~oologicalJoumalofthe Linnean Socie[y (1999), 126: 355-364. With 4 figures
Article ID: zjls 1998.0178, available online at http://ulYw.idealihrary.com on
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Redescription of Spirobranchus gaymardi
(Quatrefages, 1866) (Polychaeta: Serpulidae) from
the Indo-Pacific with remarks on the
Spirobranchus giganteus complex
DIETER FIEGE*
Forschungsinstitut Senckenbq, Senckenbergzmlage 25, 0-60325 Frankfurt, Germany
HARRY A. TEN HOVE
Instituut uoor Systematiek en Populatiebiologie/<oologisch Museum, Uniuersiteit van Amsterdam,
Mauritskade 61, NL-1090 G T Amsterdam, T h e Netherlands
Received October 1997; accepted for publication June 1998
As a contribution to unravel the confusion of the Spimbranchus comiculatus complex a detailed
redescription is given of S.gaymardi, together with an overview of the S.&anteus complex.
The operculum of S.gaymardi in its most typical form is clearly distinguished from that of
any other Spirobranchus species by the presence of two short, broadened, and abraded dorsal
tines which usually fuse or touch above the midline of the operculum.
0 1999 The Linnean Society of London
ADDITIONAL KEY WORDS: - SPirobranchus gtganteus complex
-
systematics - taxonomy.
CONTENTS
Introduction . . . . . . . . . . . . . . . . . . . . . . .
Systematics . . . . . . . . . . . . . . . . . . . . . . .
Spimbranchus gaymardi (Quatrefages, 1866) . . . . . . . . . . . .
Acknowledgements
. . . . . . . . . . . . . . . . . . . .
References . . . . . . . . . . . . . . . . . . . . . . .
356
356
356
362
363
INTRODUC’IION
The serpulid genus Spirobranchus Blainville, 1818 comprises c. 20 species mainly
occurring in the subtropics and tropics, many of them living in close association
* Corresponding author. E-mail: [email protected]
0024-4082/99/070355
+ 10 $30.00/0
355
0 1999 The Linnean Society of London
356
D. FIEGE AND H. A. TEN HOVE
with hermatypic corals. Their major identification character is the shape of the
operculum. This character, however, is often quite variable and species identification
is difficult, especially as older descriptions are not sufficiently detailed.
Spirobranchusgaymardi was described in 1866 by Quatrefages and originally placed
in the genus Cymospira. Since that time this species has never been reported again
although specimens identified by Pruvot as Pomatoceros gaymardi, but synonymized by
Fauvel with Spirobranchusgiganteus (in Pruvot & Fauvel, 1930), probably represent S.
gaymardi. Pomatoceropsis roxasi Holly, 1935 (later renamed Pseudopomatoceros roxasi Holly,
1936) of which the syntypes recently have been refound in the collections of the
Naturhistorisches Museum W e n (NHMW) proved to be synonymous with S. gaymardi.
In his revision of the genus Spirobranchus ten Hove (1970) listed C. gaymardi among
other species as a synonym of the Indo-Pacific subspecies Spirobranchus giganteus
corniculatus. More recently Smith (1985) reported a Spirobranchus sp. (‘type B’) from
the Great Barrier Reef which fits very well with the description of C. gaymardi. In
subsequent discussions Smith and ten Hove agreed on the need to evaluate the
validity of the names used for the Spirobranchus gganteus complex as given in the
synonymy of ten Hove (1970). Forms like those depicted by Quatrefages (1866: pl.
20, fig. 13) had been collected from various areas in the Indo-Pacific. Based on this
material the taxon gaymardi is redescribed and placed in the genus Spirobranchus. The
re-establishing of this long forgotten species as a valid taxon is part of an attempt
to unravel the confusion of the S. corniculatus complex (see ten Hove, 1994) at least
for those forms where a distinction based on morphological characters is possible
(Fiege & ten Hove, 1997).
SYSTEMATICS
Spirobranchusgaymardi (Quatrefages, 1866)
(Figs 1-3)
Cymospira Gaymardi Quatrefages, 1866: 539-540, pl. 20, fig. 13.
Spirobranchus giganteus: Pruvot & Fauvel, 1930: 88-89.
Pomatoceropsis roxasi Holly, 1935: 99-100, figs 1-6.
Pseudopomatoceros roxasi Holly, 1936: 223 [new genus name only].
Spirobranchusgiganteus corniculatus: ten Hove, 1970: 24-32, 50-5 1 figs 63-73, partim.
Pseudopomatoceros roxasi: Fauchald, 1977: 146.
Pseudopomatoceros roxasi: Uchida, 1978: 72-73.
Spirobranchus gaymardi: Fossii & Nilsen, 1996: 140, 141 [name only].
Spirobranchus corniculatus: Nishi, 1996: 3 14-3 15, figs 1b-i, k [probably not j] .
Material examined
?Expedition ASTROLABE 1826- 1829, coll. Quoy & Gaymard (1 spm., MNHN
356a).
JAPAN: Okinawa Island, Zampa Cape, subtidal reefs, leg. E. Nishi, 17.2.1994 (4
spms, ZMA V.Pol. 402 1).
South China Sea, Hainan Island: Yezhu Island, off Hainan Island, 24.1 1.1990,
6-10m (1 spm., SMF 5339; SEM stubs 237,238). Yezhu Island south side, 22.3.1992,
357
Figure 1. Spirobranchus gymardi. A, Lizard Island, larger specimen (SMF 6087). Thoracic region,
dorsal side. B, Tukang Besi, smallest specimen (SMF 85 13). Thoracic region, dorsal side. Operculum
partly overgrown. C, Hainan Island (SMF 5336). Tube. Scale bars A-C = 3mm.
10m, leg. D. Fiege. Occurring together with specimens of Spirobranchus gardineri
(Pixell, 1913) in the same coral block (3 spms, SMF 5336).
PHILIPPINES: Mindoro Island, Puerto Galero (types of P roxasi, 2 spms, 1
without operculum, NHMW A 118).
INDONESIA: Indonesian-Dutch Snellius I1 Expedition, leg. H.A. ten Hove: Sta.
4.003, Ambon, Ambon Bay, near Seilale, 3"46'S, 128"7'E, reefflat to 1 m deep,
358
D. FIECE AND H. A. TEN HOVE
Figure 2. Skimbranchus gymardi; Hainan Island (SMF 5339; SEM stubs 237, 238) A, B, operculum.
C, collar chaetae. D, thoracic uncini. E, abdominal uncini. Scale bars: A = 1000pm; B = 300 pm; C =
30pm; D, E = 10pm.
REDESCRIPTION OF SPIROBMVCHUS GAYMARDI
359
u
A
B
C
G
Figure 3. Spimbrunchus ggyrnardi. A, Lizard Island, larger specimen (SMF 6087). Operculum, dorsal
side. B, Tukang Besi, larger specimen (SMF 8513). Operculum, dorsal side. C, same from lateral.
D-F, antler-like spines of opercula, seen from top. D, E, Ambon, 2 specimens (ZMA V.POL. 4023).
F, Okinawa (ZMA V.POL. 4021). G, ?Astrolabe specimen (MNHN 356A). Operculum, seen from
top. Scale bar A-G = 3 mm.
23.8.1984 (6 spms, from coral Porites ?lobata, together with S. corniculatus s.s., S. ?
gardineri and S. paumotanus, ZMA V.Po1.4023, 4024-4026); Sta. 4.030 A, Tukang
Besi Islands, Binongko, 5"55'S, 123"59'E, gently sloping reef, 3-6m (15 spms,
360
D. FIECE AND H. A. TEN HOVE
National Natural History Museum, Leiden (NNM) 18370, SMF 8513 incl. SEM
stubs 438, 439); Sta. 4.096, Komodo Island, NE cape, 8"29'S, 119"34.1'E, edge of
narrow coastal reef sloping down to sandy bottom at 30m, reef patches in sand at
3m, 19./20.9.1984 (1 spm., together with S. corniculatus s.s., SMF 5759); Sta. 4.120,
N of Sumbawa, Bay of Sanggar, 8"20.5'S, 1 18' 15.7'E, nearly horizontal reef, 1-3 m,
23.9.1984 (3 spms from Porites, ZMA V.Pol. 4020); Sta. 4.145A, NE Taka Bone
Rate (Tiger Isld.), NE of Tarupa Besar, 6"27'S, 121"8'E, sloping reef, 6-8m,
26.9.1984 (3 spms from Porites Lichen, with S. corniculatus s.s., NNM 18371).
INDIA: Laccadive Islands, Kavaratti Is., 5.1.1980, leg. A. Misra; Agatti Is. 4.4.84,
leg. A. Misra (3 spms, Zool. Survey of India, Reg. No. 1326/ 1).
AUSTRALIA: Dampier Archipelago, Kendrew Island, 20"28.7'S, 1 16'30'E. In
Porites, reef front, 20 ft. deep, leg. Crown of Thorns Survey, 14.5.1973 (5 spms,
ZMA V.Pol. 3574; exchange W.A. Museum. no. 223-75); Queensland, Great Barrier
Reef, Carter Reef, 10-15m, leg. P. Hutchings, 10.3.1986 (1 spm., AM W24045);
Lizard Island, Palfrey Island, S of light house, coral heads on sandy bottom, 7m,
leg. H.A. ten Hove et al., Stn.17, 2.3.1986 (3 spms AM W24046, 2 spms SMF 6087);
Lizard Island, S South Island, sloping silty reef, little coral cover, 18-20m, leg. H.A.
ten Hove et al., Stn. 21, 6.3.1986 (5 spms, ZMA V.Pol. 4027).
Description
Branchial crown: two lobes in spiral (up to 8 whorls, mean 6, n= 16). Radioles
connected by interradiolar membrane for c. f to of their length (Fig. 1A).Membrane
with smooth edge between radioles, no lappets or frills. Radioles with short terminal
filament. In most populations white pigment (remnants of ocellar clusters?) observed
at either side of the base of the terminal filament (e.g. missing in populations from
Komodo and Ambon). Length of crown up to 13mm, proportionally shorter in
larger specimens. Pinnules almost same length along radiole, only those at tip a
little shorter. Radioles square-shaped in cross-section, outer side flat (Fig. 1A, B).
Stylodes absent. Kidney-shaped compound eye present at base of dorsalmost radiole
on each side, as figured by Smith (1984, fig. 10: be, branchial eyes; 1985, figs 44B,
C, 45).
Mouthparts: two smooth lips; left side of dorsal lip with bean-shaped membrane
folded backwards, right side possibly with tiny mouthpalp (difficult to observe, found
in three specimens only).
Peduncle: smooth dorsoventrally flattened, triangular in cross-section. Inserted just
left of median line. Broadly winged at least for distal f to of entire length. Wings
with smooth edges, ending in broadly rounded 'tips', proximally not set off by a
notch (Fig. IA, B).
Operculum: with inverted fleshy cone, distally topped by oval concave calcareous
plate. Length of plate 4.8-7.1-1 1.0mm (SD= 1.5, n= 16), width 44.7-7.3mm
(SD = 0.9, n = 16). Two or three antler-like spines originating from a short common
stem. Two evenly curved dorso-lateral spines, round in cross-section, with 2-3
secondary spinules terminating in pointed tips. Spines may raise at an angle of
almost 60" but there is a tendency towards levelling off in plane parallel to opercular
plate (Figs lA, B, 2A, B [calcareous plate broken], 3A-C). Each spine with short,
flattened dorsal tine which broadens distally; left and right tine most commonly fuse
or touch above midline of operculum. Usually extremities of these dorsal tines blunt,
abraded (Quatrefages, 1866: 540 'brusquement tronquk') probably as a result of
REDESCRIPTION OF SPIROBRRNCHUS GAYMARDI
36 1
passage against walls of tube (Fig. 2A, B). However, unworn tines may terminate in
three or more short acute spinules (Fig. 3D-F).
Development of the medio-ventral spine more variable. It may be completely
absent (n= 10) (Figs lA, 3A, F) or present only as slight protuberance (n= 12) but
it may also be several millimeters in length and bifurcate at its tip ( n = 5 ) (Figs 2A,
B, 3D, E, G) (see ten Hove, 1970: figs 35-45 for similar situation in Spirobranchus
gzganteus s.s.). Opercular plate usually pink, especially along the edge, sometimes
extending to the spines, otherwise white (decolorized by preservative?); often overgrown (Fig. 1B).
Collar and thoracic membranes: collar one fifth of length of radioles, but up to
half the length in small specimens; divided into one big triangular ventral lobe
corresponding to spine over entrance of tube, and two smaller lateral lobes (Fig.
1B) continuing into thoracic membranes forming short ventral apron. Tonguelets
present between lateral and ventral lobes, as usual in the genus. Notch between
ventral and lateral collar lobes extending as groove across ventral side of thorax
marking base of collar; as long as, and running parallel to, first row of uncini.
Thorax: seven thoracic chaetigers, six uncinigerous. Collar chaetae of ‘Spirobranchustype’, i.e. with many small teeth at the base of the ‘blade’. Chaetae made up of
fibrils densely packed on one side, the axis, and more loosely on the other side, the
‘teeth and blade’ of the literature, thus appearing feathery (Fig. 2C). Few chaetae
without teeth at base of blade, smoother except for horizontal incisions at regular
intervals, possibly representing preparation artefact or juvenile character (Fig. 2C:
left). Collar chaetae sometimes deeply retracted into collar membrane. Thoracic
chaetae finely striated (‘limbate’), arranged in bundles with three rows of slightly
different length. Thoracic uncini with 15-16 teeth in a single row, anteriormost
tooth gouge-shaped (Fig. 2D). Triangular depression between tori as in generic
diagnosis (ten Hove & Nishi, 1996).
Abdomen: number of abdominal chaetigers 1 6 7 - 2 m 3 13 (SD = 37, n = 12). Abdominal chaetae compressed trumpet-shaped with long tip, 2-5 in one fascicle.
Abdominal uncini with 1 1-16 teeth in a single row (Fig. 2E).
Tube: very massive, embedded in living corals (mostly Porites) leaving almost
nothing visible except the opening. Dorsal crest projecting into spine pointing over
opening (Fig. 1C). Tubes may be up to 13mm wide, internal diameter 6-7mm,
wall-thickness 2-4mm around opening, 0.5-1 mm inside coral. Interior lining of
tube salmon colored, occasionally tinted blue on substrate-orientated side.
Measurements
Length 28-53-85mm (SD = 16.8, n= 16); width of thorax (3.5)4.3-6.2-7.8mm
(SD= 1.05, n= 16). Specimen with thoracic width of 3.5mm is smaller than half
the maximum size, and not included in statistics.
Distribution
Japan (Okinawa), South China Sea (Hainan Island), Philippines (Mindoro Island),
Indonesia (Sunda Sea, Banda Sea), India (Laccadive Islands), New Caledonia,
Australia (Dampier Archipelago; Great Barrier Reef).
Remarkr
Usually Spirobranchusgaymardican easily be recognized by the characteristic shape of
its operculum, more specifically the arrangement and shape of the two dorsal tines.
362
D. FIEGE AND H. A. TEN HOVE
They are short and broad with blunt, abraded tips and meet above the midline. Abraded
spines and tines are very rare in other members of the S. corniculatus complex. In S.
corniculatus S.S. the opercular spines are less elaborate and the tines, if present, not broad
but narrow. However, in some specimens of S. gaymardi we observed the dorsal tines
not abraded, but with 3-7 pointed tips (Fig. 3E, F).This situation may also be found
in S. spec. ‘type C’ of Smith (1985), but that has the tines thinner and spaced more
widely apart, not touching across midline. The overall appearance of both types of
opercula is very similar. O n the other hand, the spines may be almost completely
abraded in S. gaymardi, thus making identification uncertain. Although the crimson
coloration of the operculum is a fairly constant feature for S. gaymardi we also observed
a few smaller specimens with a white operculum (e.g. Nishi & Asakura, 1996: fig. 3), a
colour more frequently occurring in S. sp. ‘type C’ of Smith (e.g. ten Hove, 1970: figs
68, 69) and S. corniculatus S.S. In conclusion, it is not always possible to attribute single
specimens, especially if without tube, to one of these three species.
As far as we know, the groove marking the base of the collar has not been
described before, nor the flap at the base of the mouth-lips. We observed the groove
also in a number of specimens of S. gardineri, S. giganteus S.S. and S.incrassatus Morch,
1863, all belonging to the S. giganteus s.1. complex, but could not find it in a number
of specimens of S. tetraceros Schmarda, 1861 and S. decoratus (Imajima, 1982) both
belonging to the S. tetraceros complex. The lip-associated membrane was present in
S. gardineri and S. gzganteus S.S. as well. Both characters may be more widely spread
in the S. giganteus complex s.1.
Type material of S. gaymardi appears to be lost. The specimen from the MNHN
(No. 356a) had been labelled ‘holotype’ probably assuming that Quatrefages based
his description on this single individual present in the collection of the MNHN.
However, measurements of this specimen do not conform with those given by
Quatrefages (length c. 60mm for 130-140 segments, width 7mm, 4 branchial
whorls), the present specimen being much smaller with length: 32mm for 174
chaetigers, width: 5.5mm; 3 branchial whorls.
In his revision of the genus Spirobranchus ten Hove (1970) listed Cymospira gaymardi
among other species as a synonym of the Indo-Pacific subspecies Spirobranchusgganteus
corniculatus (Grube, 1862). Since this revision was based on limited material from
Indo-West Pacific areas with many mistaken identifications, ten Hove (1994) changed
his opinion and deemed his ‘subspecies’ of 1970 to be full species complexes. In the
S. corniculatus complex s.s., three species can be distinguished by their opercular
shape, colour of their tubes and possibly by the shape of their branchial spires
(Smith, 1985, ‘types A, B, C’), i.e. S. corniculatus s.s., S. gaymardi and S. spec. ‘type C’
of Smith, probably identical with S. crucigerus (Grube, 1862). These three species form
the Indo-Pacific part of the S. gzganteus complex, which comprises all circumtropical
Spirobranchus species that have spiralling branchiae. With regard to this character S.
niganucha (Fischli, 1900)’and S. paumotanus (Chamberlin, 1919) can also be attributed
to the S. corniculatus complex s.1. as well as S. gardineri (Pixell, 1913) and a species not
yet described (see Fig. 4,which is modified from Fossii & Nilson, 1996: 140).
ACKNOWLEDGEMENTS
We thank the following: H. Zibrowius for information on the existence of a specimen
of S. gaymardi in the collections of the MusCum National d’Histoire Naturelle, Paris
S. spinosus Moore, 1923
S. giganteus complex S.S.
S. spec 'type C of Smith, 1985
S. corniculatus complex
I
r
I
I
S. gurdineri (Pixell, 1913)
G
Indo-Pacific
Figure 4. Graph showing diversification of Spimbranchus giganteus complex (adapted from Fossg & Nilson, 1996).
S. polycerus complex
I
Atlantic and Pacific Coast of
North America
Spirobrunchus giganteus
complex s.1.
W
w
m
k
364
D. FIEGE AND H. A. TEN HOVE
(MNHN); J.-C. Dauvin and Mme. d’Hondt for the loan of this specimen; H.
Sattmann of the Naturhistorisches Museum Wien (NHMW) for unearthing two
‘undetermined’ specimens which turned out to be syntypes of Pomatoceropsis roxasi;
E. Nishi of the Natural History Museum and Institute, Chiba, Japan; A. Misra,
Zoological Survey of India, Calcutta, for the loan of additional material. Ruth
Barnich skilfully made the drawings and critical remarks on the manuscript. Last
and certainly not least we thank Richard Smith. His thesis and discussions between
him and the second author started a re-evaluation of the validity of nominal species
within the S. giganteus complex. The Max Planck Society, the Senckenberg Society
of Natural History and the Institute of Oceanology, Chinese Academy of Sciences,
supported fieldwork on Hainan Island, South China Sea; the Trustees of the
Australian Museum, Sydney, supported fieldwork in Australia. The Netherlands
Marine Science Foundation enabled participation of HAtH in the Indonesian-Dutch
Snellius I1 Expedition.
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