Content uploaded by Monika Bryce
Author content
All content in this area was uploaded by Monika Bryce on Jun 10, 2015
Content may be subject to copyright.
160
Accepted by E. Pante: 2 Apr. 2015; published: 26 May 2015
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2015 Magnolia Press
Zootaxa 3963 (2): 160
–
200
www.mapress.com
/
zoota xa
/
Article
http://dx.doi.org/10.11646/zootaxa.3963.2.2
http://zoobank.org/urn:lsid:zoobank.org:pub:86A305B7-2B9C-403E-8FC0-3420EFB13F52
Digitate and capitate soft corals (Cnidaria: Octocorallia: Alcyoniidae) from
Western Australia with reports on new species and new Australian geographical
records
MONIKA BRYCE
1, 4,5
, ANGELO POLISENO
2
, PHILIP ALDERSLADE
3
& SERGIO VARGAS
2
1
Western Australian Museum, Locked Bag 49, Welshpool DC, Perth, Western Australia 6986, Australia.
E-mail: Monika.bryce@museum.wa.gov.au
2
Department of Earth & Environmental Sciences, Paleontology & Geobiology
Ludwig-Maximilians-University Munich, Richard-Wag-
ner Strasse 10, 80333 Munich,Germany.
3
CSIRO Marine and Atmospheric Research, GPO Box 1538, Hobart, Tasmania 7001. E-mail: phil.alderslade@csiro.au
4
Queensland Museum, PO Box 3300, South Brisbane BC, Queensland 4101, Australia.
5
Corresponding author
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162
Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163
Systematics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163
Family Parasphaerascleridae McFadden & Ofwegen, 2013 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163
Genus Parasphaerasclera McFadden & Ofwegen, 2013 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163
Parasphaerasclera grayi (Thomson & Dean, 1931) new record . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163
Parasphaerasclera kimberleyensis sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166
Family Alcyoniidae Lamouroux, 1812 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169
Genus Eleutherobia Pütter, 1900. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169
Eleutherobia australiensis sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169
Eleutherobia imaharai sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
Eleutherobia somaliensis Verseveldt & Bayer, 1988 new record . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175
Eleutherobia splendens (Thomson & Dean, 1931) new record . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180
Genus Sphaerasclera McFadden & Ofwegen 2013 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
Sphaerasclera flammicerebra (Williams, 2003) new record . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .182
Genus Paraminabea Williams & Alderslade, 1999 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
Paraminabea aldersladei (Williams, 1992) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184
Paraminabea cf. aldersladei Williams & Alderslade, 1999 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .186
Molecular Phylogeny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198
Abstract
We report on digitate and capitate Octocorallia within the genera Parasphaerasclera McFadden & Ofwegen, 2013,
Eleutherobia Pütter, 1900, Sphaerasclera McFadden & Ofwegen, 2013, and Paraminabea Williams & Alderslade, 1999
from tropical Western Australian waters. Three new species (Parasphaerasclera kimberleyensis, Eleutherobia australien-
sis, Eleutherobia imaharai) are described, with a discussion of their taxonomic placement in the light of a recent treatment
of the genus Eleutherobia and related taxa by McFadden & Ofwegen (2013). In addition, range extensions for three spe-
cies are reported, Parasphaerasclera grayi (Thomson & Dean, 1931) known from Indonesia and the Pacific Ocean,
Eleutherobia somaliensis Verseveldt & Bayer, 1988 from Somalia, and Eleutherobia splendens (Thomson & Dean, 1931)
recorded from Indonesia and the Philippines. Additionally, one new Australian geographical record (Sphaerasclera flam-
micerebra) (Williams, 2003) with a known distribution from Palau to Mauritius, has been included. We complement mor-
Zootaxa 3963 (2) © 2015 Magnolia Press
·
161
SOFT CORALS OF WESTERN AUSTRALIA
phological taxonomy with molecular data (mtMutS, 28S rDNA) to analyse and clarify phylogenetic placement of these
species. The mitochondrial mtMutS phylogeny supported Eleutherobia, Paraminabea, Parasphaerasclera and Sphaeras-
clera as distinct monophyletic genera. Phylogenetic analyses based on 28S rDNA lacked resolution and were largely un-
resolved. Additionally, the molecular data corroborated our proposed morphological hypothesis of the placement of the
new species P. kimberleyensis sp. nov. with no anthocodial armature in the genus Parasphaerasclera, and the assignment
of the new species, E. australiensis sp. nov. and E. imaharai sp. nov., with distinct polyps sclerites in the genus Eleuther-
obia.
Key words: Eleutherobia, Paraminabea, Parasphaerascleridae, Parasphaerasclera, Sphaerasclera, Kimberley, Indian
Ocean
Introduction
The tropical marine environment of Western Australia extends northwards from the Tropic of Capricorn at 23.44°
S and encompasses several marine bioregions, which reflect the region’s diverse macro-scale habitat structure. The
soft coral fauna of this large area, while largely unknown, is represented in many of the region’s habitats, including
inlets, estuaries, coastal and off-shore reefs and islands. Deep water soft coral communities are even less well
known having received little collecting effort. Since 2009 the Western Australian Museum has been undertaking
comprehensive biodiversity surveys off the Kimberley coast, in the state’s far north (Bryce & Sampey 2014).
Examination of soft coral species from these recent collections, paired with historical material from the Western
Australian Museum’s collection, has provided a base-line dataset on soft coral species occurrence and community
composition.
In this account we focus on small, digitate and capitate species of the genera Eleutherobia, Parasphaerasclera,
Sphaerasclera and Paraminabea; genera that are often represented by one or few species (Eleutherobia 11,
Parasphaerasclera 6, Sphaerasclera 1, Paraminabea 10), and often have very narrow geographic distributions
(Table 1; McFadden & Ofwegen 2013). To date only three relevant species have been recorded from Australian
waters; Eleutherobia rubra (Brundin, 1896) was described from the north west coast of the continent (Verseveldt &
Bayer 1988), Parasphaerasclera zanahoria (Williams, 2000) was more recently recorded from the north east coast
on the Great Barrier Reef by one of us (Alderslade, unpublished) and Paraminabea aldersladei Williams, 1992
from the north east and north west coasts of Australia (Williams 1992; Williams & Alderslade 1999). More than
half of the described species of these genera were collected during two expeditions, the Siboga Indonesia
Expedition in 1899 (Thomson & Dean 1931); and the German deep-sea expedition in 1906 (Kükenthal 1906a).
Considering the limited survey effort in remote areas in comparison to more readily accessible areas it is
hypothesized that the number of recorded deep-water species will increase with further collecting effort. The same
holds true for the small species found in shallower habitats, which have adapted to a cryptic lifestyle preferring low
light areas, such as overhangs and caves (Williams & Alderslade 1999; Williams 2000, 2001, 2003). Sampling in
these rather inaccessible, high energy areas, in combination with the small colony size and apparent low
abundance, makes them difficult to find. This present contribution describes three new species of soft coral within
the genera Parasphaerasclera and Eleutherobia. The species, P. kimberleyensis sp. nov. was collected under an
overhang at Long Reef, north Kimberley. E. australiensis sp. nov. was collected from deep-water off the Dampier
Archipelago and E. imaharai sp. nov. off North West Cape, which is in the Pilbara region of Western Australia.
Further, we report on range extensions of a number of species of Eleutherobia, Parasphaerasclera, Sphaerasclera
and Paraminabea, and discuss the taxonomic placement of all included species which we establish using an
integrative taxonomic approach (see Dayrat, 2005; Will et al. 2005; McFadden et al. 2014), combining
morphological examinations with molecular phylogenetic analyses derived from two independent markers (i.e.
mtMutS and 28S rDNA).
Abbreviations
WAM Western Australian Museum, Locked Bag 49, Welshpool DC, WA 6986, Australia.
QM Queensland Museum.
AIMS The Australian Institute of Marine Science.
BRYCE ET AL.
162
·
Zootaxa 3963 (2) © 2015 Magnolia Press
CSIRO The Commonwealth Scientific and Industrial Research Organization.
ZMB Museum für Naturkunde, Berlin.
Material and methods
Material was collected by SCUBA off the Kimberley region in Western Australia (Fig. 1). Upon collection,
specimens were photographed and preserved in 70 % ethanol prior to further examination. Historical trawled
material from the WAM collection was also examined. Sclerites were prepared for both light and scanning electron
microscopy (SEM) by cutting small pieces of the specimen from five different regions (polyps, surface of the polyp
region, surface of the base, interior of the polyp region, interior of the base) and dissolving them in sodium
hypochlorite (13 % available chlorine). After the organic material had dissolved, the loose sclerites were rinsed
with water and dried on a glass slide for further investigation. Durcupan ACM was used as mounting media for
permanent slides (Fabricius & Alderslade 2001: 40). Twenty-five sclerites were measured per sclerite type. SEM
images were taken using a Hitachi TM-1000. Images of the specimens were made using an Olympus SZ-CTV
dissecting microscope with TSview software (TUCSEN) and a NIKON D 300 camera. Type material has been
registered and deposited in the Western Australian Museum, Perth.
FIGURE 1. Location of the Kimberly region and collection sites.
DNA extraction of ethanol-preserved specimens was done using Macherey-Nagel NucleoSpin
®
Tissue kit
(M&N, Duren, Germany). DNA quality was examined on a 1.5 % agarose gel and the quantity (ng/µL) was
measured on a Nanodrop 1000. The 5’ end of the mitochondrial gene mtMutS and a partial fragment of the 28S
nuclear ribosomal gene were amplified according to McFadden et al. (2011) and McFadden & Ofwegen (2012) or
Zootaxa 3963 (2) © 2015 Magnolia Press
·
163
SOFT CORALS OF WESTERN AUSTRALIA
internal primers were used to amplify smaller, overlapping fragments spanning the region of interest in both genes.
For mtMutS, we used the following internal primers: MSH-met1F (5’-ATGAGCCARATACCTATGC-3’),
MSH3010F (5’-GGATAAAGGTTGGACTATTATAG-3’; (Thoma et al. 2009) and MSH3101R (5’-
GATATCACATAAGATAATTCCG-3’; Sanchez et al. 2003). PCR programs used were adapted from previous
protocols published elsewhere (Sanchez et al. 2003; McFadden et al. 2004; Brugler & France 2008; Vargas et al.
2014). PCR products were purified by precipitation, adding one volume of 20 % (w/v) polyethylene glycol 8000 in
2.5molL
-1
NaCl and sequenced in both directions using the same primers used for PCR. Sequences were deposited
at the European Nucleotide Archive under accession numbers HG970065-970091. DNA sequences were aligned
against other octocoral sequences available in GenBank using MUSCLE (Edgar, 2004) with default options in
GENEIOUS 6.0.5 (Drummond et al. 2012). The programs RAxML 7.2.8 (Stamatakis, 2006) and MrBayes 3.1.2
(Ronquist & Huelsenbeck 2003) were used to infer Maximum Likelihood (ML) and Bayesian phylogenetic trees,
respectively. For ML analyses we use GTRGAMMA model with bootstrap analyses including 1000 pseudo-
replicates, the support values were obtained with the rapid bootstrap algorithm (Stamatakis, 2008). Rate variation
was modeled using a discrete gamma distribution with 4 categories (Yang, 1994). For the Bayesian, the best-fit
model was selected using the Akaike Information Criterion (AIC) implemented in jModeltest 2.1.3 (Darriba et al.
2012). The analyses ran for 10,000,000 generations under the best fitted model (i.e. GTR + I + G) with a sample
frequency of 500 and the first 25 % of sampled trees were discarded as burn-in. Convergence between the runs
(stationarity of parameters) was assessed using the standard deviation of the split frequencies. We assumed
convergence was achieved when this value reached 0.01. The 28S tree was re-rooted considering the stoloniferans
Cornularia pabloi and Cornularia cornucopiae as outgroups. Due to the lack of mtMutS sequences for C.
cornucopiae, the mitochondrial tree was re-rooted using C. pabloi as the only outgroup. All the nodes with
bootstrap values < 70 % and a posterior probability < 0.95 were collapsed into polytomies using TreeGraph 2.3.0-
425 beta (Stöwer & Müller 2010).
Results
Systematics
Family Parasphaerascleridae McFadden & Ofwegen, 2013
Diagnosis. Soft corals with a digitiform, digitate or lobate growth form. Usually with a bare stalk, which can be
indistinct. Polyps monomorphic, retractile, producing small, coenenchymal mounds when retracted. Permanent
calyces absent. Sclerites of colony surface and interior predominantly radiates and/or tuberculate spheroids,
occasionally along with rodlets and crosses. Sclerites permanently coloured. Polyp sclerites absent.
Azooxanthellate. (adapted from McFadden & Ofwegen 2013).
Genus Parasphaerasclera McFadden & Ofwegen, 2013
Type species. Alcyonium rotiferum Thomson, 1910 by original designation.
Diagnosis. As for the family.
Parasphaerasclera grayi (Thomson & Dean, 1931) new record
(Figs. 2A,B, 3, 4; Tabs. 1, 2)
Nidalia grayi Thomson & Dean, 1931: 37, Pl. 2, Fig. 2.
Eleutherobia grayi Verseveldt & Bayer 1988: 33–34, Figs. 24, 25; Williams 2001: 210–216, Figs. 1–10. Benayahu et al. 2004:
550 (recorded only); Dautova & Savinkin 2009: 4–10, Figs. 3–7.
Parasphaerasclera grayi McFadden & Ofwegen 2013: 70, 71, 78.
Material examined. WAM Z54774, six whole specimens, Station 69/K11, unnamed outcrop NW Black Rocks,
BRYCE ET AL.
164
·
Zootaxa 3963 (2) © 2015 Magnolia Press
close to White Island, NW Australia, 14.9741° S, 124.3974° E, SCUBA, depth 12 m, coll. M. Bryce, 17 October
2011.
FIGURE 2. A, B, Parasphaerasclera grayi WAM Z54774; C–E Parasphaerasclera kimberleyensis sp. nov. C, D, holotype
WAM Z59789; E, paratype WAM Z67195.
FIGURE 3. Parasphaerasclera grayi, WAM Z54774, sclerites from the surface of the polyparium.
Description. In total six upright, unbranched colonies were collected. They are digitiform, symmetrical with a
wide base tapering distally towards the rounded apex of the polyparium (Fig. 2A,B) and were always growing in
pairs, arising from a common, polyp-free, one mm thick, encrusting holdfast. The colonies vary in size, but are in
general very similar in shape, colour, arrangement of polyps and sclerite composition. The polyp-free basal portion
of each colony is very short and occupies only 10 % of the total colony length. Polyps are monomorphic, large,
Zootaxa 3963 (2) © 2015 Magnolia Press
·
165
SOFT CORALS OF WESTERN AUSTRALIA
retractile, quite numerous, evenly distributed over the colony and are translucent when fully extended and the
coenenchymal mounds associated with the polyps are narrow. Pair one consists of one colony that is 31 mm in total
length, with an apex 4 mm in diameter and a base 10 mm in diameter, together with a smaller colony that is 5 mm
in total length with an apex 4 mm in diameter and a base 9 mm in diameter. Pair two consists of one colony that is
25 mm in total length, with an apex 5 mm in diameter and a base 10 mm in diameter, together with a smaller colony
that is 13 mm in total length with an apex 3 mm in diameter and a base 6 mm in diameter. Pair three consists of one
colony that is 19 mm in total length, with an apex 5 mm in diameter and a base 8 mm in diameter, together with a
smaller colony that is 9 mm in total length with an apex 5 mm in diameter and a base 7 mm in diameter.
FIGURE 4. Parasphaerasclera grayi, WAM Z54774, sclerites: A, surface of the stalk; B, interior of the polyparium; C,
interior of the stalk.
BRYCE ET AL.
166
·
Zootaxa 3963 (2) © 2015 Magnolia Press
In the surface of the polyparium, including the coenenchymal mounds, the majority of sclerites are 6, 7- and 8-
radiates and rodlets 0.04–0.08 mm long and crosses 0.03–0.07 mm in diameter (Fig. 3). The smaller radiates are
tuberculate capstan-like forms and the longer rodlets are essentially 8-radiates with more distant prominences and a
smooth, often long, shaft.
The stalk surface is densely spiculated with irregular radiates 0.05–0.08 mm long, crosses 0.05–0.08 mm long
and rodlets (Fig. 4A).
The interior sclerites of the polypary are 6- and 7-radiate capstans 0.04–0.09 mm long, crosses 0.06–0.09 mm
in diameter, and elongated rod-like sclerites 0.10–0.11 mm long (Fig. 4B). The interior of the stalk coenenchyme is
populated with sclerites consisting of rod-like forms 0.10–0.11 mm long with a smooth shaft and large warty
prominences, and robust crosses around 0.09 mm in diameter are also occasionally present (Fig. 4C).
Polyp sclerites are absent.
Colour. In situ the colonies were uniformly bright rusty orange with small, bright, light orange, rounded
coenenchymal mounds. The polyp-free encrusting holdfast was uniformly rusty red. The polyps were transparent
with bright white tentacles. The colour did not change on deck or in alcohol. In preserved specimens the surface
and internal coenenchymal sclerites are brownish dark red to pale-ochre, the sclerites of the polyp mounds are pale-
ochre, and the interior coenenchyme is pink.
Habitat. Steep, forward reef slope extending up to the reef crest at 14 metres. On the crest of the slope were
small coral outcrops and the underlying base rock was covered with encrusting corals and soft coral communities,
dominated by Sinularia and Sarcophyton as well as several species of gorgonian. The slope was dissected by
narrow surge grooves with vertical sides marked by caves and ledges extending to 20 metres. A small group of
Parasphaerasclera grayi colonies was found attached to the wall in an overhang at 12 metres depth amongst other
scattered soft corals.
Remarks. This species has a wide distribution in the Indo-Pacific region, but it is a new record in Australia
(Table 1). More recently Benayahu et al. (2004) reported P. grayi as a new record from Taiwan, and in 2013 it was
collected by WAM from Christmas Island (Richards, unpublished). McFadden & Ofwegen (2013) described
material from Palau as P. aff. grayi and also discussed in detail the high variability of shape and size of the sclerites
of specimens of P. grayi that have been described in the literature, stating the possibility that multiple species might
be involved. They also pointed out the similarities between their P. aff. grayi and the P. grayi of Williams (2001)
from the Solomon Islands, as well as the differences between their material and the P. grayi from Vietnam
described by Dautova & Savinkin (2009) and the lectotype of P. grayi described by Verseveldt & Bayer’s (1988).
Specimens from Palau and the Solomon Islands have tuberculated rods that lack a smooth waist. Our material
agrees best with the well-illustrated and detailed re-description of the Vietnamese material and with the description
of the lectotype based on the presence of distinctive smooth rod-like forms.
Parasphaerasclera kimberleyensis sp. nov.
(Figs. 2C–E, 5, 6; Tabs. 1, 2)
Material examined. Holotype: WAM Z59789, Station 44/K10, Long Reef, northern Kimberley, NW Australia,
13.88867° S, 125.74942° E, SCUBA, depth 10 m, coll. M. Bryce, 20 October 2010. Paratype: WAM Z67195, 1
specimen, same data as the holotype.
Description. The holotype is a digitiform colony, with a wide base, which tapers distally towards the rounded
apex of the polyparium (Fig. 2C–D). It is 15 mm tall, 2 mm in diameter at the apex and 4 mm in diameter across the
holdfast. The colony has no real stalk as polyps occur almost to the very base. The polyp-free basal portion
occupies less than 20 % of the total colony length. The polyps are rather sparse and evenly distributed. They are
monomorphic, large, and when completely retracted, leave a prominent coenenchymal mound on the surface of the
polyparium. All polyps are tightly retracted—as they were at the time of collection—and occupy most of the
interior of the polyparium.
The majority of the sclerites from the upper part of the polyparium are tuberculate capstans, some slightly
club-shaped, 0.05–0.13 mm long (Fig. 5), along with crosses 0.06–0.08 mm diameter and a few rodlets (see Fig.
5a). An apparent triradiate sclerite (see Fig. 5b) has been included to point out where errors are possible, as this is
actually the broken end of a sclerite with the fracture side down. The authors believe that the sclerites shown in Fig.
24b–f for E. grayi in Versevedt & Bayer (1987), especially d–f, are most probably of this nature.
Zootaxa 3963 (2) © 2015 Magnolia Press
·
167
SOFT CORALS OF WESTERN AUSTRALIA
FIGURE 5. Parasphaerasclera kimberleyensis sp. nov., holotype WAM Z59789, sclerites: surface of the polyparium (a =
rodlet; b = broken end of a sclerite with the fracture side down).
BRYCE ET AL.
168
·
Zootaxa 3963 (2) © 2015 Magnolia Press
FIGURE 6. Parasphaerasclera kimberleyensis sp. nov., holotype, WAM Z59789, sclerites: A, surface of the stalk (a = club);
B, interior of the stalk (b = cross).
The sclerites of the coenenchymal mounds of P. kimberlyensis n. sp. are similarly shaped to those of the
polyparium. There are no sclerites in the interior of the polyparium.
The base of the colony is densely spiculated with small radiates and tuberculate rods 0.02–0.10 mm long (Fig.
6A). Rare clubs 0.08–0.13 mm long are also represented in the base of the colony (see Fig. 6Aa). The interior of the
base is less densely populated with sclerites, and contains mainly spindles, clubs and irregular forms 0.04–0.15 mm
long (Fig. 6B). Crosses 0.09 mm in diameter are also present in the interior of the base (see Fig. 6Bb). The
tubercles of all sclerites are ornamented with granules that tend to be elongate and arranged in rows or united to
form ridges. This is most obvious in the interior stalk sclerites. Polyp sclerites are absent.
Colour. In situ the colonies were uniformly bright yellow-orange with bright red rounded coenenchymal
mounds. The polyp-free base was uniformly bright yellow and the polyps were white. The colour did not change on
Zootaxa 3963 (2) © 2015 Magnolia Press
·
169
SOFT CORALS OF WESTERN AUSTRALIA
deck or in alcohol. In preserved specimens the surface and internal coenenchymal sclerites are pale-yellow to
colourless, the sclerites of the polyp mounds are red, and the interior coenenchyme is white.
Etymology. Named for the type locality, Kimberley, Western Australia.
Habitat. A steep and fractured fore-reef slope with a near vertical wall ascending from 20 metres to a depth of
4 metres. The wall is heavily pocketed with small caves and deep, steep-sided fissures. At 20 metres there are large
rocky outcrops forming long reef-gullies, which are almost devoid of life, probably due to the heavy siltation.
Sediment between the rocky outcrops is very fine and smothering. The Parasphaerasclera kimberleyensis colonies
were found in a small group attached to the wall in an overhang at ten metres depth amongst other scattered soft
corals.
Variability. The paratype is very similar to the holotype in shape, colour, arrangement of polyps and sclerite
composition, but is smaller. It is also digitiform, with a wider base and tapering distally towards the rounded apex
of the polyparium. It is 9 mm tall, 3 mm in diameter at the apex and 4 mm in diameter across the holdfast (Fig. 2E).
Remarks. Parasphaerasclera kimberleyensis sp. nov. does not demonstrate a close resemblance to any
currently described species within the genus. Morphologically, it is distinguished by the shape of the colony and
the shape and colour of the sclerites, and also by the colony colour, the bright yellow colony being offset by the
bright red polyp mounds, ,but there is some superficial resemblance to P. zanahoria (Williams, 2000), which was
originally described from Tonga in the South Pacific. More recently P. zanahoria was recorded from the Great
Barrier Reef by one of us (Alderslade, unpublished). Like P. kimberleyensis sp. nov., P. zanahoria, which is
uniformly orange, is characterised by having some sclerites in the form of large crosses, but in that species the
crosses have finely tapered and acutely tipped rays. In comparison P. kimberleyensis sp. nov. has crosses with
thorny, rounded rays, and, in addition, all the radiates are distinctly more thorny than those in P. zanahoria.
Family Alcyoniidae Lamouroux, 1812
Diagnosis. Membranous or more or less fleshy, massive colonies. The latter with a basal part, the stalk, and a distal
part bearing the anthocodiae, the polyparium. Sclerites are often less than one mm long, but they can be over 10
mm. They include spindles, clubs, radiates, and double heads. Polyps monomorphic or dimorphic and the
autozooids are retractile. This family has a more or less global distribution. It is especially speciose in the Indo-
Pacific, but so far absent in the shallow waters of the tropical western Atlantic (Williams 2003; Fabricius &
Alderslade 2001).
Genus Eleutherobia Pütter, 1900
Type species. Eleutherobia japonica Pütter, 1900 by monotypy; = E. rigida (Pütter, 1900).
Diagnosis. Colonies usually small when contracted with symmetrical, conical to cylindrical polyparium, but
branched, lobate, or clavate growth forms may occur. Polyps are monomorphic with large anthocodiae, which are
retractile into low rounded to conspicuous coenenchymal mounds. The polyp-free basal portion is usually short,
but can occupy up to half of the colony. Found mostly in deep water or restricted to caves and overhangs in shallow
waters, and often in small groups. Sclerites include radiates, capstans, double heads, spindles, spheroids; rod-like
forms or crosses are sometimes present. Anthocodial sclerites present, arranged in points or collaret and points
(McFadden & Ofwegen 2013; Fabricius & Alderslade 2001; Verseveldt & Bayer 1988).
Eleutherobia australiensis sp. nov.
(Figs. 7A–C, 8, 9; Tabs. 1, 2)
Material examined. Holotype: WAM Z31488, one sectioned colony, Station PF06/S1-200/R2, 190 km NW of
Dampier, Pluto Gas Field, NW Australia, 19.9352°–19.9308°S, 115.3261°–115.3288° E, epibenthic sled, depth,
200 m, coll. B.F. Cohen, 8 December 2005. Paratype: WAM Z66778, one whole specimen, same data as holotype.
Description. The holotype is a tapering digitiform colony, with a small lobe (Fig. 7A–C) and it is attached to a
BRYCE ET AL.
170
·
Zootaxa 3963 (2) © 2015 Magnolia Press
piece of hard coral skeleton. It is 50 mm tall, 4 mm in diameter at the apex and 10 mm in diameter across the
holdfast. The polyp-free basal portion is very short and occupies only 10 % of the total colony length (Fig. 7A).
Polyps are large, monomorphic, evenly distributed over the polyparium and are completely retracted leaving
distinct, rounded coenenchymal mounds on the surface.
In the surface of the polyparium the majority of the sclerites are spindles, 0.10–0.35 mm long, with pointed
ends, and ovals 0.08–0.22 mm long (Fig. 8A). Sclerites of the coenenchymal mounds are small, spindle-like bodies
0.10–0.15 mm long (Fig. 8B). The sclerites of the interior coenenchyme of the polyparium are few, being, thin
spindles up to 0.4 mm long (Fig. 8C).
FIGURE 7. A–C, Eleutherobia australiensis sp. nov., holotype, WAM Z31488; D, Eleutherobia imaharai sp. nov., holotype,
WAM Z13252; E, (photo courtesy Y. Imahara) Eleutherobia dofleini (Kükenthal, 1906) “Type”, ZMB 6524; F, Eleutherobia
somaliensis, WAM Z31487; G, Eleutherobia splendens, WAM Z23988-1; H. Sphaerasclera flammicerebra, WAM Z31480.
The stalk surface is densely spiculated mainly with warty ovals, short, plump spindles 0.10–0.30 mm long with
tapering ends and some irregular bodies (Fig. 9A). The sclerites of the interior coenenchyme of the stalk are thin
spindles, 0.10–0.35 mm long, plump spindles, 0.10–0.25 mm long, with the ends tapering to a point and a few
irregular, somewhat triangular forms of similar size (Fig. 9B). All of the above sclerites have high, prickly,
complex warts
The armature of the polyps consists of a collaret eight to ten rows deep and points of six to eight pairs of
curved flattened spindles 0.25–0.45 mm long (Fig. 8D). The tentacles contain spikey rods up to 0.07 mm long (Fig.
8E; only smaller sclerites up to 0.05 mm are figured due to SEM preparation difficulties). No introvert sclerites are
present.
Colour. The preserved colonies are light grey with small orange, rounded coenenchymal mounds. The wider,
polyp-free encrusting base is uniformly light grey. The interior coenenchyme is light brown. Surface and internal
coenenchymal sclerites are colourless, while the sclerites of the coenenchymal mounds are pale-ochre to reddish.
Zootaxa 3963 (2) © 2015 Magnolia Press
·
171
SOFT CORALS OF WESTERN AUSTRALIA
FIGURE 8. Eleutherobia australiensis sp. nov., holotype, WAM Z31488, sclerites: A, surface of the polyparium; B,
coenenchymal mounds; C, interior of the polyparium; D, polyps; E, tentacles.
Etymology. The species is named for being collected in Australia.
Habitat. The specimens were sampled from 200 m along the continental slope utilising an epibenthic sled, but
the exact habitat is unknown.
Variability. The paratype is digitiform, with a wide base, and tapering distally towards the rounded apex of the
polyparium. It is 45 mm in total length and 3 mm in diameter at the apex. The holdfast is laterally flattened, 15 mm
wide and 4 mm thick. The specimen is attached to a piece of hard coral skeleton.
Remarks. The new species is characterised by coenenchymal mounds which are orange because of the
coloured sclerites, anthocodial armature of flattened spindles, distinct oval sclerites and long warty spindles in the
polyparium surface, and the occurrence of plump spindles with tapered, pointed ends in the lower part of the
colony. The ovals and the plump, tapering spindles found in the new species are very like those found in E. dofleini
(Kükenthal, 1906), however, in that species these sclerites occur in both the lower and upper parts of the colony.
Also, the shape of the colony is very different (see Verseveldt & Bayer 1988: Fig. 19a) and the warts on the
sclerites are lower and far less prickly.
Eleutherobia imaharai sp. nov.
(Figs. 7D; 10, 11; Tabs. 1, 2)
Material examined. WAM Z13252, one whole specimen, Station 1031302, North West Cape, 190 km north west
of Dampier, NW Australia, 21.2802° S, 114.0606° E, epibenthic sled, depth 200–250 m, coll. AIMS NW Cape
survey, 13 March 2001.
Description. The colony is essentially two digitiform lobes with an irregular surface, arising from a common
BRYCE ET AL.
172
·
Zootaxa 3963 (2) © 2015 Magnolia Press
base, each slightly tapering to a rounded summit (Fig 7D). The largest lobe is about 4 mm wide at its base and
projects 16 mm above the holdfast, and the smaller lobe extends 3 mm from the base of the larger lobe before it
bends upwards and extends a further 9 mm. The aspect of the holdfast seen in Fig. 7D is about 15 mm wide. Apart
from the holdfast, the numerous monomorphic polyps are evenly distributed over most of the colony, and they are
all retracted within low to moderate coenenchymal mounds.
FIGURE 9. Eleutherobia australiensis sp. nov., holotype, WAM Z31488, sclerites: A, surface of the base; B, interior of the
base.
Zootaxa 3963 (2) © 2015 Magnolia Press
·
173
SOFT CORALS OF WESTERN AUSTRALIA
FIGURE 10. Eleutherobia imaharai sp. nov., holotype, WAM Z13252, sclerites: A, surface of polyparium; B, interior of
polyparium; C, collaret and points; D, tentacle rachis; E, pinnules; F, tentacle rachis sclerites in situ; black arrow showing
position of pinnules.
BRYCE ET AL.
174
·
Zootaxa 3963 (2) © 2015 Magnolia Press
FIGURE 11. Eleutherobia imaharai sp. nov., holotype, WAM Z13252, sclerites: A, surface of the base (a = capstan); B,
interior of the base.
In the surface of the polyparium the sclerites include spindles, up to 0.29 mm long, with acute ends, and ovals
0.08–0.16 mm long; some of latter may have very large, complex warts and one or two sharply tapering ends (Fig.
10A). The sclerites of the interior of the polyparium are markedly narrow, acute spindles 0.20–0.40 mm long (Fig.
10B).
The polyp armature is formed of slightly curved spindles 0.20–0.55 mm long (Fig. 10C). The collaret is about
eight to ten rows deep and the points contain five to six pairs of obliquely arranged sclerites. The tentacles contain
densely packed rods with a curved end (Fig. 10D). They are up to 0.30 mm long, their length becoming smaller
towards the tentacle tip, and they are obliquely arranged in two rows covering the aboral and lateral faces of the
tentacles (Fig. 10F). The pinnules contain a few small spindles, up to 0.12 mm long, with relatively simple
tubercles (Figs. 10E, F(arrowed)). A few introvert spindles of a similar size and shape as the pinnule sclerites are
also present.
Zootaxa 3963 (2) © 2015 Magnolia Press
·
175
SOFT CORALS OF WESTERN AUSTRALIA
The surface of the common base is densely spiculated, mainly with warty ovals, 0.10–0.20 mm long (Fig.
11A). Short, plump spindle-like forms, up to 0.25 mm in length, with sharply tapering ends are also present at the
surface of the base, as are a few crosses and capstans (see Fig. 11Aa). The majority of the sclerites in the interior of
the common base are spindles up to 0.30 mm long, which commonly have one or both ends acute (Fig. 11B). Ovals
and a few irregular forms of similar size, that may have pointed processes, also occur.
Colour. The preserved colony has a cream interior and exterior and the sclerites are colourless.
Habitat. The specimens were sampled from a muddy-rubble environment between 200–250 m depth along the
continental slope utilising an epibenthic sled.
Remarks. The sclerites of the surface and interior of the new species are of the same form as those of
Eleutherobia dofleini as described and illustrated by Verseveldt & Bayer (1988: 29, Figs. 19, 20), by Imahara et al.
(2014: 89, Figs. 24, 25) in their new book on the octocorals of Sagami Bay, and to a lesser extent by Utinomi
(1954: 45, Fig. 2) when describing colonies collected off Minabe. However, the colony form of E. dofleini, which
is only known from Sagami Bay to Tosa Bay, Japan, is quite different to that of the new species, having a narrow,
commonly branched polypary that generally has pronounced polyp mounds of sufficient size to be termed
“Kelche” or “Polypenkelche” by Kükenthal and “calyces” by Imahara et al. and Verseveldt & Bayer. There is also
a difference in the polyp body armature of the new species. The latter lacks any intermediate sclerites between the
points while E. dofleini has two (Verseveldt & Bayer 1988: Fig. 20A1).
In order to check the exact nature of the sclerites from the tentacles and the interior of the polypary in E.
dofleini, neither of which were figured by Verseveldt & Bayer, we approached Dr Leen van Ofwegen, the Naturalis
Biodiversity Center, Leiden and Dr Yukimitsu Imahara, Biological Institute of Kuroshio, for assistance. Dr van
Ofwegen kindly examined the microscope slides used by Verseveldt and Bayer (1988) but found that there were no
preparations present of either character. Dr Imahara examined Utinomi’s specimen of E.dofleini from Tanabe Bay,
Kii Pininsula, Japan (Utinomi 1960) and found that the tentacles contain small, somewhat scale-like, curved,
flattened spindles with scalloped edges and not curved bars as in the tentacles of the new species, and that the
sclerites of the interior of the polypary are not at all long and thin but are shorter and somewhat stouter, like those
illustrated in Fig. 25C of the new book on Sagami Bay (Imahara et al. 2014). He also confirmed the difference in
colony form by sending an image of colony ZMB 6524 labelled “Nidalia dofleini Kük. Type” (see Fig. 7E) and
added the information that the pinnule sclerites of a colony from Tanabe Bay used by Utinomi (1960) are slender
rods that are quite unlike those of the new species.
Etymology. It is with pleasure that we name this new species after our colleague, Dr Yukimitsu Imahara, in
recognition of his assistance with our enquiries regarding E. dofleini, his continuing work on Japanese octocorals,
and especially the publication (with Drs Fumihito Iwase & Hiroshi Namikawa) of the extremely valuable book on
the Sagami Bay representatives of this faunal group.
Eleutherobia somaliensis Verseveldt & Bayer, 1988 new record
(Fig. 7F; 12; Tabs. 1, 2)
Eleutherobia somaliensis Verseveldt & Bayer, 1988: 39–40, Figs. 18g,h; 33b; 34.
Material examined. WAM Z31487, two whole specimens, grooved, digitate, 25 and 41 mm tall, Station PF06/S1-
200/R2, 190 km NW of Dampier, Pluto Gas Field, NW Australia, 19.9308°–19.9352° S, 115.2261°–115.2288° E,
epibenthic sled, depth 200 m, coll. B.F. Cohen, 8 December 2005; WAM Z12201, one whole specimen, branched
with two side branchlets, 65 mm tall, Station 1031302, North West Cape, 21.4666° S, 114.1016° E, epibenthic sled,
depth 200–250 m, coll. AIMS North West Cape survey, 13 March 2001.
Description. Two of the colonies are unbranched, and the third is branched. The colonies are stiff, grooved,
with a wider base and taper distally towards the rounded apex of the cylindrical polyparium (Fig. 7F). The colonies
are between 25–65 mm tall, 3 mm in diameter at the apex and between 6–11 mm in diameter across the holdfast.
No distinct stalk is present. Polyps are monomorphic, numerous and mostly irregularly distributed over the colony.
In some areas of the base the polyps are arranged in longitudinal rows, in other places they are absent. In the
preserved specimens all polyp bodies are retracted.
The sclerites in the surface of the polyparium (Fig. 12A) are mostly 8-radiate derivatives, between 0.08–0.10
BRYCE ET AL.
176
·
Zootaxa 3963 (2) © 2015 Magnolia Press
mm long, with a medial waist, the larger ones approaching clubs, together with some longer spindles up to 0.38
mm long. The sclerites of the interior of the polyparium are spindles and needles, 0.23–0.50 mm long, (Fig. 12B).
The surface of the base is densely spiculated with small rods and crosses, 8-radiate derivatives, sub-spheroidal
forms and plump spindles with sharply tapering ends. The sclerites are about 0.10-0.2 mm long (Fig. 12C). The
sclerites in the interior of the base are warty spindles up to 0.20-0.30 mm long (Fig. 12D). They are similar to the
sclerites in the interior of the polyparium, but are slightly shorter and wider. All of the polyparium and basal
sclerites have very prickly warts.
The polyp armature consists of crown and points and is formed of slightly spiny spindles around 0.03 mm long
(Fig. 12E). The tentacles contain flattened sclerites, the larger ones curved, up to 0.25 mm long (Fig. 12F).
FIGURE 12. Eleutherobia somaliensis WAM Z31487, sclerites: A, surface of polyparium; B, surface of the base; C, interior of
the polyparium; D, interior of the base; E, polyps; F, tentacles.
Zootaxa 3963 (2) © 2015 Magnolia Press
·
177
SOFT CORALS OF WESTERN AUSTRALIA
TABLE 1. Distribution and occurrences of former species and currently valid species of Eleutherobia. Data in this table are based on records from the literature. (NR): new
record in Australia.
Species Distribution Depth (m) Source Literature
Alcyonium Linnaeus, 1758
A. studeri (Thomson, 1910) South Africa 42–121 Verseveldt & Bayer 1988; Williams 1992; Williams & Little 2001;
Williams 2003
A. variabile (Thomson, 1921) South Africa 13–468 Williams 1986; Williams 1992; Williams 2003
Eleutherobia Pütter, 1900
E. australiensis sp. nov. NW Australia 200
E. dofleini (Kükenthal, 1906) Japan 69–250 Kükenthal, 1906; Thomson & Dean 1931; Utinomi 1954; Utinomi
1957; Williams 2003; Verseveldt & Bayer 1988; Imahara et al. 2014
E. imaharai sp. nov. NW Australia 200–250
E. duriuscula (Thomson & Dean, 1931) Indonesia 69–204 Thomson & Dean 1931; Verseveldt & Bayer 1988; Williams 2003
E. flava (Nutting, 1912) Japan 174 Nutting 1912; Verseveldt & Bayer 1988; Williams 2003; Imahara et
al 2014
E. grandiflora (Kükenthal, 1906) Japan shallow Kükenthal 1906; Utinomi 1957; Verseveldt & Bayer 1988; Williams
2003
E. rigida (Pütter, 1900) Japan 24–150 Utinomi 1954; Utinomi 1957; Verseveldt & Bayer 1988; Williams
2003; Imahara et al. 2014
E. rubra (Brundin, 1896) Japan; USA; NW Australia 80–128 Thomson & Dean 1931; Utinomi 1957; Verseveldt & Bayer 1988;
Williams 2003
E. somaliensis Verseveldt & Bayer, 1988 Somalia; NWAustralia (NR) 70–200 Verseveldt & Bayer 1988; Williams 2003
E. splendens (Thomson & Dean, 1931) Indonesia; Philippines; Australia
(NR)
204–511 Thomson & Dean, 1931; Verseveldt & Bayer 1988; Williams 2003
……continued on the next page
BRYCE ET AL.
178
·
Zootaxa 3963 (2) © 2015 Magnolia Press
TABLE 1. (Continued)
Species Distribution Depth (m) Source Literature
E. unicolor (Kükenthal, 1906) Japan 70–73 Kükenthal 1906; Utinomi 1957; Verseveldt & Bayer 1988; Williams
2003
E. sumbawaensis Verseveldt & Bayer, 1988 Indonesia 69 Verseveldt & Bayer 1988; Williams 2003
E. vinadigitaria Williams & Little, 2001 South Africa 52–86 Williams & Little 2001
Parasphaerasclera gen.n. (McFadden & van Ofwegen, 2013)
P. albiflora (Utinomi, 1957) Japan 45 Utinomi 1957; Williams 2003
P. aurea (Benayahu & Schleyer, 1995) South Africa 24–36 Benayahu & Schleyer 1995; Williams & Little 2001; Williams 2003;
McFadden & Ofwegen 2013
P. grayi (Thomson & Dean, 1931) New Guinea; Solomon Islands;
Indonesia; Japan, Taiwan;
Vietnam; Australia (NR);
5–73 Thomson & Dean 1931; Verseveldt & Bayer 1988; Williams 2001;
Benayahu et al. 2004 Williams 2003; McFadden & Ofwegen 2013
P. kimberleyensis sp. nov. NW Australia 10
P. nezdoliyi (Dautova & Savinkin, 2009) Vietnam 12–27 Dautova & Savinkin 2009
P. rotifera (Thomson, 1910) South Africa 27–120 Verseveldt & Bayer 1988; Williams & Little 2001; Williams 1992;
Williams 2003; McFadden & Ofwegen 2013
P. zanahoria Williams, 2000 Tonga; NE Australia 25–30 Williams 2000; Williams 2003
Sphaerasclera gen.n. (McFadden & van Ofwegen, 2013)
S. flammicerebra (Williams, 2003) Palau; New Caledonia;
Mauritius; Australia (NR)
142–450 Williams 2003; McFadden & Ofwegen 2013
Zootaxa 3963 (2) © 2015 Magnolia Press
·
179
SOFT CORALS OF WESTERN AUSTRALIA
TABLE 2. Morphological comparison of the Western Australian digitate and capitate species of Parasphaerasclera, Eleutherobia, Sphaerasclera and Paraminabea.
*Plump spindles with sharply tapering ends. N.R.: new record; Sp. nov: new species.
Species Status in
Australia
Colour in EtOH Growth form Introvert
sclerites
Collaret
sclerites
Tentacle
sclerites
Polyparium
surface (mm)
Polyparium
interior (mm)
Stalk
surface (mm)
Base
interior (mm)
I. Parasphaerasclera
P. grayi N.R. rusty orange with bright
orange coenenchymal
mounds
digitiform absent absent absent 6–, 7, – 8-radiates,
rodlets, crosses:
0.03–0.08
6–, 7–radiates,
crosses: 0.04–0.09;
rods: 0.010–0.011
irregular radiates,
crosses, rods: 0.05–
0.08
crosses, rods:
0.10–0.11
P. kimberleyensis Sp. nov. bright yellow with red
coenenchymal mounds
digitiform absent absent absent capstans: 0.05–0.13;
crosses 0.06–0.08;
rodlets: 0.05
absent radiates, rods:
0.02–0.10
radiates, crosses:
0.07–0.09; spindles,
clubs, irregular bodies:
0.04–0.15
II. Eleutherobia
E. australiensis Sp. nov. grey with orange
coenenchymal mounds
digitiform absent present present ovals : 0.08–0.22;
spindles: 0.10–0.35
thin spindles up to
0.40
ovals, *spindles:
0.10–0.30
thin spindles: 0.10–0.35;
*spindles: 0.10–0.25
E. imaharai Sp. nov. cream digitiform to
digitate
few or
absent
present present ovals: 0.08–0.16
*spindles 0.20–0.29
spindles up to 0.40 ovals, rare crosses:
0.10–0.20;
*spindles up to 0.25
ovals, *spindles, irregular
bodies:
0.20–0.30
E. somaliensis N.R. light brown digitiform to
digitate
present present present 8–radiates: 0.08–0.10;
spindles: 0.18–0.38
narrow spindles and
needles: 0.23–0.50
8-radiates,
capstans, ovals,
*spindles:
0.1–0.2;
crosses present
long narrow spindles
0.20–0.30
E. splendens N.R. cream to light orange
with white to light red
coenenchymal mounds
digitiform to
digitate
present present present 8–radiates, clubs:
0.08–0.15
narrow spindles:
0.34–0.47;
rods: 0.16–0.20
8–radiates, clubs:
0.07–0.16
spindles: 0.25–0.35
III. Sphaerasclera
S. flammicerebra N.R. orange with red
coenenchymal mounds
capitate absent absent absent 8–radiates: 0.05–0.08;
spheroids: 0.12–0.20
radiates, double-
heads, barrels,
spheroids:
0.13–0.20
radiates: 0.05–0.08
spheroids:
large 0.15–0.20
medium 0.10–0.13
predominately
spheroids: 0.17–0.21
IV. Paraminabea
P. aldersladei recorded orange digitiform absent absent absent 8–radiates, double-
heads: 0.04–0.09
8-–radiates, double-
heads: 0.04–0.09
8–radiates, double-
heads: 0.05–0.11
8–radiates, double-heads:
0.05–0.11
P. cf. aldersladei recorded orange digitiform absent absent absent 8–radiates, double-
heads: 0.04–0.09
8–radiates, double-
heads: 0.04–0.09
8–radiates, double-
heads: 0.06–0.09
8–radiates, double-heads:
0.06–0.09
BRYCE ET AL.
180
·
Zootaxa 3963 (2) © 2015 Magnolia Press
Colour. The preserved colonies are uniformly cream.
Habitat. The specimens were sampled from a depth range between 200–250 m along the continental slope
utilising an epibenthic sled, but the exact habitat is unknown.
Remarks. Eleutherobia somaliensis has only been described from off the coast of Somalia, Africa (Table 1),
and this is the first record for Australia. Although Verseveldt & Bayer’s (1988) original description of the holotype
did not include illustrations of the sclerites from the polyps or the interior of the base, we think the sclerites of our
specimens bear a very close resemblance to those that were illustrated and to the descriptions of those that were
not. The notable differences are the more prickly nature of the sclerite warting and the lack of well defined 8-
radiates as shown in Verseveldt & Bayer’s Fig 34a–d.
Eleutherobia splendens (Thomson & Dean, 1931) new record
(Fig. 7G; 13; Tabs. 1, 2)
Nidalia splendens Thomson & Dean, 1931: 38, Pl. 1, Fig. 7; Pl. 6, Fig. 9; Pl. 25, Fig. 8.
Eleutherobia splendens Verseveldt & Bayer, 1988: 40–41, Figs. 18a, 33a, 35, 36c.
Material examined. WAM Z23988, two whole specimens, cylindrical, colony 1 unbranched, 45.3 mm tall,
attached to a mollusc shell, colony 2 branched with three branches in one plane, 51.6 mm tall, Station SO1/84/055,
Lacepede Archipelago, NW Australia, 19.9500°–19.9833° S, 120.7338°–120.7350° E, CSIRO FRV "Soela" cruise
VI, trawl, depth 297 m, coll. S.M. Slack-Smith, 10 February 1984; WAM Z54996, one whole specimen,
unbranched, 31.5 mm tall, Station SO1/84/056, Kimberley, Beagle Bay, NW Australia, 16.9297° S, 122.5411° E,
CSIRO FRV "Soela" cruise VI, trawl, depth 301 m, coll. S.M. Slack-Smith, 11 February 1984. NTM C002899, one
whole specimen, 19.3338° S, 115. 6836° E, FRV “Soela” cruise 0184, trawl, depth , 306–308 m, coll. A.J. Bruce,
29 January 1984; NTM C013059, one whole specimen, Station SS1005 130-015, off Red Bluff, 23.9908° S, 112.
3547° E, RV "Southern Surveyor", beam trawl, depth 411 m, K. Gowlett-Holmes, 8 December 2005.
Description. The colonies are erect, cylindrical, with large bodied anthocodiae up to 4 mm long (Fig. 7G). The
colonies are unbranched or branched in one plane and are up to 51.6 mm tall. Some specimens are attached to a
mollusc shell.
In the surface of the polyparium the majority of the sclerites are thorny clubs, up to 0.22 mm long but mainly
between 0.08–0.15 mm , with the warts below the head arranged in girdles (Fig. 13A). There are also a few 8-
radiates present. The majority of sclerites of the base are thorny 8-radiates up to about 0.10 mm long, but there are
also a few larger sclerites are up to 0.16 mm that are club-shaped (13B). The sclerites of the interior of the
polyparium are markedly narrow, needle-like forms, 0.34–0.47 mm long, with girdles of high spines (Fig. 13C).
The sclerites in the interior of the base are spindles up to 0.35 mm long. They are similar to the sclerites in the
interior of the polyparium, but are slightly shorter and wider (Fig. 13D).
The tentacles contain densely packed stout, flattened rods with a curved end up to 0.45 mm long (Fig. 13E).
The polyp armature is strongly developed. It consists of collaret and point and is formed of slightly curved, spiny
spindles around 0.05 mm long (Fig. 13F).
Colour. The preserved colonies are cream with the distal part of the coenenchymal mounds being sometimes
of the same colour, but usually are a distinct pink to red (Fig. 7G). The tentacles are white. The introvert contains
brick-red sclerites.
Habitat. The specimens were sampled from a depth range between 297–411 m along the continental slope
utilising otter and beam trawls, but the exact habitat is unknown.
Remarks. Previously this species has only been described from Indonesia and the Philippines (Table 1),
making this the first record for Australia. Our specimens agree well with the holotype colony described and figured
in the original report of Thomson and Dean, but it is impossible to make any worthwhile comparisons with the rest
of their brief description. The notable difference between the characters of our material and the comparable
features reported by those authors is the more prickly warting of the sclerites (as was the case with E. somaliensis
above). Unfortunately, the redescription did not include illustrations of the polyp sclerites, so we asked Dr Leen
van Ofwegen to make a comparison using Verseveldt & Bayer’s microscopic slides of the holotype sclerites of E.
splendens held in the Naturalis Biodiversity Centre, Leiden, and he was able to confirm they are of the same form.
Zootaxa 3963 (2) © 2015 Magnolia Press
·
181
SOFT CORALS OF WESTERN AUSTRALIA
FIGURE 13. Eleutherobia splendens WAM Z23988-1, sclerites: A, surface of polyparium; B, surface of the base; C, interior of
the polyparium; D, interior of the base; E, tentacle rachis; F, collaret and points.
Genus Sphaerasclera McFadden & Ofwegen 2013
Type species. Eleutherobia flammicerebra Williams, 2003, by original designation
BRYCE ET AL.
182
·
Zootaxa 3963 (2) © 2015 Magnolia Press
Diagnosis. Colonies with capitate growth form, with distinct, spherical polyparium raised on a bare stalk.
Polyps are monomorphic and form rounded coenenchymal mounds over the entire surface of the capitulum.
Sclerites are coloured spheroids and smaller radiates. Polyp sclerites are absent. Species are found mostly in deep
water. Azooxanthellate. (adapted from McFadden & Ofwegen 2013).
Sphaerasclera flammicerebra (Williams, 2003) new record
(Fig. 7H; 14; Tabs. 1, 2)
Eleutherobia flammicerebra Williams, 2003: 423–434, Figs. 1 a–c, 2–8.
Sphaerasclera flammicerebra McFadden & Ofwegen 2013: 66–67, Fig. 2–3.
Material examined. WAM Z31480, three whole specimens, 21.6–26.3 mm tall, stalk length 12.0–20.4 mm,
polyparium diameter 10.2–11.2 mm, Station PF06/S1-200/R2, 190 km NW of Dampier, Pluto Gas Field, NW
Australia, 19.9352°–19.9308° S, 115.2261°–115.2288° E, epibenthic sled, depth 200 m, coll. B.F. Cohen, 7
December 2005; WAM Z31465, one small colony, 16.2 mm tall, stalk length 9.4 mm, polyparium diameter 7.9
mm, Station PF06/S1-200/R2, 190 km of NW of Dampier, Pluto Gas Field, 19.9352°–19.9308° S, 115.2261°–
115.2288° E, epibenthic sled, depth, 200 m, coll. B.F. Cohen, 7 December 2005; WAM Z13065, Station 1031302,
North West Cape, 21.4672° S, 114.1016° E, epibenthic sled, depth 200–250 m, coll. P. Alderslade, J. Fromont and
L.M. Marsh, 21 March 2002; WAM Z55265, one whole colony, 22.0 mm tall, stalk length 13.2mm, polyparium
diameter 15.2 mm, Station SO1/84/055, Lacepede Archipelago, 19.9500°–19.9833°S, 120.7683°–120.7350° E,
CSIRO FRV "Soela" cruise VI, trawl, depth 297 m, coll. S.M. Slack-Smith, 10 February 1984; WAM Z54995, one
whole specimen, 32.4 mm tall, stalk length 20.9 mm, polyparium diameter 16.1 mm, Station SO1/84/056,
Kimberley, Beagle Bay, 16.9297° S , 122.5411° E, CSIRO FRV "Soela" cruise VI, trawl, depth 301 m, coll. S.M.
Slack-Smith, 1 February 1984; WAM Z55271, one whole specimen, 20.4 mm tall, stalk length 11.3 mm,
polyparium diameter 14.9 mm, Station SO1/84/059, Kimberley, Beagle Bay, 15.1500°–15.1833° S, 121.0833°–
120.0500° E, CSIRO FRV "Soela" cruise VI, trawl, depth 449 m, coll. S.M. Slack-Smith, 11 February 1984. All
colonies are attached to a fragment of hard coral skeleton.
Description. All colonies are attached to hard corals and have a capitate growth form, with a rounded
capitulum with a scrolled lower margin and a straight or slightly bent, distinct stalk (Fig. 7H). The polyps are
monomorphic, numerous and evenly distributed over the entire surface of the capitulum. In the preserved
specimens all polyp bodies are retracted forming low, round coenenchymal mounds at the surface. The stem is
polyp free. Colonies are between 16.2–32.3 mm tall and the length of the stalks ranges between 9.40–20.9 mm. The
height of the polypariums range between 7.0–11.4 mm and have diameters between 7.9–16.1 mm. The diameter of
the stalks ranges at base between 6.3–13.1 mm and between 4.2–11.5 mm at the distal end.
The surface of the polyparium is densely spiculated with large tuberculated spheroids between 0.12–0.20 mm
long and smaller, oval 8-radiates 0.05–0.08 mm long (Fig. 14A). The sclerites of the interior of the polyparium are
predominately round to oval-shaped tuberculated spheroids between 0.13–0.20 mm in length (Fig. 14B). Some
robust 8-radiates around 0.17 mm in length are also present. The sclerites of the surface of the base is densely
spiculated with large tuberculated spheroids 0.15–0.20 mm in length, medium-sized tuberculated spheroids mainly
between 0.10–0.13 mm long, and smaller, oval 8-radiates 0.05–0.08 mm long (Fig. 14C). The sclerites in the
interior of the stalk are predominately large tuberculated spheroids 0.17–0.21 mm in length and very robust
radiates mainly around 0.10 mm long (14D). Polyp sclerites are absent.
Colour. The polyparium of the preserved colonies are cream to light orange with dark orange spots
representing the protuberances. One colony has a uniformly light pink capitulum. The stalks are cream to orange.
Sclerites are colourless and orange to around the protuberances.
Habitat. The specimens were sampled by epibenthic sled and trawl from a depth range between 200–449 m
along the continental slope, but the exact habitat is unknown.
Remarks. Sphaerasclera flammicerebra (Williams, 2003) was first described from Palau, Pacific Ocean.
McFadden & Ofwegen (2013: 62, 66, 67, Table 1, Fig. 2, 3) included museum material of S. flammicerebra
collected from New Caledonia in 2008 and Mauritius collected in 1929 in their phylogenetic and morphological
analyses. This material of S. flammicerebra agrees with the original description by Williams (2003: 423–430, Fig.
1–8) in most characters. The main difference are the rather spiky 8-radiates of our material in comparison to more
Zootaxa 3963 (2) © 2015 Magnolia Press
·
183
SOFT CORALS OF WESTERN AUSTRALIA
rounded 8-radiates of Williams (2003) material. As the colonies agree with the original description of S.
flammicerebra by being monomorphic, in the absence of polyp sclerites, the size and shape of the colonies, the size
and distribution of sclerites types, and the genetic similarity, we consider the difference in sclerite shape of the
smaller 8-radiates as a intraspecific variation (Fig. 7, 14, 19, 20).
FIGURE 14. Sphaerasclera flammicerebra, WAM Z31480, sclerites: A, surface of polyparium; B, interior of the polyparium;
C, surface of the base; D, interior of the base.
Genus Paraminabea Williams & Alderslade, 1999
Type species. Bellonella indica Thomson & Henderson, 1905 by subsequent designation (Williams & Alderslade
1999).
Diagnosis. Colonies usually small, unbranched, with symmetrical, cylindrical polyparium. Growth forms such
as dome-shaped, digitiform, hemispherical, or digitate-lobate can occur. Dimorphic polyps are evenly distributed
over the polyparium and are devoid of sclerites. Autozoids are large and completely retractile. Siphonozooids
scarce, small to minute, distributed between the autozooids. The polyp-free basal portion is variable in length.
Mostly in deep water or restricted to caves and overhangs in shallow waters. Sclerites of polyparium mostly
barrels, double heads and 6- or 8-radiates. Also, radiates, tuberculate spheroids, 7-radiates or double stars and
spindle-like forms derived from radiates may occur.
(Williams & Alderslade 1999).
BRYCE ET AL.
184
·
Zootaxa 3963 (2) © 2015 Magnolia Press
Paraminabea aldersladei (Williams, 1992)
(Figs. 15A,B,C; 16A–D; Tab. 2)
Bellonella indica (non Thomson & Henderson, 1905) Bayer 1974: 261; Faulkner and Chesher 1979: 267, Pl. 22; Minabea
aldersladei Williams, 1992: 3–9, Figs 1b, 4, 5.
FIGURE 15. A–C, Paraminabea aldersladei, WAM Z59775: A, in situ; B, a preserved specimen and fresh specimens on deck
after collection; C, longitudinal section showing gastric cavities of autozooids and siphonozooids and detail of the colony
surface, black arrow pointing to siphonozooids; D–F, Paraminabea cf. aldersladei, WAM Z59783: D, in situ; E, a preserved
specimen and fresh specimens on deck after collection; F, longitudinal section showing gastric cavities of autozooids and
siphonozooids and detail of the colony surface, black arrow pointing to siphonozooids.
Material examined. WAM Z59775, five whole specimens, Station 43/K10, Long Reef, Kimberley, NW Australia,
13.92155° S, 125.73268° E, scuba, depth 12–20 m, coll. M. Bryce, 20 October 2010; WAM Z66736, six whole
specimens, Station 115/K11, Heritage reef, Kimberley, NW Australia, 14.30367° S, 115.20915° E, scuba, depth
12–20 m, coll. M. Bryce, 22 September 2011; WAM Z67010, four whole specimens, Station 130/K13, Ashmore
Reef, NW Australia, 12.18848° S, 123.12887° E, scuba, depth 12–20 m, coll. M. Bryce, 29 September 2013; WAM
Z67106, 33 whole specimens, Station 144/K13, Hibernia Reef, NW Australia, 11.97404° S, 123.32208° E, scuba,
depth 14 m, coll. M. Bryce, 5 October 2013; WAM Z67241, 24 whole specimens, Station 152/K14, Clerke Reef,
NW Australia, 17.25188° S, 119.38378° E, scuba, depth 16 m, coll. M. Bryce, 3 October 2014; WAM Z67271, 1
whole specimens, Station 156/K14, Clerke Reef, NW Australia, 17.29298° S, 119.37819° E, scuba, depth 16 m,
coll. M. Bryce, 5 October 2014; WAM Z67366, 1 whole specimens, Station 170/K14, Clerke Reef, NW Australia,
17.31697° S, 119.38378° E, scuba, depth 15 m, coll. M. Bryce, 10 October 2014; WAM Z67385, 1 whole
Zootaxa 3963 (2) © 2015 Magnolia Press
·
185
SOFT CORALS OF WESTERN AUSTRALIA
specimens, Station 173/K14, Clerke Reef, NW Australia, 17.31753° S, 119.31216° E, scuba, depth 15 m, coll. M.
Bryce, 11 October 2014; WAM Z67404, 2 whole specimens, Station 178/K14, Mermaid Reef, NW Australia,
17.16154° S, 119.6471° E, scuba, depth 15 m, coll. M. Bryce, 13 October 2014.
Description. All colonies are digitiform, slightly laterally flattened, tapering toward the rounded apical end
and have a short, polyp-free base (Fig. 15A,B,C). They are between 10–53 mm tall, 4–18 mm in diameter at the
apex and 4–25 mm in diameter across the holdfast. The polyps are dimorphic, and arranged uniformly over the
surface of the colonies. In all colonies the autozooid polyps are completely retracted and surrounded by minute
pores representing the siphonozooids.
Sclerites from the polyparium surface are predominantly eight radiates with double heads, that appear to be
derived from 6- and 8-radiates, 0.04–0.09 mm long (Fig. 16A). Those from the interior of the polyparium are more
robust barrels and sub-sheroidal forms of similar size with a very short waist (Fig. 16B). Sclerites from the base
surface and interior are similar in shape to those of the polyparium, but slightly longer, 0.05–0.11 (Fig. 16C,D).
Polyp sclerites are absent.
FIGURE 16. Paraminabea aldersladei, WAM Z59775, sclerites: A, surface of the polyparium; B, interior of the polyparium;
C, surface of the base; D, interior of the base.
Colour. The colonies were uniformly bright orange in situ, on deck and in alcohol. The polyps in the preserved
specimen are cream. Sclerite colour is orange.
Habitat. Station 43/K10: steep outer reef wall descending to 20 meters where the bottom is of fine silt. From
there the honeycombed wall extends upwards with encrusting corals and soft corals, which rapidly increase in
coverage. Specimens of Paraminabea aldersladei were found attached to the wall between 12 and 20 metres.
Station 115/K11: fore-reef slope with a vertical wall descending down to a coral rubble bottom. The reef crest
has a high diversity of hard and soft corals, sea fans, and sponges. The vertical wall has small caves and ledges with
little benthic invertebrate life. Specimens of Paraminabea aldersladei were attached to the wall between 12 and 20
metres.
Station 130/K13: a very steep and fractured fore-reef slope descending to a depth of 20 metres. The slope is
deeply incised with caves and small ledges which are well covered with encrusting invertebrates. The small group
of Paraminabea aldersladei colonies was found attached to a wall in an overhang at 20 metres depth.
Station 144/K13: the fore-reef cemented slope consists of separate "hillocks" that are joined at the base with
deep "gullies" between. There is a high diversity of soft and hard corals and an abundant occurrence of Paraminabea
aldersladei. The colonies were attached to the walls under overhangs amongst other scattered soft corals.
BRYCE ET AL.
186
·
Zootaxa 3963 (2) © 2015 Magnolia Press
Remarks. Paraminabea aldersladei is a well-known species from the Indo-Pacific (Williams 1992; Table 2).
In Australia P. aldersladei has been reported from the Great Barrier Reef and also from the Rowley Shoals, north-
western Australia. The characters of this material agree well with the original description by Williams (1992,
although no yellow colonies were encountered.
Paraminabea cf. aldersladei Williams & Alderslade, 1999
(Figs. 15D,E,F, 17A,B; 18A,B; Tab. 2)
FIGURE 17. Paraminabea cf. aldersladei, WAM Z59783, sclerites: A, surface of the polyparium; B, interior of the
polyparium.
Zootaxa 3963 (2) © 2015 Magnolia Press
·
187
SOFT CORALS OF WESTERN AUSTRALIA
FIGURE 18. Paraminabea cf. aldersladei, WAM Z59783, sclerites: A, surface of the base; B, interior of the base.
Material examined. WAM Z59783, five whole specimens, Station 44/K10, Long Reef, Kimberley, NW Australia,
13.92155° S, 125.73268° E, scuba, depth 10–20 m, coll. M. Bryce, 20 October 2010.
Description. The colonies are either uniformly cylindrical from the base to the apical tip or are tapering toward
the rounded apical end (Fig. 15D,E,F). They have a short, slightly laterally flattened polyp-free base. The polyps
are dimorphic, and arranged uniformly over 60–90 % of the surface of the colonies. In all colonies the autozoids
are completely retracted. The surface of the preserved specimens is extremely contracted and appears convoluted,
which makes the siphonozooids difficult to locate. The gastric cavities of the autozooids form curved tubes (Fig.
15F). Colonies are between 16–24 mm tall, 3–7 mm in diameter at the apex and 5–7 mm in diameter across the
holdfast.
In the surface of the polyparium the sclerites are mostly 8-radiates and some cylindrical to oval forms, 0.04 to
0.09 mm long (Fig. 17A), and in the interior the sclerites are double stars or double heads and irregular forms, some
approaching crosses, from about 0.05 to 0.07 mm long (Fig. 17B). The sclerites from the base are similar to those
of the polyparium, consisting mostly of 8-radiates in the surface, from 0.06 to 0.08 mm long (Fig.18A), and double
heads or double-stars, with a long waist, together with irregular forms approaching crosses, up to 0.09 mm long, in
the interior (Fig. 18B). Polyp sclerites are absent.
Colour. The colonies were uniformly dark red in situ and on deck. In alcohol the specimens are dark orange.
The polyps in the preserved specimens are cream. The colour of the sclerites is red.
Habitat. Very steep and fractured fore-reef slope ascending from 20 up to four metres depth. The slope is
heavily pocketed with small caves and deep, steep-sided fissures. At 20 metres there are large rocky outcrops
forming long-reef gullies, which are almost devoid of life, probably due to the heavy siltation. Sediment between
the rocky outcrops is very fine and smothering. A small group of Paraminabea cf. aldersladei was found attached
BRYCE ET AL.
188
·
Zootaxa 3963 (2) © 2015 Magnolia Press
to the wall together with a group of Eleutherobia kimberleyensis sp. nov. in an overhang at ten meters depth
amongst other scattered soft corals.
Remarks. Paraminabea cf. aldersladei has similarities to P. aldersladei, P. indica, and P. robusta. It resembles
P. aldersladei in growth form, but differs in type and distribution of the sclerites. P. aldersladei is bright orange and
has mainly orange 8-radiates, double-heads, robust barrels and subspheroidal forms, while this species has red
sclerites consisting predominately of radiates, double heads and complex cross-like forms. Paraminabea indica
differs from Paraminabea cf. aldersladei by being dichromatic, the restriction of the polyps to less than half of the
colony length, and the lack of tuberculated spheroids. It also appears to be restricted to deep water. The main
differences between P. cf. aldersladei and P. robusta are colour, colony length and the morphology and distribution
of the siphonozooids.
Molecular Phylogeny
The mitochondrial marker mtMutS was successfully amplified for all analyzed specimens, except for E. splendens
(Table 3). In contrast, the ~800 bp fragment of the 28S nuclear ribosomal gene was only successfully amplified for
five specimens. For Parasphaerasclera grayi, Parasphaerasclera kimberleyensis sp. nov. and Eleutherobia
somaliensis only partial sequences were recovered and amplification was not possible for Sphaerasclera
flammicerebra and Eleutherobia splendens. Both mtMutS and 28S phylogenies resulted in a polyphyletic family
Alcyoniidae (Figs. 19, 20). The mitochondrial phylogenetic tree separates the genera Eleutherobia, Paraminabea,
Parasphaerasclera and Sphaerasclera in four monophyletic groups. Apart from the clade including the genus
Parasphaerasclera, supported only in the ML analysis, the others were well supported by both methods. The
sampled specimens belonging to these genera were distributed in three clades in the mitochondrial phylogeny. In
particular the new species Parasphaerasclera kimberleyensis was included in a clade with five other species of
Parasphaerasclera (P. a ure a , P. gr a y i , P. aff. grayi, P. rotifera and P. valdiviae). P. grayi and P. aff. grayi were
respectively sister to P. aurea and P. kimberleyensis, while the relationship between P. rotifera and other members
of Parasphaerasclera was unresolved. S. flammicerebra was sister to Paraminabea aldersladei, P. cf. aldersladei
and a clade including the scleraxonians Paragorgia wahine, Corallium laauense and the alcyoniid Anthomastus
ritteri. The nuclear phylogenetic tree recovered a monophyletic Paraminabea and Eleutherobia, but did not clarify
the relationships within Parasphaerasclera, as many of the nodes including those species were collapsed into
polytomies due to their low support values (Figs. 19, 20). The species belonging the genus Paraminabea (i.e. P.
aldersladei and
P. cf. aldersladei) formed a well supported clade, but their relationships with other alcyoniids
remained largely unresolved. Notably, both the mitochondrial and nuclear phylogenies recovered the three sampled
species belonging the genus Eleutherobia (i.e. E. australiensis sp. nov., E. imahari sp. nov. and E. somaliensis) as
monophyletic, and this clade nested in a larger clade composed of Alcyonium species.
Discussion
General summary. The present account is the first comprehensive inventory of digitate and capitate soft corals of
the genera Eleutherobia, Parasphaerasclera, Sphaerasclera and Paraminabea of tropical Western Australia. It
reports on three new species (Parasphaerasclera kimberleyensis, Eleutherobia australiensis, Eleutherobia
imaharai), three range extensions (Parasphaerasclera grayi, Eleutherobia somaliensis, Eleutherobia splendens),
and one new geographical record (Sphaerasclera flammicerebra), and discusses their taxonomic placement. Given
the limitations of traditional morphological taxonomy, we used an integrative approach for the Western Australian
material, combining morphological examinations with molecular phylogenetic analyses to clarify the status and
placement of the specimens.
The literature detailing species of the genus Eleutherobia prior to 2013 is plagued with confusion (Williams
1986). Verseveldt & Bayer revised the genus Eleutherobia in 1988, but a new taxonomic revision of this genus was
overdue as more species had subsequently been added. Benayahu & Schleyer (1995) mentioned the possible poly-
or paraphyly of Eleutherobia, as the genus incorporated species with highly variable sclerites and body plans
resulting in confusion over its generic characters. In 2013 McFadden & Ofwegen used morphology together with
Zootaxa 3963 (2) © 2015 Magnolia Press
·
189
SOFT CORALS OF WESTERN AUSTRALIA
molecular investigation of some nominal species of the genus, to provide sufficient evidence to propose the new
family Parasphaerascleridae and the new genera Parasphaerasclera and Sphaerasclera. Six nominal species of
Eleutherobia without polyp sclerites were assigned to the new genus Parasphaerasclera (Table 1, 2), the only
capitate nominal species of Eleutherobia with spheroids (E. flammicerebra) was assigned to the new genus
Sphaerasclera and the capitate E. variabile was re-assigned to Alcyonium. Our molecular results confirm the
separation of Eleutherobia, Parasphaerasclera, and Sphaerasclera as different genera and provide further
evidence that the genus Eleutherobia is monophyletic (Figs. 19, 20; Table 2).
FIGURE 19. Phylogenetic tree of mtMutS. Numbers at the nodes represent: on the left ML bootstrap values and on the right
Bayesian posterior probabilities (pp). Nodes with bootstrap value < 70 % and pp < 0.95 were collapsed. Triangles represent
collapsed clades with strong support (pp = 1). Specimens analyzed belonging to Parasphaerasclera, Eleutherobia, and
Sphaerasclera are in bold. Cornularia pabloi was used as the outgroup.
Former species of Eleutherobia and those species of the genus currently considered valid, incorporate possible
rare species with an Indo-Pacific biogeography, ranging from South Africa to the central west Pacific, Japan and
Australia (Table 1) (Williams, 1999; 2001), but the known geographic range of some of the species has been
expanded. Two species, Eleutherobia somaliensis and Sphaerasclera flammicerebra, have now been reported from
areas far removed from their type locality, and Benayahu et al. (2004) reported P. grayi as a new record from
Taiwan. The latter authors suggest that the position of Taiwan, between the West Pacific Ocean, the East China Sea
and the crossroad of the Philippine-Japan Island Arc forms a stepping stone for the dispersal of shallow reef
organisms. The finding of new species and geographical records in this paper may indicate that species diversity,
and associated distributional ranges of Eleutherobia and Parasphaerasclera, may have been underestimated in
remote areas, such as along the Western Australian coast.
BRYCE ET AL.
190
·
Zootaxa 3963 (2) © 2015 Magnolia Press
FIGURE 20. Phylogenetic tree of the 28S ribosomal gene. Numbers at the nodes represent: on the left ML bootstrap values and
on the right Bayesian posterior probabilities (pp). Nodes with bootstrap value < 70 % and pp < 0.9 were collapsed. Triangles
represent collapsed clades with strong support (pp ≥ 0.95). Specimens analyzed belonging to Parasphaerasclera, Eleutherobia,
and Sphaerasclera are in bold. Cornularia cornucopiae and Cornularia pabloi were used as outgroup.
One new species, Parasphaerasclera kimberleyensis sp. nov., was found together with Paraminabea
aldersladei at Long Reef. These species found in shallower habitats have adapted to a cryptic lifestyle, preferring
darker areas, such as overhangs and caves. A dark red specimen identified herein as Paraminabea cf. aldersladei
was also found at this site. Despite these two specimens of Paraminabea, Paraminabea aldersladei and
Paraminabea cf. aldersladei, being exposed to the same environmental conditions, colour and sclerite shape are
distinctly different. Nevertheless, the genus is known for its high intraspecific phenotypic variability. Using sclerite
shape as the only determining factor for species identification is problematic (Williams & Alderslade 1999). The
lack of molecular differences between the dark red Paraminabea cf. aldersladei specimen and that of P. aldersladei
point to them being colour and morphological variants of the latter species, rather than separate species. However,
we acknowledge that the lack of divergence in the mtMutS marker does not necessarily mean conspecificity, given
the slow rates of molecular evolution reported for the mitochondria of anthozoans (Shearer et al. 2002). Further
studies and the analysis of rapidly evolving molecular markers seem necessary to clarify this matter (see Pante et
al. 2014).
Molecular considerations. The combination of morphological characters, geographic distribution and the
compatibility with the generic diagnoses provide enough evidence to propose the placement of the new species P.
kimberleyensis sp. nov. in the newly erected genus Parasphaerasclera and the second and third new species, E.
australiensis sp. nov. and E. imaharai sp. nov., in the genus Eleutherobia.
Zootaxa 3963 (2) © 2015 Magnolia Press
·
191
SOFT CORALS OF WESTERN AUSTRALIA
TABLE 3. Octocoral taxa used for phylogenetic analysis with voucher and GenBank accession numbers. New specimens here analysed are in boldface. RMNH = Naturalis
Biodiversity Center; ZSM = Zoologische Staatssammlung München; ZMTAU = Zoological Museum, Tel Aviv University; WAM = Western Australian Museum; NTM =
Museum and Art Gallery of the Northern Territory; CSM = C.S. McFadden Laboratory; MNHN = Museum National d’Histoire Naturelle, Paris; SBMNH: Santa Barbara
Museum of Natural History; QM = Queensland Museum; USNM = National Museum of Natural History (Smithsonian Institute); CRCNI = Palau International Coral Reef Center;
UF = Florida Natural History Museum; NIWA = National Institute of Water and Atmospheric Research (New Zealand); OCDN = Coral Reef Research Foundation (Palau). *see
Pante et al. (2012).
GenBank accession numbers
Order Suborder Family Species Voucher mtMutS 28S rDNA
Acrophytum claviger RMNH Coel. 40222 JX203770 JX203655
Alcyonium acaule AY607775
Alcyonium aurantiacum DQ302806
Alcyonium bocagei RMNH Coel. 39672 GU355960 KF728088
Alcyonium coralloides AY607772 JX203640
Alcyonium digitatum AY607777 JX203641
Alcyonium glomeratum RMNH Coel. 39666 AY607776 KF728091
Alcyonium haddoni ZSM 20061191 GU355974 JX203642
Alcyonium hibernicum RMNH Coel. 39661 AY607771 KF728089
Alcyonium palmatum GQ342467 JX203643
Alcyonium sidereum GU355972 KF728090
Alcyonium roseum ZSM 20061195 GQ342468 JX203644
Alcyonium variabile KF728095 JX203645
Alcyonium verseveldti ZMTAU CO33097 GU356012 JX991219
Anthomastus ritteri RMNH Coel. 40802 DQ302816 JX203761
Alcyonium bocagei AY607774
Cladiella sp. WAM Z59835 HG970087 HG970073
Dampia pocilloporaeformis WAM Z59725 HG970088 HG970074
Discophyton rudyi CSM-DIRU15 DQ302808 JX203659
Elbeenus lauramartinae DQ536320
Eleutherobia australiensis, s
p. nov. W
AM Z31488 HG970078 HG970065
Eleutherobia imaharai, s
p. nov. W
AM Z13252 HG970080 HG970067
Eleutherobia somaliensis WAM Z12201 HG970079 HG970066
Alcyonacea Alcyoniina Alcyoniidae
Klyxum sp. WAM Z59659 HG970089 HG970076
......continued on the next page
BRYCE ET AL.
192
·
Zootaxa 3963 (2) © 2015 Magnolia Press
TABLE 3. (Continued)
GenBank accession numbers
Order Suborder
Family Species Voucher
mtMutS 28S rDNA
Lampophyton planiceps RMNH Coel. 40201 GQ342477 JX203656
Lobophytum cf. altum WAM Z59839 HG970075
Lobophytum compactum NTM-C011566 DQ280559
Lobophytum legitimum NTM-C013980 DQ280571
Lobophytum pauciflorum UF 2856 JX203649
Lobophytum ransoni NTM-C013929 DQ280578
Lobophytum sarcophytoides RMNH-Coel.33065 DQ280582
Lobophytum strictum NTM-C011271 DQ280584
Malcacanthus capensis RMNH Coel. 40801 DQ302811 JX203660
Paraminabea aldersladei NTM C14895 JX203767 JX203763
Paraminabea aldersladei WAM Z66736 HG970083 HG970070
Paraminabea aldersladei WAM Z59775 HG970084 HG970071
Paraminabea cf. aldersladei WAM Z59783 HG970085 HG970072
Rhytisma sp. NTM-C001942 DQ302812
Sarcophyton ehrenbergi NTM C14455 JX203650
Sarcophyton elegans UF2637 DQ280520
Sarcophyton gemmatum ZMTAU CO34091 GU356017
Sarcophyton mililatensis RMNH-Coel.33080 DQ280541
Sarcophyton trocheliophorum NTM C14854 JX203651
Sarcophyton trocheliophorum AB759314
Sarcophyton trocheliophorum NTM-C014469 DQ280549
Sinularia abrupta ZMTAU CO33623 KC542822
Sinularia brassica NTM C13507 FJ621379
Sinularia brassica NTM C14185 FJ621380
Sinularia brassica WAM Z59651 HG970090
Sinularia brassica RMNH Coel. 41306 KF915494
Sinularia brassica RMNH Coel. 41309 KF915493
Sinularia compressa ZMTAU CO34142 FJ621387
......continued on the next page
Zootaxa 3963 (2) © 2015 Magnolia Press
·
193
SOFT CORALS OF WESTERN AUSTRALIA
TABLE 3. (Continued)
GenBank accession numbers
Order Suborder
Family Species Voucher
mtMutS 28S rDNA
Sinularia digitata RMNH Coel.40841 KC542830
Sinularia dura NTM C13808 FJ621402
Sinularia erecta ZMTAU CO34144 FJ621404 KC542835
Sinularia finitima RMNH Coel.38728 FJ621407
Sinularia finitima RMNH Coel.41332 KF915495
Sinularia gardineri ZMTAU CO34097 KC542819
Sinularia hirta ZMTAU CO34103 FJ621429
Sinularia hirta ZMTAU CO34100 KC542820
Sinularia leptoclados ZMTAU CO35308 KC542836
Sinularia mammifera NTM C14198 FJ621444
Sinularia maxima NTM C14512 FJ621448 KC542839
Sinularia polydactyla ZMTAU CO34138 FJ621466
Sinularia polydactyla RMNH Coel.41339 KF915515
Sinularia querciformis ZMTAU CO34096 FJ621469 JX203652
Sinularia robusta NTM C14518 KC542843
Sinularia sp. WAM Z59808 HG970091
Sinularia sp. WAM Z59770 HG970077
Sinularia terspilli ZMTAU CO34156 KC542821
Sphaerasclera flammicerebra MNHN-IK-2012-
12004
JX203765 JX203638
Sphaerasclera flammicerebra WAM Z31480 HG970086
Thrombophyton coronatum SBMNH 145123 DQ302814 JX203661
Nephtheidae Capnella imbricata DQ302817
Dendronephthya sinaiensis ZMTAU CO34163 JX124349
Dendronephthya sp. NTM-C012655 DQ302818
Dendronephthya sp. A RMNH Coel. 40907 KF915354
Eunephthya thyrsoidea RMNH Coel. 40182 JX124364 JX124340
......continued on the next page
BRYCE ET AL.
194
·
Zootaxa 3963 (2) © 2015 Magnolia Press
TABLE 3. (Continued)
GenBank accession numbers
Order Suborder
Family Species Voucher
mtMutS 28S rDNA
Lemnalia sp. RMNH Coel. 40818 JX203802 JX203726
Leptophyton benayahui GQ342507
Nephthea acuticonica ZMTAU CO34070 GU356023
Nephthea sp. NTM-C012400 DQ302822
Nephthea sp. JN383340
Nephthea sp. B RMNH Coel. 40972 KF915417
Neospongodes sp. NTM-C013130 DQ302823
Paralemnalia digitiformis DQ302824
Paralemnalia digitiformis RMNH Coel. 40941 KF915418
Paralemnalia eburnea ZMTAU CO34081 GU356030
Paralemnalia thyrsoides ZMTAU CO34087 JX203727
Scleronephthya corymbosa ZMTAU CO34159 GQ342511 JX124350
Scleronephthya sp. NTM-C011489 DQ302825
Scleronephthya sp. A RMNH Coel. 41039 KF915486
Stereonephthya cundabiluensis ZMTAU CO34204 JX124351
Stereonephthya sp. NTM-C011307 DQ302826
Stereonephthya sp. B RMNH Coel. 41061 KF915541
Umbellulifera sp. NTM-C011063 DQ302827
Chironephthya sp. NTM-C012426 DQ302830
Nidaliidae
Chironephthya sp. ZMTAU CO34203 GQ342513 JX203730
Chironephthya sp. 1 ZMTAU CO34223 GU356025
Chironephthya sp. 2 ZMTAU CO34222 GU356024
Chironephthya sp. 3 ZMTAU CO34226 GU356026
Chironephthya sp. A RMNH Coel. 40893 KF915337
......continued on the next page
Zootaxa 3963 (2) © 2015 Magnolia Press
·
195
SOFT CORALS OF WESTERN AUSTRALIA
TABLE 3. (Continued)
GenBank accession numbers
Order Suborder
Family Species Voucher
mtMutS 28S rDNA
Chironephthya sp. B RMNH Coel. 40889 KF915339
Chironephthya sp. C RMNH Coel. 40890 KF915343
Chironephthya sp. D RMNH Coel. 40900 KF915348
Chironephthya sp. D RMNH Coel. 40897 KF915347
Nephthyigorgia sp. NTM-C011345 DQ302831
Nephthyigorgia sp. RMNH Coel.40819 JX203804 JX203732
Nidalia sp. NTM C014876 DQ302828 JX203729
Pieterfaurea khoisanianum CSM-SAF183 GQ342510 JX203657
Siphonogorgia sp. NTM-C011159 DQ302832
Siphonogorgia sp. A RMNH Coel. 41044 KF915529
Siphonogorgia godeffroyi RMNH Coel. 40833 JX203803 JX203731
Ceeceenus quadrus UF2858 GQ342514 JX124346 Paralcyoniidae
Paralcyonium spinulosum RMNH Coel. 40820 DQ302833 JX124347
Parasphaerasclera aurea RMNH Coel. 40799 JX203766 GQ377456
Parasphaerasclera grayi WAM Z54774 HG970081 HG970068
Parasphaerasclera aff. grayi NTM C14902 DQ302809
Parasphaerasclera kimberleyensis, sp. nov. WAM Z59789 HG970082 HG970069
Parasphaerasclera rotifera UF3890 GQ342472 JX203639
Parasphaerasclera valdiviae RMNH Coel. 41532 KF728097 KF728085
Parasphaerascleridae
Parasphaerasclera valdiviae RMNH Coel. 41534 KF728096 KF728084
Anthelia glauca ZMTAU CO34183 JX203812 JX203753
Asterospicularia randalli DQ302836
Asterospicularia randalli RMNH Coel. 41521 KF915316
Cespitularia erecta OCDN-8504C JX203813 JX203755
Cespitularia sp. NTM-C013542 DQ302837
Xeniidae
Efflatounaria sp. NTM-C012311 DQ302838
Ovabunda obscuronata ZMTAU CO34077 GU356027
......continued on the next page
BRYCE ET AL.
196
·
Zootaxa 3963 (2) © 2015 Magnolia Press
TABLE 3. (Continued)
GenBank accession numbers
Order Suborder
Family Species Voucher
mtMutS 28S rDNA
Sansibia sp. NTM-C012955 DQ302840
Sarcothelia edmondsoni CSM-SKB JX203757
Sarcothelia sp. NTM-C015151 DQ302841
Sarcothelia sp. KM201434
Xenia hicksoni ZMTAU CO34072 GQ342529 JX203759
Chrysogorgiidae *Stephanogorgia faulkneri NTM C14927 GQ342485 JX203718 Calcaxonia
Ellisellidae Viminella sp. RMNH Coel. 40032 JX203794 JX203703
Corallidae Corallium laauense USNM 1071433 GQ293301 GQ293265 Scleraxonia
Paragorgiidae Paragorgia wahine NIWA 3326 GQ293314 GQ293263
Clavulariidae Clavularia sp. RMNH Coel.40809 JX203778 JX203678
Cornularia cornucopiae PLG 2012 JX203760
Cornulariidae Cornularia pabloi RMNH Coel.40197 JX203791
Cornularia pabloi USNM 1178390 JX203792 JX203699
Telestula cf. spiculicola GU563311
Stolonifera
Telestidae
Telestula sp. NTM C14984 JX203697
Helioporacea Helioporidae Heliopora coerulea CRCNI 577 DQ302872 JX203716
Echinoptilidae Actinoptilum molle RMNH Coel. 40822 GQ342491 JX203738
Halipteridae Halipteris finmarchica NTM C14596 DQ302868 JX203741
Pennatulidae Gyrophyllum sibogae NOR89/53 JX203740
Gyrophyllum sp. NTM-C014392 DQ302869
Pennatulacea
Subsessiliflorae Virgulariidae Virgularia schultzei RMNH Coel. 40823 GQ342527 JX203743
Zootaxa 3963 (2) © 2015 Magnolia Press
·
197
SOFT CORALS OF WESTERN AUSTRALIA
From a molecular perspective, in the mitochondrial tree, Parasphaerasclera kimberleyensis sp. nov. is sister to
Parasphaerasclera grayi, both being collected from shallow water habitats of the Kimberley, and is included in a
supported clade together with Parasphaerasclera rotifera, Parasphaerasclera aurea, and Parasphaerasclera
valdiviae. These species are characterized by lacking sclerites in their polyps. In contrast, the 28S rDNA phylogeny
resulted in Parasphaerasclera kimberleyensis sp. nov. being separated from the co-generic species. However, the
relationships shown in the 28S rDNA phylogeny are poorly resolved in this part of the tree, with several deep
nodes collapsed into polytomies due to their low support values. In addition, it is worth noting that only a partial
28S rDNA sequence (~250 bp) was recovered from Parasphaerasclera kimberleyensis sp. nov. and the recovered
fragment includes a highly conserved region of the 28S rDNA gene. Thus the phylogenetic placement of
Parasphaerasclera kimberleyensis sp. nov. should be treated as uncertain in the 28S rDNA phylogeny.
Until now the only published records of Eleutherobia from the west coast of Australia were for E. rubra
(Brundin, 1896) from Port Headland (Verseveldt & Bayer, 1988). Colonies of this species are slightly flattened and
have dome-shaped polyp mounds with the openings offset to the top of the colony. Our molecular analyses
supported a clade including E. australiensis sp. nov., E. somaliensis and E. imaharai sp. nov. and in both
phylogenies these taxa nested in a clade including Alcyonium species. These results are in agreement with
McFadden & Ofwegen (2013) who reassigned two species of Eleutherobia with capitate growth form to the genus
Alcyonium and stated that all species of Eleutherobia, ‘with polyps with distinct collaret and points of spindles;
radiates, spindles and club-like sclerites in the colony surface; and spindles in the interior coenenchyme’ likely
belong to Alcyonium sensu stricto. These species include the new species E. australiensis sp. nov. and E. imaharai
sp. nov., as well as the remaining Eleutherobia species E. dofleini, E. duriuscula, E. flava, E. grandiflora, E.
rigida, E. rubra, E. splendens, E. somaliensis, E. sumbawaensis, E. unicolor, and E. vinadigitaria.
In the interest of efficiency and clarity over time we believe that all remaining Eleutherobia species and the
newly described ones will move into the
Alcyonium clade once further molecular information is forthcoming.
However, to prevent confusion the authors prefer at this stage that this action is better undertaken as a single
change at a future date.
The distinct, curved bars found in the tentacles of E. imaharai sp. nov. are different from the tentacle sclerites
found in all other species of Eleutherobia where they are known. There are, however, some similarities to the
flattened, curved spindles of E. australiensis sp. nov., which themselves may be similar to those of E. somaliensis
described by Verseveldt & Bayer (1988: 39) as “two rows of distally diverging, flat, marginally toothed
sclerites…”.
E. splendens DNA did not amplify and therefore no genetic or phylogenetic information exists on this species.
Finally, the capitate Sphaerasclera flammicerebra shows as being related to species of Paraminabea in the
mitochondrial tree.
Implications. We would also like to highlight the role of comprehensive surveys in remote areas, such as the
surveys of Kimberley marine environments by the Western Australian Museum. These surveys provide a clearer
understanding of soft coral taxonomy, systematics and distributional patterns in tropical marine environments. This
has implications for marine area protection (Fabricius 2008; Schleyer & Benayahu 2008; Keesing et al. 2011) as
collections of new and known species are growing and new records of soft corals can be expected from this poorly
studied area of the eastern Indian Ocean. Molecular techniques clarifying soft coral taxonomy are rapidly
advancing and becoming a reliable tool for understanding their taxonomic relationships (McFadden et al. 2006;
Breedy et al. 2012; McFadden et al. 2013). Future integrative studies will increase our knowledge and result in a
more reliable system for the Octocorallia. This work underpins the hypothesis that the west coast of Australia is a
unique ecosystem supporting a rich and diverse marine fauna (Marsh 1993; Bryce 2009; Masini et al. 2009; Wilson
2013), and further investigations will enhance our understanding of these unique environments.
Acknowledgements
Invertebrate collections were made as part of the Census of Marine Life (CReefs Program) funded by the
Australian Government under auspices of the Australian Institute of Marine Science, the Great Barrier Reef
Research Foundation and BHP Billiton. This research is part of Australian Biological Resources Study (ABRS)
National Taxonomic Research Grant (number 209-05) “Taxonomy of tropical Australian Octocorallia (Anthozoa:
Coelenterata) (J.N.A. Hooper & P.N.A. Alderslade).
BRYCE ET AL.
198
·
Zootaxa 3963 (2) © 2015 Magnolia Press
The work was made possible due to logistical support from the WAM funded by Woodside Energy, as a part of
the Kimberley Woodside Collection Project (2008–2011). The authors would like to gratefully acknowledge the
help provided by the collection manager of the QM, Merrick Ekins, as well as the scientific staff and ship's
company aboard the Kimberley Quest II. Angelo Poliseno acknowledges financial support from the Deutsche
Akademische Austauschdienst (DAAD). We also thank Prof. Gert Wörheide for providing access to the laboratory
facilities of the Department of Earth & Environmental Sciences, Paleontology & Geobiology. Leen P. van
Ofwegen, Naturalis Biodiversity Center, is thanked for providing information concerning the holotype of E.
splendens. Gary C. Williams is also acknowledged for taxonomic information concerning Paraminabea. Michael
Verrall (CSIRO, Perth) is thanked for SEM assistance.
References
Alderslade, P. (1998) Revisionary systematics in the gorgonian family Isididae, with descriptions of numerous new taxa
(Coelenterata: Octocorallia). Records of the Western Australian Museum, Supplement No. 55, 1–359.
Bayer, F.M. (1974) A new species of Trichogorgia and records of two other octocorals new to Palau Islands. Micronesica, 10,
257–271.
Benayahu, Y. & Schleyer, M.H. (1995) Corals of the south-west Indian Ocean II. Eleutherobia aurea spec. nov. (Cnidaria,
Alcyonacea) from deep reefs on the KwaZulu-Natal Coast, South Africa. South African Association for marine biological
research, Oceanographic Research Institute, Investigational Report, 68, 1–12.
Benayahu, Y., Jeng, M.S., Perkol-Finkel, S. & Dai, C.F. (2004) Soft Corals (Octocorallia: Alcyonacea) from Southern Taiwan. II.
Species Diversity and distributional patterns. Zoological Studies, 43(3), 548–560.
Breedy, O., van Ofwegen, L. & Vargas, S. (2012) A new family of soft corals (Anthozoa, Octocorallia, Alcyonacea) from the
aphotic tropical eastern Pacific waters revealed by integrative taxonomy. Systematics and Biodiversity, 10, 351–359.
http://dx.doi.org/10.1080/14772000.2012.707694
Brugler, M.R. & France, S.C. (2008) The mitochondrial genome of a deep-sea bamboo coral (Cnidaria, Anthozoa, Octocorallia,
Isididae): genome structure and putative origins of replication are not conserved among octocorals. Journal of Molecular
Evolution, 67, 125–136.
http://dx.doi.org/10.1007/s00239-008-9116-2
Bryce, C. (Ed.) (2009) Marine Biodiversity Survey of Mermaid Reef (Rowley Shoals), Scott and Seringapatam Reef. Records of
the Western Australian Museum, 77, 1–255.
Bryce, M. & Sampey, A. (2014) Kimberley Marine Biota: Soft corals and sea fans (Octocorallia). Records of the Western
Australian Museum 84 (in press).
Dautova, T.N. & Savinkin, O.S. (2009) New data on soft corals (Cnidaria: Octocorallia: Alcyonacea) from Nha Trang Bay, South
China Sea. Zootaxa, 2027, 1–27.
Darriba, D., Taboada, G.L., Doallo, R. & Posada D. (2012) jModelTest 2: more models, new heuristics and parallel computing.
Nature Methods 9 (8), 772.
http://dx.doi.org/10.1038/nmeth.2109
Dayrat, B. (2005) Towards integrative taxonomy. Biological Journal of the Linnean Society, 85, 407–415.
http://dx.doi.org/10.1111/j.1095-8312.2005.00503.x
Edgar, RC. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research
32, 1792–1797.
http://dx.doi.org/10.1093/nar/gkh340
Fabricius, K. & Alderslade, P. (2001) Soft corals and sea fans - a comprehensive guide to the tropical shallow water genera of
the central-west Pacific, the Indian Ocean and the Red Sea. Australian Institute of Marine Science, Townsville, 264 pp.
Fabricius, K. (2008) A brief photo guide to the shallow-water octocorals of the Rowley Shoals, Western Australia. Report to the
Department of Environment and Conservation, Government of Western Australia, 39 pp.
Faulkner, D. & Chesher, R. (1979) Living corals. Clarkson N. Potter, Inc., New York, 311 pp.
Imahara, Y., Iwase, F. & Namakawa, H. (2014) The octocorals of Sagami Bay. Tokyo University Press, 398 pp.
Keesing, J.K., Irvine, T.R., Alderslade, P., Clapin, G., Fromont, J., Hosie, A.M., Huisman, J.M., Phillips, J.C., Naughton, K.M.,
Marsh, L.M., Slack-Smith, S.M., Thomson, D.P. & Watson, J.E (2008) Marine benthic flora and fauna of Gourdon Bay
and the Dampier Peninsula in the Kimberley region of north-west Australia. Journal of the Royal Society of Western
Australia, 94, 285–301.
Kükenthal, W. (1906a) Alcyonaria. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition, 13 (1), 1–111.
Kükenthal, W. (1906b) Japanische Alcyonaceen. Abhandlungen der königlich Bayerischen Akademie der Wissenschaften. II.
KL. Suppl. Bd. 1 Abt. (Beitraege zur Naturgeschichte Ostasiens), pp. 1–86.
Lamouroux, J.V.F. (1812) Extrait d’un mémoire sur la classification des polypiers coralligènes non entièrement pierreux.
Nouveaux Bulletin Des Sciences par la Société. Philomathique de Paris, 3, 181-188.
Marsh, LM. (1993) Cnidaria, other than reef-building corals, of Ashmore Reef and Cartier Island. In: Berry, P.F. (Ed.) Marine
Zootaxa 3963 (2) © 2015 Magnolia Press
·
199
SOFT CORALS OF WESTERN AUSTRALIA
Faunal Survey of Ashmore Reef and Cartier Island, North-western Australia. Records of the Western Australian Museum,
44, 1–91.
Masini, R, Sim, C.B. & Simpson, C.J. (2009) Protecting the Kimberley. A synthesis of scientific knowledge to support
conservation management in the Kimberley region of Western Australia. Department of Environment and Conservation,
Western Australia, 45 pp.
McFadden, C.S., Tullis, I.D., Hutchinson, M.B., Winner, K. & Sohm, J.A. (2004) Variation in coding (NADH dehydrogenase
subunits 2, 3 and 6) and non-coding intergenic spacer regions of the mitochondrial genome in Octocorallia (Cnidaria:
Anthozoa). Marine Biotechnology, 7 (4), 405–406.
http://dx.doi.org/10.1007/s10126-005-1000-0
McFadden, C.S., France, S.C., Sanchez, J.A. & Alderslade, P. (2006) A molecular phylogenetic analysis of the octocorallia
(Cnidaria: Anthozoa) based on mitochondrial protein–coding sequences. Molecular Phylogenetics and Evolution, 41,
513–527.
http://dx.doi.org/10.1016/j.ympev.2006.06.010
McFadden, C.S., Benayahu, Y., Pante, E., Thoma, J.N, Nevarez, P.A. & France, S.C. (2011) Limitations of mitochondrial gene
barcoding in the cnidarian sub-class Octocorallia. Molecular Ecology Resources, 11, 19–31.
http://dx.doi.org/10.1111/j.1755-0998.2010.02875.x
McFadden, C.S. & van Ofwegen, L.P. (2012) Stoloniferous octocorals (Anthozoa, Octocorallia) from South Africa, with
descriptions of a new family of Alcyonacea, a new genus of Clavulariidae, and a new species of Cornularia
(Cornulariidae). Invertebrate Systematics, 26, 331–356.
http://dx.doi.org/10.1071/IS12035
McFadden, C.S. & Ofwegen, L.P. (2013) Molecular phylogenetic evidence supports a new family of octocorals and a new genus
of Alcyoniidae (Octocorallia, Alcyonacea). ZooKeys, 346, 59–83.
http://dx.doi.org/10.3897/zookeys.346.6270
McFadden, C.S., Brown, A.S., Brayton, C., Hunt, C.B. & Ofwegen, L.P. (2014) Application of DNA barcoding in biodiversity
studies of shallow-water octocorals: molecular proxies agree with morphological estimates of richness in Palau. Coral
Reefs, 33, 275–286.
http://dx.doi.org/10.1007/s00338-013-1123-0
Nutting, C.C. (1912) Description of the Alcyonaria collected by the U.S. Fisheries steamer “Albatross”, mainly in Japanese
waters, during 1906. Proceedings of the United States National Museum 43, 1–104.
http://dx.doi.org/10.5479/si.00963801.43-1923.1
Pante, E., France, S.C., Couloux, A., Cruaud, C., McFadden, C.S, Samadi, S. & Watling, L. (2012) Deep-Sea origin and in-situ
diversification of Chrysogorgiid octocorals. PLOS ONE 7 (6), 1–14.
http://dx.doi.org/10.1371/journal.pone.0038357
Pante, E., Abdelkrim, J., Viricel, A., Gey, D., France, S., Boisselier, M.C. & Samadi, S. (2014) Use of RAD sequencing for
delimiting species.
http://dx.doi.org/10.1038/hdy.2014.105
Pütter, A. (1990) Alcyonaceen des Breslauer Museum. Zoologische Jahrbücher (Systematik), 13, 443–462.
Ronquist, F. & Huelsenbeck, J. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19,
1572–1574.
http://dx.doi.org/10.1093/bioinformatics/btg180
Schleyer, M.H. & Benayahu, Y. (2008) Soft coral biodiversity and distribution in East Africa: Gradients, function and
significance. Proceedings of the 11
th
International Reef Symposium, Ft. Lauderdale, Florida, 7–11 July 2008, pp. 1388–1391.
Sanchez, J.A., McFadden, C.S., France, S.C. & Lasker, H.R. (2003) Molecular phylogenetic analyses of shallow-water
Caribbean octocorals. Marine Biology, 142, 975–987.
Shearer, T.L., van Oppen, M.J.H., Romano, S.L. & Wörheide, G. (2002) Slow mitochondrial DNA sequence evolution in the
Anthozoa (Cnidaria). Molecular Ecology, 11, 2475–2487.
http://dx.doi.org/10.1046/j.1365-294X.2002.01652.x
Stöver, B.C. & Müller, K.F. (2010) TreeGraph 2: Combining and visualizing evidence from different phylogenetic analyses.
BMC Bioinformatics, 11, 7.
http://dx.doi.org/10.1186/1471-2105-11-7
Stamatakis, A. (2006) RAxML–VI-HPC: maximum likelihood based phylogenetic analyses with thousands of taxa and mixed
models. Bioinformatics, 22, 2688–2690.
http://dx.doi.org/10.1093/bioinformatics/btl446
Stamatakis, A. Hoover, P. & Rougemont, J. (2008) A rapid bootstrap algorithm for the RAxML Web servers. Systematic
Biology, 57, 758–771.
http://dx.doi.org/10.1080/10635150802429642
Thoma, J.N., Pante, E., Brugler, M.R. & France, S.C. (2009) Deep-sea octocorals and antipatharians show no evidence of
seamount-scale endemism in the NW Atlantic. Marine Ecology Progress Series 397, 25–35.
http://dx.doi.org/10.3354/meps08318
Thomson, J.S. (1910) The Alcyonaria of the Cape of Good Hope and Natal. Alcyonacea. Transactions of the Royal Society of
Edinburgh, XLVII, 549–589.
BRYCE ET AL.
200
·
Zootaxa 3963 (2) © 2015 Magnolia Press
Thomson, J.A. & Henderson, W.D. (1906) An account of the Alcyonarians collected by the Royal Indian Marine survey ship
Investigator in the Indian Ocean. Printed by order of the trustees of the Indian Museum, 128 pp.
http://dx.doi.org/10.5962/bhl.title.26574
Thomson, J.A. & Dean, L.M.I. (1931) Alcyonacea of the Siboga Expedition with an addendum to the Gorgonacea. Siboga-
Expedition Reports, 13d, 1–227.
Utinomi, H. (1954) Some alcyoniid octocorals from Kii coast, middle Japan, with descriptions of two new species. Publications of
Seto Marine Biological Laboratory, 6 (2), 147–168.
Utinomi, H. (1957) The Alcyonarian genus Bellonella from Japan, with description of two new species. Publications of Seto
Marine Biological Laboratory, 6 (2), 24–44.
Utinomi, H. (1960) Noteworthy octocorals collected off the southwest coast of Kii Peninsula, middle Japan. 1. Stolonifera and
Alcyonacea. Publications of Seto Marine Biological Laboratory, 8 (1), 1–26.
Utinomi, H. & Harada, E. (1958) A list of bottom animals collected by a trawler “Kaiun-maru” off the southwest coast of Kii
Peninsula. Publications of Seto Marine Biological Laboratory, 6 (3), 385–395.
Vargas, S., Guzman, H.M., Breedy, O. & Wörheide, G. (2014) Molecular phylogeny and DNA barcoding of tropical eastern
Pacific shallow-water gorgonian octocorals. Marine Biology, 161 (5), 1027–1038
http://dx.doi.org/10.1007/s00227-014-2396-8
Verseveldt, J. & Bayer, F.M. (1988) Revision of the genera Bellonella, Eleutherobia, Nidalia and Nidaliopsis (Octocorallia:
Alcyoniidae and Nidalliidae), with description of two new genera. Zoologische Verhandelingen Leiden, 245, 1–131.
Will, K.P., Mishler, B.D. & Wheeler, Q.D. (2005) The perils of DNA Barcoding and the need for integrative taxonomy.
Systematic Biology, 4, 844–851.
http://dx.doi.org/10.1080/10635150500354878
Williams, G.C. (1986) Morphology, systematics and variability of the southern African soft coral Alcyonium variabile (J. Stuart
Thomson, 1921) (Octocorallia, Alcyoniidae). Annals of the South Africa Museum, 96, 241–270.
Williams, G.C. (1992) Revision of the soft coral genus Minabea (Octocorallia: Alcyoniidae), with new taxa from the Indo-West
Pacific. Proceedings of the California Academy of Sciences, 48, 1–26.
Williams, C.G. (1992) The Alcyonacea of southern Africa. Stoloniferous octocorals and soft corals (Coelenterata, Anthozoa).
Annals of the South African Museum, 100, 249–358.
Williams, G.C. (2000) A new species of the soft coral genus Eleutherobia Pütter, 1900 (Coelenterata: Alcyonacea) from the Tonga
Islands. Proceedings of the California Academy of Sciences, 52, 159–169.
Williams, G.C. (2001) First record of a bioluminescent soft coral: description of a disjunct population of Eleutherobia grayi
(Thomson & Dean, 1931) from the Solomon Islands, with a review of bioluminescence in the Octocorallia. Proceedings of
the California Academy of Science, 52, 209–225.
Williams, G.C. (2003) Capitate taxa of the soft coral genus Eleutherobia (Octocorallia: Alcyoniidae) from Palau and South Africa:
a new species and a new combination. Zoologische Verhandelingen Leiden, 345, 419–436.
Williams, G.C. & Alderslade, P. (1999) Revisionary systematics of the western Pacific soft coral genus Minabea (Octocorallia:
Alcyoniidae), with descriptions of a related new genus and species from the Indo-Pacific. Proceedings of the California
Academy of Sciences, 51, 337–364.
Williams, G.C. & Little, S.A (2001) A new species of the soft coral genus Eleutherobia Pütter, 1900, (Octocorallia: Alcyoniidae)
from South Africa. Proceedings of the California Academy of Sciences, 52, 195–208.
Wilson, B. (2013) The biogeography of the Australian North West Shelf: Environmental change and life’s response. Elsevier,
413 pp.
Yang, Z. (1994) Maximum–likelihood phylogenetic estimation from DNA-sequences with variable rates over sites–
approximate methods. Journal of Molecular Evolution, 39, 306–314.
http://dx.doi.org/10.1007/BF00160154
