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Molecular systematics of the speciose Indo-Pacific soft coral genus, Sinularia
(Anthozoa: Octocorallia)
Catherine S. McFadden,
1,a
Leen P. van Ofwegen,
2
Elizabeth J. Beckman,
1
Yehuda Benayahu,
3
and Phil Alderslade
4
1
Department of Biology, Harvey Mudd College, Claremont, California, USA
2
National Natural History Museum (Naturalis), Leiden, The Netherlands
3
Department of Zoology, University of Tel Aviv, Ramat Aviv, Tel Aviv, Israel
4
CSIRO, Marine & Atmospheric Research, Castray Esplanade, Hobart, Tasmania, Australia
Abstract. The speciose tropical soft coral genus Sinularia traditionally has been divided into
five intrageneric taxonomic groups based on variation in a single morphological character:
the shape of the club sclerites (calcite skeletal elements) embedded in the surface tissues of the
colony. To test the phylogenetic utility of this system of classification, we used a 735-bp
fragment of the octocoral-specific mitochondrial msh1 gene to construct a molecular phy-
logeny that included 80 of the B150 recognized morphospecies of Sinularia. The msh1 phy-
logeny recovered five well-supported clades, but they were not congruent with the traditional
intrageneric taxonomic groups. Mapping of characters onto the tree suggested that the five
major clades plus several additional sub-clades of Sinularia can be distinguished based on a
suite of four morphological characters; these include the presence of sclerites in the tentacle,
collaret, and point regions of the polyps, in addition to the shape of the club sclerites in the
surface tissues. The overall growth form of the colony also distinguishes some clades. Polyp
sclerites have for the most part been overlooked taxonomically in Sinularia, and as a result
information on these characters is lacking or is incorrect in many species descriptions. As has
been the case in other recent studies of lower metazoan groups, construction of a molecular
phylogeny has led us to recognize the phylogenetic and taxonomic importance of previously
overlooked morphological characters. A revised taxonomic key that includes these characters
is already improving our ability to discriminate species boundaries, and facilitating descrip-
tion of new Sinularia species.
Additional key words: Alcyoniidae, molecular phylogenetics, taxonomy, msh1
Use of molecular systematic approaches continues
to revolutionize our understanding of evolutionary
relationships within and among higher taxa in the
lower metazoan phyla, in particular, the cnidarians
and sponges. Demosponges, scleractinian corals, and
octocorals are three major groups in which molecular
data are necessitating a radical reappraisal of ordi-
nal- and family-level taxonomy (Romano & Palumbi
1996; Fukami et al. 2004, 2008; Boury-Esnault 2006;
McFadden et al. 2006b). Historically, taxonomic
classification and phylogenetic inference in these
groups has been challenging because of their relative
morphological simplicity and a consequent paucity
of characters suitable for analysis (e.g., Bayer 1956).
Many of the morphological characters tradition-
ally used for higher level classification in these
taxa have now been shown to be discordant with
molecular phylogenetic evidence, and likely repre-
sent homoplasies.
In many cases in which there is a disagreement be-
tween classification based on traditional morphol-
ogy-based and molecular phylogenetic approaches,
careful re-examination of morphology has revealed
new sets of characters that are indeed congruent with
the molecular data. For example, new sets of phylo-
genetically informative microskeletal characters have
been discovered for the scleractinian families Favi-
idae (Fukami et al. 2004) and Mussidae (Budd &
Stolarski 2009) after molecular studies showed the
traditional taxonomy of these groups to be incorrect
Invertebrate Biology 128(4): 303–323.
r2009, The Authors
Journal compilation r2009, The American Microscopical Society, Inc.
DOI: 10.1111/j.1744-7410.2009.00179.x
a
Author for correspondence.
E-mail: mcfadden@hmc.edu
(Fukami et al. 2004, 2008). In Octocorallia, molecular
studies clearly indicate that the traditional morphol-
ogy-based sub-ordinal and family-level taxonomy
does not reflect phylogenetic relationships (McFad-
den et al. 2006b). Re-examination of some unexpected
relationships has, however, facilitated identification
of new morphological characters that are congruent
with molecular data. For example, by mapping mor-
phological characters onto a molecular phylogeny,
Sa
´nchez et al. (2003) found that the characters that
have traditionally been considered most important
for classifying octocorals belonging to the holaxon-
ian group of sea fans (axial structure and the size and
shape of surface sclerites) were not congruent with
the phylogeny, whereas several characters heretofore
considered of only minor taxonomic importance
(presence of polyp sclerites and sclerite ornamenta-
tion) were diagnostic for the major molecular clades.
Similarly, in their analysis of a cryptic genus of soft
corals revealed by molecular data, McFadden et al.
(2006a) showed that several previously overlooked
morphological characters (polyp sclerites and subtle
aspects of colony growth form) clearly separate this
genus from two other genera with which its members
had been confounded. In the present study, we use a
similar character-mapping approach to identify mor-
phological characters congruent with sub-generic
phylogenetic relationships in the speciose soft coral
genus Sinularia M
AY
1898.
Members of the genus Sinularia are among the
most widespread and commonly encountered octo-
corals on Indo-Pacific coral reefs (Benayahu & Loya
1977; Tursch & Tursch 1982; Dinesen 1983; Riegl
et al. 1995; Fabricius 1997). Their fleshy colonies of-
ten attain a large size and monopolize extensive areas
of shallow reef habitat (Benayahu & Loya 1981; Fab-
ricius 1995, 1997; Fabricius & Dommisse 2000; Bast-
idas et al. 2004), particularly in disturbed, near-shore
areas where they are sometimes considered invasive
(Fabricius 1998). Despite their ecological dominance
in some habitats, species of Sinularia have also been
impacted severely by recent coral bleaching events,
suffering mortality rates 490% in some locations
(Fabricius 1999; Bruno et al. 2001; Loya et al. 2001).
Work on the ecology of this important group has,
however, been hampered by taxonomic uncertainties
and difficulty in distinguishing species in the field,
limiting most studies to the generic level (e.g., Fab-
ricius 1995). Colony growth morphology is often
highly variable (Benayahu et al. 1998), even among
parts of the same colony (Alderslade & Shirwaiker
1991; Benayahu 1998), and examination of micro-
scopic sclerites (calcite skeletal elements) in the tis-
sues is required to distinguish species reliably. Many
species remain undescribed, while the validity of oth-
ers is uncertain (van Ofwegen 2002).
Until now, most taxonomic work on Sinularia has
followed the protocol of Verseveldt (1980), who pub-
lished the last comprehensive taxonomic revision of
the genus. The two characters that he considered the
most important for taxonomy and that are the ex-
clusive focus of his and most subsequent studies are:
(1) the morphology of the club-shaped sclerites found
in the surface layer of the colony and (2) the overall
colony growth form as assessed primarily from pre-
served material. In his revision, Verseveldt (1980)
presented a key in which he subdivided Sinularia
into five groups based on characteristics of the club
sclerites, a system of classification that taxonomists
have continued to use (e.g., Verseveldt & Benayahu
1983; Alderslade 1987; Alderslade & Baxter 1987;
van Ofwegen & Vennam 1991; Benayahu 1993; van
Ofwegen 2001). Verseveldt’s Group I is the most dis-
tinctive of these five, having club sclerites with a
characteristic shape referred to as ‘‘leptoclados-
type’’ clubs after the nominate species in the group,
Sinularia leptoclados (E
HRENBERG
1834). Group II in-
cludes species in which the sclerites are not of the
leptoclados-type, but instead have a central wart
(Fig. 1). Groups III and IV have club sclerites that
lack a central wart and are distinguished from one
another by size (Group III: clubs o0.12 mm long;
Group IV: clubs 40.12 mm long). Species in Group
V have few or no club sclerites on the colony surface.
van Ofwegen (2002) summarized the current taxo-
nomic status of Sinularia, and recognized 128 valid
species. Since then, an additional 26 species of this
Fig. 1. Illustration of an octocoral polyp indicating the
locations and types of sclerites typically found in genus
Sinularia. Modified from Bayer et al. (1983).
304 McFadden, van Ofwegen, Beckman, Benayahu, & Alderslade
Invertebrate Biology
vol. 128, no. 4, fall 2009
genus have been described (Manuputty & van Of-
wegen 2007; van Ofwegen 2008a,b), bringing the to-
tal to B150. This recent taxonomic explosion has
been fueled in part by the recognition that preserved
specimens may not retain important aspects of col-
ony growth form that can be observed easily in situ.
Species whose colony form differs markedly when
alive are often transformed by preservation such that
their differences are less apparent (van Ofwegen
2008b), and as a result many distinct species have
gone unrecognized. Careful documentation of colony
growth form in situ is now improving taxonomic res-
olution as well as facilitating identification of species
in the field (van Ofwegen 2008b).
A recent molecular phylogeny of the related alcyo-
niid genera Sarcophyton L
ESSON
1834 and Lobophytum
V
ON
M
ARENZELLER
1886 revealed unexpected clades
(including a previously unsuspected third genus), and
subsequently led to the identification of a new set of
morphological characters for taxonomy (McFadden
et al. 2006a). In particular, the presence, distribution,
and form of sclerites in the polyps, characters that
were rarely mentioned by Verseveldt (1980, 1982,
1983) in his revisions, were discovered to be phylo-
genetically and taxonomically informative characters
for these genera. In the present study, we construct a
molecular phylogeny for the genus Sinularia using
the same genetic marker (the octocoral-specific mi-
tochondrial protein-coding gene msh1) used effec-
tively in our work on Sarcophyton and Lobophytum
(McFadden et al. 2006a). We test for congruence be-
tween the molecular phylogeny and Verseveldt’s
(1980) taxonomic groups, and subsequently use the
phylogeny to identify a new suite of morphological
characters that distinguish five major clades and sev-
eral distinct sub-clades within the genus Sinularia.
Methods
Specimens of Sinularia were collected and identi-
fied by the authors from Ambon, Moluccas, Indone-
sia (November 1996); Gulf of Carpentaria, NT,
Australia (December 2003); Republic of Palau
(May 2005); and Eilat, Israel (July 2007). Taxonomic
accounts and descriptions of new species have been
published for the Gulf of Carpentaria, Moluccas, and
Palau collections (Manuputty & van Ofwegen 2007;
van Ofwegen 2008a,b). Specimens have been depos-
ited in the Museum and Art Gallery of the Northern
Territory, Australia (NTM); the Nationaal Nat-
uurhistorish Museum, Leiden (NNM, formerly
RMNH); and the Tel Aviv University Zoological
Museum, Israel (ZMTAU) (Appendix 1). Additional
species of Sinularia were obtained from collections in
the NTM and the Florida Natural History Museum
(UF), a majority of them collected throughout the
Indo-Pacific from 1995 to 2002 by the Coral Reef
Research Foundation, Republic of Palau, and iden-
tified by PA.
Extraction of DNA from ethanol-preserved tissue
samples, PCR amplification, and sequencing of the
msh1 gene followed the protocols published in
McFadden et al. (2006a). Sequence data were proof-
read using LaserGene software (DNASTAR, Inc.,
Madison, WI, USA), and aligned using Muscle v.
3.6 (Edgar 2004). PhyML (Guindon & Gascuel 2003)
was used to construct maximum likelihood trees using
aGTR1I1Gmodel of substitution with 100 boot-
strap replicates. Maximum parsimony analyses were
run using TNT (Goloboff et al. 2000) with default
parameters and 100 bootstrap replicates. Bayesian
analysis was conducted using MRBAYES v 3.1
(Huelsenbeck & Ronquist 2001) with a GTR1I1G
model run for 5 10
6
generations (burnin 510
6
gen-
erations). PAUP
(Swofford 2002) was used to com-
pute pairwise genetic distances (Kimura 2-parameter)
among taxa, and to generate maximum likelihood
scores for trees constrained to reflect monophyly of
Verseveldt’s (1980) five taxonomic groups. Con-
strained trees were compared with the best-fit maxi-
mum likelihood tree using the Shimodaira–Hasegawa
(SH) test (Shimodaira & Hasegawa 1999) with 10,000
RELL bootstrap replicates. Species of Sarcophyton,
Lobophytum, and Dampia A
LDERSLADE
1983 were in-
cluded as outgroup taxa in all analyses; molecular
analyses of relationships within Octocorallia support
these three genera as the clades most closely related to
Sinularia (McFadden et al. 2006b).
Following phylogenetic reconstruction, morpho-
logical character states were mapped onto the tree
manually. Specific characters examined included the
presence, form, and size of the sclerites in the tenta-
cles, collaret and point regions of the polyp (Fig. 1; see
Bayer et al. 1983 for further definitions of anatomical
terms); presence, shape, and size of the club sclerites
in the colony surface tissues; and overall colony
growth form. When information on particular char-
acters (especially polyp sclerites) could not be found
in published taxonomic descriptions or conflicted
with the observed phylogenetic distribution of traits,
we re-examined original material if it was available,
including type specimens and the permanent sclerite
slides prepared by Verseveldt, now kept at NNM.
Results
A total of 735 nucleotides of the msh1 sequence
were obtained for 119 specimens representing 80
Phylogeny of Sinularia 305
Invertebrate Biology
vol. 128, no. 4, fall 2009
morphospecies of Sinularia (Appendix 1). No indels
were present, and nucleotide alignments were unam-
biguous. All phylogenetic methods supported the
same tree topology (Fig. 2). We identified five major
clades of Sinularia, each supported by bootstrap val-
ues 490% and Bayesian posterior probabilities
40.95. The two largest clades (4, 5) each could be
further subdivided into three or four sub-clades; the
three sub-clades within clade 5 were supported
strongly by all analyses, whereas only one of the
four sub-clades identified within clade 4 was well sup-
ported by all phylogenetic methods (Fig. 2). All an-
alyses recovered clade 1 as the sister clade to all other
Sinularia species. The phylogenetic relationships
among the other four major clades, however, were
unresolved (Fig. 2).
Clade 1 includes four specimens: three identified as
Sinularia brassica M
AY
1898, and one as Sinularia
dura (P
RATT
1903). These two species, which differ
only in colony growth form, have been synonymized
previously (Benayahu et al. 1998), although the two
distinct haplotypes we found are suggestive of two
species (Fig. 2). The shape of the club sclerites (very
wide heads with no central wart) is unique within the
genus (Fig. 3E, b), as is the enormous variation in
colony growth form that these species exhibit (Bena-
yahu et al. 1998). Clade 1 is further distinguished
from the other four major clades by the presence of
scales in the tentacles (Fig. 3E, a). Point and collaret
sclerites are absent (Table 1).
Clade 2 is formed by Sinularia vrijmoethi V
ERSE-
VELDT
1971, Sinularia flaccida
VAN
O
FWEGEN
2008b,
Sinularia loyai V
ERSEVELDT
&B
ENAYAHU
1983, and
Sinularia grandilobata V
ERSEVELDT
1980. All four
species have in common a well-developed collaret
and points (collaret spindles to 0.25–0.4mm length
[Fig. 3D, b]; point clubs to 0.17–0.28 mm length [Fig.
3D, a]) as well as rods in the tentacles (Table 1). On
the colony surface, they all have club sclerites with a
central wart (see Fig. 1) that is somewhat obscured by
three lateral warts (Fig. 3D, c). Sinularia vrijmoethi,
S. flaccida, and S. loyai are very similar in having
long clubs (averaging 0.26, 0.29, and 0.30 mm, re-
spectively); S. grandilobata, which forms a sister
clade to the other three species, has much shorter
clubs (0.10 mm). Sinularia grandilobata also has an
encrusting colony growth form, while the other
three species are stalked.
Clade 3 is formed by Sinularia foveolata V
ERSE-
VELDT
1974, Sinularia fungoides T
HOMSON
&H
ENDER-
SON
1906, and the monotypic species Dampia
pocilloporaeformis A
LDERSLADE
1983, which falls
within genus Sinularia rather than within the other
outgroup taxa (Fig. 2). Verseveldt (1980) classified
the two former species in his Group III: ‘‘most clubs
0.06–0.12 mm long, not of the leptoclados-type, and
without central wart.’’ However, all three species
clearly have clubs with a central wart, although it is
mostly nestled within the three lateral warts below it,
which tend to obscure it and give the club head a tri-
angular shape (Fig. 3C). This unique club shape is
diagnostic for clade 3. In addition, species in this
clade have indistinct points formed by rod-like scle-
rites (see Alderslade 1983: fig. 4J–M; Alderslade
1987: fig. 9).
Clade 4 is the largest clade in the genus (Fig. 2).
Species belonging to this group have clubs with a
central wart, and lack collaret and tentacle sclerites
(but see ‘‘Discussion’’); the majority also lack point
sclerites. Clade 4 can be divided into four sub-clades
(4A–D) that vary in support values but can be dis-
tinguished morphologically (Table 1). Sub-clade 4A
is the most distinct of these groups genetically, and is
supported by bootstrap values 490%. It includes
Sinularia querciformis (P
RATT
1903) and Sinularia va-
riabilis T
IXIER
-D
URIVAULT
1945, both of which Verse-
veldt (1980) classified as Group IV (‘‘clubs with no
central wart’’), as well as Sinularia flexibilis (Q
UOY
&
G
AIMARD
1833) and Sinularia procera V
ERSEVELDT
1977, placed by Verseveldt in Group V (‘‘No [or
hardly any] clubs in the surface layer of the lobes’’).
Sinularia querciformis and S. variabilis do have clubs
with a central wart, but this central wart is often leaf-
like, obscuring the arrangement (Fig. 3B, b). Sinul-
aria procera and S. flexibilis normally have no clubs
on the surface layer of the lobes (Verseveldt 1977,
1980). van Ofwegen & Vennam (1994), however,
found a specimen of S. flexibilis from Ambon that
had surface clubs, mostly with a central wart and not
unlike those of S. querciformis and S. variabilis. All
four species in sub-clade 4A have a similar colony
growth form with a stalk.
Sub-clade 4B is formed by Sinularia he-
terospiculata V
ERSEVELDT
1970, Sinularia notanda
T
IXIER
-D
URIVAULT
1966, Sinularia humilis
VAN
O
FWE-
GEN
2008b, Sinularia numerosa T
IXIER
-D
URIVAULT
1970, Sinularia sublimis
VAN
O
FWEGEN
2008b, Sinul-
aria curvata M
ANUPUTTY
&O
FWEGEN
2007, Sinularia
ultima
VAN
O
FWEGEN
2008b, Sinularia sobolifera
V
ERSEVELDT
&T
URSCH
1979, Sinularia cruciata T
IX-
IER
-D
URIVAULT
1970, and Sinularia polydactyla
(E
HRENBERG
1834). Although support for monophyly
was lacking for this group, all of these species have in
common the presence of point sclerites (Fig. 3A, a), a
character that distinguishes them from other mem-
bers of clade 4.
Sub-clades 4C and 4D include a large number of
species (Table 1) that all lack sclerites in the polyps
306 McFadden, van Ofwegen, Beckman, Benayahu, & Alderslade
Invertebrate Biology
vol. 128, no. 4, fall 2009
0.02
*
4C
4D
4B
4A
2
5C
5B
5A
1
3
2
4
1
5
3
*
*
*
*
*
*
*
*
*
*
*
Fig. 2. Maximum likelihood tree of genus Sinularia constructed using 735 bp of the mitochondrial gene msh1. Major
clades discussed in the text are indicated by circled numbers, and sub-clades by vertical bars to the right of node labels.
Numbers above branches indicate bootstrap values from maximum likelihood (left) and maximum parsimony (right)
analyses; numbers below branches are Bayesian posterior probabilities. 100, both bootstrap values 100 and Bayesian
credibility 1.0; Asterisk, bootstrap values 470 and posterior probabilities 40.90. Roman numerals following specimen
names indicate the sub-generic taxonomic group to which a species has been assigned traditionally based on the
classification of Verseveldt (1980).
Phylogeny of Sinularia 307
Invertebrate Biology
vol. 128, no. 4, fall 2009
Fig. 3. Typical morphology of sclerites from the colony surface and polyps of Sinularia, clades 1 to 4B. A. Clade 4B:
Sinularia humilis RMNH Coel. 38737; a, point clubs. B. Clade 4A: a, Sinularia flexibilis, clubs from the base of the colony
(uncatalogued colony from Raja Ampat, Indonesia); b, Sinularia querciformis RMNH Coel. 34308. C. Clade 3: a,
Sinularia fungoides RMNH Coel. 34321; b, Dampia pocilloporaeformis RMNH Coel. 19843. D. Clade 2: Sinularia flaccida
RMNH Coel. 38731; a, point clubs; b, collaret spindle; c, surface clubs. E. Clade 1: Sinularia brassica RMNH Coel. 34304;
a, tentacle scales; b, surface clubs.
308 McFadden, van Ofwegen, Beckman, Benayahu, & Alderslade
Invertebrate Biology
vol. 128, no. 4, fall 2009
Table 1. Summary of diagnostic morphological characteristics of clades of Sinularia identified in the molecular phy-
logeny (Fig. 2). Confirmed clade members 5species included in the phylogeny. Additional species not included in the
phylogeny, but predicted to belong to each clade on the basis of morphology: 2: S. dactyloclados, S. elongata, S. licrocla-
dos, S. macrodactyla, S. manaarensis, S. parva, S. schumacheri.3:S. molokaiensis.S. megasclera, 4A: S. arborea, S. as-
terolobata, S. flabelliclavata, S. inexplicita, S. portieri.4B:?S. barcaformis, S. capillosa, S. macropodia, S. mira, S. mollis,
?S. muqeblae, S. platylobata, S. sandensis. 4C/4D: S. acetabulata, S. agilis, S. deformis, S. discrepens, S. flexuosa, S. gib-
berosa, S. higae, S. incompleta, S. inflata, S. microclavata, S. ovispiculata, S. paulae, S. tanakai, S. venusta, S. yamazatoi.
5A/B: S. crustaformis, S. granosa, S. lochmodes, S. ramosa, S. recurvata, S. slieringsi.5C:S. corpulentissima, S. dissecta, S.
exilis, S. facile, S. firma, S. fishelsoni, S. inelegans, S. jasminae, S. kavarattiensis, S. microspiculata, S. vanderlandi.
Clade Confirmed clade members Points Collaret Tentacle
sclerites
Surface clubs Other
1S. brassica M
AY
1898 No No Yes, scales Wide heads Variable
colony
shape
S. dura (P
RATT
1903)
2S. flaccida
VAN
O
FWEGEN
2008b
Yes, strong Yes,
strong
Yes, rods Central wart
obscured by
three lateral
warts
S. grandilobata V
ERSEVELDT
1980
S. loyai V
ERSEVELDT
&
B
ENAYAHU
1983
S. vrijmoethi V
ERSEVELDT
1971
3D.pocilloporaeformis
A
LDERSLADE
1983
Yes,
indistinct
No No Central wart
obscured by
three lateral
warts
S. foveolata V
ERSEVELDT
1974
S. fungoides T
HOMSON
&
H
ENDERSON
1906
4A S. flexibilis (Q
UOY
&G
AIMARD
1833)
No No No Central wart
often leaf-
like; or no
surface
clubs
Colony has a
stalk
S. procera V
ERSEVELDT
1977
S. querciformis (P
RATT
1903)
S. variabilis T
IXIER
-
D
URIVAULT
1945
4B S. cruciata T
IXIER
-D
URIVAULT
1970
Yes No No Central wart
distinct
S. curvata M
ANUPUTTY
&
O
FWEGEN
2007
S. heterospiculata V
ERSEVELDT
1970
S. humilis
VAN
O
FWEGEN
2008b
S. notanda T
IXIER
-D
URIVAULT
1966
S. numerosa T
IXIER
-
D
URIVAULT
1970
S. polydactyla (E
HRENBERG
1834)
S. sobolifera V
ERSEVELDT
&
T
URSCH
1979
S. sublimis
VAN
O
FWEGEN
2008b
S. ultima
VAN
O
FWEGEN
2008b
4C S. abhishiktae O
FWEGEN
&
V
ENNAM
1991
No No No Central wart
distinct,
warts often
leaf-like
S. babeldaobensis
VAN
O
FWEGEN
2008b
S. bremerensis
VAN
O
FWEGEN
2008a
S. capitalis (P
RATT
1903)
Phylogeny of Sinularia 309
Invertebrate Biology
vol. 128, no. 4, fall 2009
S. confusa
VAN
O
FWEGEN
2008a
S. diffusa
VAN
O
FWEGEN
2008a
S. foliata
VAN
O
FWEGEN
2008b
S. gravis T
IXIER
-D
URIVAULT
1970
S. mammifera M
ALYUTIN
1990
S. rigida (D
ANA
1846)
S. siaesensis
VAN
O
FWEGEN
2008b
S. tumulosa
VAN
O
FWEGEN
2008b
4D S. compressa T
IXIER
-
D
URIVAULT
1945
No No No Central wart
distinct
S. conferta (D
ANA
1846)
S. crassa T
IXIER
-D
URIVAULT
1945
S. crebra
VAN
O
FWEGEN
2008b
S. finitima
VAN
O
FWEGEN
2008b
S. gaweli V
ERSEVELDT
1978
S. linnei
VAN
O
FWEGEN
2008a
S. luxuriosa
VAN
O
FWEGEN
2008b
S. nanolobata V
ERSEVELDT
1977
S. ornata T
IXIER
-D
URIVAULT
1970
S. papula
VAN
O
FWEGEN
2008a
S. pavida T
IXIER
-D
URIVAULT
1970
S. polydactyla (E
HRENBERG
1834)
S. scabra T
IXIER
-D
URIVAULT
1970
S. uniformis
VAN
O
FWEGEN
2008b
S. verruca
VAN
O
FWEGEN
2008b
S. woodyensis
VAN
O
FWEGEN
2008a
5A S. gardineri (P
RATT
1903) Yes Yes Yes, scales Central wart
distinctS. halversoni V
ERSEVELDT
1974
S. humesi V
ERSEVELDT
1968
S. mauritiana V
ENNAM
&
P
ARULEKAR
1994
5B S. cruciata T
IXIER
-D
URIVAULT
1970
Yes Yes Yes, rods Central wart
distinct
S. depressa T
IXIER
-D
URIVAULT
1970
S. hirta (P
RATT
1903)
S. lamellata V
ERSEVELDT
&
T
URSCH
1979
S. terspilli V
ERSEVELDT
1971
Table 1. (cont’d).
Clade Confirmed clade members Points Collaret Tentacle
sclerites
Surface clubs Other
310 McFadden, van Ofwegen, Beckman, Benayahu, & Alderslade
Invertebrate Biology
vol. 128, no. 4, fall 2009
and have clubs with a distinct central wart. Maxi-
mum likelihood analysis supported the monophyly of
sub-clade 4C (bootstrap value 590%), but other
phylogenetic methods did not support the distinction
of sub-clades 4C and 4D (Fig. 2). Nonetheless, there
is often a detectable, albeit subtle, difference in the
typical shape of the club sclerites between these two
groups, with those in sub-clade 4C having more leaf-
like warts (Fig. 4D,E).
Two very different msh1 haplotypes were found in
specimens identified as S. polydactyla: one belonging
to sub-clade 4B and the other to 4D (Fig. 2). Spec-
imens collected from the Red Sea (type locality of the
species) that were present in the NNM and identified
as S. polydactyla show the normal point clubs of sub-
clade 4B, while a re-examination of RMNH-Coel.
38442 found no polyp sclerites whatsoever, consis-
tent with its placement in sub-clade 4D.
Clade 5 is another large clade that can be divided
into three sub-clades, each with high support values
(Fig. 2). All of the species in clade 5 have a collaret
and point sclerites present in the polyps. Sub-clade
5A includes Sinularia gardineri, Sinularia halversoni
V
ERSEVELDT
1974, Sinularia mauritiana V
ENNAM
&
P
ARULEKAR
1994, and Sinularia humesi V
ERSEVELDT
1968, all of which have club sclerites with a distinct
central wart (Fig. 4C, c). Sinularia gardineri, S.
halversoni, and S. humesi also have many scales in
5C S. abrupta T
IXIER
-D
URIVAULT
1970
Yes Yes ?rods Leptoclados-
type clubs
S. acuta M
ANUPUTTY
&
O
FWEGEN
2007
S. bisulca
VAN
O
FWEGEN
2008b
S. compacta T
IXIER
-
D
URIVAULT
1970
S. cristata T
IXIER
-D
URIVAULT
1969
S. densa (W
HITELEGGE
1897)
S. digitata
VAN
O
FWEGEN
2008b
S. erecta T
IXIER
-D
URIVAULT
1945
S. gaveshaniae A
LDERSLADE
&
S
HIRWAIKER
1991
S. intacta T
IXIER
-D
URIVAULT
1970
S. leptoclados (E
HRENBERG
1834)
S. longula M
ANUPUTTY
&
O
FWEGEN
2007
S. maxima V
ERSEVELDT
1971
S. molesta T
IXIER
-D
URIVAULT
1970
S. muralis M
AY
1899
S. parulekari A
LDERSLADE
&
S
HIRWAIKER
1991
S. robusta M
ACFADYEN
1936
S. rotundata T
IXIER
-
D
URIVAULT
1970
S. verseveldti O
FWEGEN
1996
Different specimens of S. polydactyla and S. cruciata exhibit msh1 haplotypes characteristic of two different clades or
sub-clades.
Table 1. (cont’d).
Clade Confirmed clade members Points Collaret Tentacle
sclerites
Surface clubs Other
Phylogeny of Sinularia 311
Invertebrate Biology
vol. 128, no. 4, fall 2009
Fig. 4. Typical morphology of sclerites from the colony surface and polyps of Sinularia, clades 4C to 5C. A. Clade 5C; a–
c, Sinularia digitata
VAN
O
FWEGEN
2008b RMNH Coel.38727; a, point clubs; b, collaret spindle; c, surface clubs; d,
Sinularia acuta RMNH Coel. 38432. B. Clade 5B: Sinularia depressa RMNH Coel. 38399; a, point clubs; b, collaret
spindle; c, surface clubs. C. Clade 5A: Sinularia humesi RMNH Coel. 38418; a, point clubs; b, collaret spindle; c, surface
clubs; d, tentacle scales. D. Clade 4D: Sinularia uniformis
VAN
O
FWEGEN
2008b RMNH Coel. 38753. E. Clade 4C:
Sinularia foliata
VAN
O
FWEGEN
2008b RMNH Coel. 38732.
312 McFadden, van Ofwegen, Beckman, Benayahu, & Alderslade
Invertebrate Biology
vol. 128, no. 4, fall 2009
the tentacles (Fig. 4C, d), the latter two species even
sharing a particular type of tentacle scale (see Manu-
putty & van Ofwegen 2007: fig. 13A). The tentacle
scales of S. gardineri are less pronounced. Sinularia
mauritiana has been described as lacking polyp scle-
rites (Vennam & Parulekar 1994), but we were unable
to verify that character state for the specimen in-
cluded here.
Sub-clade 5B includes Sinularia lamellata V
ERSE-
VELDT
&T
URSCH
1979, S. cruciata NTM-C13505,
Sinularia hirta (P
RATT
1903), Sinularia depressa T
IX-
IER
-D
URIVAULT
1970, and Sinularia terspilli V
ERSE-
VELDT
1971. Like sub-clade 5A, these species all
have club sclerites with a distinct central wart, and
the polyps have a collaret and points (Fig. 4B). No-
tably, however, the tentacular sclerites are rods
rather than scales, a character that distinguishes
this sub-clade from 5A (Table 1). Two distinct msh1
haplotypes were found for S. cruciata: one in sub-
clade 5B and the other in sub-clade 4B. Microscope
slides of the type show characters consistent with
placement in sub-clade 4B, and suggest that the spec-
imen identified as S. cruciata in sub-clade 5B (NTM-
C13505) is a different species.
Sub-clade 5C makes up the majority of clade 5 and
includes species with leptoclados-type clubs in the
colony surface (Fig. 4A), as well as collaret and point
sclerites in the polyps. A majority of the species in
this sub-clade (Sinularia abrupta T
IXIER
-D
URIVAULT
1970, Sinularia parelukari A
LDERSLADE
&S
HIRWAIKER
1991, Sinularia densa (W
HITELEGGE
1897), Sinularia
leptoclados, Sinularia rotundata T
IXIER
-D
URIVAULT
1970, Sinularia intacta T
IXIER
-D
URIVAULT
1970, Sin-
ularia verseveldti O
FWEGEN
1996, Sinularia compacta
T
IXIER
-D
URIVAULT
1970, Sinularia acuta M
ANUPUTTY
&O
FWEGEN
2007, Sinularia longula M
ANUPUTTY
&
O
FWEGEN
2007, Sinularia molesta T
IXIER
-D
URIVAULT
1970, Sinularia maxima V
ERSEVELDT
1971, Sinularia
muralis M
AY
1899, and Sinularia erecta T
IXIER
-D
URI-
VAULT
1945) have previously been placed in Group I
of Verseveldt (1980) (leptoclados-type clubs). The
sub-clade also includes Sinularia robusta M
ACFADYEN
1936 and Sinularia cristata T
IXIER
-D
URIVAULT
1969,
neither of which has been assigned to this group pre-
viously (Group III; Verseveldt 1980). In these two
species, however, some leptoclados-type clubs are
present although they are not well developed. Insuffi-
cient information is available to determine whether
or not all of the species in sub-clade 5C share the
presence of tentacle rods with sub-clade 5B.
The tree shown in Fig. 2 had a significantly greater
log likelihood (ln L52999.22) than an alternative
tree in which the five taxonomic groups of Verseveldt
(1980) were constrained to be monophyletic (ln L5
3528.84) (SH test, po0.001). Among our five major
clades, clade 1 included species placed in Verseveldt’s
Group IV, and clade 3 only consisted of Group III
species, but each of the other three clades comprised
a mix of species from two or more different taxo-
nomic groups (Fig. 2).
Within clades, the mean pairwise genetic distan-
ces (Kimura 2-parameter) between species ranged
0.002–0.029 (Table 2), and some morphospecies
shared the same msh1 sequence, reflecting the typical
lack of variability in anthozoan mitochondrial genes
(Shearer et al. 2002). The mean genetic distances be-
tween clades 2, 3, 4, and 5 ranged 0.028–0.072, com-
parable to the mean distance between the genera
Sarcophyton and Lobophytum at msh1 (McFadden
et al. 2006a). Clade 1, however, differed from the
other four clades by 0.085–0.112, distances that were
slightly greater than those between each of the other
Sinularia clades and the outgroup taxa, Sarcophyton
and Lobophytum (Table 2).
Discussion
The five major clades and sub-clades of Sinularia
distinguished by msh1 can all also be recognized and
separated from one another based on a suite of just
four primary morphological characters: presence of
sclerites in the (a) tentacle, (b) collaret, and (c) point
regions of the polyp, and (d) shape of the club scle-
rites in the colony surface tissues (Table 1, Fig. 1).
Table 2. Average pairwise genetic distances (Kimura 2-parameter) between species within (bold) and among the five
major clades of Sinularia. Clades are defined as in Fig. 2. Values are means (SD).
Outgroup Clade 1 Clade 2 Clade 3 Clade 4 Clade 5
1 0.121 (0.027) 0.005 (0.004)
2 0.113 (0.007) 0.112 (0.003) 0.029 (0.023)
3 0.080 (0.006) 0.085 (0.001) 0.052 (0.002) 0.002 (0.001)
4 0.091 (0.008) 0.100 (0.003) 0.059 (0.005) 0.028 (0.004) 0.015 (0.010)
5 0.105 (0.008) 0.112 (0.003) 0.072 (0.004) 0.042 (0.003) 0.048 (0.005) 0.011 (0.007)
Phylogeny of Sinularia 313
Invertebrate Biology
vol. 128, no. 4, fall 2009
Colony growth form and type of sclerites in the tenta-
cles further distinguish several clades. Identification of
morphological characters diagnostic for each of the
clades and sub-clades allows us to predict the phyloge-
netic placement of a majority of the Sinularia species
that were not included in our molecular phylogeny
(Table 1), thereby providing a hypothetical framework
for further tests of the phylogenetic utility of these
characters. The only species whose phylogenetic place-
ment we were unable to predict because of unclear spe-
cies descriptions and lack of access to type material
were Sinularia andamanensis (T
HOMSON
&S
IMPSON
1909), Sinularia anomala V
ERSEVELDT
&B
ENAYAHU
1983, Sinularia pedunculata T
IXTER
-D
URIVAULT
1945,
Sinularia schleyeri B
ENAYAHU
1993, and Sinularia white-
leggei L
U
¨TTSCHWAGER
1914, and the species described
by L
I
(1982) (Sinularia corpulenta L
I
1982, Sinularia
fibrillosa L
I
1982, Sinularia monstrosa L
I
1982, Sinularia
papillosa L
I
1982, and Sinularia tenella L
I
1982).
There was strong support for the monophyly of
genus Sinularia, provided that Dampia pocilloporae-
formis is regarded as a species of Sinularia.Inhis
original description of Dampia, Alderslade (1983)
discussed the similarities between it, Sinularia foveo-
lata, and Sinularia fungoides, the two other species
with which it groups in clade 3. All three of these
species have calyx-like structures surrounding the
polyps, although they are developed to a much
greater degree in Dampia. Fabricius & Alderslade
(2001) also expressed the opinion that D. pocillopor-
aeformis may be simply an aberrant species of Sinul-
aria; it bears a close resemblance to Sinularia
triangula T
IXIER
-D
URIVAULT
1970, and may in fact
be that species.
Although the phylogeny presented here supports
monophyly of Sinularia, it should be noted that in
some molecular analyses based on different genes
and including a wider range of outgroup taxa, the
species in clade 1 (Sinularia brassica and Sinularia
dura) fall outside of Sinularia (C.S. McFadden, un-
publ. data). Clade 1 was recovered here as sister to all
remaining Sinularia species, but the genetic distances
separating it from the other Sinularia clades are as
great or greater than those among the genera Sinul-
aria, Sarcophyton, and Lobophytum (Table 2). If fu-
ture molecular phylogenetic analyses confirm a
paraphyletic relationship between clade 1 and the
other four Sinularia clades, clade 1 might merit a ge-
neric status. Because S. brassica is the type species of
Sinularia, however, such a revision would make Sin-
ularia a monotypic genus and require that a new ge-
nus be established for the other B150 species.
The only one of Verseveldt’s (1980) five taxonomic
groups that was congruent with the molecular phy-
logeny and formed a monophyletic group was his
Group I (our sub-clade 5C). All of the species in this
sub-clade have the leptoclados-type club sclerites that
distinguish them clearly from other clades. Only Sin-
ularia robusta and Sinularia cristata, in both of which
the leptoclados-type clubs appear to be rare, were not
included in Group I by Verseveldt (1980). Species
belonging to his other four taxonomic groups are
distributed throughout the other nine sub-clades in
the msh1 phylogeny, with some sub-clades (e.g., 4D)
comprising species from three different groups (Fig.
2). In particular, species from Verseveldt’s Group II
(club sclerites with a central wart) and Groups III/IV
(club sclerites without a central wart) often fall into
the same clade, and our re-examination of type ma-
terial suggests that his classification of club sclerites
as having a central wart or not was often incorrect.
For instance, species in clade 3 were classified by
Verseveldt as lacking a central wart and were there-
fore placed in his Group III, but careful examination
of these clubs reveals the presence of a small central
wart that is often obscured (Fig. 3C). In fact, the only
Sinularia species whose clubs truly lack a central wart
are those in clade 1 (S. brassica/dura) and those with
leptoclados-type clubs (5C).
Three of the four characters that we found to be
the most informative phylogenetically for Sinularia
were the presence/absence of sclerites in three distinct
regions of the polyp (tentacles, collaret, and points;
Fig. 1). Characters associated with the polyps, how-
ever, have not traditionally been considered impor-
tant in taxonomic work on this genus. As discussed
previously (McFadden et al. 2006a), the methods
used by Verseveldt (1980) to prepare microscope
slides of sclerites may inadvertently have resulted in
the loss of many of the smallest sclerites, particularly
those from the tentacles. His and many subsequent
descriptions of Sinularia species do not include infor-
mation on tentacle sclerites, as a consequence of
which we are unable to verify their presence/absence
in all members of some of the clades identified here
(Table 1). In addition to omissions, in a number of
cases, our subsequent re-examination of type mate-
rial has revealed errors in original taxonomic descrip-
tions. For instance, Sinularia curvata (sub-clade 4B)
was described recently by Manuputty & van Ofwegen
(2007) as having no sclerites in the polyps, but a care-
ful re-examination of type material revealed some
point sclerites. No polyp sclerites have been described
previously for Sinularia sobolifera (4B) either, but the
specimen identified here (RMNH-Coel. 38748)
clearly showed them. Re-examination of permanent
microscope slides of the type material for Sinularia
cruciata and Sinularia numerosa also showed point
314 McFadden, van Ofwegen, Beckman, Benayahu, & Alderslade
Invertebrate Biology
vol. 128, no. 4, fall 2009
sclerites to be present in those species. In contrast,
Vennam & van Ofwegen (1996) erroneously reported
the presence of polyp sclerites in Sinularia gravis
T
IXIER
-D
URIVAULT
1970, mistaking immature surface
clubs for those typical of the points. Further exam-
ination of type material for species whose reported
morphological character states appear to be incon-
gruent with the molecular phylogeny is likely to re-
veal additional omissions or errors in the primary
taxonomic literature. For example, both Sinularia
mauritiana and Sinularia jasminae A
LDERSLADE
&
S
HIRWAIKER
1991 have been reported to lack polyp
sclerites (Alderslade & Shirwaiker 1991; Vennam &
Parulekar 1994), and yet both species belong to a
clade (5) whose other members have them, a discrep-
ancy that should motivate a future re-evaluation of
the type material.
Assessment of species boundaries
Although msh1 effectively distinguishes the major
clades and sub-clades within Sinularia, it should be
noted that this gene is not variable enough to distin-
guish all species of this genus from one another.
Within each clade, there are numerous examples of
distinct morphospecies that nonetheless share the
same msh1 sequence. The traditional morphological
differences that distinguish some of these genetically
similar species from one another are discussed further
in van Ofwegen (2008b). Analysis of a more rapidly
evolving gene region will be necessary to detect mo-
lecular differences among these closely related Sinul-
aria species. Indeed, preliminary results from the
more variable nuclear ribosomal ITS genes show
clear genetic differences among species such as Sin-
ularia diffusa
VAN
O
FWEGEN
2008a, Sinularia bre-
merensis
VAN
O
FWEGEN
2008a, Sinularia mammifera
M
ALYUTIN
1990, and Sinularia confusa
VAN
O
FWEGEN
2008a, four species in sub-clade 4C that share iden-
tical msh1 sequences (C.S. McFadden, unpubl. data).
Because msh1 sequences often do not differ among
closely related species, cases where individuals iden-
tified to the same species do have quite different msh1
sequences may suggest either cryptic species or mis-
identification of specimens. The most obvious exam-
ples of this in the msh1 tree are Sinularia polydactyla
and S. cruciata, for which specimens identified to the
same species using traditional morphological charac-
ters fell into entirely different phylogenetic clades.
Other examples of genetic differences among speci-
mens of a magnitude that suggests the possibility of
cryptic species can be seen for Sinularia grandilobata
(clade 2), Sinularia variabilis (sub-clade 4A), Sinul-
aria heterospiculata (sub-clade 4B), Sinularia com-
pressa T
IXIER
-D
URIVAULT
1945 (sub-clade 4D), and
Sinularia leptoclados (sub-clade 5C). Indeed, a sub-
sequent taxonomic re-evaluation of S. leptoclados
based on morphological characters has revealed this
taxon to be a complex of several different species
(L.P. van Ofwegen & Y. Benayahu, unpubl. data). In
addition, Sinularia rotundata, Sinularia intacta, and
Sinularia molesta, three species in the leptoclados
group that had been synonymized (van Ofwegen
2001), have msh1 sequences that are sufficiently
different to suggest that they do represent distinct
species, as is also the case for the previously syn-
onymized S. brassica and S. dura (clade 1). Clearly,
much work still needs to be carried out to verify spe-
cies boundaries in Sinularia in cases where a signifi-
cant genetic variation is present within a recognized
taxon, or, conversely, where morphospecies are in-
distinguishable genetically.
Although the morphological characters identified
here (Table 1) distinguish clades rather than species,
consideration of these characters in future taxonomic
studies will also improve our ability to discern species
boundaries (see van Ofwegen 2008b). This point is
best illustrated by the cases we have identified in
which specimens that were assigned to the same spe-
cies on the basis of traditional taxonomic characters
actually belong to very different clades. For example,
by examining the polyp sclerites, it was possible to
distinguish morphologically between genetically dis-
parate specimens of S. polydactyla and S. cruciata,
and in retrospect assign them to the correct clades.
Examination of polyp sclerites a priori (coupled with
recognition of the phylogenetic importance of those
characters) would have prevented these specimens
from ever having been assigned to the same species.
Recent evidence for hybridization among species of
Sinularia may, however, complicate our ability to as-
sess species boundaries and to reconcile morphologi-
cal with molecular data. Slattery et al. (2008)
document the ability of Sinularia maxima and S. poly-
dactyla to hybridize successfully in the laboratory, and
identify a putative naturally occurring hybrid zone be-
tween these species in Guam. Sinularia maxima be-
longs to the distinctive leptoclados-club sub-clade (5C)
while S. polydactyla belongs to clade 4, characterized
by club sclerites with a distinct central wart. Hybrid
offspring contain a mixture of both parental types of
club sclerites (Slattery et al. 2008). This case is partic-
ularly instructive because it demonstrates (1) that suc-
cessful hybridization can occur between Sinularia
species from very different genetic groups and (2)
that hybrid individuals can exhibit morphological
traits diagnostic for more than one clade. ITS poly-
morphisms that are shared among species also suggest
Phylogeny of Sinularia 315
Invertebrate Biology
vol. 128, no. 4, fall 2009
the possibility of ongoing or past hybridization be-
tween some members of sub-clades 4C and 4D (C.S.
McFadden, unpubl. data). Much additional work
needs to be carried out, however, before any conclu-
sions can be drawn about the prevalence or evolution-
ary ramifications of hybridization in Sinularia.
In summary, as has also been shown recently for a
variety of other cnidarian groups (Sa
´nchez et al. 2003;
Fukami et al. 2004, 2008; McFadden et al. 2006b), we
have demonstrated that the morphological characters
traditionally considered most important for inferring
taxonomic relationships within the genus Sinularia (i.e.,
the form and size of club sclerites) are incongruent with
molecular phylogenetic data. However, other charac-
ters whose phylogenetic importance has until now been
overlooked (i.e., polyp sclerites) recover relationships
that are congruent with the phylogeny. Recognition of
the utility of these new characters will necessitate a re-
working of the established taxonomy and taxonomic
protocols in Sinularia, an effort that is already under-
way (van Ofwegen 2008b). In octocorals, the character-
mapping approach used here continues to be useful for
reconciling genus- and family-level taxonomy with mo-
lecular phylogenetics (Sa
´nchez et al. 2003; McFadden et
al. 2006a). In future, we hope to be able to use a similar
method to identify morphological characters that are
congruent with and diagnostic for even higher level (or-
dinal and sub-ordinal) clades within Octocorallia, a
problem that historically has and continues to bedevil
the taxonomy of this group (McFadden et al. 2006b).
Acknowledgments.
We thank Vanessa Brisson for
laboratory assistance, John Starmer for material from
Guam, and the Coral Reef Research Foundation (Pat
and Lori Colin) for specimens collected over the years as
part of the US National Cancer Institute shallow water
marine collections and taxonomy program. Partial support
for this study was provided by the Assembling the
Cnidarian Tree of Life project, NSF grants EF-0531570
to C.S. McFadden and EF-0531779 to Paulyn Cartwright.
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Phylogeny of Sinularia 317
Invertebrate Biology
vol. 128, no. 4, fall 2009
Appendix 1. Specimens of Sinularia and outgroup taxa included in molecular phylogenetic analysis. NTM, Museum and
Art Gallery of the Northern Territory, Darwin, Australia; RMNH, Nationaal Natuurhistorisch Museum, Leiden,
ZMTAU, Zoological Museum, Tel Aviv University; UF, Florida Natural History Museum.
Species Museum Cat. No. Collection location Latitude Longitude Date GenBank no.
S. abhishiktae RMNH Coel. 38720 Palau, Koror,
Mutremdiu
7116.25.2
00
N 134131
0
26.8
00
E 2005 FJ621373
S. abrupta NTM C13799 Palau, Neco Channel 7112.34
0
N 134122.32
0
E 1995 FJ621374
S. acuta RMNH Coel. 38432 Indonesia, Ambon,
near Morela
3133
0
S 128112
0
E 1996 FJ621375
S. acuta RMNH Coel. 38721 Palau, Koror,
Wonder Channel
7110
0
53.3
00
N 134121
0
38.6
00
E 2005 FJ621376
S. babeldaobensis RMNH Coel. 38723 Palau, Babeldaob,
Toagel Mlungi
Channel
7132
0
33
00
N 134128
0
06.6
00
E 2005 FJ621377
S. bisulca RMNH Coel. 38724 Palau, Koror,
Wonder Channel
7110
0
53.3
00
N 134121
0
38.6
00
E 2005 FJ621378
S. brassica NTM C13507 Malaysia, Sabah,
Semporna I.
4139.48
0
N 118146.94
0
E 1999 FJ621379
S. brassica NTM C14185 Papua New Guinea,
Cape Nelson, Tufi
9104.89
0
S 149119.10
0
E 2002 FJ621380
S. brassica NTM C13660 Australia, WA,
Ashmore Reef
12114.29
0
S 123100.77
0
E 2002 FJ621381
S. bremerensis NTM C14488 Australia, Gulf of
Carpentaria,
Bremer I.
12105.66
0
S 136147.75
0
E 2003 FJ621382
S. capitalis NTM C14530 Australia, Gulf of
Carpentaria,
Bremer Is.
12105.66
0
S 136147.75
0
E 2003 FJ621383
S. compacta RMNH Coel. 38433 Indonesia, Ambon,
Latuhalat
03146
0
S 128106
0
E 1996 FJ621384
S. compressa RMNH Coel. 38420 Indonesia, Ambon,
Ambon Bay
03143
0
S 128104
0
E 1996 FJ621385
S. compressa ZMTAU CO 34140 Red Sea, Israel, Eilat,
Nature Reserve
29130.6
0
N34155.35
0
E 2007 FJ621386
S. compressa ZMTAU CO 34142 Red Sea, Israel, Eilat,
Nature Reserve
29130.6
0
N34155.35
0
E 2007 FJ621387
S. compressa ZMTAU CO 34150 Red Sea, Israel, Eilat,
Nature Reserve
29130.6
0
N34155.35
0
E 2007 FJ621388
S. conferta NTM C13972 Mauritius 20117.60
0
S57121.06
0
E 1999 FJ621389
S. confusa NTM C14456 Australia, Gulf of
Carpentaria,
Bremer I.
12105.66
0
S 136147.75
0
E 2003 FJ621390
S. crassa RMNH Coel. 38430 Indonesia, Ambon,
Ambon Bay
03140
0
S 128110
0
E 1996 FJ621391
S. crebra RMNH Coel. 38726 Palau, Koror,
Uchelbeluu Reef
7116
0
25.2
00
N 134131
0
27
00
E 2005 FJ621392
S. cristata NTM C13818 Palau, Ngiwal 07131.43
0
N 134137.66
0
E 1996 FJ621393
S. cruciata NTM C13505 Malaysia, Sabah,
Semporna I.
04138.06
0
N 118142.58
0
E 1999 FJ621394
S. cruciata ZMTAU CO 34152 Red Sea, Israel, Eilat,
Nature Reserve
29130.6
0
N34155.35
0
E 2007 FJ621395
S. curvata RMNH Coel. 38436 Indonesia, Ambon,
W of Sahuru
03140
0
S 128109
0
E 1996 FJ621396
S. densa NTM C1993 Australia, GBR, Qld,
Rib Reef
1980 FJ621397
S. depressa RMNH Coel. 38428 Indonesia, Ambon,
Latuhalat
03146
0
S 128106
0
E 1996 FJ621398
318 McFadden, van Ofwegen, Beckman, Benayahu, & Alderslade
Invertebrate Biology
vol. 128, no. 4, fall 2009
S. diffusa NTM C14457 Australia, Gulf of
Carpentaria,
Bremer I.
12105.66
0
S 136147.75
0
E 2003 FJ621399
S. diffusa NTM C14464 Australia, Gulf of
Carpentaria,
Bremer I.
12105.66
0
S 136147.75
0
E 2003 FJ621400
S. digitata RMNH Coel. 38727 Palau, Babeldaob,
Toagel Mlungi
Channel
7132
0
33
00
N 134128
0
06.6
00
E 2005 FJ621401
S. dura NTM C13808 Philippines, Davao,
N of Talikud I.
7157.72
0
N 125140.94
0
E 1996 FJ621402
S. erecta NTM C12557 Persian Gulf, Iarak I. 26152
0
N56125
0
E 1997 FJ621403
S. erecta ZMTAU CO 34144 Red Sea, Israel, Eilat,
Nature Reserve
29130.6
0
N34155.35
0
E 2007 FJ621404
S. finitima RMNH Coel. 38729 Palau, Koror,
Toachel Mid
7120
0
21.9
00
N 134131
0
03.8
00
E 2005 FJ621405
S. finitima RMNH Coel. 38730 Palau, Koror,
Wonder Channel
7110
0
53.3
00
N 134121
0
38.6
00
E 2005 FJ621406
S. finitima RMNH Coel. 38728 Palau, Babeldaob,
Ngaregabal Reef
7124
0
50.4
00
N 134126
0
38.2
00
E 2005 FJ621407
S. flaccida RMNH Coel. 38731 Palau, Koror, Big
Drop Off
7106
0
48.8
00
N 134115
0
36.4
00
E 2005 FJ621408
S. flexibilis RMNH Coel. 38378 Indonesia, Ambon,
Latuhalat
3146S 128106
0
E 1996 FJ621409
S. foliata RMNH Coel. 38732 Palau, Koror,
Mutremdiu
7116
00
25.2
0
N 134131
0
26.8
00
E 2005 FJ621410
S. foveolata NTM C13966 Mauritius 20114.38
0
S57122.80E 1999 FJ621411
S. fungoides NTM C13910 Palau, Ngerm 1 7133.47
0
N 13410.03E 1998 FJ621412
S. gardineri NTM C14050 Vanuatu, Santo,
Tutuba
15133.07
0
S 167116.65E 2000 FJ621413
S. gardineri ZMTAU CO 34097 Red Sea, Israel, Eilat,
Nature Reserve
29130.6
0
N34155.35
0
E 2007 FJ621414
S. gardineri ZMTAU CO 34146 Red Sea, Israel, Eilat,
Nature Reserve
29130.6
0
N34155.35
0
E 2007 FJ621415
S. gaveshaniae RMNH Coel. 38733 Palau, Koror,
Ngederrak Reef
7117.05
0
N 134129
0
20
00
E 2005 FJ621416
S. gaweli UF3498 Guam, Agat Bay 1998 FJ621417
S. gaweli UF3181 Guam, Double Reef 1998 FJ621418
S. gaweli RMNH Coel. 38734 Palau, Koror, Big
Drop Off
7106
0
48.8
00
N 134115
0
36.4
00
E 2005 FJ621419
S. grandilobata NTM C13796 Palau, Lighthouse
Channel marker 3
1995 FJ621420
S. grandilobata NTM C14114 American Samoa,
Pago Pago Harbor
14106.82
0
S 170140.07
0
W 2001 FJ621421
S. gravis RMNH Coel. 38736 Palau, Peleliu,
Ngermoket
6158
0
36.9
00
N 134113
0
20
00
E 2005 FJ621422
S. halversoni NTM C13848 Fiji, Vanubalavu
Lagoon
17110.27
0
S 179101.30
0
W 1996 FJ621423
S. halversoni NTM C14058 Vanuatu, Santo,
Malo Seamount
15138.10
0
S 167118.72
0
E 2000 FJ621424
S. heterospiculata NTM C13968 Mauritius 20114.38
0
S57122.80
0
E 1999 FJ621425
S. heterospiculata NTM C14003 Vanuatu, Konanda
Reef
17145.17
0
S 168117.28
0
E 2000 FJ621426
S. hirta RMNH Coel. 38399 Indonesia, Ambon,
Manuala Beach
3135
0
S 128105
0
E 1996 FJ621427
Appendix 1. (cont’d).
Species Museum Cat. No. Collection location Latitude Longitude Date GenBank no.
Phylogeny of Sinularia 319
Invertebrate Biology
vol. 128, no. 4, fall 2009
S. hirta ZMTAU CO 34100 Red Sea, Israel, Eilat,
Nature Reserve
29130.6
0
N34155.35
0
E 2007 FJ621428
S. hirta ZMTAU CO 34103 Red Sea, Israel, Eilat,
Nature Reserve
29130.6
0
N34155.35
0
E 2007 FJ621429
S. hirta ZMTAU CO 34148 Red Sea, Israel, Eilat,
Nature Reserve
29130.6
0
N34155.35
0
E 2007 FJ621430
S. humesi UF3500 Guam, Piti Bay 1998 FJ621431
S. humilis RMNH Coel. 38737 Palau, Koror,
Wonder Channel
7110
0
53.3
00
N 134121
0
38.6
00
E 2005 FJ621432
S. intacta NTM C12479 India, Gulf of
Mannar
9116
0
N79112
0
E 1991 FJ621433
S. lamellata NTM C13824 Palau, E Babeldaob,
Ngchesar, RRII
Outlet
07125.07
0
N 134135.98
0
E 1996 FJ621434
S. lamellata NTM C13931 Papua New Guinea,
Milne Bay, Alotau
4
10122.63
0
S 150123.05
0
E 1998 FJ621435
S. leptoclados NTM C5421 Australia, WA,
Broome, Roebuck
Bay
1987 FJ621436
S. leptoclados NTM C14492 Australia, Gulf of
Carpentaria,
Bremer I.
12105.66
0
S 136147.75
0
E 2003 FJ621437
S. leptoclados NTM C14519 Australia, Gulf of
Carpentaria,
Bremer I.
12105.66
0
S 136147.75
0
E 2003 FJ621438
S. leptoclados ZMTAU CO 34095 Red Sea, Israel, Eilat,
Nature Reserve
29130.6
0
N34155.35
0
E 2007 FJ621439
S. linnei NTM C14480 Australia, Gulf of
Carpentaria,
Bremer I.
12105.66
0
S 136147.75
0
E 2003 FJ621440
S. longula RMNH Coel. 38439 Indonesia, Ambon,
Seri Bay
3145
0
S 128109
0
E 1996 FJ621441
S. loyai ZMTAU CO 34154 Red Sea, Israel, Eilat,
Nature Reserve
29130.6
0
N34155.35
0
E 2007 FJ621442
S. luxuriosa RMNH Coel. 38742 Palau, Koror,
Toachel Mid
7120
0
21.9
00
N 134131
0
03.8
00
E 2005 FJ621443
S. mammifera NTM C14198 Vanuatu, Konanda
Reef
17145.17
0
S 168117.28
0
E 2000 FJ621444
S. mauritiana NTM C13852 Fiji, Vanua Levu,
Viani Bay
16145.50
0
S 179155.10
0
E 1996 FJ621445
S. maxima NTM C14001 Vanuatu, Konanda
Reef
17145.17
0
S 168117.28
0
E 2000 FJ621446
S. maxima NTM C14255 India, Gulf of
Mannar
19110
0
N09120
0
E 2001 FJ621447
S. maxima Guam, Piti Bay 1998 DQ302813
S. maxima NTM C14512 Australia, Gulf of
Carpentaria,
Bremer I.
12105.66
0
S 136147.75
0
E 2003 FJ621448
S. molesta RMNH Coel. 38440 Indonesia, Ambon, E
of Cape Nusanive
3148
0
S 128106
0
E 1996 FJ621449
S. muralis NTM C13978 Mauritius 19158.44
0
S57138.89
0
E 1999 FJ621450
S. nanolobata RMNH Coel. 38441 Indonesia, Ambon,
near Morela
3133
0
S 128112
0
E 1996 FJ621451
Appendix 1. (cont’d).
Species Museum Cat. No. Collection location Latitude Longitude Date GenBank no.
320 McFadden, van Ofwegen, Beckman, Benayahu, & Alderslade
Invertebrate Biology
vol. 128, no. 4, fall 2009
S. notanda NTM C14555 Australia, Gulf of
Carpentaria, W.
Woody I.
12111.10
0
S 136140.29
0
E 2003 FJ621452
S. numerosa NTM C13790 Palau, Lighthouse
Channel marker 3
7117.21
0
N 134127.77
0
E 1995 FJ621453
S. ornata NTM C13971 Mauritius 20119.23
0
S57122.01
0
E 1999 FJ621454
S. ornata NTM C14138 American Samoa, W
end Ofu I.
14100.95
0
S 169140.85
0
W 2001 FJ621455
S. papula NTM C14527 Australia, Gulf of
Carpentaria,
Bremer I.
12105.66
0
S 136147.75
0
E 2003 FJ621456
S. parulekari NTM C13989 Mauritius 19156.75
0
S57137.24
0
E 1999 FJ621457
S. parulekari NTM C14249 India, Gulf of
Mannar
19110
0
N9120
0
E 2001 FJ621458
S. pavida RMNH Coel. 38744 Palau, Koror,
Uchelbeluu Reef
7116
0
25.2
00
N 134131
0
27
00
E 2005 FJ621459
S. peculiaris NTM C13959 Palau, Angauri 2 6153.94
0
N 134107.35
0
E 2000 FJ621460
S. peculiaris NTM C14092 Micronesia, Yap,
Ulithi Atoll
9159.73
0
N 139140.05
0
E 2000 FJ621461
S. polydactyla NTM C14142 American Samoa,
Ofu, Ofu Harbor
14100.138
0
S 169140.86
0
W 2001 FJ621462
S. polydactyla NTM C14173 Papua New Guinea,
Normanby I.
9143.93
0
S 150144.77
0
E 2002 FJ621463
S. polydactyla RMNH Coel. 38442 Indonesia, Ambon,
Ambon Bay, E of
Erie
3145
0
S 128108
0
E 1996 FJ621464
S. polydactyla ZMTAU CO 34106 Red Sea, Israel, Eilat,
Nature Reserve
29130.6
0
N34155.35
0
E 2007 FJ621465
S. polydactyla ZMTAU CO 34138 Red Sea, Israel, Eilat,
Nature Reserve
29130.6
0
N34155.35
0
E 2007 FJ621466
S. procera RMNH Coel. 38386 Indonesia, Ambon,
W of Sahuru
3140
0
S 128109
0
E 1996 FJ621467
S. querciformis NTM C14019 Vanuatu, Efate, Paul
Reef
17139.87
0
S 168110.85
0
E 2000 FJ621468
S. querciformis ZMTAU CO 34096 Red Sea, Israel, Eilat,
Nature Reserve
29130.6
0
N34155.35
0
E 2007 FJ621469
S. querciformis ZMTAU CO 34191 Red Sea, Israel, Eilat,
Princess Beach
29129.77
0
N34154.53
0
E 2007 FJ621470
S. rigida NTM C13937 Papua New Guinea,
E Fields 02
10100.66
0
S 145139.90
0
E 1998 FJ621471
S. rigida NTM C14141 Am. Samoa, Olosega
I., SE Asaga Strait
14100.95
0
S 169137.67
0
W 2001 FJ621472
S. robusta NTM C14518 Australia, Gulf of
Carpentaria,
Bremer I.
12105.66
0
S 136147.75
0
E 2003 FJ621473
S. robusta NTM C14526 Australia, Gulf of
Carpentaria,
Bremer I.
12105.66
0
S 136147.75
0
E 2003 FJ621474
S. rotundata NTM C3755 Australia, Cobourg
Peninsula, Orontes
Reef
11104
0
S 132109
0
E 1982 FJ621475
S. scabra NTM C14043 Vanuatu, Santo,
Tutuba
15133.32
0
S 167116.59
0
E 2000 FJ621476
S. siaesensis RMNH Coel. 38746 Palau, Koror, Siaes
Reef
7110
0
56.5
00
N 134121
0
38.7
00
E 2005 FJ621477
Appendix 1. (cont’d).
Species Museum Cat. No. Collection location Latitude Longitude Date GenBank no.
Phylogeny of Sinularia 321
Invertebrate Biology
vol. 128, no. 4, fall 2009
S. siaesensis RMNH Coel. 38747 Palau, Koror, Siaes
Reef
7110
0
56.5
000
N 134121
0
38.7
00
E 2005 FJ621478
S. sobolifera RMNH Coel. 38748 Palau, Koror,
Wonder Channel
7110
0
56.5
00
N 134121
0
38.7
00
E 2005 FJ621479
S. sublimis RMNH Coel. 38750 Palau, Koror, Siaes
Reef
7110
0
56.5
00
N 134121
0
38.7
00
E 2005 FJ621480
S. terspilli ZMTAU CO 34156 Red Sea, Israel, Eilat,
Nature Reserve
29130.6
0
N34155.35
0
E 2007 FJ621481
S. tumulosa RMNH Coel. 38751 Palau, Peleliu,
Ngermoket
6158
0
36.9
00
N 134113
0
20
00
E 2005 FJ621482
S. ultima RMNH Coel. 38752 Palau, Koror,
Uchelbeluu Reef
7116
0
25.2
00
N 134131
0
27
00
E 2005 FJ621483
S. uniformis RMNH Coel. 38753 Palau, Babeldaob,
Toagel Mlungi
Channel
7132
0
33
00
N 134128
0
06.6
00
E 2005 FJ621484
S. variabilis NTM C14134 American Samoa,
Ofu, W of Nuutele
I.
14100.46
0
S 169141.16
0
W 2001 FJ621485
S. variabilis NTM C14164 Papua New Guinea,
Woodlark I.
9112.51
0
S 151155.28
0
E 2002 FJ621486
S. verruca RMNH Coel. 38754 Palau, Koror, Siaes
Reef
7110
0
56.5
00
N 134121
0
38.7
00
E 2005 FJ621487
S. verseveldti NTM C13520 Malaysia, Sabah,
Semporna I.
4136.40
0
N 118146.63
0
E 1999 FJ621488
S. vrijmoethi NTM C14095 Micronesia, Yap,
Ulithi Atoll
10100.79
0
N 139147.33
0
E 2000 FJ621489
S. woodyensis NTM C14557 Australia, Gulf of
Carpentaria, W.
Woody I.
12111.10
0
S 136140.29
0
E 2003 FJ621490
D. pocilloporae-
formis
NTM C5805 Australia, WA,
Rowley Shoals
17107.70
0
S 119120.20
0
E 1987 DQ280593
Sar. ehrenbergi NTM C11208 Indonesia, Central
Java Sea, Jepara
6135.00
0
S 110139.00
0
E 1992 DQ280512
Sar. troche-
liophorum
NTM C14469 Australia, Gulf of
Carpentaria,
Bremer I.
12105.66
0
S 136147.75
0
E 2003 DQ280549
L. compactum NTM C11566 Australia, GBR,
Orpheus I.
18133
0
S 146130
0
E 1992 DQ280559
Appendix 1. (cont’d).
Species Museum Cat. No. Collection location Latitude Longitude Date GenBank no.
322 McFadden, van Ofwegen, Beckman, Benayahu, & Alderslade
Invertebrate Biology
vol. 128, no. 4, fall 2009
Appendix 2. Key to the clades of Sinularia.
Clade 1: Polyps with tentacle scales but without collaret or points; clubs with extreme
wide heads (see Verseveldt 1980: fig. 2D)
Clade 2: Polyps with collaret, points and tentacle rods; clubs with central wart indistinct
because the central wart and the three warts below it are close together
Clade 3: Polyps with indistinct points andno collaret; clubs with small central wart giving
the clubs a triangular form
Clade 4: Polyps without collaret or tentacle sclerites; clubs with central wart
4A: Polyps without sclerites; clubs with central wart indistinct because of leaf-like
processes, or clubs absent
4B: Polyps with point sclerites; clubs with central wart distinct, or clubs absent
4C, 4D: Polyps without sclerites; clubs with central wart distinct
Clade 5: Polyps with collaret and point sclerites
5A: Polyps with tentacle scales; clubs with central wart distinct
5B: Polyps with tentacle rods; clubs with central wart distinct
5C: Leptoclados-type clubs
1. No collaret or point sclerites ..................................................... 2
Collaret and/or points present .................................................... 4
2.Scalesintentacles,extremelywideclubheads ........................................ Clade1
Rods or no sclerites in tentacles................................................... 3
3.Centralwartwithleaf-likeprocesses,orclubsabsent................................... Clade4A
Centralwartdistinct........................................................... Clade4C(4D)
4. Only point sclerites present ...................................................... 5
Collaretandpointspresent...................................................... 6
5. Points indistinct, point sclerites rod-like, clubs triangular................................ Clade3
Points distinct, point sclerites club-like.............................................. Clade4B
6.Centralwartindistinct.......................................................... 7
Centralwartdistinct........................................................... 8
7. Central wart indistinct, clubs not leptoclados-type ..................................... Clade2
Leptoclados-typeclubs.......................................................... Clade5C
8.Scalesintentacles............................................................. Clade5A
Rods in tentacles . . ............................................................ Clade5B
Phylogeny of Sinularia 323
Invertebrate Biology
vol. 128, no. 4, fall 2009
