First record of Dipolydora blakei (Annelida: Spionidae) from Europe: Greece, Mediterranean Sea

Abstract

The spionid polychaete Dipolydora blakei is identified from benthic samples collected in the Aegean Sea off Chalkis and the Ionian
Sea off Kalamitsi, both on the coast of Greece. Adults of D. blakei are characterized by having the falcate spines of chaetiger 5 with a large
lateral tooth and bristles on the convex side of a long and pointed main fang, awl-like spines in posterior notopodia, branchiae beginning
from chaetiger 7 and a pygidium with two lateral lobes. This is the first report of the species from the Mediterranean Sea and European
waters and the second record (after one from Brazil) outside its type locality in deep-water off New England, north-west Atlantic Ocean.
The morphology of the Greek specimens is described and illustrated and diagnostic characters of the species are discussed. The origin of
these specimens in the coastal waters of Greece is uncertain and the species is considered as cryptogenic in the Mediterranean Sea. An
identification key to Dipolydora species with bristle-topped falcate spines in chaetiger 5 known from the Mediterranean Sea and European
waters is provided.

Full text

Medit. Mar. Sci., 14/1, 2013, 19-23 19
First record of Dipolydora blakei (Annelida: Spionidae) from Europe:
Greece, Mediterranean Sea
V. I. RADASHEVSKY1,2 and N. SIMBOURA3
1 A.V. Zhirmunsky Institute of Marine Biology, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok 690059, Russia
2 Far Eastern Federal University, Vladivostok 690091, Russia
3 Hellenic Centre for Marine Research, Institute of Oceanography, 19013, Anavissos, Attica, Greece
Corresponding author: radashevsky@gmail.com
Received: 25 Arpil 2012; Accepted: 3 August 2012; Published on line: 19 February 2013
Abstract
The spionid polychaete Dipolydora blakei is identied from benthic samples collected in the Aegean Sea off Chalkis and the Ionian
Sea off Kalamitsi, both on the coast of Greece. Adults of D. blakei are characterized by having the falcate spines of chaetiger 5 with a large
lateral tooth and bristles on the convex side of a long and pointed main fang, awl-like spines in posterior notopodia, branchiae beginning
from chaetiger 7 and a pygidium with two lateral lobes. This is the rst report of the species from the Mediterranean Sea and European
waters and the second record (after one from Brazil) outside its type locality in deep-water off New England, north-west Atlantic Ocean.
The morphology of the Greek specimens is described and illustrated and diagnostic characters of the species are discussed. The origin of
these specimens in the coastal waters of Greece is uncertain and the species is considered as cryptogenic in the Mediterranean Sea. An
identication key to Dipolydora species with bristle-topped falcate spines in chaetiger 5 known from the Mediterranean Sea and European
waters is provided.
Keywords: polychaete, morphology, Aegean Sea, Ionian Sea, Mediterranean, cryptogenic species.
Short Communication
Mediterranean Marine Science
Indexed in WoS (Web of Science, ISI Thomson) and SCOPUS
The journal is available on line at http://www.medit-mar-sc.net
Introduction
In total, 398 species of polychaetes were identied in
samples collected from 188 stations in the Aegean and
Ionian Seas during ecological surveys along the coast
of Greece (Simboura et al., 2000). One sample from
the Ionian Sea, off Kalamitsi on the western coast
of Greece (Fig. 1), yielded small immature spionid
polychaetes which were reported by Zenetos et al. (1997)
as Polydora caulleryi Mesnil, 1897. One specimen of
the same species was also collected from the Aegean
Sea, Evvoikos Gulf, off Chalkis on the eastern coast
of Greece. These specimens are here described and re-
identied as Dipolydora blakei (Maciolek, 1984). This is
the rst report of the species from the Mediterranean Sea
and European waters.
Materials and Methods
Field collections were made in the Aegean and Ionian
Seas, Greece, eastern Mediterranean, on soft substrata
at depths ranging from 3 to 380 m between 1982 and
1993 (see Simboura et al., 2000). Sediment samples
were collected using grabs and screened using a 1 mm
mesh sieve. The residue was xed in 10% formaldehyde
solution, rinsed in fresh water, and then transferred to
70% ethanol. Polychaetes were removed from the residue
under a stereomicroscope at the Hellenic Centre for
Marine Research, Hellas, Greece. Photographs were taken
using a Zeiss Axio Imager 7.2 microscope equipped with
a digital camera; before photography, specimens were
stained with a solution of methylene green in alcohol.
The examined material was deposited at the Museum of
the Institute of Marine Biology (MIMB), Vladivostok,
Russia. Information about the samples is given below
along with description of the specimens. The number of
specimens in each sample is given in parentheses after
the museum abbreviation and registration number.
Type specimens of Polydora blakei from off New
England, U.S.A. deposited in the Allan Hancock
Foundation Polychaete collection of the Natural History
Museum of Los Angeles County, Los Angeles, U.S.A.
(LACM-AHF), and specimens of D. blakei from off the
states of Rio de Janeiro and São Paulo, Brazil deposited
in the E.F. Nonato Polychaete Collection of the Institute
of Biology of the Federal University of Rio de Janeiro,
Rio de Janeiro, Brazil (IBUFRJ), Museum of Zoology of
the University of São Paulo, São Paulo, Brazil (MZUSP),

20 Medit. Mar. Sci., 14/1, 2013, 19-23
and the Museum of Zoology of the State University
of Campinas, Campinas, Brazil (ZUEC) were also
examined.
Taxonomic Account
Spionidae Grube, 1850
Dipolydora blakei (Maciolek, 1984)
Polydora blakei Maciolek, 1984: 124–127, gs 1, 2.
Dipolydora blakei: Blake, 1996: 194. Radashevsky
& Paiva, 2010: 44–45, textg. 44.
Polydora caulleryi: Zenetos et al., 1997: 445, 449.
Not Mesnil, 1897.
Material
Greece: Mediterranean, Aegean Sea, Evvoikos Gulf,
off Chalkis, sta. XA1, 38°28.9´N, 23°35.9´E, 18 m, muddy
sand with coral debris, coll. N. Simboura, 8 Mar 1991,
MIMB 27141 (1). Ionian Sea, off Kalamitsi, sta. C4b,
38°58.155´N, 20°42.264´E, 63 m, sand with shell debris,
coll. N. Simboura, 26 Apr 1991, MIMB 27123 (11).
Supplementary material
U.S.A.: New England Continental Slope, Gay
Head-Bermuda Transect, R/V Atlantis, cruise 283, sta.
slope 2, 40°1.8´N, 70°42´W, 200 m, anchor dredge, coll.
H.L. Sanders, 28 Aug 1962, LACM-AHF POLY 1387
(holotype), 1388 (600 paratypes).
Brazil: Rio de Janeiro Continental Slope, northern
Bacia de Campos, Oceano Profundo II Project: sta.
50A, 22°02.847´S, 39°52.402´W, box core layer 2–5
cm, 1048 m, 29 Jun 2003, IBUFRJ 1284 (1); sta. 50,
22°04.567´S, 39°52.083´W, box core layer 5–10 cm,
1030 m, 30 Jun 2003, IBUFRJ 822 (1). São Paulo
Continental Platform, REVIZEE/Score Sul - Benthos
Project, sta. 6661, 24°07.637´S, 45°51.895´W, 147 m, 9
Jan 1998, ZUEC POL 669 (11); sta. 6665, 24°20.844´S,
44°09.913´W, 258 m, 10 Jan 1998, ZUEC POL 670 (4);
sta. 6686, 25°36.988´S, 45°13.571´W, 380 m, 13 Jan
1998, ZUEC POL 668 (1); REVIZEE/Score Central -
Benthos Project, sta. Y1-4, 22°19.083’S, 40°48.7’W, 60
m, 11 Jun 2002, MZUSP 81 (1).
Fig. 1: Map showing world-wide records of Dipolydora blakei and collecting sites off Greece. 1 – continental slope off New
England, U.S.A. (Maciolek, 1984); 2 – continental platform and slope off the states of São Paulo and Rio de Janeiro, Brazil
(Radashevsky & Paiva, 2010); lled triangle – Aegean Sea, off Chalkis; lled circle – Ionian Sea, off Kalamitsi (present study).
Fig. 2: Dipolydora blakei adult morphology (Aegean and
Ionian Seas, Greece, MIMB 27123, 27141). Relationship
between distribution of branchiae (number of the last
branchiate chaetiger) and total number of chaetigers in worm
(empty triangles), and anterior position of awl-like spines in
notopodia (number of the rst spine-bearing chaetiger) and
total number of chaetigers in worm (lled circles).

Medit. Mar. Sci., 14/1, 2013, 19-23 21
Adult morphology (Greek specimens)
Up to 4 mm long and 0.2 mm wide for 37 chaetigers.
Pigmentation absent. Prostomium anteriorly bilobed,
posteriorly extending to end of chaetiger 2 as a low
caruncle. Occipital antenna and eyes absent. Nuchal
organs as wide ciliary bands on sides of caruncle. Palps
as long as 10–15 chaetigers, with frontal longitudinal
groove lined with ne cilia.
Chaetiger 1 with short capillaries and small
postchaetal lamellae in both rami. Posterior notopodia
from chaetigers 17–28 to the end of the body with up to
15 awl-like spines gathered in cone-shaped bundles and
10 slender capillaries (Fig. 3F); spines gradually lost with
age from anterior notopodia and starting in more posterior
chaetigers in bigger individuals (Fig. 2). Spines with
distal tips pointed and basal parts blunt, conspicuously
protruding out of the body, slightly thinner and fewer at
rst appearance than in posterior chaetigers; old spines
thinner and shorter than new spines in same bundle.
Chaetiger 5 twice as long as chaetigers 4 and 6, with
up to 5 dorsal superior winged geniculate capillaries (Fig.
3C), 4 heavy falcate spines alternating with 1–2 thin
companion capillaries (Fig. 3D), and 6 winged ventral
capillaries (Fig. 3B); postchaetal lamellae absent. Dorsal
superior capillaries fewer and shorter than those on
chaetigers 4 and 6; ventral capillaries fewer but similar
in size to those on chaetigers 4 and 6; companion chaetae
alimbate or with narrow wing, situated between newly
developed falcate spines in posterior part of row. Falcate
spines each with long, distally pointed main fang and
large lateral tooth; convex side of main fang, except tip,
covered with ne bristles (Fig. 3E).
Hooded hooks in neuropodia from chaetiger 7, up
to 5 in a series, accompanied by 1–2 inferior capillaries
throughout and 1–2 alternating capillaries in 5–10
posterior chaetigers (Fig. 3A). Hooks bidentate; shaft
slightly curved, without constriction; newly developed
hooks bigger, situated in upper part of hook row (Fig. 3A,
ho). Inferior capillaries winged in anterior neuropodia,
very thin and alimbate in posterior neuropodia (Fig. 3A,
ic). Alternating capillaries hair-like, alimbate, situated
between upper hooks in row (Fig. 3A, ac).
Branchiae from chaetiger 7 to chaetiger 12, fewer
in small individuals (Fig. 2); attened, with surfaces
oriented parallel to body axis, basally fused to notopodial
postchaetal lamellae.
Pygidium small and eshy, bilobed, divided by
dorsal gap and ventral furrow into two rounded lateral
lobes, whitish due to numerous spindle-shaped glandular
cells with striated contents (Fig. 3F).
Glandular pouches in neuropodia from chaetiger 6,
large in chaetigers 7 and 8 and then gradually diminishing
in size.
Digestive tract without gizzard-like structure.
Habitat
In the Ionian Sea, adults of D. blakei were found off
the Kalamitsi coast, inhabiting silty tubes in clusters of
empty calcareous serpulid tubes on a sandy seabed with
shell debris, at a depth of 63 m. In total, 313 specimens
were found in two Ponar grab samples, each with a
sampling surface area of 0.045 m2. In the Aegean Sea,
the only individual of D. blakei was found off the Chalkis
Fig. 3: Dipolydora blakei adult morphology (Ionian Sea, Greece, MIMB 27123). A – chaetae from a posterior neuropodium, left
lateral view, dorsal side is to the left and the posterior end is up, showing alimbate capillary chaeta (ac) alternating upper (bigger,
newly developed) hooks in a row, four bidentate hooded hooks (ho), and alimbate inferior capillary chaeta (ic). B–E – chaetae from
notopodium of chaetiger 5: B, ventral capillary chaeta, C, short geniculate dorsal superior capillary chaeta, D, slender companion
capillary chaeta (anterior-row notochaeta), E, heavy falcate spine with large lateral tooth and ne bristles on convex side of
main fang (posterior-row notochaeta). F – posterior end, dorsal view, showing awl-like spines (sp) and capillary chaetae (ca) in
notopodia, and pygidium (py) composed by two lateral lobes containing large glandular cells.

22 Medit. Mar. Sci., 14/1, 2013, 19-23
coast, on a muddy sand seabed with fragments of a stony
coral Cladocora caespitosa (Linnaeus, 1767).
Distribution
West Atlantic Ocean, off North America (USA)
and South America (Brazil); rst record from European
waters, Mediterranean, Greece, presented here (Fig. 1).
Discussion
Dipolydora blakei was originally described by
Maciolek (1984, as Polydora blakei) based on material
collected at depths of 140–200 m along the east coast
of the United States, off New England, in the north-west
Atlantic Ocean. After its original description, the species
was reported from off the coast of the state of Rio de
Janeiro, Brazil, south-west Atlantic Ocean, at depths of
1030–1048 m (Radashevsky & Paiva, 2010) (Fig. 1). It
has not been reported from the east Atlantic Ocean or
the Mediterranean Sea despite intensive studies around
Europe. Type specimens of P. blakei from the continental
slope off New England, U.S.A. and specimens of D. blakei
from the continental slope and platform off the states of
Rio de Janeiro and São Paulo, Brazil were examined and
no morphological difference was found between them
and specimens from Greece. Both Brazilian and Greek
specimens are therefore referred to D. blakei. The origin
of the specimens in coastal waters around Greece remains
uncertain and the species is considered as cryptogenic in
the Mediterranean Sea.
Dipolydora blakei is a small gonochoristic species
up to 7 mm long and 0.5 mm wide for 40 chaetigers;
females produce small number of oocytes each at least
241 µm in diameter (Maciolek, 1984). Worms occur
rarely but may form local dense aggregations as a result of
asexual reproduction; specimens with signs of architomy
were found off the Brazilian coast (Radashevsky V.I.,
unpublished data). The details of the reproduction and
larval development of D. blakei are unknown.
With the present study, four Dipolydora species
with bristle-topped falcate spines in chaetiger 5 have
been reported from the Mediterranean Sea and European
waters. These are D. armata (Langerhans, 1880), D.
blakei, D. caulleryi (Mesnil, 1897), and D. quadrilobata
(Jacobi, 1883). Adults of these species are characterized
by having the prostomium incised to bilobed on the
anterior margin, branchiae beginning from chaetiger 7
and basally fused to notopodial postchaetal lamellae,
and awl-like spines in posterior notopodia. Dipolydora
armata adults bore into various calcareous substrata
while adults of the other species inhabit tubes on soft
substrata. A key for their identication is provided below.
Adults of D. blakei appear similar to those of D.
caulleryi in that the heavy falcate spines of chaetiger 5
have dense bristles on the convex side of the long, pointed
main fang. They differ, however, in having the falcate
spines of chaetiger 5 with a large lateral tooth, rather than
a main fang only and in having a bilobed pygidium with
two lateral lobes, rather than a quadrilobate pygidium with
one pair of dorsal and one pair of ventral lobes (see Blake,
1971). Dipolydora caulleryi is a common inhabitant
of soft bottom communities in the Mediterranean Sea
and European waters, thus D. blakei might have been
confused with this species. Reassessment of Dipolydora
specimens with bristle-topped falcate spines in chaetiger
5 from European waters would clarify the origin of D.
blakei in the Mediterranean Sea.
Key to Dipolydora species with bristle-topped falcate
spines in chaetiger 5 from the Mediterranean Sea and
European waters
1. Boring into shells, corals and coralline algae. Falcate
spines of chaetiger 5 each with large lateral tooth and
an apical transverse ange on the convex side of the
main fang. Pygidium cup-shaped to bilobed armata
- Inhabiting tubes on soft sediments. Falcate spines of
chaetiger 5 without apical transverse ange; lateral
tooth present or absent. Pygidium with two or four
lobes 2
2. Falcate spines of chaetiger 5 distally bifurcated, each
with two short massive unequal teeth and ne bristles
between them. Pygidium with four lobes
quadrilobata
- Falcate spines of chaetiger 5 each with a long pointed
main fang bearing dense bristles on the convex side.
Pygidium with two or four lobes 3
3. Falcate spines of chaetiger 5 without lateral teeth.
Pygidium four-lobed, with one pair of dorsal lobes
and one pair of ventral lobes caulleryi
- Falcate spines of chaetiger 5 each with large lateral
tooth. Pygidium bilobed, with one pair of lateral
lobes blakei
Acknowledgements
Our sincere thanks are to Leslie Harris, Tim Worsfold
and anonymous reviewers for valuable comments and
edits to the manuscript. Financial support was provided
by the Russian Foundation for Basic Research (Project
12-04-00609) through the A.V. Zhirmunsky Institute
of Marine Biology, and the Government of the Russian
Federation (Project 11.G34.31.0010) through the Far
Eastern Federal University.
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