Table Of ContentPROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
115(2):350-358. 2002.
Contribution to the present status of Spiochaetopterus costarum:
description of Spiochaetopterus koreana, a new species of
Chaetopteridae (Polychaeta) from the West Coast of Korea
M. R. Bhaud, B. S. Koh, and J. S. Hong
—
(MRB, BSK) Observatoire Oceanologique de Banyuls, Universite P. et M. Curie CNRS, BP 44,
66650 Banyuls-sur-mer, Cedex, France, [email protected];
(JSH) Department of Oceanography, Inha University, Incheou 402-751, Korea, [email protected]
—
Abstract. Examination of a collection of Chaetopteridae (Annelida: Poly-
chaeta) from the Yellow Sea (West coast of South Korea) led to the erection
of a new species, Spiochaetopterus koreana. Its main characteristics are the
presence of a ventral shield in two contrasting black and white parts, visible
through the transparent wall of the tube; a pair of black spots at the base of
the A2 setae; Bl and B2 with unilobed neuropodia; the hindmost ventral se-
cretion area extending well beyond the setae of A9; upper oblique plan of the
specialized seta on A4, with two sub-equal lobes. The erection of this new
species confirms, in spite of the probable existence of a long larval life which
is accepted as a means of countering local differentiation, the extensive differ-
entiation within this genus.
Many marine species with wide distri- verse parts of the world's oceans and out-
butions, the so-called cosmopolitan species, side its actual specific area. Consequently,
are sibling species-complexes and are, in recent descriptions of new species of Spi-
fact, more subdivided than originally ochaetopterus in areas as different as Bra-
thought (Hilbish 1996; Knowlton 1993). zilian coasts (Bhaud & Petti 2001), Persian
The broad geographic distribution attribut- Gulf (Bhaud & Martin 2002) and the Yel-
ed to many species is often a taxonomic low Sea (the present paper), are explained.
artifact and needs to be reassessed. This
general statement is particularly applicable Materials and Methods
to the polychaetous annelids (Mackie &
Pleijel, 1995; Williams, 1984), and more Specimens were collected on the west
specifically to the Chaetopteridae, (genus and south coasts of South Korea in three
Spiochaetopterus and species S. costarum). groups of stations: areas A, B and C (Fig.
Reported throughout the world's oceans, S. 1). The type locality is Station 7 (Fig. 1) in
m
costarum was at first believed to be a cos- the Sorae Bight, 14 depth relative to low
mopolitan species (Okuda 1935; Day, 1967; tide (37°20'20 N; 126°36'80 E). The sta-
Blake, 1996). However, Bhaud (1998a & tions of the three areas have similar char-
1998b) showed that different species ofSpi- acteristics; they all lie in sheltered zones
ochaetopterus are found on the western and and the bottom is a silty-sand sediment. The
eastern coasts of the North Atlantic Ocean total number in the collection was 82, dis-
despite the probable ability oflarvae to dis- tributed as follows: area A: 35 specimens
seminate throughout the Atlantic Ocean. sampled in Nov 1998 and Feb 1999, area
Since then, the same author has been very B: 9 specimens sampled in Sep 1999 and
attentive to reports of Spiochaetopterus un- Area C: 38 specimens sampled in Sep 1999.
der the species name of S. costarum in di- Observations were made in several ways.
VOLUME 115, NUMBER 2 351
124° 125 127° 128°
120km
East Sea
(Sea of Japan)
126''35'E 127°14'E 127»22'E
Fig. 1. Map ofthe collection areas. The principal map indicates the location ofthree districts. A, B and C
ofthe Korean coasts, in which specimens were collected. In the lower part, for each district, detail oflocation
of stations is given. In district A, the arrow indicates the type locality.
352 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. A-E, orientation ofthe animal is the same: anteriorparton the right side. A, Anteriorpart (Aregion)
observed with a scanning electron microscope. Because color cannot be shown the different parts identified on
VOLUME 115, NUMBER 2 353
For observations by SEM, setae were dis- ochaetopterus species. Each A4 parapodi-
sected from preserved specimens, im- um has a single modified, enlarged cutting
KOH
mersed in (diluted 1:2) for 15 min, seta at the distal end. This obliquely trun-
rinsed three times in distilled water, 30 min cated distal end is approximately heart-
each, processed through a graded alcohol shaped in end view, with two equally de-
series and stored in 70% ethanol until ob- veloped lobes. The shaft of the seta has a
servation. Immediately prior to viewing in subcircular cross section throughout its
a Hitatchi S.520 SEM (University of Per- length. Region A has 9 setigerous segments
pignan. Centre ofElectron Microscopy), se- {n = 37). At the level of setal bundles 6
tae were transferred to absolute alcohol, air- and 7 of region A, the ventral face exhibits
dried, attached to a stub and coated with a shield with two contrasting parts. The
gold palladium. Fragments oftube were ob- hindmost ventral secretion area extends
served by a Diaplan Leitz light microscope; well beyond the setae of A9. Region B has
photographs were taken with a Leica-Wild more than 2 segments (maximum observed:
MPS 32. 12). The neuropodia on the first two seg-
Following the suggestion by Crossland ments have only one lobe. The tube is un-
(1904), the anterior, middle and posterior branched, smooth, transparent, without dis-
body regions of chaetopterides are herein tinct articulations by with areas regularly
designated A, B, and C, respectively; A4 disposed with folded tube wall. Tube di-
thus refers to setigerous segment4 inregion ameter: 1200 |—JLm.
A, B2 to setigerous segment 2 in region B, Description. Body size and colour: The
mm
etc. anterior part reached 6 in length. All
the A segments plus 10 B segments extend-
Systematics ed to 17 mm. The general color on pre-
served specimens was pink for the anterior
Family Chaetopteridae
region, and creamy white for the middle
Spiochaetopterus Sars, 1856
part. Fragments of the posterior region,
Spiochaetopterus koreana, new species
were partly greenish brown. The ventral
—
Type material. Holotype: National Mu- part carried a contrasting black and white
suem of Natural History, Paris N° MNHN- shield.
POLY 70. Paratypes: National Musuem of Region A: On all examined specimens (n
Natural History, Paris N° MNHN- POLY = 37), 9 setigerous segments were present
71a and 71b. Other specimens were depos- in region A. The horseshoe-shaped peris-
ited in the personal collections of the au- tomium allowed sufficient room for the
thors. Photographs ofspecimens fixed in al- blunt prostomium whose front edge was al-
cohol were ke—pt by the authors. most flat. There were two black ocular
Diagnosis. This is a large, oculate Spi- spots anterior to the tentacles on the lateral
the ventral face have been delimited by a thin line. As for the whole family, the letters A, B and C indicatethe
three regions of the body. B. Anterior part (A region) photographed with a dissecting microscope in order to
observe the contrasting ventral shields. C. Structure ofneuropodia (N) in B1 and B2 segments formed withone
lobe. D. Structure of all notopodia formed with three lobes (1-3); structure of neuropodia in B3 segment and
following segments formed with two lobes Nl and N2. E. Structure ofnoto and neuropodia in a segment ofC
region. F, G, Specialized seta ofA4 segment. H, I. Uncini observed in B1 segment. J, K. Structure ofthe tube
with an articulation, a transversal septum applied on the tube wall and eggs (J), with longitudinal serration (K).
Abbreviations: Ar, Aregion; Br, B region; Al toA9, setigeroussegments 1 to9ofAregion; BC: blackcrescent;
WCl to WC4, white crescents 1 to 4. Regularly arranged secretory pores occupy the first partWCl. Scalebars:
A, B = 860 |xm; C-F = 100 jjim; G = 50 jj-m; H, I = 4 |xm; J, K = 600 \im.
354 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
edge of the prostomium. In all the exam- part and a large unilobed outer part (total:
ined specimens, a long shield in two con- three lobes; Fig. 2D). The insertion point of
trasting colors characterized the ventral sur- this lobe was far from the internal lobes but
face: black for the anterior part and white close to the main neuropodial lobe. The
for the posterior one (Fig. 2A, B). The me- base of these notopodia was preceded by a
dio-ventral part of the short black crescent- pronounced, raised transverse ring on the
shaped fore-shield (BC or black crescenton first B segments, diminishing on later seg-
Fig. 2A, B) extended between the bundles ments. Neuropodia had only one lobe on
on A6 and A7 (Fig. 2B). The two points of segments 1 and 2 (Fig. 2C), and 2 lobes,
the crescent extended posterior to the A7 always vertically extended, on the other
bundles. The crescent was thus oblique. segments (Fig. 2D). As in other species
&
Alongside the ventral posterior border of (Bhaud Petti 2001) the uncial plates of
the black shield, the white shield, on setig- the neuropodial lobes were inserted in such
erous segments 8 and 9, reached well be- a way that their respective sets of teeth
yond the A9 setal bundle. This white shield faced each other, pointing posteriorly on the
was heterogeneous: a narrow crescent just anterior lobe (progressive position) and an-
after the A7 setae (WCl or white crescent teriorly on the posterior lobe (retrogressive
1) and easily identified by the presence of position). Two brick-red lines extended the
regularly arranged oval pores, a homoge- length of the ventral face of region A and
neous white area ending at the middle of then disappeared around Bl or B2. A cili-
the A8 setae (WC2) and a white area punc- ated groove, with a pink band on each side,
tuated by brown spots on the posterior part extended the length of the dorsal face.
(WC3 and WC4) with a marked constric- Region C was incomplete (maximum of
tion at the level ofthe A9 setal bundle. The eight segments on specimens with A+B re-
region anterior to the black crescent was gions); some fragments of C region ob-
brick coloured; it was delimited at its for- served with pygidium. The structures of
ward end by a transversecrescent atthelev- noto- and neuropodia were typical for the
el ofsegment 1 and two longitudinal curved genus (Fig. 2E): notopodia were cylindrical,
lines from the Al setae to the A7 setae not inflated or slightly inflated at the distal
where they come into oblique contact with part, without or with one seta projecting
both black and white crescents. These lines outside the terminal lobe. These short no-
were deepest around A4. Ventral to them topodia were inserted at the bottom of the
the tegument was brown to dark pink and intersegmental constrictions. Neuropodia
dorsal to them it was pale pink. On several were composed of two fleshy lobes with
specimens, the color ofthe ventral face, and uncini: a short upper, anteriorly oriented
ventrally to the longitudinal lines was not lobe, and a vertically extended, lower, pos-
homogeneous: there was a wide, transverse teriorly oriented lobe. These lobes were es-
dark pink belt at the level of A2 and A3. A corted with prominent tegumentary crests.
black spot immediately ventral to each A2 Pygidium without special characteristics; a
seta constituted the last specific criterion. dorsal notch separated two lateral lobes.
On some preserved specimens these two The morphology of the A4 specialized
black spots were linked together by a thin seta was specific. Setigerous segment A4
transverse black ventral line. carried a single modified seta on each ra-
Region B: It had a maximum of 12 se- mus, straw-colored to yellowish, with a
tigerous segments {n = 14), with uniramous swollen, obliquely truncated distal end. The
neuropodia on B1 and B2, and biramous upper oblique plane was cordate with the
neuropodia on the remaining setigerous horizontal ventral edge hollowed out. The
segments. The notopodia, as in many other two lobes of the upper oblique plane were
species, carried a bilobed Y-shaped inner equal or subequal. This weak assymmetry
VOLUME 115, NUMBER 2 355
of lobes was only observed when the seta to 1997 with a simultaneous increase in the
is viewed face on. The overhang ofthe ven- number of tubes per m^. The proportion of
tral edge was clearly marked. The cross sec- sand, silt and clay varied over the same pe-
%
tion of the shaft was regularly sub-circular riod from 64, 33 and 3 respectively (in
%
with no longitudinal furrow (Fig. 2F, G) on 1996) to 34, 58 and 8 respectively (in
the dorsal side as well as the ventral side. 1997). In the same time, the tube density
Uncini were particularly small (length: (N/m2) increased from 3 to 74 (Lee 1999).
20 |JLm), triangular in shape; the small teeth Proportion of silt and number of individu-
were clearly individualized at 100 magni- als, depended on the current flow rate
fication. Nine teeth were counted on the which, if low, increased particle deposition
straight median section of 10 jjim (Fig. 2H, and decreased larval dispersion. During the
I). period 1996-1997 the dynamics of both
The tube unbranched, smooth and trans- currents and sediments were altered by the
parent, without distinct articulations building of several jetties and the dredging
reached a diameter of 1200|xm. The trans- of a large coastal area, thus explaining the
parent wall had regular folded areas, not variation in the number of specimens.
immediately visible (Fig. 2J). A comparison Variation in space was also observed. In
with other described species (Bhaud 1998a) this collection the tube structure varied with
indicated that the number oflayers secreted the district where the specimen was found.
by the ventral shield of area A was limited. In Cll (district C, station 11 in Fig.l) the
The difference was that in the present case, tubes were all straw-yellow but much dark-
this thin wall was observed all along the er than in A10 (district A, station 10 in
collected tubes (8 cm), and not only on the Fig.l). This difference correlated with the
2-3 cmprotruding outofthe sediment. This number of layers forming the tube wall,
tube was probably the upper part of a lon- was probably linked to the very high sedi-
ger tube because we never observed the mentation in station A10 correlated with the
whole body. Septa were present and had a proximity of the Yellow Sea and of the es-
A
single perforation. transverse serration at tuary of the near river. This sedimentation
the tube opening, as described in Bhaud et forced the animals to construct their tubes
al. (1994) and Bhaud (1998a), and other continually as they were covered over. In
transverse serrations further posteriorly on station C11, a zone characterized by a low-
the tube, representing previous tube open- er sedimentation, tube extension could be
ings, were also observed. A longitudinal slower which allowed the animal to con-
rupture, which extends lengthwise, follow- struct a tube consisting of a higher number
ing approximately the same meridian, was of mucopolysaccharide layers.
also present (Fig. 2K). It showed one or Sea surface temperature varied from
several series of parallel indentations, -0.5°C in February to 4-28.5°C in August
which always occurred in pairs because (Koh et al 1997, Park & Park 2000).
they were the result of the rupture of some For conditions of reproduction, some
or all of the wall layers, related to body specimens displayed mature segments in re-
growth in the—width of the tube. gion B. In addition, eggs were observed in
Etymology. This species name is taken one tube. The spherical shape reached a di-
from the country name Koreaplus theLatin ameter of 76—80 |JLm.
suffix -anus—(=belonging to)
Ecology. In the collection area, the bot- Discussion
tom is a silty-sand sediment. However, the
proportion of silt and sand was variable in This new species has already been ob-
time. At station 10 ofdistrict A, an increase served in the studied area under different
&
in the silt fraction was observed from 1996 designations: Spiochaetopterus sp (Hong
356 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Lee 1983), Telepsavus costarum (Paik specific area. This latter hypothesis is prob-
1989) Spiochaetopterus costarum (Lee ably correct (Bhaud 2000). The putative
1999; Lim & Hong 1997; Lim et al 1992). cosmopolitism ofS. costarum was theresult
In the nearby Japanese area, before the of both inadequate morphological exami-
work ofNishi et al (1999) which began the nation and the use of questionable ecolog-
revision of Chaetopteridae in the area, only ical arguments linked to larval dispersal.
2 Spiochaetopterus were known at the sub- Thus, larval dispersal was often usedtojus-
species level: S. costarum okudai and S. tify a wide geographic distribution for a
costarum costarum reported by Nishi species. It was also said that the gene flux
&
(1996) and Nishi Arai (1996). These sub- inside the area counteracted any differenti-
species have now been promoted to the lev- ation that might have arisen. However, Vrba
el ofspecies: S. okudai and S. okinawaensis (1983) argued that genetic mixing does not
and they have been joined by the new spe- contribute as much to the integrity ofa spe-
cies S. sesokoensis (Nishi & Bhaud 2000). cies as selection in a particular environ-
These Japanese species are different from ment. Selection is a firstrate cohesive force.
the one described in the present note. The Mobility acts by reducing a population's
two equal or subequal lobes of the upper chances of isolating itself, but it also acts
oblique plane of the specialized seta differ- by augmenting the probability of encoun-
entiate the Korean species from the two tering a new environment that is not nec-
Japanese species: S. okinawaensis and S. essarily favorable. In this case, the more the
sesokoensis Nishi & Bhaud 2000. In addi- individuals are mobile, the greater is the
tion, this new species is differentiated from likelihood that they will experience a new
S. sagamiensis and S. okudai by the clear selection factor, and speciation becomes
division of the upper oblique plane of the more probable.
specialized seta into two lobes. A provi-
sional summary for the Pacific Ocean in- Acknowledgements
cludes the following species: S. pottsi (E.
Berkeley 1927), S. monroi (Monro 1933), We wish to thank Dr R. H. Jung, (Na-
S. okudai (Okudai 1935); S. okinawaensis tional Fisheries Research and Development
Nishi & Bhaud 2000, S. sesokoensis Nishi Institute), who gave us a large number of
& Bhaud, 2000, S. sagamiensis Nishi et al, specimens of the Korean Spiochaetopterus.
1999. A total of6 species with, in addition, We also wish to thank Dr D. Martin, Centre
the one described in the present paper. In d'Estudis Avancats de Blanes (C.S.I.C) co-
the Atlantic Ocean, five species were re- ordinator ofthe INTAS 97-0916 project en-
A
defined (Bhaud 1998a): S. typicus Sars titled 'The Sibling Species Problem:
1856, S. bergensis Gitay 1969, S. costarum Morphological, Ecological, and Genetic
(Claparede 1870), S. oculatus Webster Approach Using Polychaete Worms' in
1879, and S. solitarius (Rioja 1917) and a which the present work was developed.
new species recently described S. nonatoi
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—
VOLUME 115, NUMBER 2 357
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