SCIENTIA MARINA
SCI. MAR., 67 (1): 99-105
2003
Spiochaetopterus creoceanae, a new species of
Chaetopteridae (Polychaeta) from the Persian Gulf
belonging to the costarum complex*
MICHEL BHAUD1, DANIEL MARTÍN2,‡ and JOÃO GIL2
1
Observatoire Océanologique de Banyuls, Université P. et M. Curie - CNRS, BP 44,
66650 Banyuls-sur-mer, Cedex, France.
2 Centre d’Estudis Avançats de Blanes (CSIC), Carrer d'accés a la Cala Sant Francesc 14, 17300 Blanes (Girona),
Catalunya, Spain. E-mail: dani@ceab.csic.es
SUMMARY: Examination of a collection of Chaetopteridae (Annelida: Polychaeta) from the Iranian coast of the Persian
Gulf led to the erection of a new species of Spiochaetopterus: S. creoceanae sp. nov. This is a small, oculate Spiochaetopterus with a single specialised, enlarged cutting seta on each A4 parapodium; this seta is roughly heart-shaped (cordate) in end view and with the outer lobe of the oblique section more developed than the inner one. The new species is mainly characterised by the presence of 10 segments in the anterior region, while all other species described to date have only 9.
The description of this new species of Spiochaetopterus is a contribution to the growing evidence supporting the hypothesis that Spiochaetopterus costarum, formerly considered to be a cosmopolitan species, has in fact been used for several distinct species. In addition, the habitat preferences of the new species are also discussed.
Key words: new species, chaetopterid, Annelida, species complex, distribution, habitat preferences.
RESUMEN: SPIOCHAETOPTERUS CREOCEANAE, UNA NUEVA ESPECIE DE CHAETOPTERIDAE (POLYCHAETA) DEL GOLFO PÉRSICO,
PERTENECIENTE AL COMPLEJO COSTARUM. – El estudio de una colección de Chaetopteridae (Annelida: Polychaeta) procedente de la costa iraniana del Golfo Pérsico nos ha llevado a describir una especie nueva de Spiochaetopterus: S. creoceanae
sp. nov. Se trata de una especie pequeña, provista de ojos y con una única seda especializada en cada parápodo A4. dicha
seda, vista apicalmente, es acorazonada y presenta el lóbulo externo de la sección oblicua más desarrollado que el interno.
La característica principal de la nueva especie es, sin embargo, la presencia de 10 segmentos en la región anterior, mientras
que las especies descritas con anterioridad poseen sólo 9. La descripción de esta nueva especie es una contribución adicional al constante incremento de pruebas que evidencian que Spiochaetopterus costarum, anteriormente considerada como una
especie cosmopolita, se ha venido usando para denominar lo que, de hecho, han resultado ser especies distintas. Asimismo,
se discuten las características del hábitat de la nueva especie.
Palabras clave: especie nueva, quetoptérido, Annelida, complejo de especies, distribución, preferencias de hábitat.
INTRODUCTION
The chaetopterid polychaete Spiochaetopterus
costarum (Claparède, 1868) was traditionally con‡
Corresponding author.
*Received November 28, 2001. Accepted October 8, 2002.
sidered a cosmopolitan species (Okuda, 1935; Day,
1967; Blake, 1996). Recently, however, Bhaud
(1998a) showed the existence of similar but clearly
different species on both western and eastern coasts
of the North Atlantic Ocean, where previous studies
had only reported different subspecies of S.
costarum (Gitay, 1969). Accordingly, Bhaud
A NEW SPIOCHAETOPTERUS SPECIES FROM THE PERSIAN GULF 99
(1998a) pointed out the limited geographical distribution of adult populations, in spite of the possible
ability of their planktonic larvae to disseminate
throughout the whole ocean.
Careful observations have been addressed to the
available specimens corresponding to the multiple
worldwide reports of S. costarum. As a consequence, marked morphological differences have
been identified, supporting the existence of distinct
species of Spiochaetopterus in geographical areas as
diverse as Japan (Nishi et al., 1999; Nishi and
Bhaud, 2000), Brazil (Bhaud and Petti, 2001), the
Yellow Sea (Bhaud et al., 2002) and the Persian
Gulf (present paper).
Based on the previous knowledge of the relevant
morphological characters distinguishing the different species within the genus Spiochaetopterus, the
present paper describes a new species, S. creoceanae, from the Iranian coast of the Persian Gulf.
Additional data on the habitat preferences of the
new species at the study site are also provided.
MATERIALS AND METHODS
Samples were taken by the French company
CREOCEAN with a van Veen grab of about 0.1 m2,
as part of an environmental baseline study carried
out off Asaluyeh and Nakhl e Taqi (Iranian shoreline
of the Persian Gulf) during August 1998. Among the
sediment samples collected for this study, 7 contained specimens of the new species (Table 1). The
density of worms was estimated on the basis of a
total sampling area of 0.3 m2 (i.e. three grabs). The
grab contents were mixed in a sufficiently large container, and then sieved out on board by pouring the
contents through a 1 mm mesh sieve. The retained
sediment was then transferred to a plastic bag, fixed
with a 4% formaldehyde/seawater solution, stained
with Rose Bengal and stored until sorted. An initial
sorting was done under a stereomicroscope (Zeiss
Stemi 2000-C) to separate the tubes containing Spiochaetopterus worms, which were then counted to
express their densities as number of individuals per
m2, from the empty tubes.
For the morphological observations, more than
50 worms were carefully extracted by cutting their
tubes under the stereomicroscope. Size measurements were estimated for the 20 most complete
worms. For light microscope observations (LM), the
specimens were placed on slides with a solution of
glycerine and distilled water. LM drawings were
made with a Wild M3Z compound microscope
equipped with a camera lucida. LM micrographs of
uncini were made by dissecting a fragment from a
parapodium and squashing it directly between the
slides. The LM micrograph of a region-C parapodium was obtained after simple dissection. LM micrographs of a tube were made on empty fragments.
LM micrographs were made with a Leica-Wild MPS
32 through a Diaplan Leitz LM or through a Wild
M3Z compound microscope.
For SEM observations of the specialized,
enlarged cutting setae (referred to as “the specialized setae”), an A4 parapodium was dissected from
the body and isolated in a glass dish containing a
nearly saturated solution of potassium peroxide.
Once transparent (about 12 hours later), the parapodia were rinsed with distilled water, dissected and
the specialised seta was removed by slight pressure
with fine forceps under a compound microscope.
The isolated setae were then rinsed three times in
distilled water (30 min each), run through a series of
ethanol concentrations, and stored in 90% ethanol
until observation. Immediately prior to viewing in a
Hitachi S.520 SEM (University of Perpignan, Centre of Electron Microscopy), they were transferred to
100% alcohol, critical-point dried, attached to a
stub, and coated.
The morphological description of the new
species is structured in three different sections corresponding to the three body regions that charac-
TABLE 1. – Spiochaetopterus creoceanae sp. nov. Geographical location of the sampling stations and main environmental parameters of the
inhabited sediments. The asterisk indicates the Type Locality for the new species.
Station
8
9
13
14
18*
19
24
North
Coordinates
East
Depth
(m)
Gravel
(%)
Silt
(%)
52°33.574’
52°33.323’
52°33.831’
52°33.591’
52°33.244’
52°34.003’
52°34.168’
27°30.229’
27°30.064’
27°29.948’
27°29.677’
27°29.406’
27°29.156’
27°28.797’
19
21
15
19
16
19
18
45.1
41.6
45.9
52
46.2
34.9
22.4
7.84
7.17
5.96
4.41
3.9
5.81
6.85
100 M. BHAUD et al.
Organic Matter Pore water Density
(%)
(%)
(ind. m-2)
3.4
4
3.7
3.6
3.2
3.3
2.7
24.5
23.6
20.8
26.3
19.7
26.3
30.4
350
3
520
427
2693
423
3
terise the genus Spiochaetopterus: Anterior (regionA), mid-body (region-B) and posterior region
(region-C) following Bhaud et al. (1994). Thus, for
instance, A4 = setiger 4 in region-A and 10A = 10
setigers in region-A. As all specimens were incomplete, the potential length for the maximum number
of setigers at region-C was estimated on the basis of
the only 4 specimens having measurable segments.
Potential length for region C is then given as an
average.
RESULTS
Systematic account
Family CHAETOPTERIDAE
Spiochaetopterus Sars, 1853
Spiochaetopterus creoceanae sp. nov.
(Figs. 1, 2)
Material examined. A total of 1325 specimens were collected by E.
Dutrieux off Asaluyeh and Nakhl e Taqi, north Nay Band Bay, 250
km south of Bandar Bousher, Iranian shoreline along the east coast
of the Persian Gulf (Table 1). Among the 7 stations containing the
new species, the most densely populated one, station 18, is designated the type locality.
Type material. Holotype: Museo Nacional de Historia Natural,
Madrid, number MNCN 16.01/9023a; paratypes: Museo Nacional
de Historia Natural, Madrid, number MNCN 16.01./9023b. The
remaining specimens are in the authors’ collections.
Etymology. The specific name refers to CREOCEAN (France), the
company responsible for the environmental survey, which collected
the samples with the new species.
Species diagnosis. A large, oculate Spiochaetopterus with 10A (invariate) + up to 27B + at
least 43C (all specimens incomplete), measuring up
to 28 mm long and 0.76 mm wide for over 79 segments. A4 specialised seta enlarged asymmetrically
(cordate), heart-shaped in distal view; shaft with two
grooves, a ventral one all along its length and another on dorsal face, just below the more developed
lobe. Neuropodia uniramous on B1 and biramous on
all remaining setigers. Tube unbranched, smooth,
transparent, with distinct articulations, 750-800 µm
in diameter.
Description:
Region-A (Fig. 1A). All examined specimens
with 10A (n = 53). Region-A measuring 0.69 ±
0.063 mm wide and 3.89 ± 0.41 mm long (n = 20).
Peristomium horseshoe, surrounding the prostomi-
FIG. 1. – Spiochaetopterus creoceanae sp. nov. A, Anterior end
drawn from SEM picture (the A4 specialized setae may be seen in
apical view), including a frontal view of pr (prostomium with the
two eyespots) and pe (peristomium) and a dorsal view of t (tentacles); B, Notopodia of region-C; C-D, Specialized A4 seta in ventral (C) and dorsal (D) views; E-F, Uncinal plates of segment B1.
Scale bars: A = 1 mm; B = 30 µm; C, D = 50 µm; E, F = 8.5 µm.
um ventrally and laterally. Prostomium blunt, relatively broad frontally. Both prostomium and peristomium finish at same level. Two well-separated
black ocular spots are located facing the tentacles,
on the lateral edge of the prostomium. Ventral
shield long, divided into several sections, each with
a different colour on preserved specimens. Glandular tissues occasionally pink to reddish due to the
use of Rose Bengal during the sample fixation
process. The ventral surfaces of both peristomium
and A1 have a whitish glandular tissue; a white,
non-glandular circle is present at the level of A1
A NEW SPIOCHAETOPTERUS SPECIES FROM THE PERSIAN GULF 101
and A2; from A2 to A7, the surface is whitish and
glandular again. These glandular areas can only be
distinguished from non-glandular ones in stained
worms; otherwise, all of them look whitish. At the
level of A7, there is a brownish crescent-like structure, extending just after A8. The height of this
structure is variable: low in the anterior part (at the
level of A7), maximum at the level of A8 and
decreasing towards the posterior part. The ventral
surface is then occupied by a series of white areas,
which may be divided into two parts: a nacreouswhitish crescent up to A9 and a pale white crescent
up to A10. The brown ventral shield is indistinct to
the naked eye, unlike the white region which is
clearly distinguishable. A longitudinal line on each
side of region-A, extending from A1 to A8, defines
the ventral limit between the muscular masses,
which are responsible for setal functioning, and the
ventral shield. This line describes an arch with the
ventral surface.
Region-B (Fig. 1A). This region always has more
than 2 segments, with a maximum of 27 (average
11.5 ± 2.4 mm in length, n = 18), each with two
rami. The notopodia, like in many other species,
have a bilobed Y-shaped inner part and a small
unilobed foliaceous outer part. The neuropodia are
unilobed on B1 and bilobed on B2 and following
segments, always transversally oriented.
Region-C (Fig. 1B). A maximum of 43 segments
were observed in this region, with a potential maximum length of 14.2 ± 2.9 mm (n = 4). The structure
of both noto- and neuropodia is the typical one of
the genus. The notopodia are erect, slightly inflated
distally, with no or 1 particularly evident lancet-like
notoseta projecting outside the terminal lobe. The
neuropodia have two fleshy lobes bearing uncini: an
upper, short, anteriorly directed lobe and a lower,
posteriorly oriented, vertically extended lobe, as on
neuropodia posterior to segment B2. Mature segments were observed in region-C.
Specialised seta of A4 (Fig. 1C, D). Each A4
parapodium has a single straw-coloured to yellowish
specialised seta, with a swollen, obliquely truncated
distal end, which is roughly heart-shaped (cordate)
in distal view. The outer lobe of the oblique section
is more developed than the inner one. The setal shaft
has a ventral groove all along its length and another
groove on the dorsal face just below the more developed lobe.
102 M. BHAUD et al.
FIG. 2. – Spiochaetopterus creoceanae sp. nov. Tube. A, Part of the
tube with marked joints, observed at low magnification; B, A onehole septum removed by the worm and then attached to the tube
wall; C, Serrated opening at border between two successive tube
sections; D, Lengthwise tear; E, Traces of the two directions of
mucus-layer deposition, observed at high magnification: arrow indicates top direction. Scale bars: A = 400 µm; B = 100 µm; C, D = 115
µm; E = 80 µm.
Uncini (Fig. 1E, F). Uncini elongate-triangular,
particularly small (22 µm long, 12-13.5 µm wide at
B1; 20 µm long, 11.8-12 µm wide on last segments
of region-C). The number of uncini vary between
340 at B1 and 20-30 at the last setigers of region-C
(but being only 4-10 in pre-pygidial setigers). The
teeth are small but visible under 100x magnifications. There are 15 to 17 teeth, 8 of which are located on the straight median section (of 10 µm in
length).
Tube (Fig. 2A–E). Tube unbranched, smooth, transparent, with distinct articulations and 750-800 µm in
diameter. Several free transverse partitions with a
single, median hole are observed inside the tube.
The differences observed in tube transparency are
directly related to the number of mucopolysaccharide layers (i.e. fewer in the more transparent tubes).
Longitudinal and transverse serration marks may be
observed between two successive tube sections.
Biological remarks
Some specimens have intra-coelomic oocytes
(80-85 µm in diameter), but neither the extent of the
reproductive period nor the current phase of the
cycle can be determined due to the brevity of the
collecting period. For the same reason, it is not
known whether the species reproduces asexually or
not. However, there is only one worm inside each
tube and no signs of regeneration are observed.
Ecological remarks
The characteristics of the water column over the
type locality during the sampling period (i.e.
August) may be summarised as follows: temperature
ranging from 35°C (bottom) to 36.8°C (surface),
water salinity always around 37‰, water turbidity
ranging from 2.5 mg (surface) to 6.5 mg (bottom) of
suspended solids per litre. The density of S. creoceanae sp. nov. (average of 630 ind. m-2) was characterised by a patchy distribution, with a maximum
of more than 2600 ind. m-2 at a single station, surrounded by areas with intermediate densities (430 ±
70 ind. m-2), and a few sites far from the main station
having less than 5 ind. m-2 (Table 1). Along the Iranian coast, hard bottoms are often present from the
beach shoreline down to 12-15 m in depth. In most
cases, the limestone rock is covered with dead coral
colonies (Price et al., 1993). At the study site, the
population of S. creoceanae sp. nov. was mainly
located seaward of this dead coral reef barrier. The
presence of hard bottoms near the shoreline prevented the worms from occurring in shallower bottoms
(i.e. less than 15 m deep). Thus, the population was
mainly concentrated at medium-depth bottoms
(around 17.6 ± 1.9 m in depth). These bottoms were
characterised by having compact sediments with
low contents of silt, pore water and organic matter
(average of 5.6 ± 1.6%, 23.5 ± 3.1% and 3.4 ± 0.2%
respectively) and intermediate values for gravel content (average of 44.8 ± 6.2%) (Table 1). The worms
were absent in all samples below 20 m in depth.
DISCUSSION
The Persian Gulf has been considered as a concentration basin (Tchernia, 1978), in which the
evaporation processes dominate the evaporation/
rainfall ratio (Sheppard et al., 1992). Seawater thus
tends to be salty and warm. The hydrological spatial
and temporal gradients are usually marked and
sometimes complex, often showing extreme temperatures and salinities (high or low depending on the
season). Although the local environmental data registered during this study cannot be extrapolated to
the whole year, they certainly correspond to ecological conditions that may be considered as extreme
for marine life. The water column over the area
inhabited by the studied population of Spiochaetopterus creoceanae sp. nov. was characterised by a very low oxygen concentration (Eric
Dutrieux, pers. comm.), as well as by high salinity
rates and extreme seasonal thermal oscillations
(which are particularly high in summer). The
observed bathymetrical differences in temperature
and salinity (both being slightly higher at the surface) cannot be attributed to a real water-mass stratification. As occurs in the whole Gulf, the influence
of a high irradiation level is probably a better explanation. The only significant difference with depth
was the high content in suspended matter near the
bottom. As in the whole Gulf (Sheppard, 1993),
continental inputs seldom occur in the study site
during summer (E. Dutrieux, pers. comm.). Thus,
the presence of particulate matter can probably be
attributed to hydrographical processes causing a
slight but significant increase of re-suspension near
the bottom.
Among the corresponding habitat preferences of
Spiochaetopterus creoceanae sp. nov., the low organic matter content in the sediments was particularly
relevant. Recent studies carried out in a Galician
estuary (NW Atlantic coast of the Iberian peninsula)
on a population refering to Spiochaetopterus
costarum postulated that the species could be considered as an indicator of organically polluted sediments (López-Jamar, 1985; Nozais, 1991). The main
reason was the observation of a gradient of decreasing density towards the mouth of the estuary, in parallel with decreasing silt and organic matter content
in the sediment. A more reliable explanation, however, could be a reduction of larval losses inside the
estuary. In fact, some topographical conditions particularly favourable to the existence of a hydrographical regime may generate retention structures, as
reported in other species with a theoretically high
dispersal capability linked to long-living planktonic
larvae (Bhaud, 1998b). Although our study did not
allow us to define the existence of retentive hydrographical structures, the presence of a nearby harbour at Nakhl e Taqi, as well as the particular position of the dead coral reefs facing the studied shoreline, may support their possible presence. More likely, the main concentration of worms could be
explained by the combination of an adequate hydrographical regime, suitable sediments and protection
by the hard bottoms and coral reefs.
A NEW SPIOCHAETOPTERUS SPECIES FROM THE PERSIAN GULF 103
FIG. 3. – Comparison of A4 specialized setae of the small-sized Spiochaetopterus species most closely resembling S. creoceanae sp.
nov. (A): B, S. solitarius; C, S. koreana; D, S. bergensis.
At a larger geographical scale, the circulation
pattern in the Persian Gulf is characterised by a
main current coming from the Strait of Ormuz and
drifting along the Iranian coastline towards the
inner regions of the Gulf. The current then turns
back and runs along the Arabian coast before again
reaching the strait (Sheppard et al., 1992). This
pattern may certainly contribute to submitting the
studied Iranian population to some degree of isolation inside the Gulf.
In fact, the morphological differences between
Spiochaetopterus creoceanae sp. nov. and the previously known species of the genus cannot be
underestimated and undoubtedly support the erection of the new taxon. The main difference
between S. creoceanae and the other known
species is the existence of 10 segments in the
region-A. The new species belongs to the group of
104 M. BHAUD et al.
small-sized Spiochaetopterus, among which it
closely resembles S. solitarius (Rioja, 1917), S.
bergensis Gitay, 1969 and S. koreana Bhaud et al.,
2002. A second relevant character, the morphology of the A4 specialized setae, confirms the validity of the new species. Effectively, S. creoceanae
has specialised setae roughly heart-shaped in distal view, with the outer lobe of the oblique section
being more developed than the inner one. S. solitarius and S. koreana both have two similar-sized
lobes while S. bergensis has three similar-sized
lobes (Fig. 3).
The presence of a new species of Spiochaetopterus in the Persian Gulf is an additional
evidence supporting the existence of phyletic radiations within the genus (Bhaud, 1998a). In fact, Spiochaetopterus costarum, which was considered as a
single cosmopolitan species by several authors,
turned out to be a complex species with a still undefined number of members (but probably more than
10-15 known to date). The main reasons that supported S. costarum as a cosmopolitan species were
the existence of a long-living planktonic larva having a large dispersal ability (Scheltema, 1974), and
too many authors willing to ignore specific differences. As was recently pointed out by Bhaud
(1998b) for other polychaete species, the increasing
number of descriptions of new species of Spiochaetopterus from all around the world (e.g.
Japan, Brazil, Yellow Sea, Persian Gulf) strongly
supports the existence of a discontinuity between
the existence of long-living swimming larvae and
the specific area of adult distribution. Thus, it seems
evident that careful studies using different
approaches (from classical but fine morphological
observations to genetic analyses) should be carried
out prior to attributing a cosmopolitan status to a
given species.
ACKNOWLEDGEMENTS
The authors wish to thank Eric Dutrieux who was
responsible for the field survey carried out by the
French company CREOCEAN. The authors are also
grateful to Meritxell Codina from the Departament
de Biologia Animal (Facultat de Biologia, Universitat de Barcelona) for the help with the preliminary
macrofaunal sorting. The study was partly financed
by a research contract between the CEAB (CSIC)
and CREOCEAN. This paper is a contribution to the
research project INTAS–OPEN–97–0916.
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A NEW SPIOCHAETOPTERUS SPECIES FROM THE PERSIAN GULF 105