ZooKeys 350: 75-90 (20 | 3) A peer-revi iewed open-access journa I

doi: 10.3897/zookeys.350.61 17 RESEARCH ARTICLE #ZooKey

WWwWwW.ZOO keys.o rg Launched to accelerate biodiversity research

Two new species of gorgonian octocorals from the
Tropical Eastern Pacific Biogeographic Region
(Cnidaria, Anthozoa, Gorgoniidae)

Odalisca Breedy!?, Gary C Williams**, Hector M Guzman*#

I Centro de Investigacién en Ciencias del Mar y Limnologta, Universidad de Costa Rica. Centro de Investi-
gacién en Estructuras Microscépicas, RO. Box 11501-2060, Universidad de Costa Rica, San José, Costa Rica
2 Department of Invertebrate Zoology and Geology, California Academy of Sciences, 55 Music Concourse Dri-
ve, San Francisco, California, 94118, USA 3 Smithsonian Tropical Research Institute, PO. Box 0843-03092,
Panama, Republic of Panama

T Attp://zoobank.org/BCOD5F1C-C101-4C51-B70B-0C6460B14732
* Attp://zoobank.ore/4BD094B7-F9C6-4FB5-8 9F4-3BAAFF46DDI6
§ Attp://zoobank. org/AD7868EC-2998-4E7E-8BE0-E35BEB31A7FO

Corresponding author: Odalisca Breedy (odaliscab@gmail.com)

Academic editor: L. van Ofwegen | Received 18 August 2013 | Accepted 20 October 2013 | Published 14 November 2013
http://zoobank.ore/C7545ECA-1F50-42FB-99AB-3D40B03B 1 DCF

Citation: Breedy O, Williams GC, Guzman HM (2013) Two new species of gorgonian octocorals from the Tropical
Eastern Pacific Biogeographic Region (Cnidaria, Anthozoa, Gorgoniidae). ZooKeys 350: 75-90. doi: 10.3897/
zookeys.350.6117

Abstract

The gorgoniid Eugorgia is exclusively an eastern Pacific genus. It has a wide geographic and bathymetric
range of distribution, found from California to Pert and extends down to 65 m deep. Two new species are
herein described. The morphological characters were analyzed and illustrated by light and scanning elec-
tron microscopy. Eugorgia beebei sp. n. can be distinguished by its white, ascending, sparse colony growth.
Eugorgia mutabilis sp. n. can be distinguished by its white colony that changes color after collection, and
the conspicuous sharp-crested disc sclerites. From a morphological point of view the new species are
related to the daniana-group, the rubens-group and the siedenburgae-group of Eugorgia; their affiliations,
and the proposal of a new group are discussed. These new species increases the number of species in the

genus to 15, and contribute to the knowledge of the eastern Pacific octocoral biodiversity.

Keywords

Eugorgia, eastern Pacific, gorgonian, soft corals, taxonomy, white species

Copyright Odalisca Breedy et al. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0
(CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

76 Odalisca Breedy et al. / ZooKeys 350: 75—90 (2013)

Introduction

Eugorgia is a gorgonian octocoral (family Gorgoniidae) with 13 valid species. The ge-
nus is considered to be exclusively eastern Pacific and is distributed from southern
California to Pert, and found in oceanic islands. It presents a wide bathymetric range
of distribution, found in shallow waters (down to 40 m), and in the mesophotic re-
gion (down to 65 m) (Breedy and Guzman 2013). Eugorgia is characterized by having
flabellate to bushy colonies with one or multiple planes. Branching is lateral, par-
tially dichotomous, or pinnate-like, often bushy, and combinations of them; branch
anastomosis is absent, but pseudo-anastomosis frequently occurs (Breedy et al. 2009).
Colonies could be attached to hard substrates, debris, and coarse sand or muddy sedi-
ments. Polyps are fully retractile into the coenenchyme in slightly raised to promi-
nent mounds arranged in series of longitudinal rows, or evenly distributed on the
branches. Coenenchymal sclerites are of various types: spindle, disc-spindle, capstan,
and the most dominant form that defines the genus is the characteristic double disc.
Double discs could be incomplete, when the sclerite tubercles have a partial fusion,
or complete, when the fused tubercles of the sclerites form wheels like flying saucers.
Anthocodial sclerites are rarely found. The color of the sclerites is variable: brownish,
orange, red, violet, white, yellow or combinations of these (Breedy et al. 2009). The
colonies are orange, pink, purple, red, white, or yellow, some have with colored rings
around the polyp mounds. ‘They are produced by the arrangement of darker or lighter
color sclerites around the polyp aperture, in some cases they are not surrounding the
polyps, just sparsely distributed giving a sprinkled appearance to the branches. Accord-
ing to the morphological features, the species are proposed to form three groups, the
daniana-group, the amp/a-group and the monospecific rubens-group (see Breedy et al.
2009). A new group characterized by bushy, irregularly pinnate, bicolored colonies
has been proposed for the recently described species Eugorgia siedenburgae Breedy &
Guzman, 2013.

Eugorgia is recognised for their bright colored colonies. The white color has been
reported only for one species, Eugorgia alba Bielschowsky, 1929 in the ampla-group
(Breedy et al. 2009), although white specimens have been observed either in collec-
tions or in the field. Herein we describe two new species that were previously recorded
as color varieties (Breedy et al. 2009, E. Deichmann as a museum label).

Materials and methods

Repository abbreviations

CAS California Academy of Science, California, USA

UCR Museo de Zoologia, Universidad de Costa Rica

STRI = Smithsonian Tropical Research Institute, Panama
USNM_ National Museum of Natural History, Washington, USA

Two new species of gorgonian octocorals from the Tropical Eastern Pacific... Fel.

The specimens used in this study belong to the octocoral collections of the above
cited museums.

Morphological analysis. Preserved specimens were photographed for later de-
tailed observation. Sclerites were obtained by dissolving tissues from branches with
3.5% sodium hypochlorite (household bleach). Sclerites were rinsed many times with
distilled water then 100% ethanol, dried, and mounted on stubs for scanning electron
microscopy (SEM), and coated with 60-80 nm Pt/Pd. They were observed and pho-
tographed using an Hitachi 3700 SEM operated at 15kV. For light microscopy, clean
sclerites were mounted in water or glycerin and observed and photographed using an
Olympus LX 51 inverted stereoscope.

We followed Verrill (1868) and Breedy et al. (2009) for characters assessment. The
terminology used in descriptions mostly follows Bayer et al. (1983), Breedy and Guz-
man (2002), and Breedy et al. (2009).

Morphological characters of colonies and the most abundant sclerite types of the
species examined here are presented in Table 1. The most abundant sclerites in these
species are disc-spindles and double discs that present various degrees of tubercle fu-
sion. The illustrations of the sclerites are presented in different planes to provide a bet-
ter idea of their architecture (Figs 3, 5). Comparison is made with the closest morpho-
logical groups, in this case, the daniana-, siedenburgae-, and rubens-groups (Table 1).

Taxonomy

Class Anthozoa Ehrenberg, 1831
Subclass Octocorallia Haeckel, 1866
Order Alcyonacea Lamouroux, 1812
Family Gorgoniidae Lamouroux, 1812
Genus Exugorgia Verrill, 1868

Eugorgia beebei sp. n.
http://zoobank.org/8B75AC49-5089-4BEF-BE80-B76198B9DOE8
http://species-id.net/wiki/Eugorgia_beebei

Figs 1-3

Eugorgia rubens var. beebei (species name suggested by E. Deichmann in a museum

label, unpublished)

Material examined. Holotype. CASIZ 75783, ethanol preserved, Los Frailes, Baja
California sur, México, 52 m, coll. R. Adcock, 18 June 1979.

Paratypes. MCZ 36106, dry, Paita, Piura, Peru, no more data available. USNM
56879, ethanol preserved, El Alto, Piura, Peri, 1860-1815 m but depth data dubious
(F. M. Bayer’s note on label: “specimen probably from previous shallow station’), An-

ton Bruun Cruise , 18B, Sta. 766, 4°10'S., 81°27'W, 9 September 1966.

78 Odalisca Breedy et al. / ZooKeys 350: 75—90 (2013)

Type locality. Baja California sur, México.

Diagnosis. Ascending colony sparse growing, branching irregularly pinnate, and
multiplanar, subdividing up to 11 times, some pseudo-anastomosis present. Promi-
nent polyp-mounds up to 0.70 mm tall, dome-shaped, arranged irregularly, and close-
ly placed on branchlets, and very distant on thick branches. Colony and sclerites white.
Spindles and disc-spindles up to 0.14 mm in length, double discs up to 0.07 mm long,
and 0.05 mm wide. Anthocodial rods absent.

Description. Holotype 24 cm tall, and 20 cm wide, ascending, sparse grow-
ing, (Fig. 1A). Branching irregularly pinnate, and multiplanar, several pseudo-
anastomosis occurs in branchlets and branches (Fig. 1A—B). Main stem 4 mm
diameter at base, slightly compressed, and short, about 80 mm long arising from a
fragment of holdfast, 0.6 mm diameter. Main stem gives off several branches and
stumps. The three main branches, 3.0-4.0 mm in diameter, emerging at angles of
45—90°and producing secondary branches subdividing and giving off thin branch-
lets, up to 2.5 mm diameter, including polyp-mounds. Branchlets irregularly ar-
ranged, separated 5-16 mm, and giving off 2 or 3 lateral, secondary branchlets, of
same thickness and arrangement. Colony branching up to 11 times. Unbranched
terminal twigs blunt, and reaching up to 50 mm long (Figs. 1A—B). Polyp-mounds
prominent, up to 0.7 mm height and 1 mm in diameter, dome-shaped, with slit-
like apertures, arranged irregularly, close together along the branchlets, and very
distantly distributed or absent along the thick branches (Fig. 1B). Holdfast de-
void of polyps. Colony white (Fig. 1A—B). Sclerites of coenenchyme white, mostly
double discs (Fig. 1C). Spindles and disc-spindles, up to 0.14 mm long and 0.04
mm wide, with 4 or 5 whorls of warty tubercles, the ends mostly blunt (Fig. 3A).
Double discs up to 0.07 mm long, and 0.05 mm wide (Fig. 3B). Crosses about
0.08x0.06 mm, scarce on samples (Fig. 3C). No anthocodial sclerites present in
the samples.

Variability. Paratype MCZ 36106 reaches up to 34 cm tall, and 31 cm wide, the
main stem 0.7 mm diameter, slightly compressed, and short, about 1.0 cm long arising
from an oval holdfast 3.2 cm diameter, and 0.2 cm thick (Fig. 2A—B). Sclerites as in
the holotype (Fig. 2C). The other examined specimens are smaller, but very consistent
in all aspects with the holotype.

Discussion. The morphology of the colony, i.e., irregular-pinnate branching
and prominent polyps, immediately segregates the new species from the ampla-
group, and suggest a similarity with daniana- , rubens- and siedenburgae-groups.
Eugorgia beebei and E. siedenburgae differ from the species in the daniana-group,
including Eugorgia mutabilis sp. n. (described below), firstly, in the colony growth,
which is sparse and ascending in E. beebei sp. n. but bushy and profuse in E£.
siedenburgae, not flabellate as it is in the daniana-group species. Secondly, it dif-
fers in the branching patterns because branchlets in the daniana-group form flat
pinnate fronds with pinnae projecting in the same plane. That is not the case in
E. beebei and E. siedenburgae where the secondary branchlets stick out in several,
irregular planes.

Two new species of gorgonian octocorals from the Tropical Eastern Pacific...

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Figure |. Eugorgia beebei (CASIZ 75783) holotype. A entire colony B detail of branches C S

tH

scleri

tes.

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80 Odalisca Breedy et al. / ZooKeys 350: 75—90 (2013)

Figure 2. Eugorgia beebei (MCZ 36106) paratype. A entire colony B detail of branches; SEM sclerites.

Two new species of gorgonian octocorals from the Tropical Eastern Pacific... 81

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Figure 3. Eugorgia beebei (CASIZ 75783) holotype, SEM coenenchymal sclerites. A spindles and disc-
spindles B double discs C cross.

Eugorgia siedenburgae and E. rubens form monospecific groups, they differ espe-
cially in the colony growth. The rubens-group have pink, sparse and laterally branched
colonies, and the siedenburgae-group, have bushy, bicolored colonies (Breedy and
Guzman 2013).

Eugorgia beebei and E. siedenburgae are very similar in sclerite content (Table 1),
but they are different especially in the growth form and in the color. The conspicuous
bushy colony immediately distinguished it from E. beebei; additionally, E. beebei has

Odalisca Breedy et al. / ZooKeys 350: 75—90 (2013)

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Two new species of gorgonian octocorals from the Tropical Eastern Pacific... 83

thicker branches and branchlets than EF. siedenburgae; the polyp mounds are pointed
and higher in the latter, and are more rounded in £. beebei. Branchlets in E. beebei are
longer than in E. siedenburgae (Table 1).

Remarks. We found the paratype in the MCZ (36106), labelled in Elisabeth
Deichmann’s handwriting (Ardis Johnston, pers. comm.) as a variety of E. rubens,
however, she certainly had not published anything on this genus, thus the variety or
the species was never established. The specimen was part of an MCZ public exhibition,
the only data we have are the locality. It is probable that this specimen examined by
Deichmann came from the Zaca expedition of 1937 and 1938. We do not consider
E. beebei as a variety of E. rubens because they differ in the traits that have been shown
to be informative to separate species in the genus: color, branching pattern, colony
growth and sclerite content (see Table 1), as mentioned above.

Etymology. This new species of Eugorgia is named for explorer/naturalist Wil-
liam Beebe (1877-1962) who studied the marine fauna at numerous locations along
the west coast of Central America from Mexico to Columbia during the Templeton
Crocker Zaca expedition between 1937 and 1938. Beebe subsequently wrote the book,
Book of Bays, which chronicles the five month expedition (Gould 2004).

Distribution. Presently known from Piura, Pert and Baja California, but it is very
likely that it exist along all along the geographic range. The depth range is 50-60 m,
it is possible the range could extends deeper, but not as deep as reported for paratype
USNM 56879, which is probably a mistake, as was remarked by F. M. Bayer (former
USNM curator).

Eugorgia mutabilis sp. n.
http://zoobank.org/B552B4F0-50F4-4E16-BC00-5A3CB2C565C5
http://species-id.net/wiki/Eugorgia_mutabilis

Figs 4—6

Material examined. Holotype. UCR 2297, ethanol preserved, Burbujas, between
Los Potreros and Playa Arenitas, Puerto Jiménez, Golfo Dulce, Costa Rica, 11 m, O.
Breedy, 9 May 2013. Paratypes. UCR 2298, same data as the holotype. UCR 2272,
2276, fragments, ethanol preserved, Burbujas, 12 m, C. Sanchez, May 2012. UCR
2299, fragment, ethanol preserved, Roca Matapalo, Cabo Matapalo, Golfo Dulce, 20
m, O. Breedy, 6 February 2009. STRI 408, dry, Isla Seca Grande, Gulf of Chiriqui,
Panama, 20 m, H. Guzman, 26 August 2002. STRI 444, dry, Isla Jicarita, Gulf of
Chiriqui, 20 m, H. Guzman, 29 August 2002. STRI 511, dry, Isla Ladrones, Gulf of
Chiriqui, 15 m, H. Guzman, 14 April 2003. STRI 1073, Santa Cruz, Coiba Island,
Panama, 15 m, H. Guzman, 27 April 2007. STRI 1076, Twin Peaks, Coiba Island, 15
m, H. Guzman, 27 April 2007. STRI 1168, fragment, ethanol preserved, Bajo Hacha,
Coiba Island, 20 m, O. Breedy, 16 April 2009. STRI 1122, ethanol preserved, La
Blanca, Oxaca, Mexico, 46 m, R. Abeytia, 29 August 2004.
Type locality. Puerto Jiménez, Golfo Dulce, Costa Rica, 11 m.

84 Odalisca Breedy et al. / ZooKeys 350: 75—90 (2013)

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Figure 4. Eugorgia mutabilis (UCR 2297) holotype. A entire colony B detail of branches C SEM sclerites.

Powe es

Diagnosis. Broad, stout, flabellate colony, main branches sinuous, branching ir-
regularly pinnate, subdividing 5—7 times, no anastomosis present. Prominent polyp-
mounds closely spaced and irregularly distributed around branches and branchlets
(Figs. 4, GA-C). Colony white, pale pink when alive (Fig. 6A—C), dark grayish when
dry or ethanol preserved. Change in color after collection very conspicuous. Longi-
tudinal grooves evident along branches and branchlets. Sclerites white. Spindles and
disc-spindles up to 0.15 mm long, double discs mostly 0.05—0.08 mm long. Sclerite
discs sharp, serrated and prominent. No anthocodial rods found.

Description. Holotype 30 cm tall, and 47 cm wide; colony broad, flabellate, very
flexible. Branching irregularly pinnate. Main stem 6 mm diameter, laterally flattened,
and short, 14 mm long. Holdfast oval, 40 mm diameter without polyps. Main stem

Two new species of gorgonian octocorals from the Tropical Eastern Pacific... 85

subdividing in 5 sinuous main branches. Main branches slightly flattened on plane of
colony, 3—4 mm in diameter emerging at angles of about 45°, bifurcating and diverg-
ing producing five flat pinnate fronds of long pinnate branchlets (Fig. 4A—B). Pinnae
short, up to 8 mm long, and 1.5—2 mm diameter including polyp-mounds, close to-
gether 1-3 mm apart. Branching up to 7 times. Unbranched terminal twigs blunt, and
reach up to 8 mm long (Fig. 4A—B). Longitudinal grooves distinct along branches and
pinnate branchlets, evident in living and preserved specimens. Polyps white (Fig. 4B).
Polyp-mounds prominent, up to 0.7 mm height and 0.8 mm in diameter, arranged
mostly in lateral rows along the branchlets and separated by the longitudinal grooves,
and more sparsely and irregularly distributed along the thicker branches (Fig. 4B).
Colony white to pale pink when alive, gray to dark grayish in ethanol/dry preservation
(Fig. 4A). Change in color very conspicuous possibly by liberation of black pigments
after collection. Sclerites of coenenchyme white. Sclerite discs conspicuous mostly
sharp, serrated and prominent (Fig. 4C, 5A—B). Disc-spindles 0.08—0.12 mm long,
and up to 0.06 wide with 4—5 whorls of discs (Fig. 5B); spindles and disc-spindles,
longer and thinner up to 0.15 mm long and 0.05 mm wide, with 5—7 whorls of warty
tubercles, the ends acute, blunt, or both (Fig. 5A). Double discs up to 0.08 mm long,
and 0.05 mm wide with prominent discs (Fig. 5B), some almost complete (Fig. 5B).
No crosses, capstans or anthocodial sclerites present in samples.

Variability. The specimens present some variation in sclerite color, white sclerites
being dominant, but some pale yellow hues could be observed in the samples. In all
other aspects they agree with the holotype, including the change of color from bright
white alive or recently collected to grayish when fixed. It is interesting that after collec-
tion, the specimens discharge a black pigment that turns the water black or the alcohol,
and the colony becomes gray. The specimens from Mexico represent the deeper record;
they have been observed down to 50 m, meaning that the range of depth extends from
40 to 50 mas it is for E. rubens, E. siedenburgae, and E. beebei, but the morphology
of the colony and sclerites remain the same described for E. mutabilis. Current flow
and depth are some of the environmental factors that could influence interspecific
variability in octocorals (Fabricius and Alderslade 2001), but in this case no effect was
observed. The morphology of the colony and sclerite content are persistent along the
depth range.

Remarks. The species was mentioned before as a variety of E. daniana: ‘a white va-
riety has been observed in shoals in Costa Rica and Mexico occurring together with the
red form’ (Breedy et al. 2009). However, after examination of many specimens from
various localities and depths, we found enough differences with respect to E. daniana,
and to the other species in the group, especially in the color and the conspicuous scle-
rites, to establish E. mutablilis as a new species.

Habitat. The new species is found on rocky substrates, in general with other spe-
cies of gorgonians, including FE. daniana, but in some places, it is the only Eugorgia
present (Fig. GA—C). Other gorgonians found normally inhabiting the same localities
are Pacifigorgia irene, P. stenobrochis, Leptogorgia alba, and Carijoa riseii, which were
very abundant in the type locality. A variety of associated invertebrates were found

86 Odalisca Breedy et al. / ZooKeys 350: 75—90 (2013)

Figure 5. Eugorgia mutabilis (UCR 2297) holotype. SEM coenenchymal sclerites. A, B spindles and
disc-spindles C double discs.

87

Two new species of gorgonian octocorals from the Tropical Eastern Pacific...

Pore
(isp

Figure 6. Exvgorgia mutabilis, in situ colonies. A, B Burbujas, Puerto Jiménez, Golfo Dulce, Costa Rica,

12 m deep (photographs by C. Sanchez) C Montafia Rusa, Islas Contreras, Panama, 32 m deep.

88 Odalisca Breedy et al. / ZooKeys 350: 75—90 (2013)

on the holotype and paratype UCR 2298, including ophiuroids, Ophiotrhix sp., and
crustaceans, shrimps, Periclimenaeus sp. and abundant crabs, Orthochela sp.

Etymology. The specific epithet is from Latin, mutablilis, changeable, in allusion
to the change in color after collecting.

Discussion. Eugorgia mutabilis belongs to the daniana-group with a characteristic
flabellate colony composed of flat pinnate fronds, and irregular pinnate-branching pat-
tern, and prominent polyp mounds. ‘The white color of the colony and sclerites of EF.
mutabilis separates it from the rest of the group. However, the new species is similar to
E. daniana in some features, e.g. maximum number of branches, branchlet distance,
polyp distribution (see Table 1), but the sclerite composition is very different. The
dominant sclerites in FE. mutabilis have very sharp crested discs that are very consistent
in all specimens revised from Mexico, Costa Rica and Panama, and along the depth
range. These type of sclerites are distinct also from the ones in E. beebei, and in E.
siedenburgae.

Distribution. Records from Costa Rica, México and Panama suggest a wide dis-
tribution, at least from Mexico to Panama, but this has to be further explored. ‘The
deepest record in Panama is 35 m, in Costa Rica 25 m, and in Mexico 50 m. Thus,
the occurrence of this species from 11 to 50 m deep also suggests a large bathymetric
range of distribution.

Final remarks

There are not many morphological characters to differentiate species in octocorals,
normally the combination of growth form, and size and color of colony and scler-
ites are the features used for identification. However, as it was acknowledged above,
colony shape can vary within species in response to environmental conditions, light
availability, wave exposure and currents (Fabricius and Alderslade 2001). Thus, species
delimitation are sometimes difficult to draw. For this reason, the study of a reasonable
amount of samples and habitats will be of aid to decide about morphological species
and varieties, especially when accessibility to collection sites is possible. However, in
some cases, only old museum specimens are available but identification and recogni-
tion for biodiversity accounts is imperative.

Molecular studies have been taken to understand boundaries between species and
interspecific or intraspecific phylogenetic relationships; however, a complete molecular
phylogeny has not been achieved due to the lack of molecular markers with adequate
resolution to distinguish species (or sometimes genera) (Cairns and Bayer 2005, Mc-
Fadden et al. 2006, McFadden et al. 2010, Williams 2013, Wirshing et al. 2005).

Morphological phylogenetic studies in Eugorgia (Breedy et al. 2009) have shown
that color of the colonies and sclerites, similar colony morphology, and ecological
habits are significant characteristics to separate clades. These characters alone are not
informative enough. The combination of the characters used to separate the new spe-

Two new species of gorgonian octocorals from the Tropical Eastern Pacific... 89

cies that are analyzed here, showed consistency when compared with other related taxa
in the genus.

Four groups of Eugorgia have been proposed for the eastern Pacific, four species in
the daniana-group, one in the rubens-group, one in the siedenburgae-group, and eight
in the ampla-group.

Although, £. beebei sp. n. was first mentioned as a variety of E. rubens, we have
demonstrated that it is a different species that does not even fit in the rubens-group, or
in the other related groups (daniana-, siedenburgae-). Thus, a new group is here pro-
posed, the deebei-group characterised by white colonies and sclerites, and with ascend-
ing, sparse colony growth.

The diagnostic characters of the daniana-group are herein modified adding the
white color for colony and sclerites, to include E. mutabilis sp. n. in the group. It
is important to mention that occurrence of complete double discs sclerites in these
species-groups is scant, and the closest example to this type are the ones in the new
species. Actually, after examined many specimens especially, in the daniana-group,
the occurrence of neat complete double discs is not frequent. It seems a more com-
mon character of the amp/a-group. The recently described Eugorgia ahorcadensis Soler-
Hurtado & Lépez-Gonzalez, 2012 from Ecuador should be placed in the latter group.
However, there is not enough evidence to separate this species from Eugorgia nobilis
Verrill, 1868, from which it represents a morphological variety, basically with longer
branches and darker sclerites. Therefore, herein we synonymize it with E. nobilis.

Presently, a total of 15 valid species are recorded for the eastern Pacific but this num-
ber should increase when more geographic areas and bathymetric ranges are explored. ‘This
research is a contribution to the knowledge of the eastern Pacific octocoral biodiversity.

Acknowledgements

We are grateful to anonymous reviewers for critical improvements of the manuscript.
We thank Adam Baldinger (MCZ) and Stephen Cairns (USNM) for specimen loans,
and Ardis Johnston (former collection manager MCZ) for providing information
about the MCZ collection, and Rita Vargas (UCR), and Carlos Guevara (STRI) for
laboratory and field work. We thank Rosalinda Abeytia for collecting paratype STRI
1122, and Celeste Sanchez for providing the submarine pictures in Figure 3. We thank
Federico Bolafios, director of the Field Biology and Conservation in the Tropics Pro-
gram (Universidad de Costa Rica/ University of Manitoba, Canada) for funding the
2013 collection trip to Golfo Dulce.

This project was partially funded by the Vicerrectoria de Investigacion, Universi-
dad de Costa Rica, and Smithsonian Tropical Research Institute.

Specimens from Costa Rica were collected under the MINAE 0890 permit, and
from Panama under the permits: DAPVS-02-2007, DAPVS-01-2008, SE/A-71-12,
SE/A-16-13 of the Panama Environmental Authority.

90 Odalisca Breedy et al. / ZooKeys 350: 75—90 (2013)

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