jolla, in the Torrey Pines kelp bed, 5
miles north of Scripps Institution); Newport Beach: 0–12 m; San Pedro: 0–15 m;
Santa Barbara: 5–8 m; Naples Reef: 5–11 m; Islands: Coronados Islands: 2–12 m; San
Clemente Island: 21 m; Santa Catalina Island: 5–18 m.
Biology. Seen more commonly in southern areas of California kelp beds (Ricketts,
4th Ed. 1968); also, oshore pilings. Seen as well in lowest intertidal zone, outer Los
Angeles Harbor; one of the most common species in southern California, in 15–30 m
depths, Point Conception to Baja California (Grigg 1977; Gotshall 1994, 2005). Both
this and M. californica seem to prefer subtidally occurring solid substrata at depths
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Figure 47. Muricea fruticosa, SBMNH 265945, light microscopy arrays. A 4× magnication, showing
large spindles with pronounced conical spinules; central, vertically aligned one, measures 650 µm B 10×
magnication of similar large spindles. Sclerite denoted by arrow is 0.875 mm in length, with a maximum
length for this spindle form ~1.0 mm.
between 1.0–30 m (Grigg 1977). Lissner and Dorsey (1986) reported that while this
species is common around the California Channel Islands and rocky areas of the main-
land, it is conspicuously absent on the Tanner and Cortes Banks and the Santa Rosa –
Cortes Ridge. Grigg (anecdotal communication) reported that populations of Muricea
may be limited by cold water and/or poor dispersal abilities of the larvae. Grigg (1974)
stated that this species is ca. one-tenth as abundant as M. californica o La Jolla. Based
on work done by Grigg, it was estimated that a colony 30 cm high is ~20 years old. As
very few colonies are seen larger than that, few colonies likely exceed this age. Mortality
attributed mostly to abrasion, occurring when particulate matter is suspended in the
water during periods of high waves and by smothering coming from accumulations of
sand. One untitled and unpublished identication guide stated that colonies are able
to survive in some of the most polluted near shore waters, as well as uncontaminated
oshore waters. It appears this species is immune to encrustations known to cause
mortality in M. californica. (SBMNH 422390, collected by MacGinitie in Newport
Bay, if indeed this species–original identication indicated it was, based on white pol-
yps, but sclerites do not support the identication–does have several galls produced by
an acorn-type barnacle, on bare axis as well as on an area covered by coenenchyme.)
Apparently fed upon by only one species of sh, the Garibaldi Hypsypops rubicunda.
Grigg (1974) calculated that between 10% and 15% of annual growth of this species
and M. californica is cropped by Garibaldi predation.
Remarks. Gift of FH Bradley; collected below low, low-water mark. Panama and
(?) Pearl Islands, 6–8 fm (11–15 m) (Verrill 1869a). Identied as Muricea fruticosa
var. typica (?).
ere are two specimens at NMNH identied as this species, with SEM images:
USNM 57171, SEM 237-238 (label reads ‘Albatross’ 28-29) and also, “Muricea fruticosa
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Figure 48. Muricea fruticosa, SBMNH 265945, SEM image. A Possible anthocodial sclerites B Spindles
from the axial sheath C An unusual quadriradiate spindle from coenenchyme D Calycular/coenenchymal
unilateral spinous spindles E Prominent calycular/coenenchymal tardigrade-type spindles.
Verrill,” collected by Limbaugh, from St. (Cape) Lucas Canyon, SEM image 999.
NMNH has an additional lot (USNM 52486), collected at Point Vicente, California,
along with material in the “Limbaugh” Collection, which could well be this species,
having been collected from the following locations (multiple lots): from California, at
Huntington Beach Gas & Electric Steam Plant discharge pipe; from Baja, at Turtle Bay.
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For all colonies of Muricea found in the California Bight, there is the possibility
of other species in the genus not previously reported as appearing in the Bight, which
may have very similar shape, etc. to the commonly recognized species described here.
Perhaps there are new, undescribed species. It may well be that previously described
species other than the standards make appearances in the Bight, and do so more often
than previously thought. is possibility is supported by the fact that climatic factors
can greatly expand ranges, even if only temporarily. A number of Mexican species may
occasionally (or more often) make an appearance in the California Bight during certain
substantial climatic/atmospheric events, such as an El Niño.
is conclusion has some support; Hardee and Wicksten (1996) in their closing
paragraphs state “(a)lthough our material could be identied as (simply) M. fruticosa
and M. californica, it is possible that other species of Muricea occur in southern Cali-
fornia.” ey recommended that a comparison of a series of specimens be made. is
would help to clarify which previously described species are valid and which are “merely
growth forms.” I agree with that recommendation. Fresh Muricea colonies, collected in
a very intentional manner, from south to north, both within the Bight and to the south
outside the Bight, must be done; that collection process is underway, with the help of
the Santa Barbara Museum of Natural History’s Sea Center sta, sta of the Aquarium
of the Pacic in Long Beach, California and sta of both the Los Angeles (LACSD)
and Orange County (OCSD) Sanitation Districts. With the dramatic weather events
we have recently experienced here in California, it will be interesting to see whether
other species of the genus are making an appearance, for any length of time.
Genus Placogorgia Studer, 1887
Placogorgia Studer, 1887: 56 [without species]. Wright and Studer 1889: 113. Nutting
1912: 83. Kükenthal 1919: 841; 1924: 209–210. Deichmann 1936: 141–142.
Bayer 1956a: F206; 1959b: 54–55. Grassho 1977 (pars): 26. Muzik 1979: 80–81.
? nec Placogorgia Nutting (part), 1910a: 76 [= Discogorgia Kükenthal].
Clematissa Studer, 1887: 106–107.
Pseudothesea Kükenthal, 1919: 843.
Discomuricea Gordon, 1926: 521.
Type species. Placogorgia atlantica Wright & Studer, 1889; SM Wright and Studer
1889 (= Pseudothesea Kükenthal, 1919).
Diagnosis. Colonies usually branched laterally in one plane; main stem generally
long; primary branches with tendency to curve upwards; primary branches tend to run
parallel with main stem, tips button-shaped, prominent swellings. Polyp height mod-
erately low, on all sides of branches; especially dense at branch tips. Calyces truncated,
cone shaped, armed with spindles (thorn scales). With crown (collaret) and points
arrangement (= operculum of Paramuricidae): each of eight points composed of two-
three pointed, convergent spindles in triangular arrangement above collaret of spinous
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rods, latter forming spiny transverse ring; fairly large triangular space free from scler-
ites between each point, situated in tentacle base. orn scales of calyx typically large,
coarse, thick; wider than tall, each with broad, abundantly branched basal root (broad,
at), and a (usually) short, stout, more or less laciniated but usually strong, blunt
spine; these sclerites overlap like roof tiles. Coenenchymal sclerites diverse spindles,
simple, branched, often attened, occasionally with one or more projections. Outer
coenenchyme with long, often bent, sclerites (spindles), blunt points on both ends; at
calyx base these form enclosing annular ring.
Placogorgia species A
Figures 49, 50A, B, 51A–C, 52A–G
Type locality and type specimens. Until there is species conrmation, no information
can be provided.
Material examined. ~5 lots (see Appendix 1: List of material examined). While
labeled Placogorgia material was examined at NMNH, nothing resembled in any way
the specimens in the SBMNH collection.
Description. Colony (Figures 49, 50A) generally branched in one plane; rarely, few
reticulate; colony height (base to tip of upper-most branch) approximately 20–21 cm.;
long, generally dichotomous branches and branchlets, moderately thick, cylindrical;
branch diameter thickness averages 5.0 mm (including calyces); meandering sinuously,
branches bent upward, parallel with main stem (not always obvious); tips of branches
and branchlets swollen, to rounded 8.0-mm diameter (Figure 50B). Main stem bifur-
cates (sometimes), some distance (±10 cm) from base; branches then again bifurcate
at ~1.0->5.0 cm from rst division. Further branching asymmetrical. Not all branches
subdivide; of those that do, distance from previous subdivision varies. Polyps distribut-
ed over entire surface, sparsely placed at base, becoming progressively most crowded at
branch tips. Color of freshly collected specimens, via video and still image (Figure 49),
pale pinkish tan with conspicuous, uy, cotton candy-pink polyps; in preserved (dry)
specimens, color dull tan-brown (Figure 50A, B); axis color slightly darker brown.
Calyces moderately low, ~1.0 mm tall, 2.0 mm across, 2.5 mm apart; blunt/rounded,
conical, armed with sclerites of various shape. Coenenchyme (relatively thin) contains
long, blunt-ending spindles, often bent; largest bent spindles 0.3–0.6 mm L, 0.05–0.1
mm H (average 0.5 mm L × 0.08 mm H), often with strong external spines (Fig-
ure52D). Distinctive sclerites often like thorn-scales or thorn-stars; small, spinulate or
laciniate ones predominantly calycular (Figures 52E–G). Anthocodial sclerites dicult
to extract; sclerites of collaret also blunt-ended, bent spindles, at base of polyp, tend-
ing to circular arrangement. A few as large, heavy, tapered spindles, sometimes with
several heavy, rough spines projecting from one side; some few (the calycular sclerites)
as branched torches (Figure 52B); also, crutch-types (Figures 51A, 52C); some few as
crosses and irregular ones, most as unilateral spiny shapes (Figure 51B, C). Color of
sclerites cream to very light tan, at least in specimens that are long dry.
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Figure 49. Placogorgia species A. In situ image, as seen in the Santa Barbara Channel. Image 6435B_
Snook_018, taken by L. Snook. Image courtesy of Milton Love, UCSB.
Distribution. For genus (based on material found/examined at NMNH and other
institutions, such as CAS) from Point Conception (California Bight) to Gulf of Califor-
nia (eastern North Pacic Ocean); western Pacic from Hawaii south to Philippines and
Indonesia. NMNH also has specimens in this genus collected from o the coast of Chile
in the southeastern Pacic Ocean; these look very dierent from the one described here.
Biology. May be considered a subtidal to deep-sea genus; this based on collection
data for known species.
Remarks. Bayer’s review of the genus (1959b), and description of a new species
from Florida, was invaluable for understanding the SBMNH specimens; a thorough
discussion of both the calycular thorn scales and the cortical sclerites was provided.
For specimens described here, the calycular thorn scales were dicult to extract; most
of the material studied was quite dry, but an examination of wet specimens (SBMNH
422968 and 422970) revealed that the form labeled “E” in Bayer (1959) is the closest
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150
t to what is seen in these specimens. Cortical sclerites in these specimens best t those
labeled “K,” “L,” and “M” (latter, in part; Bayer 1959). As well, sclerites seen in these
specimens also corresponded as follows to Diechmann’s (1936) illustrations for several
species in the genus Placogorgia from the Atlantic Ocean: pl 15, gs 19–20 best match
those identied as cortical thorn scales, pl 15, gs 23–24 best match those likely caly-
cular (branched torch) thorn scales, while pl 15, gs 26 and 32 match the blunt-ended
spindles seen here. Nearly all sclerite shapes in this unidentied species are far broader
in their root than they are tall; based on interpretation of the key in Bayer’s review I
have made a tentative placement of this species in the genus Placogorgia.
ere are multiple species of Placogorgia from the Atlantic, as well as a number
of species from the South Pacic and Indian Oceans (Kükenthal 1924). Only a few
records were found from the southeastern Pacic: unidentied species found west of
southern Chile (USNM 80162 [wet] and 98863 [wet]), and a number of unidentied
specimens from the western North Pacic (Hawaii), such as USNM 75077; none were
from the eastern North Pacic. ere was one reference by Harden (1979) indicating
that the species, using one of the same specimens currently discussed here (SBMNH
422969), is Placogorgia ramosa (Wright and Studer 1889), stating that it is synony-
Figure 50. Placogorgia species A, SBMNH 422970. A Colony measures 28 cm tall (excludes missing
base) × 9.0 cm broad B Magnication of branch tips; one tip malformed due to presence of barnacle gall.
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 151
mous with Paramuricea ramosa (Wright & Studer, 1889). Cordeiro et al. (2018i) does
not list a species P. ramosa in the list of recognized Placogorgia species in the WoRMS
Database; Harden’s designation does not inform the identication of the SBMNH
specimen in any way. As well, I could not nd any other indication that Paramuricea
ramosa is synonymous with Placogorgia ramosa.
Figure 51. Placogorgia species A. SBMNH 422970, Light microscopy arrays, 10× magnication. ASome
of the more unusually shaped sclerites B–C Several highly ornamented and distinctive sclerites. Crutch
form ~80 µm tall, thin spindles measure in the range of 150–191 µm in length, larger, thicker spindles are
~250 µm long, with unusual forms, such as B, the tall, spikey sclerite measuring some 300–350 µm tall
and that appearing as a very large quadriradiate (C) measuring 280 µm across.
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Figure 52. Placogorgia species A. SBMNH 422970, SEM image. A Small capstan form B Chunky
torch-type spindle C Crutch-type spindles D Long spindles E orn scales typical of the genus F Small,
developing thorn scale, dorsal view G More developed thorn scales, dorsal view.
Muzik (1979) stated that coenenchyme in species of Placogorgia is lled with diverse
spindles that are simple, branched and often attened, occasionally with one or more
projections; this is descriptive of what was seen here. Muzik (1979) also stated that the
calycular thorn scales are wider than tall, the projection usually short, with the base broad
and at. e Hawaiian species described by Muzik (1979) with their distinctive features
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 153
were of interest: the presence of 1) broad calycal thorn scales and 2) very branched, at-
tened sclerites of the coenenchyme. It seemed best to place the examined specimens in
the SBMNH collection within the genus Placogorgia based on the appearance of the
sclerites, but other genera are under consideration. However, any link between an eastern
Pacic locality and described members of this genus, or any other possible genera, can-
not be made (Ekman 1935: 66, 1953: 40; Bayer and Deichmann 1960).
Should specimens in the SBMNH collection represent a new species, this would be
the rst description of a species from this genus in the northeastern Pacic Ocean; if not a
new species, then this is the rst record of a known species of Placogorgia (seen elsewhere)
from the northeastern Pacic Ocean; further study of specimens is currently underway.
Conclusions
e SBMNH research collection, including substantial material from the Allan Han-
cock Foundation’s ‘Velero’ Expeditions, provides a good representation of the fami-
lies Gorgoniidae and Plexauridae (and many species included therein) from the area
known as the California Bight. e collection eectively displays the variation (and
dierences) of species found in the California Bight as compared with other locations
which may also harbor the same species. While many species from the collection match
well with species collected in other locations, there are some intriguing dierences seen
in many specimens taken from within the Bight that reect, perhaps, the dynamic
environmental system that is the California Bight.
As the family Plexauridae is so well represented in the SBMNH collection, two of
the remaining genera in the family, Swiftia and esea, will be discussed more thorough-
ly in Part III of this work, completing the full and comprehensive review of all species
recorded to date as appearing in the California Bight, based on the SBMNH research
collection, and specically on the many specimens that came to SBMNH through the
Allan Hancock Foundation’s historic ‘Velero’ Expeditions collection events.
References
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Appendix 1
List of material examined – Part II
(Material examined = whole colony study plus multiple sclerite preparations; all
with light microscopy, plus selected colonies under SEM, shown in gures associ-
ated with text)
Adelogorgia phyllosclera Bayer, 1958
Material examined. ~25 lots. USA, California – fragments, plus full colony; San Die-
go County, San Diego, Point Loma, 32°41'28"N, 117°16'53"W, station PL-13, no
depth recorded; coll. SCCWRP, Sea Quest, 10 September 1975; LACoMNH Marine
Biodiversity Center and SBMNH 422894 [wet]. – 1 fragment; Los Angeles County,
o the southern California Channel Islands, seaward of Newport Beach, 33°34'02"N,
118°02'20"W, 96 m; coll. M Love via submersible ‘Delta’, 6 October 2005; (his num-
ber: A6651), SBMNH 422893 [wet]. –1 colony; Los Angeles County, Catalina Is-
land, Bird Rock, 33°27'00"N, 118°29'14"W, 50 m; coll. R Given, July 1977; SBMNH
51252 [wet]. –1 colony; Los Angeles County, Santa Catalina Island, .33 miles, 180° T
from Ship Rock Light, 33°27'19"N, 118°29'34"W, 58 m; coll. R/V ‘Velero IV’, sta-
tion 2132-52, 22 July 1952; SBMNH 422308 [wet]. –1 colony; Los Angeles County,
Santa Catalina Island, 4 miles, 140° T from Ship Rock Light, at Isthmus Cove, by
dredge, 33°27'36"N, 118°28'58"W, 76 m; coll. R/V ‘Velero IV’, station 12400-68,
17 October 1968; SBMNH 423087 [dry]. –1 colony; Los Angeles County, o Santa
Monica, South Banks, 33°53'00"N, 118°31'00"W, by trawl, 82–100 m; coll. unknown,
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 165
10 October 1978; SBMNH 422892 [wet]. MEXICO, Baja California Sur (Pacic
Coast) – multiple fragments; southwest of Punta San Eugenio, 8.5 miles S of Canal de
Dewey, 27°42'15"N, 115°05'02"W, 89 m; coll. R/V ‘Velero III’, station 1259-41, 27
February 1941; SBMNH 422309; [wet]. –3 colonies; Isla Natividad, 7.5 miles SSW of
island, 27°44'17"N, 115°14'40"W, 115–120 m; coll. R/V ‘Velero III’, station 1258-41,
bottom of loose rocks, coral, 27 February 1941; SBMNH 422403 [dry]. MEXICO,
Baja California Norte (Pacic Coast) – multiple colonies; 8 miles W of Isla Cedros,
28°05'45"N, 115°31'35"W, 117–118 m; coll. R/V ‘Velero III’, station 1253-41, 26
February 1941; SBMNH 422310 [wet]. –multiple colonies/fragments; 1.5 miles o
north end of Isla Cedros, 28°23'20"N, 115°11'52"W, 100–109 m; coll. R/V ‘Velero
III’, station 1264-41, 28 February 1941; SBMNH 422311 [wet]. CENTRAL AMER-
ICA – 1 colony, (no base); Panamá, Veraguas, o Bahía Honda, rock, sand and mud,
07°45'35"N, 81°35'35"W, 55–91 m; coll. R/V ‘Velero III’, station 863-38, 01 March
1938; SBMNH 423058 [dry]. SOUTH AMERICA – multiple fragments; northern Pa-
cic Ocean, Ecuador, Galápagos Islands, O Bindloe (=Marchena Island), 00°18'20"N,
90°31'00"W, 27 m; coll. R/V ‘Velero III’, Station 310-35, 3 December 1934; SBMNH
422312 [wet]. –1 colony (bleached); southern Pacic Ocean, Galápagos Islands, Santa
Cruz Island, o Gordon Rocks, 00°33'30"S, 90°09'45"W, 46–55 m; coll. R/V ‘Velero
III’, station 317-35, 8 December 1934; SBMNH 422891 (‘Velero’ label written in er-
ror, 317-37) [wet]. Note: Specimens listed from Central/South America possibly one or
more of several new species recently described by Breedy and Guzmán 2018.
Other material examined. USA, California – 1 fragment; San Diego County,
taken o shore, BI 53-4, SIO 61-517, 32°38'33"N, 117°15'34"W, 30 m, with dredge;
coll. unknown, 19 November 1953 (0900); SIO-CO 457 [wet]. –2 fragments; San
Diego County, SI 50, from Canyon Rim, 80 m; coll. unknown, 2 October 1965; SIO-
CO 1683 (two specimens in the one lot) [wet]. –San Diego County, Canyon Rim, SL
49-b. 80 m; coll. R Grigg, 2 October 1965 (1200); SIO-CO 1605 [wet]. –San Diego
County, o La Jolla, Station 8, La Jolla Canyon, 114-40 m, rock dredge–rock, sand,
clay; coll. unknown, 7 January 1954; SIO-CO 1588 [wet]. –multiple colonies; San
Diego County, La Jolla, La Jolla Canyon, South Wall, 64 m; coll. C Limbaugh and
party, 11 September 1957; [NMNH collection, no catalog number, dry]. –Los Angeles
County, Santa Monica, 33°52'08"N, 118°34'05"W, 73 m; coll. unknown, 29 March
1974; SIO-CO 1579 [wet].
Other material, not examined. USA, California – San Diego County, San Die-
go, 32°45'06"N, 117°30'00"W, 9–24 m; coll. C Limbaugh, Station 1166, 3 July 1906;
USNM 57456 [wet]. –2 colonies; USA, California, San Diego County, La Jolla, 400
miles west of Point La Jolla, 18–21 m; coll. C Limbaugh, 14 December 1953; Para-
type–USNM 50187 [dry]. –6 colonies; USA, California, San Diego County, La Jolla,
La Jolla Canyon, 46–52 m; coll. C Limbaugh, 8 October 1955; Paratype–USNM
50188 [dry]. –1 colony; USA, California, San Diego County, La Jolla, La Jolla Can-
yon, 40–43 m; coll. R Ghilardi and party, 19 January 1956; Paratype–USNM 50635
[dry]. –USA, California, San Diego County, La Jolla, La Jolla Point, 32°51'01"N,
117°21'02"W, 104–130 m; coll. R/V ‘Albatross’, Station 4328, 8 March 1904; USNM
51320 [dry].
Elizabeth Anne Horvath / ZooKeys 860: 67–182 (2019)
166
Eugorgia daniana Verrill, 1868
Material examined. ~10 lots MEXICO, Baja California Sur (Pacic Coast) – 1
colony; 2 miles, 220° T from Punta San Eugenio, 27°49'30"N, 115°05'10"W, 38
m; coll. R/V ‘Velero IV’, station 11514-67, 15 June 1967; SBMNH 422895 [wet].
MEXICO, Baja California Norte (Pacic Coast) – 1 colony; Isla Cedros, coast side,
28°13'00"N, 115°10'00"W; coll. Pacic BioMarine, received from Carmelita Freeman,
26 April 1974; SBMNH 422897 [DH 423 = SBMNH-10]; [dry]. –1 colony; Isla Ce-
dros, 10 miles S of Punta Norte on east side of island, 28°13'00"N, 115°10'00"W, 9 m;
coll. R McPeak, 22 February 1983; SBMNH 422404 (B1545) [dry]. MEXICO, Baja
California Norte (Gulf of California) – 1 colony; Bahía de Los Angeles, Isla Mit-
lan, 29°04'00"N, 113°31'00"W, 6 m; coll. R McPeak, 12 September 1980; SBMNH
422405 [dry]. MEXICO (Gulf of California) – 1 colony; Sonora, Bahía de Cholla
(= Choya), 31°20'46"N, 113°38'14"W; coll. J Wilkins, 23 February 1959; SBMNH
422896 [DH 433 = SBMNH-20] [dry].
Other material examined – fragment; northern Pacic Ocean, Mexico, Baja
California Sur, oshore island of Isla Socorro, outside Punta Tosca, ~18/19°00'00"N,
112°00'00"W, 21 m; coll. unknown, 4 February 1964; SIO CO 1686 [wet]. –1 col-
ony; northern Pacic Ocean, Mexico, Baja California Sur, Puerto (Bahía) San Barto-
lomé (Tortugas or Turtle Bay); coll. unknown, 14 March 1961; USNM 52484 [dry].
–fragment; USA, California, San Diego County, Loma Sea Valley, due west of Point
Loma, by otter trawl 1, ST-908, 326–351 m; coll. Fager party, 21 January 1965; SIO
CO 1684 (Acc. No. BI 65-33) [wet]. –1 colony; California, San Diego County, San
Diego, Escondida Bay; coll. unknown, 14 November 1868; USNM 57302 [dry].
Eugorgia rubens Verrill, 1868
Material examined. ~ 50 lots. USA, California – 1 colony; California Channel Is-
lands, on a specimen shell of Haliotis sorenseni (SBMNH 349096); coll. Leon Bray,
1964 [dry]. –1 fragment; Orange County, 1 mile S of Newport Harbor, 33°36'30"N,
117°54'30"W, vertical south face of shale reef, 17 m; coll. R Given, by hand, SCU-
BA, 05 October 1957; SBMNH 422313 [wet]. –1 colony (stripped); Santa Catalina
Island, 1.5 miles, 130° T from Long Point Light, urchins, crinoids, dead brachio-
pod, diversied, very rich, 33°23'48"N, 118°21'16"W to 33°22'57"N, 118°19'54"W,
84 m; coll. R/V ‘Velero IV’, Station 2226-53, 28 February 1953; SBMNH 422314
[wet]. –1–2 colonies; Los Angeles County, Santa Catalina Island, between Avalon
and Long Point, 33°23'30"N, 118°21'50"W, 55–80 m; coll. R/V ‘Velero III’, station
1167-40, 08 August 1940; SBMNH 422407; SBMNH-14 [dry]. –colony fragments;
Los Angeles County, .33 miles, 180° T from Santa Catalina Island, Ship Rock Light,
dredge–brachiopods, rocky sand bottom, 33°27'19"N, 118°29'34"W, 58 m; coll. R/V
‘Velero IV’, station 2132-52, 22 July 1952; SBMNH 422315[wet]. –multiple frag-
ments; Los Angeles County, 7.35 miles, 89° T to Manhattan Beach Pier, 33°53'18"N,
118°32'45"W, 58 m; coll. R/V ‘Velero IV’, station 24493-76, 10 March 1976; (with
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 167
label– AHF 24493, BLM Ref. station 360; Leg. Mo. 08); SBMNH 422099 [wet]. –
fragment; Los Angeles County, o Rocky Point, close to Point Vicente, S end of Santa
Monica Bay, ~33°44'33"N, 118°25'28"W; coll. by C Limbaugh; 5 October 1941;
SBMNH 422898 [wet]. –multiple fragments; Los Angeles County, 10.75 miles W.
of Point Dume, on loose rock, sponge, 34°00'00"N, 119°01'20"W, 87 m; coll. R/V
‘Velero III’, station 1276-41, 23 March 1941; SBMNH 422324 [wet]. –1 colony;
Ventura County, Rincon, 34°22'27"N, 119°28'27"W, 98 m; coll. P Brophy, com-
mercial otter trawl, 01 May 1968; SBMNH 422901 [DH 421 = SBMNH-08; dry].
–1 colony; Ventura County, Anacapa Island, 34°01'00"N, 119°21'43"W, 11 m; coll. B
Scronce and M Conboy, by hand, SCUBA, 29 January 1964; SBMNH 422900 [DH
432 = SBMNH-19; dry]. –1 colony; Ventura County, Anacapa Island, 34°01'00"N,
119°21'43"W, reef and cove, northeast end of island, 11 m; coll. B Scronce and party,
by hand, SCUBA, 29 January 1964; SBMNH 45561 [ex MacGinitie collection; wet].
–1 colony; Ventura County, Anacapa Island, 34°01'00"N, 119°21'43"W, reef and
cove, northeast end of island, 17–18 m; coll. B Scronce and party, 29 January 1964;
SBMNH 45562 [ex MacGinitie collection; wet]. –1 fragment; Ventura County, An-
acapa Island, windward side, 34°00'36"N, 119°24'02"W, 15–24 m; coll. M Hamann
and S Bobzin, 21 February 1998; SBMNH 345323 [wet]. –1 colony; Santa Barbara
County, Santa Cruz Island, 34°01'14"N, 119°41'05"W, seaward side; 15–18 m; coll.
W Hary, February 1967; SBMNH 422411 [dry]. –1 colony fragment; Santa Barbara
County, basin o Santa Cruz Island (Santa Barbara Basin/Channel), 15.5 mi., 139°T,
from Santa Barbara Light, Santa Barbara, CA, 34°20'15"N, 119°37'45"W, 51 m; coll.
T Phillips, R/V ‘Velero IV’, station 13617-69, 13 November 1969; SBMNH 422409
[DH 428 = SBMNH-15; dry]. –small colony and fragments; Monterey County, Point
Sur, 36°16'07"N, 121°55'13"W, 65 m; coll. T Laidig, NMFS-FED, Santa Cruz, 13
October 2007; SBMNH 235537 [dry]. MEXICO, Baja California Sur (Pacic
Coast) – several fragments; 28 miles, 199° T from Abreojos Light, S of Punta Abreo-
jos, dredge–brachiopods, #12 coarse sand and rock, 26°17'15"N, 113°41'45"W (end),
100 m; coll. ‘Velero IV’, station 1709-49, 7 March 1949; SBMNH 422327 [wet]. –1
colony; 6.5 miles, 167° T from Punta Abreojos, dredge–coarse mud and sand, mud
bottom, 26°35'27"N, 113°31'45"W (end), 51 m; coll. R/V ‘Velero IV’, station 1954-
50, 29 April 1950; SBMNH 422316 [wet]. –2 colonies; SE of Isla Asunción, pinnacles
near the “6 fathom spot,27°06'28"N, 114°17'30"W, 23–27 m; coll. RH McPeak, sta-
tion B, 11-14 Nov. 1981; SBMNH 422412 [dry]. –2 colonies; 8.5 miles south of Ca-
nal de Dewey, sand, broken shell, gravel, 27°42'15"N, 115°05'02"W (end), 89 m; coll.
R/V ‘Velero III’, station 1259-41, 27 Feb. 1941; SBMNH 422323 [wet]. MEXICO,
Baja California Norte (Pacic Coast) – 1 colony; Islas San Benitos, 28°18'10"N,
115°33'20"W, in Phyllospadix, 5 m; coll. Santa Catalina Island Lab, 15 August 1977;
SBMNH 422413 [dry]. –1 colony; 8 mi. SW of Isla Cedros, green, ne sand, coral,
28°00'00"N, 115°29'00"W (end), 115–118 m; coll. R/V ‘Velero III’, station 1254-41,
26 Feb. 1941; SBMNH 422318 [wet]. –several colonies; 8 miles W of Isla Cedros,
gravel, loose rock, 28°05'45"N, 115°31'35"W (end), 116–118 M; coll. R/V ‘Velero
III’, station 1253-41, 26 Feb. 1941; SBMNH 422317 [wet]. –1 colony; 1.5 miles o
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N end of Isla Cedros, ne sand, broken shell, 28°22'18"N, 115°11'00"W (end), 82–
100 m; coll. R/V ‘Velero III’, station 1263-41, by small dredge, 28 Feb. 1941; SBM-
NH 422329 [wet]. –1 fragment; 1.5 miles o N end of Isla Cedros, shale, pebbles,
28°23'20"N, 115°11'52"W (end), 100–109 m; coll. R/V ‘Velero III’, station 1264-41,
by small dredge, 28 Feb. 1941; SBMNH 422328 [wet]. –multiple fragments; 6.5 miles
SW of Punta San Carlos, dredge, rocky bottom, 29°33'30"N, 115°35'28"W (end),
36 m; coll. R/V ‘Velero IV’, station 1944-50, 25 April 1950; SBMNH 422325 [wet].
–1 colony; 4 mi. N of Isla Todos Santos, on rock, 31°52'10"N, 116°49'25"W (end),
73 m; coll. R/V ‘Velero III’, station 1244-41, 24 Feb. 1941; SBMNH 422319 [wet].
MEXICO, Baja California Sur (Gulf of California) – 1 fragment; E of Isla San Fran-
cisco (Francisquito), sandy mud, 24°47'25"N, 110°31'20"W, 109 m; coll. R/V ‘Velero
III’, station 651-37, 9 March 1937; SBMNH 422330 [wet].
Other Material Examined. USA, California – San Diego County, San Diego, o
Point Loma, 32°35'05"N, 117°20'02"W, 91–101 m, by otter trawl; coll. R/V ‘E.B.
Scripps’ (student cruise), A Fleminger, 15 November 1980; SIO-CO 912 [wet]. –1
colony; San Diego County, Canyon Rim, 80 m; coll. R Grigg, Cruise SI 49a, 2 Oc-
tober 1965; (was SIO/BIC CO 1593(b), now SIO-CO 2087(a)); [wet]. –1 fragment;
San Diego County, Canyon Rim, 77–80 m; coll. unknown, Cruise SI 50, 02 October
1965; (was SIO/BIC CO 1683(a), now SIO-CO 2088, 2089 or 2090) [wet]. –San
Diego County (?), Sea Lab SL 49-d, 80 m; coll. R Grigg, 2 October 1965; SIO-CO
1578 [wet]. –San Diego County, La Jolla, La Jolla Canyon, 47 m; coll. R Ghilardi
and party, 4 March 1959; SIO-CO 1685; [wet]. –1 colony; San Diego County; Sta-
tion 2104, 86 m; det. M Lilly and D Pasko, City of San Diego, MBL, 26 July 1996;
[dry]. –1 colony; Los Angeles County, San Clemente Island, NE side, at Forbidden
Reef, 33°00'51"N, 118°20'37"W; coll. J Cooper, 11 June 1982; MLML CO 109
[wet]. –Los Angeles County, Santa Monica, 33°52'08"N, 118°34'05"W (end), 73 m;
29 March 1974; SIO-CO 1629 [wet]. –Ventura County, Anacapa Island, south side,
34°00'00"N, 119°26'00"W, 15–26 m; coll. FG Hochberg, 20 November 1964. LA-
CoMNH No. 64-28 [wet].
Eugorgia ljubenkovia sp. nov.
Material examined. 5 lots; two specimens of species in SBMNH collection from Mexi-
co; another 3 lots, collected by sta of OCSD in California waters, now housed at SBM-
NH. – fragment; USA, California, Orange County, o Huntington Beach, by Van Veen
grab, ~33°36'34"N, 118°02'32"W, 30 m; coll. OCSD, Survey 94–100, station 34, Rep.
1, 1994; material from J Ljubenkov; SBMNH 472234 [wet]. –numerous fragments;
USA, California, Orange County, o Huntington Beach, otter trawl, ~33°35'41"N,
117°59'34"W, 35 m; coll. OCSD, Survey 8718, station T-2, Rep. 1, Haul 1, 13 January
1987; Bottle 0011, SBMNH 472232 [wet]. –1 colony + fragment; USA, California, Or-
ange County, o Huntington Beach, otter trawl, at night, ~33°35'57"N, 118°02'47"W,
36 m; coll. OCSD, Survey 8836, station T-6, Rep. 1, 1988; Bottle 0080, SBMNH
472233 [wet]. MEXICO, Baja California Norte ySur – 1 strand; 28 miles south of
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 169
Punta Abreojos, bearing 199° T from Abreojos Light, 26°17'30"N, 113°41'45"W, 98
m; coll. R/V ‘Velero IV’, station 1709-49, 7 March 1949; SBMNH 472245 [wet].
–colony fragments; Isla Cedros, S Bay, dredge—sand and mud, Lovena (heart urchin),
sponges, 28°04'21"N, 115°17'50"W (end), 29 m; coll. R/V ‘Velero IV’, station 1703-
49, 5 March 1949; SBMNH 422333–Holotype [wet].
Leptogorgia chilensis (Verrill, 1868)
Material examined. ~25 lots. USA, California – 1 colony (partial); Los Angeles
County, Farnsworth Bank, dredge—bryozoan, 33°20'39"N, 118°30'55"W (end), 15
m; coll. R/V ‘Velero IV’, 1904-49, 7 September 1949; SBMNH 422940 [wet]. –1+
colony; Los Angeles County, 5 miles, 152° from San Pedro Breakwater, 33°38'20"N,
118°12'00"W, 31–35 m; coll. R/V ‘Velero III’, station 1232-41, 15 February 1941;
SBMNH 423066 [wet]. –1+ colony + fragments; Los Angeles County, 2.25 miles
south of San Pedro Breakwater, 33°40'30"N, 118°14'20"W, 27 m; coll. R/V ‘Velero
III’, station 1207-40, 24 November 1940; SBMNH 423067 [wet]. –two frag-
ments; Los Angeles County, Palos Verdes Estates, 33°47'18"N, 118°24'47"W; coll.
T Burch, station 40129, 22 August 1940; SBMNH 422954 [wet]. –multiple frag-
ments; Los Angeles County,7.35 miles, 89° T to Manhattan Beach Pier, 33°53'18"N,
118°32'45"W (end), 58 m; coll. R/V ‘Velero IV’, station 24493-76, 10 March 1976;
SBMNH 422943 [wet]. –2 colonies; Ventura County, Anacapa Island, 34°00'14"N,
119°23'41"W, 6 m; coll. unknown, 24 January 1963; with hydroid colony on one
branch; SBMNH 422944 [wet]. –1 colony; Ventura County, Anacapa Island, 100
yards out from Cat Rock, 34°00'13"N, 119°25'16"W, 17–18 m; coll. B Scronce and
party, 30 October 1962; SBMNH 422945 [wet]. –1 colony; Santa Barbara County,
~6 miles south and west of Summerland coast, on northwest corner of Platform
“A”, ~34°20'49"N, 119°38'25"W, ~39 m; coll. S Clark, 3 December 2010; SBMNH
265962 [wet]. –1 colony; Santa Barbara County, Goleta, Sands Beach, 34°24'00"N,
119°53'00"W, ~6 m; coll. R/V ‘Vantuna’ Cruise, 469—Mar. Bio, November 2001;
SBMNH 422953–Neotype [wet]. MEXICO, Baja California Sur (Pacic Coast)
– 1 colony + branch fragments; 6.5 miles, 167° T from Punta Abreojos, dredge–
coarse mud and sand, mud bottom, 26°35'27"N, 113°31'45"W (end), 51 m; coll.
R/V ‘Velero IV’, station 1954-50, 29 April 1950; SBMNH 422947 [wet]. –2 colo-
nies; Canal de Dewey, opposite Punta San Eugenia, coralline, rock, 27°49'40"N,
115°06'20"W (end), 38–47 m; coll. R/V ‘Velero III’, station 1260-41, 27 February
1941; SBMNH 422949 [wet]. –1 colony; 2 miles, 220° T from Punta San Eugenia,
27°49'30"N, 115°05'10"W (end), 38 m; coll. R/V ‘Velero IV’, station 11514-67,
15 June 1967; SBMNH 422950 [wet]. MEXICO, Baja California Norte (Pacif-
ic Coast) – 1 fragment; South Bay, Isla Cedros, rock, along margin of kelp bed,
28°04'45"N, 115°21'05"W, 18–27 m; coll. R/V ‘Velero III’, 287-34, 10 March
1934; SBMNH 422951 [wet]. –1 colony, incomplete; 6.5 miles SW of Punta San
Carlos, dredge–rocky bottom, 29°33'30"N, 115°35'28"W (end), 36 m; coll. R/V
‘Velero IV’, station 1944-50, 25 April 1950; SBMNH 422952 [wet].
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Other material examined. USA, California – 3 fragments; San Diego County,
Canyon Rim, 73 m; coll. R Grigg, Cruise SI 49a, 4 October 1965; SIO/BIC 1593(a)
[wet]. –1 strand; San Diego County, Canyon Rim, 70–73; coll. unknown, Cruise
SI 50, 2 October 1965; SIO/BIC CO 1683(c) [wet]. –2 lots; colony/fragments; San
Diego County, Scripps Submarine Canyon, o Scripps Institute, 15–21 m & 23–30
m; coll. C Limbaugh, December 1953; [Limbaugh Collection, NMNH, dry]. –1+col-
ony/ fragments; San Diego County, La Jolla Canyon, one quarter mile o Scripps
Institute, 46 m; coll. C Limbaugh, 23 July 1954; [Limbaugh Collection, NMNH,
dry]. –colony/ fragments; San Diego County, one quarter mile o Point La Jolla, edge
of kelp bed, 18–21 m; coll. C Limbaugh, 14 December 1953; [Limbaugh Collec-
tion, NMNH, dry]. –colony/fragments; San Diego County, 1 mile north of La Jolla,
29 m; coll. C Limbaugh, 11 February 1952; [Limbaugh Collection, NMNH, dry].
–1 colony; Los Angeles County, San Clemente Island, NE side of island, Forbidden
Reef, 33°01'26"N, 118°20'17"W; coll. J Cooper, by SCUBA, 11 June 1982; MLML
C0108 [wet]. –colony/fragments; Los Angeles County, Redondo Beach, Santa Monica
Bay, Streetcar Reef, ~33°51'00"N, 118°25'04"W, 15 m; coll. unknown, 18 July 1961;
[Limbaugh Collection, NMNH, dry]. –colony/fragments; Los Angeles County, Re-
dondo Beach, Santa Monica Bay, Streetcar Reef, 18 m; coll. unknown, 14 September
1961; [Limbaugh Collection, NMNH, dry]. –colony/fragments; Ventura County, ~10
miles south of Santa Barbara, California, Richeld Oil Island, 5–9 m; coll. unknown,
16 May 1961; [Limbaugh Collection, NMNH, dry]. –1 colony (possibly this species);
Santa Barbara County, S side of Santa Cruz Island, Smuggler's Cove, 34°01'13"N,
119°32'25"W, washed up on cobbles; coll. N Moore, det. EA Horvath, photographed
only, 10 March 2008 [dry]. MEXICO, Baja (Pacic Coast) – colony/fragments;
MEXICO, Baja California Sur, Bahia Tortugas area, ~15–18 m; coll. J Stewart, No-
vember 1959; [Limbaugh Collection, NMNH, dry]. –colony/fragments; MEXICO,
Baja California Norte, Isla Guadalupe, 42 m; coll. J Stewart and party, February 1960
[Limbaugh Collection, NMNH, dry].
Other material, not examined – 1 specimen: USA, California, Santa Barbara
County, Santa Cruz Island, Smuggler’s Cove, 21 m.; coll. B Scronce and party, by div-
ing, 24 January 1963; [DH 426 = SBMNH-13 [dry], missing from SBMNH collection.
Leptogorgia diusa (Verrill, 1868)
Material examined. 4 lots. MEXICO (Gulf of California) – 1 colony; Sonora, Cabo
Tepoca, 30°15'45"N, 112°53'20"W, on shore–rock and reef; coll. R/V ‘Velero III’, sta-
tion 1077-40, 4 February 1940; SBMNH 423089 [dry]. –multiple colonies; Sonora,
o Rocky Point, mud, sand and shell, 31°19'50"N, 113°39'10"W (end), 18–20 m;
coll. R.V ‘Velero III’, station 1072-40, 2 February 1940; SBMNH 423088 [dry]. –1
colony; Sonora, Bahia de Choya (Cholla), ~31°21'19"N, 113°37'26"W; coll./legit. JW
with # 038-1, 25 February 1959; SBMNH 423090 [dry].
Other material examined –1 colony; USA, California, San Diego County, San
Diego, o shore; coll. J Stewart, 1965; CAS IZ 97890 [dry].
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 171
Leptogorgia licrispa Horvath, 2011
Material examined. ~9 lots. USA, California – 2 fragments (1 lot); Ventura County,
o Ventura Harbor, collected with ovulid snails, 34°14'00"N, 119°19'00"W, 20–50
m; coll. P Brophy, trawled by dragnets, August 1968; SBMNH 423079 [housed in
vial, in ovulid snail collection, Neosimnia nec Simnia loebbeckeana, SBMNH 423103,
dry]. MEXICO, Baja California Sur (Pacic Coast) – multiple colonies (1 lot); o
Boca Flor de Malva, SE of Punta Tosca, 24°11'07"N, 111°21'03"W, 69–87 m; coll.
J McLean on R/V ‘Searcher’, Station 31–32, No. 71–16, 1 February 1971; SBMNH
423057–Holotype [DH 415 = SBMNH-06, dry].
Other material examined. MEXICO (Gulf of California) – several fragments;
Gulf of California, Sonora, Mexico, Guaymas, 28°15'00"N, 111°48'00"W (end),
64–75 m; coll. unknown, 21 March 1960; USNM 75099 [wet]. –numerous frag-
ments/colonies; Sonora, Cabo Tepoca, 30°20'30"N, 113°08'00"W, 56–67 m; coll. R
Parker, 31 March 1960; (with a station number: P-212-60); USNM 1106683 [dry].
MEXICO, Baja California Sur (Pacic Coast) – multiple fragments/colonies; Cabo
San Lucas Canyon, ~22°54'09"N, 109°22'45"W, on sand bottom, 30 m; coll. W
North and C Limbaugh, March 1959; USNM 1106685 [dry]. –multiple fragments;
o Boca Flor de Malva, SE of Punta Tosca, 24°11'07"N, 111°21'03"W, 69–87 m;
coll. J McLean on R/V ‘Searcher’, Station 31–32, No. 71–16, 1 February1971; LA-
CoMNH—Holoparalectotype [DH 218, SBMNH06, dry]. –multiple fragments;
o Boca Flor de Malva, SE of Punta Tosca, 24°11'07"N, 111°21'03"W, 69–87 m;
coll. J McLean on R/V ‘Searcher’, Station 31–32, No. 71-16, 1 February1971; LA-
CoMNH—Holoparalectotype [DH 415, SBMNH06, dry]. –multiple fragments;
Cabo San Lazaro, 24°48'00"N, 112°19'12"W, by shrimp trawler, 65–73 m; coll. C
Limbaugh, sometime in the 1950s; USNM 1106684 [dry]. –several fragments; Mag-
dalena Bay, ~24°38'11"N, 111°58'15"W, 86 m; coll. unknown on R/V ‘Albatross’, 8
April 1889; USNM 57144 [wet]. –multiple fragments; SW Punta San Juanico, outer
coast, ~26°12'33"N, 112°30'49"W, (no depth recorded); coll. unknown, 29 January
1971; LACoMNH [DH240, dry].
Leptogorgia exilis (Verrill, 1868)
Material examined. 5 lots. USA, California – 1 colony fragment (base missing);
Los Angeles County, San Pedro, 6.9 miles, 139° T from Point Fermin, dredge–sand,
33°37'04"N, 118°11'50"W (end), 45 m; coll. R/V ‘Velero IV’, 2042-51, 20 July 1951;
SBMNH 422941 [wet]. –2 lots; colony, fragmented; Los Angeles County, between
White's Point and Portuguese Bend, 33°42'54"N, 118°20'07"W (end), by beam trawl,
22 m; coll. R/V ‘Velero IV’, station 1644-48, 19 November 1948; SBMNH 422942
[dry & wet]. MEXICO, Baja California Sur (Pacic Coast) – fragments; 28 miles,
199° T from Abreojos Light, S of Punta Abreojos, dredge–brachiopods, beam trawl–#
12 coarse sand and rock, 26°17'15"N, 113°41'45"W (end), 100 m; coll. R/V ‘Velero
IV’, station 1709-49, 7 March 1949; SBMNH 422946 [wet]. –1 colony + multi-
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ple fragments; 2 miles, 142° T from urloe Head, 27°35'45"N, 114°49'15"W, no
depth recorded; coll. R/V ‘Velero IV’, station 11842-67, 7 December 1967; SBMNH
422948 [wet].
Leptogorgia sp. A (? = Leptogorgia tricorata Breedy and Cortés 2011)
Material examined. ~10-11 lots. USA, California – 2 colonies, 1 fragment; 9.5 miles
NW of buoy, Cortes Bank, white sand, rock, 32°33'15"N, 119°15'15"W, 91 m; coll.
R/V ‘Velero III’, station 1342-41, 10 June 1941; SBMNH 423080 [wet]. –4 colo-
nies; Los Angeles County, Santa Catalina Island, 1 mile SW of Ben Weston Point,
mud, sand, gravel, 33°20'55"N, 118°30'45(35)” W (end), 82–89 m; coll. R/V ‘Velero
III’, station 1316-41, 17 May 1941; SBMNH 422334 [wet]. –1 colony; Los Angeles
County, Santa Catalina Island, Bird Rock, ~33°27'05"N, 118°29'10"W, 46 m; coll. R
Given, July 1977; SBMNH 423083 [wet]. –1 fragment; Los Angeles County, Santa
Catalina Island, o Bird Rock, rock, coarse shell, kelp; sm. dredge boat, 33°27'20"N,
118°29'00"W (end), 56–73 m; coll. R/V ‘Velero III’, station 1187-40, 29 September
1940; SBMNH 423081 [wet]. –1 fragment; Ventura County, Santa Barbara Island,
N of island, 33°30'58"N, 119°00'50"W, on gray sand, 67-73 m; coll. R/V ‘Velero
III,’ station 1177-40, 9 September 1940; SBMNH 422903 [wet]. –2 colony frag-
ments; Los Angeles County, o Rocky Point, close in proximity to Point Vicente (S
end of Santa Monica Bay), ~33°44'33"N, 118°25'28"W; coll. C Limbaugh, 5 Octo-
ber 1941; SBMNH 423082 [wet]. –numerous colonies/fragments; 10.75 miles W of
Point Dume, on loose rock and sponge, 34°00'15"N, 119°01'30"W (end), 86-87 m;
coll. R/V ‘Velero III’, station 1276-41, 23 March 1941; SBMNH 422340 [wet]. –frag-
ment; Santa Barbara County, Channel Island area, ~34°06'56"N, 119°42'49"W, by
trawl; coll. P Brophy; SBMNH 423084 [wet].
Other material examined – fragment; Mexico, Baja California Norte (Gulf of
California), NE of Isla San Jose, Isla Las Animas, 28°42'14"N, 112°55'43"W, epizooic
on “black coral, 36–42 m; coll. Adcock and Markham, Station D-24B, 11 August
1965 (identied in error by Harden as Heterogorgia tortuosa); CAS-IZ 96757 [wet]. –1
colony; USA, California, San Diego County, near coast of San Diego, no depth record-
ed; coll. J Stewart (UCSD diver), 1965 (identied in error by Harden as H. tortuosa);
CAS-IZ 98140 [dry]. –1 strand; USA, California, San Diego County, Canyon Wall
(1200), 82 m; coll. R Grigg, Cruise SL 49c, 2 October 1965; SIO-CO 1594 [wet].
–colony fragment; USA, California, Ventura County, Channel Islands, 10 miles E of
San Nicolas Island, ~33°12'19"N, 119°43'29"W, 91 m; coll. (possible ‘Albatross’ col-
lection, according to Harden; identied in error as Heterogorgia papillosa), 29 March
1917; CAS-IZ 34636 [wet].
Other material, not examined – Japan, Shizuoka, Honshu Island, Suruga Bay,
Omae Zaki, 34°35'00"N, 138°15'00"E; coll. R/V ‘Albatross’, station nos. 3727-3735,
US Fish Commission, 16 May 1900, 62–89 m; USNM 50248 [wet].
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 173
Chromoplexaura marki (Kükenthal, 1913)
Material examined. ~60 lots. USA, California – (?)1 colony; Ventura/Los Angeles
County, 20–22 miles south of San Nicolas Island, rocky, 32°51'00"N, 119°23'45"W,
118–136 m; coll. R/V ‘Velero III’, 1344-41, 11 June 1941; SBMNH 423071 [wet].
–4 fragments (2 lots); Los Angeles County, Santa Catalina Island, 5 miles SE of
Church Rock, 33°16'00"N, 118°13'45"W (end), 215 m on sand, loose washed rock;
coll. R/V ‘Velero III’, Station 1350-41, 12 June 1941; SBMNH 265946 [wet]. –mul-
tiple fragments/colonies (two lots); Los Angeles/Ventura County, o San Nicolas Is-
land, with sponge, rock, 33°16'10"N, 119°24'30"W (end), sponge and rock, 51–56
m; coll. R/V ‘Velero III’, 1123-40, 12 April 1940; SBMNH 423061 [wet]. –3 colo-
nies, 1 fragment; Los Angeles County, 2.5 miles SE of Seal Rocks, Santa Catalina
Island, 33°17'20"N, 118°15'35"W (end), rock, lumpy gray sand, sponge, urchin,
gorgonians, hermits, 158-173 m; coll. R/V ‘Velero III’, station 1429-41, 25 Octo-
ber 1941; SBMNH 265948 [wet]. –1 colony; Los Angeles County, 5 miles, 152° T
from San Pedro Breakwater, 33°38'20"N, 118°12'10"W (end), coarse sand, dead shell,
clay, 31–35 m; coll. R/V ‘Velero III’, 1232-41, 15 February 1941; SBMNH 423066
[wet]. –1 colony; Los Angeles County, 2.25 miles south of San Pedro Breakwater,
33°40'30"N, 118°14'20"W (end), sand, pebble, broken shell, 27 m; coll. R/V ‘Velero
III’, 1207-40, 24 November 1940; SBMNH 423067 [wet]. –multiple fragments
(bearing numerous brittle stars); Santa Barbara County, Santa Cruz Island, 13 miles,
5° T to Gull Island Light, 33°44'00"N, 119°51'00"W, 155 m; coll. R/V ‘Velero IV’,
station 24329-76, 21 January 1976; SBMNH 265935 (with label reading: 24329 BFI)
[wet]. –fragment; Santa Barbara County, 16.5 miles SE X S of South Point, Santa Rosa
Island, 33°38'35"N, 119°58'05” (end) W, rocks, crinoids, brittle stars, 131–140 m;
coll. R/V ‘Velero III’, station 1397-41, 27 August 1941; SBMNH 265936 [wet]. –1
colony; Santa Barbara County, Santa Rosa Island, 6.27 miles, 341° T to Ford Point,
33°48'45"N, 119°59'20"W (end), 104 m; coll. R/V ‘Velero IV’, station 23065-75,
18 October 1975; SBMNH 265937 (with label reading: 23065 6C) [wet]. –1 col-
ony; Santa Barbara, County, San Miguel Island, 3.7 miles, 21° T to Crook Point,
33°56'48"N, 120°21'42"W (end), 118 m; coll. R/V ‘Velero IV’, station 24882-76,
28 April 1976; SBMNH 265949 (with label reading: 24882 CH) [wet]. –1 colony;
Santa Barbara County, south side of Santa Cruz Island, gray sand, shell, 33°57'45"N,
119°38'20"W (end), 67–73 m; coll. R/V ‘Velero III’, 1191-40, 30 September 1940;
SBMNH 423068 [wet]. –1 colony; 2.6 miles, 140° T to Cavern Point, Santa Cruz
Island, 34°04'30"N, 119°34'18"W (end), 76 m; coll. R/V ‘Velero IV’, station 24863-
76, 27 April 1976; SBMNH 423064 [wet]. –1 colony (fragments); Ventura County,
1 mile, 231.5° T from Ventura Pier Light, 34°10'00"N, 118°26'57"W, no depth; coll.
R/V ‘Velero IV’, station 5828-58, 21 August 1958; SBMNH 265947 [wet]. –(?)1
colony; Monterey County, o San Jose Creek Beach (N end), Carmel Submarine Can-
yon, 36°32'00"N, 121°56'00"W, 12–38 m; coll. J H McLean; 1960-1964; SBMNH
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423070 [wet] (formerly part of Los Angeles County Museum collection). –1 small
colony; Monterey County, Point Sur, 36°17'02"N, 121°57'09"W, 56 m, on large rock;
coll. D Starr, NMFS-FED, Santa Cruz, 13 October 2007; SBMNH 235536 [dry]. –1
fragment; Monterey County, southwest of Point Pinos, 36°37'25"N, 121°58'25"W,
rock and sponges, 47 m; coll. R/V ‘Velero III’, 891-38, 8 August 1938; SBMNH
423062 [wet]. –1 colony; Monterey County, Monterey Bay, 1.15 miles, 347° T from
Point Pinos Light, on rock, 36°38'54"N, 121°56'15"W; coll. R/V ‘Velero IV’, station
6437-59, 30 September 1959; SBMNH 423065 [wet]. –fragments; Monterey County,
Monterey Bay, o Point Pinos, rock, crinoids, brittle stars, 36°39'50"N, 121°58'00"W,
89–98 m; coll. R/V ‘Velero III’, station 890-38, 8 August 1938; SBMNH 423063
[wet]. –1 colony; Monterey County, Monterey, 36°47'00"N, 122°02'00"W, ~22 m;
coll. T Burch, BLM Ref. Sta. 360, 18 August 1940, with label (Burch # 40128); SBM-
NH 423060–Neotype [dry]. –1 colony + fragments; Monterey County, Monterey,
36°47'00"N, 122°02'00"W, ~22 m; coll. T Burch, possible BLM Ref. Sta. 360, 18 Au-
gust 1940, with label (Burch # 40128); SBMNH 423072 [wet]. –1 colony; Monterey
County, Monterey, 36°47'00"N, 122°02'00"W, ~30–45 m; coll. T Burch (Acc. No.
3665), BLM Ref. Sta. 360, 18 August 1940; SBMNH 422954 [wet]. –1 colony; Santa
Cruz County, Santa Cruz, 36°57'07"N, 122°00'36"W, 30 m; coll. Kinnetics Lab; Sta-
tion 1, 7 July 1978; with barnacles growing on the colony, where some branches come
o of main “stem;” SBMNH 423069 (previous no. 40612, not in Data Base; speci-
men originally labeled Psammogorgia torreyi). [wet]. USA, Oregon – 1 colony plus
fragments; Lincoln County, 15 nautical miles o coast of Newport, Oregon, Station
NH-15, 44°39'28"N, 124°24'29"W, 85 m; coll. R/V ‘Yaquina’, Cruise Y6906C, 27
June 1969; SBMNH 423073 [wet].
Other material examined. USA, California – multiple fragments; San Luis
Obispo County, 27° o Avila Beach, starting just N of bell buoy, around Souza Rock,
~35°10'35"N, 120°44'16"W, 36 m; coll. R/V ‘NB Scoeld’, Field No. 53-B-25, 9
June 1953; CAS-IZ-96742 [wet]. –1 colony; San Luis Obispo County, ~10 miles S
of Morro Bay, near Point, ~35°22'01"N, 121°02'55"W, 9 m; coll. abalone diver via
scuba (“Ed”), 1964; CAS-IZ 97951 [dry]. –1 colony; Monterey County, between Point
Lobos and Sabrina Point, ~36°31'28"N, 121°57'03"W, 364 m; coll. Capt. Alioti, on
Shrimper ‘Jackie Boy’ (lot 2), 4 March 1972; CAS-IZ 96735 [wet]. –1 colony; Monte-
rey County, Carmel Bay, o San Jose Creek Beach (Monastery Beach), ~36°31'34"N,
121°55'50"W, 38 m; coll. D Sullivan, 20 May 1962; CAS-IZ 96746 [wet]. –1 colony +
1 fragment; Monterey County, Monterey Bay, bearing S 25° E, 5.4 miles o Point Piños
lighthouse, ~36°35'49"N, 122°00'13”W 170 m; coll. USBCF ‘Albatross’, station 4543,
Haul # 816, 1904 SIO/BIC CO 361 [wet]. –1 colony/fragments; Monterey County,
Monterey Bay, bearing N 87° W, 1.74 miles o Point Piños lighthouse, ~36°38'30"N,
121°57'45"W, 64 m; coll. USBCF ‘Albatross’, station 4441, 1904SIO/BIC CO 1810
[wet]. (Both SIO/BIC CO 361 and CO 1810, are labeled P. arbuscula; this is in error,
based on location data.) –1 colony; Monterey County, Monterey Bay, Monterey Bay
Canyon Wall, ~36°52'37"N, 121°56'50"W, 500–600 m; coll. C Mah, R/V ‘Point Sur’,
28 February 1997; CAS-IZ 108905 [wet]. –2 colonies; Monterey County, Monterey
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 175
Bay, Continental Shelf, ~36°52'37"N, 121°56'50"W, 150–160 m; coll. C Mah, R/V
‘Point Sur’, 14 April 1998; CAS-IZ 113071 [wet & dry]. –colonies; Monterey County,
Monterey Bay, Monterey Canyon, canyon wall, ~36°52'37"N, 121°56'50"W, 600–700
m; coll. C Mah, R/V ‘Point Sur’, 14 April 1998; CAS-IZ 113072 [wet]. –1 colony; San
Francisco County, ? Farallon Islands, ~37°44'22"N, 123°03'13"W, no depth recorded;
coll. unknown, with exception of “Trawl no. PC1-1: sample no. 1,” 23 September
1991; CAS-IZ 96747 [wet]. –1 colony; Humboldt County, Blunt's Reef, o Cape
Mendocino, California, ~40°25'56"N, 124°27'28"W; coll. Turkington, May 1910;
det. by Bayer; USNM 51500 [dry]. USA, Oregon and Washington – 1 colony; no
specic location data, 600 m; coll. A Carey Jr., R/V ‘Acona’, Station OTB-36 16 June
1964; CAS-IZ 143903 [wet]. –1 colony and 1 fragment; Coos County, o coast, Co-
quille Bank, ~42°54'23"N, 124°51'16"W, 257 m; coll. P Etnoyer, NOAA “West Coast
Survey, Fall 2010, 3 November 2010; BS004 (CB 50001-002) and BS005 (CB 50001-
003) [wet]. –fragment; o Oregon coast, Heceta Bank, 43°55'58"N, 124°55'55"W,
140 m; coll. Heceta RB-00-05, G Hendler; RV ‘Ronald H. Brown’ (NOAA) and S/V
‘Ropos’, Dive R534-Rk-009, 24 June 2000; LACoMNH Marine Biodiversity Center
processing number 233 [wet]. –fragment; o Oregon coast, Heceta Bank, 43°56'52"N,
124°56'36"W, 189 m; coll. Heceta RB-00-05, G Hendler; RV ‘Ronald H. Brown’
(NOAA) and S/V ‘Ropos’, Dive R535-Bio-2, 3, 4, or 13, 24 June 2000; LACoMNH,
Marine Biodiversity Center processing number 379 [wet]. –1 colony; o Oregon coast,
Heceta Bank, 43°58'07"N, 124°51'10"W, hard bottom covered with invertebrates,
117.9 m; coll. A Valdés by ROV; RV ‘Ronald H. Brown’ (NOAA) and S/V ‘Ropos’,
Dive 615, 10 July 2001; LACoMNH, Marine Biodiversity Center processing number
100 [wet]. –1 colony; o Oregon coast, Heceta Bank, 43°59'16"N, 124°52'59"W,
rocky area, collected with a large rock, 81.1 m; coll. A. Valdés by ROV; RV ‘Ronald H.
Brown’ (NOAA) and S/V ‘Ropos’, Dive 613, 9 July 2001; LACoMNH Marine Biodi-
versity processing center number 85 [wet]. –1 colony; o Oregon coast, Heceta Bank,
44°09'11"N, 124°49'29"W, 81 m; coll. Heceta RB-00-05, G Hendler; RV ‘Ronald H.
Brown’ (NOAA) and S/V ‘Ropos’, Dive R530-Bio-0012, 21 June 2000; LACoMNH,
Marine Biodiversity Center processing number 205 [wet]. –1 colony; o Oregon coast,
Heceta Bank, 44°11'08"N, 124°48'24"W, rocky bottom, collected with rock, 74.3 m;
coll. R Embley by ROV; RV ‘Ronald H. Brown’ (NOAA) and S/V ‘Ropos’; Dive 608,
7 July 2001; LACoMNH, Marine Biodiversity Center processing number 56 [wet]. –2
colonies; o Oregon coast, Heceta Bank, 44°11'14"N, 124°50'03"W, rocky bottom,
collected with rock with gorgonian attached, 81.8 m; coll. N Puniwai by ROV; RV
‘Ronald H. Brown’ (NOAA) and S/V ‘Ropos’, Dive 608, 7 July 2001; LACoMNH,
Marine Biodiversity Center processing number 55 [wet]. –1 colony; o Oregon coast,
Heceta Bank, 44°13'20"N, 124°52'26"W, 98 m; coll. Heceta RB-00-05, G Hendler;
RV ‘Ronald H. Brown’ (NOAA) and S/V ‘Ropos’, Dive R538-Rk-0003, 26 June 2000;
LACoMNH Marine Biodiversity Center processing number 263 [wet]. –2 colonies;
Lincoln County, o Newport, ~44°40'10"N, 124°27'12"W, 110 m; coll. Dr JE Mc-
Cauley, R/V ‘Acona’, no collection date given; CAS-IZ 24639 [wet]. –2 colonies +
fragments; Lincoln County, 112.01 miles SW of Yaquina Head Light, 44°40'00"N,
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126°21'12"W, 2,856 m; coll. A Carey, OSU, Station OTB-30, 20 May 1964; SBM-
NH 423078 [wet]. –1 colony; Eastern N Pacic, Tillamook County, o Oregon
coast, due W of Nehalem Bay State Park, 45°38'55"N, 124°06'03"W, ~80 m; coll.
NFSC, 2006 expedition; CB 34213-093, FRAM 100105460 [wet]. –1 colony, North
Pacic, o Oregon/Washington border, Clatsop County, due W of Tillamook Head,
and mouth of Columbia River, ~46°13'30"N, 124°22'31"W, ~113 m; coll. NFSC,
2006 expedition; CB 34210-023, FRAM 100105455 [wet]. –3 colonies; North Pa-
cic, Grays Harbor County, o Washington State, due W of Quinault Beach Resort,
~47°01'33"N, 124°39'10"W, ~98 m; coll. NFSC, 2006 expedition; CB 34210-025,
FRAM 00105453 [wet]. –1 colony, North Pacic, o Washington coast, Grays Harbor
County, ~47°06'07"N, 124°32'17"W, 73 m; coll. NFSC, WCGS 2006; CB 34210-
020, FRAM 100105450 [wet].
Other material, location not known. WCGS 2006, Oregon – CB 34213-
047, FRAM 100105469 (provided by E Berntson, NOAA Fisheries Oce, Port
Orchard, WA).
Other material, not examined – several colonies; USA, N Pacic Ocean, Cali-
fornia, Monterey County, Carmel, 26 m; [YPM 8877A and YPM 8877B; notation of
“questionable form,” wet].
Muricea californica Aurivillius, 1931
Material examined. ~33 lots. USA, California – (?)1 colony; Orange County, New-
port Bay, 33°35'37"N, 117°52'51"W, on rocks of outer jetty, below extended low tide;
coll. G MacGinitie, October 1946; SBMNH 422390 [wet]. –multiple colonies/frag-
ments; Orange County, Newport Beach, “e Pipeline,” ~33°36'23"N, 117°55'50"W,
depth not recorded; coll. Aquarium of the Pacic, Long Beach (for SBMNH Sea Cent-
er), early summer, 2010; SBMNH 265938 [wet]. –1 colony; Orange County, Newport
Bay, 33°37'11"N, 117°53'41"W, depth not recorded; coll. G Sphon, 08 January 1953;
SBMNH 422920 [DH 451 = SBMNH-35; dry]. –2+ colonies; Orange County, o
Huntington Beach, with a beam trawl, 33°38'30"N, 118°00'20"W, 7–36 m; coll. R/V
‘Velero III’, 1127-40, 20 April 1940; SBMNH 422360 [1 dry, others wet]. –1 colo-
ny; Los Angeles County, San Pedro, Point Fermin, 33°41'00"N, 118°16'00"W; coll.
R/V ‘Velero IV’, station 1578-46, 20 December 1946; data from label, not Station
Log (NOTE: from August 1942-July 1948, no Station Log notes; ship not running
consistently during WWII?); SBMNH 422361[wet]. –fragment; Los Angeles County,
San Pedro, inside of Point Fermin, on shore–rock wall and loose rock, 33°42'45"N,
118°16'45"W; coll. R/V ‘Velero III’, station 1217-40, 30 November 1940; SBMNH
422357 [wet]. –multiple colonies; Los Angeles County, San Pedro, Point Fermin,
33°42'18"N, 118°17'35"W, 13 m; with data station number: 50 3 T147, 17 June
1916; SBMNH 422375[wet]. –several colonies in 3 lots; Los Angeles County, San
Pedro Breakwater, on shore, 33°42'08"N, 118°16'05"W; coll. R/V ‘Velero III’, station
1230-41, 26 January 1941; SBMNH 422358 [wet]. –1 colony; Los Angeles County,
between White's Point and Portuguese Bend, beam trawl–bryozoa, corals, sea mouse,
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 177
gorgonacea, 33°42'54"N, 118°20'07"W (end), 22 m; coll. R/V ‘Velero IV’, station
1644-48, 19 November 1948; SBMNH 422359 [wet]. –1 colony; Los Angeles Coun-
ty, between White's Point and Portuguese Bend, 33°42'54"N, 118°20'07"W, 22 m;
coll. R/V ‘Velero IV’, station 1644-48, 19 November 1948; SBMNH 422919 [dry].
–1 colony (stripped); Ventura County, Rincon Beach, 30°20'10"N, 119°24'35"W, on
beach; coll. N Moore, 2 March 2008; SBMNH 422918 [dry]. –1 colony; Santa Bar-
bara County, Anacapa Island, 34°00'13"N, 119°25'16"W, 100 yards out from Cat
Rock, 17–18 m; coll. B Scronce, and party, 30 October 1962; SMBNH 45564; [wet].
–1 colony; Santa Barbara County, Anacapa Island, 34°00'13"N, 119°25'16"W, 100
yards out from Cat Rock, 17–18 m; coll. B Scronce, and party, 30 October 1962;
SBMNH 45563; [wet]. –1 colony; Ventura County, Port Hueneme, o the north jetty,
34°09'26"N, 119°13'42"W, 11 m; coll. M Conboy and D Sprong, 16 January 1963;
SBMNH 422911 [DH 448 =SBMNH-32; dry]. –3 colonies; Ventura County, Port
Hueneme, 34°09'26"N, 119°13'42"W, 11 m; coll. M Conboy and D Sprong, 16 Janu-
ary 1963; SBMNH 422913 [DH 449 =SBMNH-33, dry]. –2 colonies; Santa Bar-
bara County, Carpinteria, several miles east of Carpinteria State Beach, 34°23'36"N,
119°31'31"W, between splash and high inter-tidal, up on rocks and exposed beach; coll.
EA Horvath, between 27–30, January 2006; SBMNH 422372 & 422373 [wet]. –2
colonies; Santa Barbara County, Carpinteria, o Carpinteria State Beach, 34°23'36"N,
119°31'31"W; coll. C Watters and J Henry, det. EA Horvath; 18 February 2008;
SBMNH 422914 [dry]. –1 fragment; Santa Barbara County, Santa Barbara, Naples
Reef, 34°26'24"N, 119°56'60"W, 15 m; coll. S Anderson and party, 11 November
1998; SBMNH 345324; [wet]. MEXICO, Baja California Sur (Pacic Coast) – 2
colonies + fragments; Cabo San Lucas, 22°52'49"N, 109°54'32"W; coll. unknown,
22 March 1936; SBMNH 422921 [dry]. –1 colony; o urloe Head, on rock with
gorgonids, 27°36'50"N, 114°50'50"W, 15–18 m; coll. R/V ‘Velero III’, station 283-
34, 9 March 1934; SBMNH 422369 [wet]. MEXICO, Baja California Norte (Pa-
cic Coast) – 1 fragment; Isla Cedros, South Bay, on rock along margin of kelp bed,
28°04'45"N, 115°21'05"W, 18–27 m; coll. R/V ‘Velero III’, station 287-34, 10 March
1934; SBMNH 422363 [wet]. –2 colonies; Baja, 6.5 miles SW of Punta San Carlos,
dredge–rocky bottom, 29°33'30"N, 115°35'28"W (end), 36 m; coll. R/V ‘Velero IV’,
station 1944-50, 25 April 1950; SBMNH 422362 [wet]. MEXICO, Baja California
Norte (Gulf of California) – 2 colonies; Roca Consag, 31°12'17"N, 114°33'39"W;
coll. L Findley, 9 July 1973; Nos. 207/208, University of Arizona; SBMNH 422376
[wet]. MEXICO (Gulf of California) – 1 colony; Sonora, South of Isla Tiburon, sand,
shell, 28°43'45"N, 112°17'50"W, 36 m; coll. R/V ‘Velero III’, 566-36, 11 March
1936; SBMNH 422384 [wet]. SOUTH AMERICA – 1 fragment; Ecuador, o Cape
San Francisco, 00°37'10"N, 80°00'30"W, 27 m; coll. R/V ‘Velero III’, station 850-38,
23 February 1938; SBMNH 422427 [dry].
Other material examined. USA, California – San Diego County, La Jolla, Quast
Rock, 21 m; coll. R Kiwala, by scuba, 7 April 1969; SIO/BIC CO 1600 [wet]. –San
Diego County, La Jolla, Quast Rock, 20 m; coll. R Grigg, December 1967; SIO/BIC
CO 1632. –San Diego County, La Jolla, Quast, 20 m; coll. unknown, October 1966;
Elizabeth Anne Horvath / ZooKeys 860: 67–182 (2019)
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SIO/BIC CO 1641. –1 lot; San Diego County, Quast Rock, 65 feet; coll. unknown,
29 October 1965; SIO/BIC CO 1945 [wet]. –2 colonies/fragments; Orange County,
just south and east of Huntington Beach, ~33°36'50"N, 117°59'13"W, 18 m; coll.
OCSD, station T0, Haul 1, 29 February 2012 [wet]. –Los Angeles County, o San
Clemente Island, 32°35'29"N, 118°19'02"W, near Station # 2, in restricted area, by
SCUBA; coll. J Cooper, 11 June 1982; MLML C0110 [wet]. –colonies/fragments;
Los Angeles County, Santa Catalina Island, just outside Blue Cavern Marine Protected
Area, near “e Quarry,” ~33°25'53"N, 118°26'49"W, depth unknown; coll. Aquar-
ium of the Pacic, Long Beach, early summer, 2010 [wet]. –Los Angeles County,
Coast Station, Palos Verdes, o Marineland of the Pacic, 33°43'58"N, 118°25'12"W,
10–15 m; large boulders with sand between; coll. R Grigg, 7 January 1967; SIO/BIC
CO 1596. MEXICO – Baja California Norte, Punta Banda, station 2 (no other data);
20 July 1968; SIOBIC CO 1943 “a1-a4”, and probably “b” as well.
All lots SIO/BIC CO 1860 – CO 1939; most of these lots probably this species,
a few mixed in likely M. fruticosa, but no specic collection location data is readily
available. In the “Limbaugh” Collection housed at NMNH, other collection locations
could include: (USA) the Richeld Oil Island (Ventura County, ~10 miles south of
Santa Barbara, California), Newport Bay (California), in the Huntington Beach Gas
and Electric Steam Plant discharge pipe (California), and from Baja: Gulf of Punta
Final, Turtle Bay and o Cape San Lucas. ere are SIO/BIC CO 1943 and CO 1945,
but these are not included in any ocial listing, and there appeared to be no data to
go with the numbers.
Other material, not examined (but photographed) or examined/photographed
(not collected) – 1 colony; N Pacic Ocean, USA, California, Los Angeles County, San
Clemente, ~33°25'21"N, 117°37'22"W, on the beach; photographer A Nelson, det. EA
Horvath; 1 March 2008. –1 colony; N Pacic Ocean, USA, California, Ventura Cali-
fornia, Santa Barbara Channel Islands, Anacapa Island, ~34°00'19"N, 119°24'55"W,
big tidal pool; photographer N Downend, det. EA Horvath; 14 April 2008.
Muricea plantaginea (Valenciennes, 1846) [= M. appressa Verrill, 1864]
Material examined. ~4 lots. MEXICO, Baja California Sur (Pacic Coast) – 1 col-
ony; Bahia de Santa Maria, 24°42'20"N, 112°14'10"W, 58 m; coll. R/V ‘Velero III’,
station 760-38, 5 January 1938; SBMNH 422418; [dry]. –1 colony; Punta Asun-
ción, 27°07'28"N, 114°17'48"W, 7 m; coll. RH McPeak, 15 October 1980; SBMNH
422417 [dry]. MEXICO, Baja California Norte (Gulf of California) –1 colony; Isla
Partida, o White Rock, 28°55'30"N, 113°05'35"W, 82 m; coll. R/V ‘Velero III’, sta-
tion 557-36, 8 March 1936; SBMNH 422416 [dry]. MEXICO (Gulf of California)
–1 colony; Sonora, Guaymas, Ensenada Carrizal, 27°52'12"N, 110°51'00"W; coll.
P LaFollette; August 1960; SBMNH 422912 [DH 450 =SBMNH-34; Also: # 0413,
and A339, dry].
Other material, not examined. USNM 52301 and USNM 52303 were both
collected in the N Pacic, USA, California, Los Angeles County, o San Pedro; no
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 179
other data given. A third, USNM 94678, may also be this form, as it is from the same
collection location. Additionally, from the Limbaugh collection of specimens housed
at NMNH, locations for possible examples of this species include: Baja California Sur
(Cabo San Lucas; Cape San Lucas Canyon; Bahia de Los Angeles); Baja California
Norte (an area near South Cedros Island; perhaps Punta Final and Turtle Bay) and
USA, California, San Diego County, La Jolla (Scripps submarine canyon). Further
sclerite examinations need to be made.
Muricea fruticosa Verrill, 1868
Material examined. ~14 lots. USA, California – 1 colony (fragmented); Orange County,
Newport Channel, Balboa, shore collected on a –1.6 tide, 33°36'02"N, 117°52'50"W;
coll. R/V ‘Velero III’, station 1224-41, 25 January 1941; SBMNH 422383 [wet]. –1
fragment (albino form or bleached–FG Hochberg); Los Angeles County, Santa Cat-
alina Island, Salta Verde Point, 15 m, 33°19'00"N, 118°25'20–27"W; coll. R Given,
1 May 1967; SBMNH 422389 [wet]. –several colonies; Los Angeles County, Santa
Catalina Island, Big Fisherman's Cove, 33°20'40"N, 118°29'07"W, 3 m; coll. R Setzer,
22 August 1968; SBMNH 422374 [wet]. –1 colony; Los Angeles County, Palos Verdes,
east of Portuguese Bend, 33°43'00"N, 118°19'57"W, 15 m; coll. unknown, 6 Novem-
ber 1949; SBMNH 422428 [dry]. –1 colony; Los Angeles County, Long Beach, Long
Beach Harbor, Back Channel, just east of Terminal Island, 33°45'41"N, 118°13'07"W,
7.6 m; coll. DB Cadien, rocky subtidal station I/S7, 31 March 1980; SBMNH
265945 [wet]. –1 fragment; Los Angeles County, Santa Monica Bay, ~33°56'07"N,
118°27'59"W; coll. Aquarium of the Pacic, Long Beach (for SBMNH Sea Center),
early summer, 2010; SBMNH 265940 [wet]. –2 colonies (1 incomplete); Santa Barbara
County, Santa Cruz Island, Smugglers Cove, 34°01'19"N, 119°32'31"W, 6 m; coll.
B Scronce and party, 24 January 1963; SBMNH 422430 [DH 477 = SBMNH-31,
dry]. –1 fragment; Santa Barbara County, Mohawk Reef, 34°23'40"N, 119°43'48"W,
4 m; coll. S Anderson, by hand, SCUBA, 01 November 1998; SBMNH 345325
[wet]. MEXICO, Baja California Sur (Pacic Coast) –1 colony; Escondido Bay area,
Isla Danzante to Punta Candeleros, 25°47'15"N, 111°15'30"W; coll. R McPeak, 30
March 1981; SBMNH 422429; [dry]. –1 colony; 2 miles, 142° T from urloe Head,
27°35'45"N, 114°49'15"W, no depth reported; coll. R/V ‘Velero IV’, station 11842-
67, 7 December 1967; SBMNH 422910; [wet]. –fragment; Scammons Lagoon, Isla
Piedras, high littoral under at sandstone, 27°46'00"N, 114°15'20"W; coll. W Wil-
liams on Kenyon-Williams Expeditions, station M12, 30 April 1946; SBMNH 422387
[wet]. MEXICO, Baja California Norte (Pacic Coast) – 1 colony; Baja, 6.5 miles
SW of Punta San Carlos, dredge–rocky bottom, 29°33'30"N, 115°35'28"W (end), 36
m; coll. R/V ‘Velero IV’, 1944-50; 25 April 1950; SBMNH 422386 [wet]. MEXICO
(Gulf of California) – 1 colony; Sonora, Guaymas, “Bahia de Conos” (El Carrisito),
27°52'00"N, 110°54'15"W, subtidal, less than 9 m, attached to rocks; coll. P LaFollette,
8-12 August 1960; SBMNH 422916 [DH 452 =SBMNH-36; ? 0336; ? A332, dry]. –1
colony/2 fragments; Sonora, Guaymas, “Bahia de Conos” (El Carrisito) 27°52'00"N,
Elizabeth Anne Horvath / ZooKeys 860: 67–182 (2019)
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110°54'15"W, less than 9 m, attached to rocks; coll. P LaFollette, 5–12 August 1960;
SBMNH 422917 [DH 455 =SBMNH-39; ? A20; ? A332; ?0336, dry]. –4 colonies,
2 other fragments; Sonora, Guaymas, Ensenada Carrizal, 27°52'12"N, 110°51'00"W;
coll. P LaFollette, 5–13 August 1960; SBMNH 422915 [DH 453 =SBMNH-37; ?
PIL-A19; ? A332; ? 0335, dry].
Other material examined. USA, California – colony; Dana Point, N of,
33°32'41"N, 117°48'36"W, 48–53 m; coll. T Matsui, with otter trawl on R/V ‘Agas-
siz’, 29 March 1974; CO/SIO 1628 (M-15, OT-8) [wet/dry?]. –CO/SIO 1939 also
identied as this species, but no other data could be found.
Placogorgia sp. A
Material examined. ~5 lots. USA, California – 1 colony, in fragments; Orange Coun-
ty, Oceanside, 33°10'29"N, 117°23'09"W, Station IP-B1046, caught in shark net, 138
m; coll. B Brophy, 1971; SBMNH 422969 [DH 844 = SBMNH-51; dry]. –mul-
tiple fragments; Los Angeles County, o Santa Monica, South Banks, 34°00'21"N,
118°29'57"W, 82–100 m; coll. Pacic BioMarine, commercial trawl, 10 October
1978; SBMNH 422970 [wet]. –1 colony fragment; Ventura County, California Chan-
nel Islands, NW edge of Hueneme Canyon, 34°02'20"N, 119°18'07"W, 145 m; coll.
NMFS Dive 32, 23 September 2011; SBMNH 235579 [dry]. –fragment; Ventu-
ra County, California Channel Islands, slightly NE of East Anacapa, 34°02'20"N,
119°18'13"W, 140 m; coll. M Love from submersible ‘Delta’, 14 October 2005; with
his number: A6689; SBMNH 422968 [wet]. MEXICO, Baja California Norte (Pa-
cic Coast) – multiple fragments; 5.5 miles S of Islas San Benito, ne green sand,
coarse grey sand, 28°13'18(55)” N, 115°33(35)’15(05)” W (end), 120–147 m; coll.
R/V ‘Velero III’, station 1251-41, 26 February 1941; SBMNH 422966 [wet].
Other material examined – 1 piece; USA, California, Orange County, north-
ern edge of San Gabriel Canyon, o Huntington Beach, trawl, ~33°33'49"N,
118°08'40"W, 113 m; coll. DB Cadien, EMAP Voucher, Survey SCBPP, southern
California Bight, Station 1476 (ID by J Ljubenkov, not collected by MEC), 29 July
1994 [wet].
A review of gorgonian coral species held in the Santa Barbara Museum of Natural History... 181
Appendix 2
Map A1. Distribution of gorgonian “red whip” forms. Includes species that are predominantly seen as a
whip rather than a branched colony as well as several species that, while usually branched, do frequently
display a slender, whip-like body type. Ranges determined from all material examined, with emphasis on
specimens housed in SBMNH collection.
Elizabeth Anne Horvath / ZooKeys 860: 67–182 (2019)
182
Appendix 3
Table A1. Contrasts and comparisons of key “red whip” species and/or species of the genus Swiftia, as
represented in SBMNH collection.
Red Whip
species
Location, S
to N
Location
Depth
Colony
Branching
Colony
Color
Polyp
Spacing
Polyp
Height
Sclerite
Color
Sclerite
Form
Sclerite
Size
Leptogorgia
exilis
Magdalena Bay,
Baja to San
Diego, CA
11 meters
to ?
in,
drooping
branches;
highly
branched
colony
Red/pink
to tan/beige
Polyps
white to
very pale
orange
No more
than
1mm
No more
than 1 mm
Bright
Salmon
Spindles &
Capstans
.03–.09
mm
Leptogorgia
chilensis
N of Magdalena
Bay to Santa
Cruz Is., CA
Approx.
15–80 m
in
branches;
moderately
branched
Orange-
red Polyps
white
1 mm Generally,
almost ush
Bright
Salmon
Spindles &
Capstans
.03-.05
mm
Red Whip
(?”Transitional/
Regional
Endemic”)
San Diego, CA
to o Oregon
coast
?20–2,000
meters
Moderate
thickness to
branches;
slightly
branched to
not branched
Orange-
red Polyps
white
Varies
from 1 to
2 mm
Generally,
from ush
to nearly 1
mm; rarely
taller
Salmon Spindles
& Dbl.
Spinds.
Approx.
0.1 mm
Red Whip
(?”Transitional/
Regional
Endemic”)
San Diego, CA
to o Oregon
coast; possible
extension to
WA coast
Approx.
12–150 m
Moderately
thin
branches,
whip-like;
slightly
branched
Orange-
red Polyps
white
(?palepink)
Varies
from 1 to
2 mm
Generally,
consistently
ush, rarely
taller; on
some,
prominent
Salmon Spindles
& Dbl.
Spinds.
Approx.
0.1 mm
“?Swiftia
Transitional/
Regional
species”
N Los Angeles
County to Point
Conception
Approx.
104–173 m
Single
branches;
also slightly
branched
(if so,
dichotomous)
Bright red
to salmon-
pink Polyps
white
Less than
1 mm
Approx. 1
mm
Salmon Spindles;
very few
Capstans,
Dbl.
Spinds. or
Rods
From
0.04 mm
to nearly
+.16
mm
Swiftia simplex N Los Angeles
County to
Alaska
200–900 m Single
branches;
sometimes
slightly
branched
Pinkish-red
(Brick-red)
Polyps
pinkish-red
No more
than
2mm
Approx. 1
mm
Pinkish-red
(Brick color)
Rods orange
Spindles,
Capstans,
Rods
and Dbl.
Spinds.
.1–.3
mm
Chromoplexaura
marki
Point
Conception
to Cape
Mendocino,
CA (?further
north to WA
state, on to
Alaska)
20–60 m;
possibly
deeper (to
600 m)
Single
branches;
sometimes
slightly to
moderately
branched
Bright red,
orange,
even
pinkish
Polyps
white or
colored
2 mm Nearly ush
to 2 mm
Salmon to
reddish
Spindles,
Capstans,
Ovals
and Dbl.
Spinds;
NO Rods
0.05 mm
to
≅
0.2
mm
Swita
spauldingi
Monterey
Bay, CA to o
Washington
coast (?further
north to Alaska)
40 to at
least 300 m
Moderate
branch
thickness;
branched to
some degree
Orange-
red Polyps
white or
very pale
pink
about
1mm
Nearly ush
to often very
conspicuous,
rounded
Salmon to
pinkish-
orange;
some yellow;
Rods orange
Spindles,
Capstans,
Rods
and Dbl.
Spinds.
About
0.1 mm
Polyp Height: includes both calyx and actual polyp.
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