Marine Ecology.

ISSN 0173-9565

ORIGINAL ARTICLE

One-year observation on the population structure

of Diopatra neapolitana Delle Chiaje (Polychaeta: Onuphidae)
in Izmir Bay (Aegean Sea, eastern Mediterranean)
Ertan Dagli, Zeki Ergen & Melih E. Cinar
Department of Hydrobiology, Faculty of Fisheries, Ege University, Bornova, Izmir, Turkey

Keywords
Diopatra neapolitana; population structure;
description; reproduction; Izmir Bay; Aegean
Sea.
Correspondence
Melih E. Cinar, Department of Hydrobiology,
Faculty of Fisheries, Ege University, 35100
Bornova, Izmir. E-mail: melih.cinar@ege.edu.tr
Accepted: 24 August 2005

Abstract
The present paper deals with the morphology and population structure of Diopatra neapolitana collected in Izmir Bay between July 1997 and June 1998. The
density of the species in the area ranged from 42 indm)2 (November 1997) to
198 indm)2 (March 1998), the biomass from 84 gm)2 (November) to
408 gm)2 (March). The coelom contained gametes in all months, except for
January, indicating that it has a very long reproductive period in the area. The
relationships among the biometrical measurements taken on specimens of Diopatra neapolitana from Izmir Bay were assessed by using regression and correlation analysis. A re-description of the species is provided.

doi:10.1111/j.1439-0485.2005.00055.x

Problem
Diopatra neapolitana Delle Chiaje 1841, which was originally described from the Mediterranean Sea (Sicily, Italy),
is known to form a dense population in the semi-polluted
zone of Izmir Bay (Ergen 1976). This species has been
sold as fish bait in fish markets in Izmir and a box of 12
specimens of this species costs approximately 1 YTL (0.58
). In addition to this species, the polychaetes Hediste
diversicolor (O. F. Muller 1776), Perinereis cultrifera
(Grube 1840) and Ophelia bicornis Savigny 1818 are also
used as fish baits in Izmir.
Diopatra neapolitana has been previously reported from
many localities in the world oceans; the Red Sea and
Indian Ocean (Fauvel 1953; Parameswaran 1973; Wehe &
Fiege 2002), the eastern Atlantic Ocean (Fauvel 1923) and
the Pacific Ocean (Choe 1960). However, Day (1967)
pointed out that the reports of D. neapolitana outside the
Mediterranean were dubious. Paxton (1993) stated that
the report of D. neapolitana from Japan by Choe (1960)
could be D. sugokai Izuka 1907. Thus, the real distributional pattern of D. neapolitana is unknown and the previous reports of this species require re-examination.

In the Mediterranean Sea, D. neapolitana was reported

to occur on vegetated or unvegetated muddy sand
bottoms in shallow waters (Fauvel 1923; Bellan 1964;
Ergen 1976; Lanera & Gambi 1993; Conti & Massa 1998;
Gambi et al. 1998). Gambi et al. (1994) also indicated its
economical importance [reported as D. cuprea cuprea
(Bosc 1802)] in Italy where almost 3.5 millions of worms
are being harvested in each year. In spite of its economical and ecological importance, no study dealing with its
population structure has been carried out up to date.
The present study provides information about the morphological feature and population structure of D. neapolitana collected from Izmir Bay (Aegean Sea) between July
1997 and June 1998.
Material and Methods
Specimens of D. neapolitana were collected monthly
between July 1997 and June 1998 on muddy sand bottom
with Cymodocea nodosa (Ucria) Ascherson off the Degas
Shipyard in the middle part of Izmir Bay, which was
regarded as a transitional or semi-polluted zone between
polluted and unpolluted waters of Izmir Bay (Kocatas

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Population structure of Diopatra neapolitana

1978) (Fig. 1). At the station, one square meter was sampled with a shovel (only one replicate), digging 15 cm
into the sediment. In the field, samples were sieved
through 2 cm mesh and the retained fauna were fixed in
4% formaldehyde.
In the laboratory, the material was washed under tap
water and specimens of D. neapolitana were separated
from other species. Each worm was removed from its
tube and weighed. Some biometrical features of each specimen such as the body length, the length of head and
first 10 chaetigers (H + 10), the width (excluding parapodia) of chaetiger 10 (W10), the tube width and length,
and the length of chaetigers carrying branchiae were
measured by using a digital compass. The total number
of chaetigers and the number of chaetigers with branchiae
were counted under a stereomicroscope. Each worm was
partly dissected to find out whether it contained gametes
or not. The diameter of eggs was measured with an
ocular micrometer.
The Regression and Pearson moment correlation analyses were performed on the data set to assess the relationship among the biometrical measurements taken. The
lengthweight relationship of unbroken specimens (a total
of 71 specimens) collected during the study period was
determined according to the allometric equation W
aLb where W is the total body wet weight (g), L the body
length (cm), and a and b are constants. The total body
lengths of broken specimens were determined by using
the linear regression equation estimated between the body
length of unbroken specimens and H + 10. The total
body weights of broken species were determined by using
the regression equation estimated between the body
weight of unbroken specimens and H + 10.
The salinity and temperature values of the area were
presented by Kaymakci (1998). The mean temperature
and salinity values of the surface water of the station
according to the seasons are as follows: Winter (14
C,

Fig. 1. The study area with the location of the sampling site.

266

33.5 psu), Spring (22
C, 34 psu), Summer (25
C,

35 psu) and Autumn (16
C, 33.5 psu).
The material was deposited at the Museum of the Faculty of Fisheries, Ege University (ESFM).

Results
Redescription of D. neapolitana from Izmir Bay

Diopatra neapolitana Delle Chiaje 1841 (Fig. 2).

Diopatra neapolitana, Clapare`de 1868: 122, pl. VI,
fig. 4; Fauvel 1923: 419420, fig. 166, ah.
Material examined: ESFM-POL/05-1, 24 March 2005,
Izmir Bay, off the Degas Shipyard, 0.5 m, on muddy sand
bottom with Cymodocea nodosa, 11 individual.
Description: Largest specimen complete, 347 mm long,
H + 10 11.42 mm, 7.74 mm wide, with 239 chaetigers.
Dorsum of anterior chaetigers dark brownish, posterior
ones pale brownish. Body iridescent. Dark brownish pigment bar present on mid-dorsal part of each chaetiger
between 2 and 41. Prostomium anteriorly pointed, with a
pair of subulate frontal lips and a pair of ventral upper
lips; oval, massive, fused at base. Styles of palps and
antennae dark brownish. Anterior part of ceratophores
dark brownish, posterior part pale yellowish. Tips of
median antenna and palps missing on the specimen
examined (Fig. 2A). Lateral antennae extending back to
chaetiger 7, with ceratophores having 1516 proximal
rings. Palps and lateral antennae arranged in rectangular
arrangement. Median antenna emerging just posterior to
line of lateral antennae. Nuchal organs subtriangular in
shape, located on posterior part of prostomium, longest
axis 528 lm long. Peristomium 1.5 times longer than
subsequent segment, with a pair of peristomial cirri, digitiform, located anteriorly, reaching back to posterior part
of peristomium (Fig. 2A).
Anterior parapodia larger than others, with rounded
prechaetal and long, subulate postchaetal lobes. Prechaetal and postchaetal lobes becoming gradually smaller,
prechaetal lobe disappearing by chaetiger 29. Ventral cirri
subulate, present on first 4 chaetigers. Ventral lobe present on chaetiger 625; most distinct on chaetigers 622.
Dorsal cirri subulate, longer on chaetigers with gills;
reaching up to mid-dorsum; gradually shortening
towards posterior end. Spiraled branchiae, greenish, from
chaetiger 5 to 65, best developed on chaetiger 7 with 14
whorls; 8 whorls on chaetiger 29; single filaments from
chaetiger 61.
Dorsal fascicle of chaetiger 1 with 2 simple chaetae and
3 pseudocompound chaetae; distal part 150 lm long;
ventral fascicle with 4 pseudocompound hooks; distal part
120140 lm long. Dorsal fascicle of chaetiger 3 with 1
limbate chaeta and 2 pseudocompound chaetae; distal

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Dagli, Ergen & Cinar

Population structure of Diopatra neapolitana

A B

F
Fig. 2. Diopatra neapolitana (ESFM-POL/05-1). A: Anterior end, B: pseudocompound hook with a modified secondary tooth, C: bidentate pseudocompound hook, D: subacicular hook, E: pectinate chaetae, F: maxillae I, II and IV. Scale bars: A: 1.5 cm, B, C: 25 lm, D: 70 lm, E: 28 lm,
F: 325 lm.

part 160 lm long. Ventral fascicle with 5 pseudocompound chaetae; distal part 150170 lm long. Pseudocompound chaetae distinctly bidentate, with long, pointed
hoods; secondary tooth sometimes broken or deformed
into very small two teeth (Fig. 2B and C). Unmodified
parapodia (chaetiger 4 onwards) with pectinate and limbate chaetae. Additional subacicular hooks clearly bidentate; proximal tooth longer than distal one, from
chaetiger 18, numbering 2 per parapodium (Fig. 2D).
Pectinate chaetae always present on dorsal fascicle, numbering 7 on chaetiger 24, 5 on chaetiger 76, 2 on chaetiger 227; arranged in oblique rows; with 610 pointed
teeth (Fig. 2E). Limbate chaetae present both dorsal and
ventral fascicles, coarsely serrated after chaetiger 40.
Pygidium with two pairs of short anal cirri.
Jaws darkly sclerotized, almost black (Fig. 2F). Mandibles with slender shafts and high calcerous cutting plates
with two weak indentations each. Maxillary formula: Mx
I 1 + 1; Mx II 8 + 5; Mx III 6 + 0; Mx IV
7 + 6; Mx V 1 + 2.
Tube composed of whitish parchment-like inner lining,
and outer covering of muddy sand and pieces of shell
fragments and algae. Fecal pellets had sand inclusions,
pieces of Cymodocea nodosa and diatom frustules.

Population structure of D. neapolitana from Izmir Bay

A total of 1294 specimens of D. neapolitana were collected

during the study period. The density of the species in the
area ranged from 42 indm)2 (November 1997) to
198 indm)2 (March 1998) (Fig. 3). As for the mean density value according to seasons, it attained the highest score
in spring (134 indm)2 32 SE), and the lowest score in
autumn (76 indm)2 8 SE). The biomass (wet weight) of
the species in the area ranged from 84 gm)2 (November)
to 408 gm)2 (March). At the seasonal base, spring had the
maximum mean biomass score (308 gm)2 50 SE),
followed by summer (283 gm)2 26 SE), winter
(201 gm)2 21 SE) and autumn (168 gm)2 43 SE).
The results of the regression analyses based on biometrical measurements of unbroken specimens are as follows:
the relationship between the body length of worms [L
(mm)] and the number of chaetigers (NC) was NC
0.53 L + 77.04 (R2 0.82, r 0.91, n 71, P < 0.05);
that between the body length of worms (L) and the width
at chaetiger 10 [W (mm)] was W 0.011 L + 3.05
(R2 0.42, r 0.53, n 71, P < 0.05); that between the
body length of worms (L) and the length of head and
first
10
chaetigers
(H + 10)
was
H + 10

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267

Dagli, Ergen & Cinar

Population structure of Diopatra neapolitana

Fig. 3. Density and biomass values of

Diopatra neapolitana at station during July
1997June 1998.

0.022 L + 5.94 (R2 0.43, r 0.53, n 71, P < 0.05);

that between the body length of worms (L) and the number of chaetigers with branchiae (NB) was NB
0.033 L + 34 (R2 0.38, r 0.53, n 71, P < 0.05);
that between the body length of worms (L) and the
length of chaetigers with branchiae [LCB (mm)] was
LCB 0.1092 L + 19.439 (R2 0.56, r 0.60, n
71, P < 0.05). The total length and number of chaetigers
of broken specimens were estimated by the formulas
given above. The correlation coefficients among parameters indicated that all of them are size dependent.
The length frequency histograms of individuals collected from July 1997 to June 1998 are shown in Fig. 4. The
longest individual was determined to be 38.9 cm in April,
the shortest to be 9.5 cm in September. The maximum
length for female was detected in June, August and October, the minimum in August; the maximum length for
male was found in August, the minimum in March and
August. Juvenile worms were observed in all months.
However, the percentage of juvenile worms in the population was higher than 10% in months from February to
September, with August (28%) and March (16%) having
the highest scores.
The lowest and highest scores for individual biomass
were found to be 0.92 and 19.75 g, respectively. The seasonal trend showed that individuals of a high biomass
value occurred in May and June, mainly due to the fact
that individuals in this months contained dense reproductive products within their coelomic cavities.
The length (cm)weight (g) relationships are presented
in Fig. 5. The regression formula for the relationship was
found to be W 0.012 L1.88, indicating that the
growth for D. neapolitana is allometric as the parameter b
(1.88) is lower than 3.
The minimum size and biomass value for males were
11 cm and 0.17 g, respectively; for females 12.5 cm and
0.77 g, respectively. The mean oocyte diameter was
268

190 2.8 SE lm (n 102, range: 50230 lm). Unripe

eggs bearing nurse cells had a mean size of 117 6.5 SE
lm (n 30, range: 50160 lm). Nurse cells (up to
10 lm in diameter) are attached to eggs with two strings,
which reach up to 35 cells and measure up to 184 lm
long. Oocytes larger than 160 lm did not bear nurse cells.
Sperms have spherical heads and grouped in capsules in
the coelomic cavity. As it can be seen in Figs 3 and 4,
D. neapolitana reproduces in all months in the area,
except for January. However, the majority of specimens
collected from April to August possessed gametes, showing that the main reproductive period for D. neapolitana
in the area took place during this period. In Figs 3 and 4,
a number of specimens were classified as unknown.
They were indeed broken specimens, which contained no
gametes on their anterior fragment available and whose
total lengths were estimated according to the regression
formulae given above. Sperm capsules and oocytes in ripe
individuals first appeared on chaetigers from 32 and 35,
respectively. On the other hand, in some specimens gametes may commence from the chaetiger 78 (female) or 85
(male). On the average, sperm capsules first appeared
from chaetiger 51 0.88 (n 100) and oocytes from
chaetiger 55 0.87. The statistical analysis showed that
there was no correlation (r 0.03) between the first
appearance of gametes on the body and, the width and
H + 10 of the worms. The overall male:female ratio was
0.75. This ratio seemed to change by months; females
particularly dominated the population in March and
April, male in May and June. In other months, the ratio
seemed near to 1. There is no significant difference
between the lengths of male and female (t-test, P 0.26).
Male and female individuals of D. neapolitana can be easily detected as the color of female becomes greenish at
maturity, while the color of males becomes light cream or
gray, mainly owing to reproductive products within the
coelomic cavity.

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Population structure of Diopatra neapolitana

Fig. 4. The length frequency histograms of individuals of Diopatra neapolitana collected from July 1997 to June 1988.

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269

Dagli, Ergen & Cinar

Population structure of Diopatra neapolitana

Fig. 5. The lengthweight relationship of Diopatra neapolitana.

Discussion
The present study examined the population structure and
morphology of D. neapolitana collected from Izmir Bay
during July 1997June 1998. The worm is of commercial
value in the area and is being used as fish bait for a number of fishes such as Gilt-head sea-bream (Sparus aurata
Linnaeus, 1758) and European sea bass [Dicentrarchus
labrax (Linnaeus, 1758)]. There is no statistical data concerning the bait worm fishery in Turkey. Diggers are collecting this species only from the area where this study
was carried out, using a sieve with the wooden frame and
a shovel. This species dominates especially the shallow
water of the area (less than 1 m depth) and rarely occurs
at 3 m or deeper. In addition to this worm, this area was
also inhabited by two other bait species, Solen marginatus
Pulteney 1799 and Upogebia pusilla (Petagna 1792).
According to an interview with the dealer of the worm
diggers, D. neapolitana also occurred in high densities off
Inciralti (see Fig. 1) in the past, but is now restricted to
the investigated area. Ten years ago each digger could collect ca. 2000 specimens within an hour, whereas now it
takes 10 h to collect the same number, indicating that the
stock has been overexploited during the time. Almost 50
persons subsist on the worm fishery in Izmir and harvest
a total of 7.5 million worms from the area in each year
earning a total of 625,000 YTL (ca. 364,000 ). The inflation rate in the prize of the worm seems to be fairly constant; a box of 12 worms cost 1.19 US$ in 1997, 1.37 US$
in 2005. All worms are consumed within Izmir, a small
amount of them are being transported to Ayvalik (north
Aegean Sea) and Mersin (Levant Sea). This worm has also
an economical importance in Italy and a total of 3.5 million of worms are being harvested each year (Gambi et al.
1994).
The density of D. neapolitana in the area had lower
scores in autumn. This could be attributed to the fact
that demands on this worm are very high during spring
270

and summer, thus the digging activity for this worm was
intense. In the winter period, it seemed that the stock was
recovering and was represented by a higher number of
individuals in March. However, a long term study is
required to assess the impact of digging activities on the
population density of this species. The maximum density
of D. neapolitana in Izmir Bay was 198 indm)2 (March
1998). However, Conti & Massa (1998) reported a very
high density (1500 indm)2) of this species in S. Gilla
Lagoon (southern Sardinia). The big difference in the
population density of this species between Izmir Bay and
S. Gilla Lagoon is complicated to evaluate, but the primary production of the areas and digging activities would
be the main factors controlling the population density of
this species.
Some authors such as Day (1967) and Paxton (1993)
pointed out that the reports of D. neapolitana outside
the Mediterranean seemed to be dubious. The species
Diopatra aciculata Knox & Cameron 1971 is very similar
to D. neapolitana in terms of general morphological appearance and chaetal type. Thus, the status about whether
D. aciculata is distinct or not was stated to be unclear
(Paxton 1993). However, we found some differences
between D. neapolitana and D. aciculata. Nuchal organs
are rounded in D. aciculata (see Paxton 1993, fig. 31B)
versus clearly sub-triangular in D. neapolitana; peristomial
cirri about three times as long as the peristomium in
D. aciculata versus the same length as the peristomium in
D. neapolitana; the ventral lobe is present from chaetiger
5 to 20 in D. aciculata versus 6 to 25 in D. neapolitana;
Mx. I is slender in D. aciculata versus medially extended
in D. neapolitana; Mx. II and III with coarse, truncated
tip in D. aciculata versus triangular, sharp in D. neapolitana. According to the differences given above, D. aciculata seems to be a distinct species. However, some
morphological differences can be expected between
distant populations of a species, thus a molecular analysis
should be carried out to reach a reliable conclusion about
the taxonomic validity of D. aciculata. It should be kept
in mind that the introduction of the species from one
area to another via ship hulls or ballast water is severe
nowadays and the introduction of D. neapolitana in the
other parts of the world could be possible.
There has been no previous gut content analysis of this
species. Our observations on remnants in digestive tracts
and fecal pellets of a number of specimens collected
showed that this species largely fed on leaves of Cymodocea nodosa and algae associated with it. We also carried
out a breeding experiment on this species to observe its
feeding behavior. This species spent most of its time
within its tube, with its tentacles protruding a few millimeters outside the aperture. Periodically, the worm extended its anterior segments outside the tube and browsed

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Dagli, Ergen & Cinar

on the bottom. It bit leaves of C. nodosa and withdrew

immediately into the tube. It also preferred eating invertebrates such as polychaetes and amphipods. It reacted
very rapidly to rotten animals, showing that it is also a
scavenger. In the absence of food, it fed on detritus on
the bottom. This also explains why this species prefers
especially organically polluted bottoms of Izmir Bay.
Mangum et al. (1968) reported that the worm tube of
Diopatra cuprea (Bosc 1802) also acted as a food-catching
device and its efficiency was governed by the volume of
water flowing past.
Our observations on the reproductive biology of
D. neapolitana showed that this species possessed gametes
within the body cavity in all months, except January. The
continuously reproducing behavior of this species, that
has been overexploited by diggers, plays an important role
to support new recruits to the area. Paxton (1993) classified four development patterns for the species of Diopatra;
brooding in the parental tube (group I); direct development in a cocoon (group II); egg masses attached to the
parental tube and direct development (group III); and
broadcast spawning with a free-swimming stage (group
IV). This species seems to belong to group IV, with an
adult reaching its maturity at the minimum length of
11 cm, an egg size of ca. 200 lm. However, Lo Bianco
(1898) reported that D. neapolitana deposited yellow eggs
in a brown, irregular, gelatinous mass (one square centimeter in size) on the distal end of its tube (group III).
However, we did not observe such a mass on tubes.
In the group IV, eggs are believed to release in a jelly
mass that immediately dissolves in seawater, as occurred
in D. cuprea (Paxton 1993). Conti et al. (2005) also
reported that this species is a free spawner and uses
external fertilization.
Conti et al. (2005) reported that the external morphology of the mature male gamete of D. neapolitana conforms to that of a typical ect-aquasperm by having a
short, rounded head with a conical acrosome, five mitochondria and a long tail. Our observation on the spermatozoo of D. neapolitana from Izmir Bay agrees with the
Contis findings.
Published information on the population structure of
Diopatra spp. is scarce. Choe (1960) performed a study
on the life history of Diopatra sugokai [this species was
given as D. neapolitana but Paxton (1993) stated that this
species might belong to D. sugokai] and provided the
reproductive and developmental biology as well as population structure of the species from Japan. He also calculated the lengthweight relationship of Japanese
specimens of D. sugokai and estimated the allometric
equation for the species as W 0.018 L1.86. We also
found an allometric growth for D. neapolitana. Peckol &
Baxter (1986) studied the population dynamics of Diopa-

Population structure of Diopatra neapolitana

tra cuprea in North River Estuary (Beaufort, North Carolina) and concluded that D. cuprea is a perennial species,
with its stable population densities in the area being
maintained by high adult survival rather than extensive
recruitment.
The present study provides information about the life
history traits of D. neapolitana from Izmir Bay, where it is
one of the polychaete species of economical importance.
Future studies to be undertaken on the population structure and biology of this species would shed more light on
its life history and functional role in the ecosystem.
Summary
The population structure of D. neapolitana was studied
during the period July 1997June 1998 at one station
located off the Degas Shipyard in the middle part of
Izmir Bay. The density of the species in the area ranged
from 42 indm)2 (November 1997) to 198 indm)2
(March 1998), the biomass from 84 gm)2 (November) to
408 gm)2 (March). The longest individual was determined to be 38.9 cm in April, the shortest to be 9.5 cm
in September. The length (cm)weight (g) relationships
were formulated as W 0.012 L1.88. The minimum size
and biomass value for males were 11 cm and 0.17 g,
respectively; for females 12.5 cm and 0.77 g, respectively.
The species, based on a single, voucher specimen from
the area, was re-described. Gut content analysis and the
experimental study showed that the species is omnivorous, a scavenger and detritus feeder. The relationships
among biometrical measurements taken on specimens
were assessed by regression and correlation analysis.
Acknowledgements
The last author (MEC) had the chance to study with Dr
Kristian Fauchald in the Smithsonian Institution (Washington DC, USA) and was greatly impressed with his deep
wisdom, knowledge about polychaetes and kind fellowship.
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Marine Ecology 26 (2005) 265272 2005 The Authors. Journal compilation 2005 Blackwell Publishing Ltd