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Redescription of Scolelepis tridentata (Southern,

1914) (Annelida: Spionidae), with description of a
new species of Scolelepis from the Black Sea

Victor Surugiu

To cite this article: Victor Surugiu (2023) Redescription of Scolelepis tridentata (Southern, 1914)
(Annelida: Spionidae), with description of a new species of Scolelepis from the Black Sea, The
European Zoological Journal, 90:2, 584-603, DOI: 10.1080/24750263.2023.2229855

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The European Zoological Journal, 2023, 584–603
Vol. 90, No. 2, https://doi.org/10.1080/24750263.2023.2229855

Redescription of Scolelepis tridentata (Southern, 1914) (Annelida:

Spionidae), with description of a new species of Scolelepis from the
Black Sea

VICTOR SURUGIU *

Facultatea de Biologie, Universitatea “Alexandru Ioan Cuza” din Iaşi, Iaşi, România

(Received 5 April 2023; accepted 18 June 2023)

Abstract
The spionid genus Scolelepis is commonly divided into the subgenera Scolelepis (s. str.) and Parascolelepis. In the Black Sea, two
species ascribed to the subgenus Parascolelepis have been reported so far – Scolelepis tridentata and Scolelepis cantabra. Both species
are quite rare and when found are usually difficult to identify because of their fragility. To ascertain the identity of specimens from
the Black Sea, the type-series of S. tridentata and non-type specimens of S. cantabra were examined. The examination of specimens
from the Black Sea identified as Scolelepis tridentata revealed that they differ from the nominal species in several characters.
Therefore, they are described herein as Scolelepis bellani n. sp., a species that most closely resembles S. korsuni from Arctic waters.
Also, the occurrence of S. cantabra in the Black Sea is confirmed.

http://zoobank.org/urn:lsid:zoobank.org:pub:78FD82BA-084F-4213-A7A0-41C1EA6ACD44
http://zoobank.org/urn:lsid:zoobank.org:act:FF4E2A3C-7607-4402-9F79-42399574CCC5

Keywords: Morphology, Polychaeta, Scolelepis, Spionidae, taxonomy

Introduction
Scolelepis Blainville, 1828, is one of the most speciose
and taxonomically difficult genera of spionid poly­
chaetes (Blake et al. 2020). Currently, the genus num­
bers 91 species and is commonly divided into two
subgenera (Sikorski & Pavlova 2015; Blake et al.
2020; Read & Fauchald 2023). The subgenus
Scolelepis Blainville, 1828, includes species with blunt
or conical, uni-, bi- or tridentate hooded hooks with
falcate or straight shafts, while the subgenus
Parascolelepis Maciolek, 1987 comprises species having
sharp, multidentate hooks with curved shafts
(Maciolek 1987). To this, Meißner and Götting
(2015) added the presence in Parascolelepis of papil­
lated palpal sheaths as opposed to smooth sheaths
fused to the palp bases in the subgenus Scolelepis (s.
str.). Even though the recent phylogenetic analysis of
the genus Scolelepis (s. lat.) made by Lee and Min
(2022b) supports the grouping of the genus into two
subgenera, the species “traditionally” defined as
Parascolelepis are scattered in both clades. Therefore,
in the present paper only the genus level is used.
Members of Scolelepis typically inhabit sandy or
mixed soft sediments in intertidal and shallow subtidal
habitats. Currently, six species of the genus Scolelepis
(s. lat.) have been reported in the Black Sea: Scolelepis
cf. cirratulus (Delle Chiaje, 1829), S. foliosa (Audouin
& Milne Edwards, 1833), S. mesnili (Bellan &
Lagardère, 1971), S. neglecta Surugiu, 2016,
S. tridentata (Southern, 1914) and S. cantabra
(Rioja, 1918) (Kisseleva 2004; Surugiu 2016; Kurt
Șahin et al. 2017; Surugiu et al. 2022). The identifica­
tion of the latter two species often proved to be diffi­
cult because of the fragmentary nature of the material
and some overlapping characters. Both species were
formerly reported either as Nerinides or as

*Correspondence: Victor Surugiu, Facultatea de Biologie, Universitatea “Alexandru Ioan Cuza” din Iaşi, B-dul Carol I, Nr. 20A 700507 Iaşi, România.
Email: vsurugiu@uaic.ro

© 2023 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group.
This is an Open Access article distributed under the terms of the Creative Commons Attribution-NonCommercial License (http://creativecommons.org/
licenses/by-nc/4.0/), which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly
cited. The terms on which this article has been published allow the posting of the Accepted Manuscript in a repository by the author(s) or with their consent.

Pseudomalacoceros, the invalid names of the genus
Scolelepis. The records of S. tridentata and S. cantabra
are confused and sometimes contradictory, not only in
the Black Sea, but also in other parts of the world.
The main morphological characters of systematic
importance of Scolelepis include: the shape of the pros­
tomium, the presence or absence of the occipital ten­
tacle, the presence or absence of notochaetae on
chaetiger 1, the degree of fusion of branchiae to noto­
podial lamellae and the chaetiger on which these start
to separate, the chaetiger on which neuropodial
hooded hooks begin, the shape and the number of
hooded hooks, the number and the arrangement of
apical teeth of hooks, the ratio of lengths of hood to
main fang, the presence or absence of a notch in
posterior neuropodial postchaetal lamellae, the shape
of the mid-body and posterior notopodial postchaetal
lamellae, and the shape of the pygidium (Pettibone
1963; Light 1977; Blake 1983; Maciolek 1987;
Imajima 1992; Sikorski 1994; Lee & Min 2022a).
Careful examination of material labeled “Nerinides
cantabra” in the collection of the Grigore Antipa
National Museum of Natural History, Bucharest,
Romania (MGAB) revealed that it actually contained
specimens belonging to two species preserved in the
same vial. One of the species was identified as
Scolelepis cantabra, while the second more closely
resembled S. tridentata. To clarify the taxonomic iden­
tity of specimens from the Black Sea, the type series of
S. tridentata (Southern, 1914), deposited in the
National Museum of Ireland, Natural History of
Dublin (NMINH), and available material of
S. cantabra has been examined. Because of some inad­
vertencies found in subsequent descriptions of
S. tridentata, the species is herein redescribed in detail
and illustrated comprehensively to avoid future confu­
sion. Due to several differences found in comparison to
S. tridentata, the material originally identified in the
Black Sea as Scolelepis tridentata is described as a new
species.
Material and methods
Morphology
For light microscopy (LM) specimens were rinsed in
distilled water and examined under stereo- or com­
pound microscopes. To contrast some characters useful
in species identification (e.g., shape of notopodial lamel­
lae, occipital tentacle, caruncle extension, palpal papil­
lae, shape of pygidium, etc.), methylene blue (MB)
temporary staining was applied. For this, specimens
were transferred first into distilled water for a while,
then dipped for a few seconds into a saturated solution
of MB in water and finally returned to distilled water for
A new species of Scolelepis from the Black Sea 585
microscopical examination (the staining fades quickly if
specimens are returned to alcohol).
Photographs of specimens were taken using a Leica
DFC450C camera coupled to a Leica M205A stereo­
microscope or to a Leica DM750 compound micro­
scope. Before photography, specimens were stained
with an aqueous solution of MB. Line drawings were
prepared based on photographs using Adobe Illustrator.
For scanning electron microscopy (SEM) studies,
selected specimens were transferred from 70% ethanol
to distilled water for a few hours to allow the ethanol
in the tissues to be replaced with distilled water. After
cutting the specimens transversally into fragments,
they were sonicated in distilled water for 0.5–3 min
at 43 kHz to remove hoods from the hooks, dehy­
drated in a graded ethanol series (50%, 70%, 80%,
90%, 96% and 99.3%, leaving them for 10 minutes in
each concentration) and then acetone, critical point
dried (CPD) in carbon dioxide, mounted on stubs,
coated with a thin layer (30 nm) of gold and exam­
ined with a scanning electron microscope at an accel­
erating voltage of 30 kV. SEM observations were
carried out using a Vega Tescan SBH in the
Electron Microscopy Laboratory, Faculty of Biology,
Alexandru Ioan Cuza University of Iaşi.
The specimens of the newly described species were
deposited in the collection of MGAB and NMINH. In
the “Material examined” sections, complete specimens
are indicated by “cs”, while anterior, middle, and pos­
terior fragments are designated as “af”, “mf”, and “pf”,
respectively. Information about samples is given below
along with descriptions of specimens.
In addition to the freshly collected material, speci­
mens deposited in various museum collections were
examined. The following abbreviations were used
for the museums and institutions that provided
loans or lodged registered specimens:
DBUA Biological Research Collection of Marine
Invertebrates, Departamento de Biologia,
Universidade de Aveiro, Portugal
MGAB Grigore Antipa National Museum of
Natural History, Bucharest, Romania
MNHN Muséum National d’Histoire Naturelle,
Paris, France
NHMD Natural History Museum of Denmark,
Copenhagen, Denmark
NHMUK Natural History Museum of United
Kingdom, London, UK
NIBR National Institute of Biological
Resources, Incheon, South Korea
NMINH National Museum of Ireland, Natural
History, Dublin, Ireland
SMF Senckenberg Museum, Frankfurt, Germany
ZMH Zoologisches Museum Hamburg, Germany
586 V. Surugiu

Morphometric analysis correlation analyses between the size of the worms (as
an independent variable) and the morphological vari­
A total of 72 specimens in optimal condition (i.e. speci­
ables, that were revealed by multivariate analysis as
mens that presented all analysed morphological char­
most useful in discriminating between species, were
acters) belonging to six species of Scolelepis (17
carried out.
individuals of S. tridentata from Ireland, Spain, and the
Mediterranean coast of France, five individuals of
S. papillosa (Okuda, 1937) from Korea (based on litera­ Taxonomic account
ture data), seven individuals of S. korsuni Sikorski, 1994
from the Atlantic coast of Norway, two individuals of Family Spionidae Grube 1850
S. quinquedentata (Hartmann-Schröder, 1965) from Genus Scolelepis Blainville, 1828, emend. Pettibone
Chile, 35 individuals of Scolelepis bellani n. sp. from 1963
the Black Sea and six individuals of S. cantabra from
Ireland, France, Portugal and the Black Sea) were ana­ Type-species: Scolelepis squamata (O.F. Müller,
lysed. The following eight variables were recorded 1806) as Lumbricus squamatus. By monotypy.
(Table S1): the ratio between the body length in milli­
metres and the number of chaetigers (L/NC); body Scolelepis tridentata (Southern, 1914)
width (W); caruncle length as the number of the chae­
tigers to which it extends back (Ca); number of chaeti­ (Figures 1–4)
gers to which branchiae are completely fused to Nerinides tridentata Southern, 1914: 98–99, pl. X,
notopodial postchaetal lamella (Br); number of chaeti­ fig. 23A–J.—McIntosh 1922: 20–21.—Fauvel
ger on which neuropodial hooded hooks start (VH); 1927: 33, fig. 10f–l.
number of apical teeth in hooded hooks (NT); max­ Scolelepis (Nerinides) tridentata.—Light 1977: 75–76,
imum number of hooks per neuropodium (MH); and fig. 3c–e (paratypes only).
the ratio of hood length to main fang height (H/F). Body Scolelepis (Parascolelepis) tridentata.—Maciolek 1987:
width refers to the distance in mm between the distal 33–34.—Hartmann-Schröder 1996: 337.
most structures on the widest chaetiger seen on the Pseudomalacoceros tridentata.—Parapar et al. 1992:
anterior end in dorsal view (including parapodia but 112, fig. 2C.
without chaetae). Maximum length and width (usually
at chaetiger 10–14) of worms were measured using Material examined.
a calibrated ocular micrometer mounted on
a stereomicroscope to the nearest 10 μm. Individuals Type material
with missing values were excluded from the multivariate
Holotype: NE ATLANTIC: IRELAND,
statistical analysis (measurements and calculations are
Blacksod Bay, Station W166, in Laminaria
available in online supplementary material, Tables S1
roots, Clare Islands Survey, coll. R. Southern,
and S2).
20 Sep. 1910, complete specimen in alcohol
To take the different data types (continuous and
(Industrial Methylated Spirits), NMINH
discontinuous variables, ratios and proportions) into
1914.325.1.—Paratypes: Same data as for holo­
account, Gower’s similarity coefficient (Gower 1971)
type, one complete specimen in alcohol (IMS),
was chosen to calculate a similarity matrix. Non-metric
NMINH 1914.325.2; one complete specimen
Multi-Dimensional Scaling (nMDS) was subsequently
broken into two fragments, NMINH
employed to display the similarities of the different
1914.325.3; one middle fragment, possibly dis­
specimens. To test for the significance of differences
sected, NMINH 1914.325.4; one anterior frag­
between species, a series of non-parametric ANOSIM
ment, NMINH 1914.325.5; one middle
(analysis of similarities) tests was performed (Anderson
fragment, NMINH 1914.325.6; one specimen
2001). Principal Component Analysis (PCA) was used
broken into an anterior end, three middle sec­
to determine variability of characters and to identify
tions (one of them was used with permission for
characters that contribute most to the species differen­
SEM) and a posterior end, NMINH 1914.325.7;
tiation. For this, the Principal Component scores were
same data as for holotype, one anterior end,
correlated (Spearman Rank Correlation Coefficient)
NHMUK 1914.12.12.29.
with the measured character values of each individual.
All multivariate statistical analyses were carried out with
Non-type material
the PRIMER v 6.1.13 software package.
To discriminate between S. korsuni from Norway IRELAND, Blacksod Bay, Station W137, on
and S. bellani n. sp. from the Black Sea, a series of shore, Clare Islands Survey, coll. R. Southern,
 A new species of Scolelepis from the Black Sea 587

c d

b e f

Figure 1. Scolelepis tridentata (Southern, 1914), syntype (NHMUK 1914.12.12.29): A. Anterior end, dorsal view; palps, left notopodial
postchaetal lamellae of chaetiger 3, right parapodia of chaetigers 1 and 2, and right notopodial postchaetal lamellae of chaetigers 3 and
4 missing. B. Anterior end, lateral view; C. Notopodial limbate capillary chaetae, chaetiger 35. D. Neuropodial hook of chaetiger 34.
E. Ventral inferior capillary chaeta, chaetiger 35. F. Alternating capillary chaeta, chaetiger 35. Scale bars: A, B = 0.5 mm; C = 0.1 mm; D–
F = 50 μm.

Mar. 1910, one complete specimen broken into two
fragments in alcohol (IMS), NMINH 1910.42.4;
Station W135, on shore, Clare Islands Survey, coll.
R. Southern, Mar. 1910, one complete specimen
broken into two fragments, NMINH 1910.42.5;
Station W168, in Laminaria roots, Clare Islands
Survey, coll. R. Southern, Sep. 1910, one dehy­
drated specimen and two middle sections,
NMINH 1910.448.3; Station W160, on shore, in
Laminaria roots, Clare Islands Survey, coll.
R. Southern, Sep. 1910, one complete specimen,
NMINH 1910.448.4.
SPAIN, NW Iberian Peninsula, Ría de Ferrol,
Galicia, Batel, coll. J. Parapar, 26 Oct. 1988,
No. 225, Ref. 225136261086-9, 4 anterior frag­
ments; SW Iberian Peninsula, Huelva, coll. J.M.
Viéitez, May 1988, 2 anterior fragments, poorly
preserved.
WESTERN MEDITERRANEAN SEA,
FRANCE, Golfe de Marseille, Carry-le-Rouet,
Sta. E 3 M C 1 (I), Corer 1, in dead Posidonia,
Carottier–KL, coll. & det. A. Willsie, 6 Sep. 1982,
1 af, SMF 16030; Carry-le-Rouet, Sta. E 3 M C 4
(II), Corer 4, in dead Posidonia, Carottier–KL, coll.
588 V. Surugiu
H. Zibrowius, det. A. Willsie, 9 Sep. 1982, 1 af,
SMF 16278; Plateau des Chèvres, Port Cros,
Sta. 1 M 5 (I), in dead Posidonia, Carottier–KL,
coll. H. Zibrowius, det. A. Willsie as Nerinides sp.,
25 Sep. 1984, 1 af, SMF 16319.
Description
Holotype complete specimen, ca. 7.5 mm long,
0.681 mm wide for about 65 chaetigers (see also
Taxonomic discussion). The longest complete para­
type ca. 19 mm long, 1.5 mm wide for ca. 65 chae­
tigers. The largest incomplete paratype 2.0 mm
wide, 5.4 mm long and with 19 chaetigers. Colour
in alcohol pale tan or opaque off-white. Body broad­
est at anterior chaetigers 8–12, tapering to both
ends. Anterior body flattened dorsally and rounded
ventrally, posterior body more or less rounded in
cross-section. Anterior segments short, mid-body
and posterior segments longer.
Prostomium sharply or bluntly tapered anteriorly,
extending posteriorly to middle of chaetiger 2 as
a depressed caruncle (Figure 1A). Short, erect,
stubby or conical occipital antenna near posterior
margin of caruncle (between palp bases), usually
directed forward. Two pairs of equal-sized,
rounded, dark-brown eyes, arranged in a nearly
straight transverse row or eyes absent. Palps missing
in all examined specimens (see also Taxonomic
discussion).
Peristomium well developed, separated from pros­
tomium by a distinct furrow, without lateral wings,
and folded longitudinally on ventral side
(Figure 1B).
Primary and secondary dorsal transverse ciliated
bands (TCB) in anterior chaetigers to around chae­
tiger 18 (Figure 1A); primary TCBs continuous with
ciliation on inner edge of branchiae; secondary
TCBs at the junction between chaetigers.
Chaetiger 1 with small, bluntly conical notopodial
lamella and small, rounded neuropodial lamella and
about 9–12 short neuropodial capillaries
(Figures 1A, B, 2B); notopodial capillary chaetae
absent (see Taxonomic discussion).
Branchiae from chaetiger 2 (Figure 2B) and con­
tinuing to last chaetiger (Figure 3A). Branchiae
meeting dorsally at around chaetigers 9–23
(Figure 1A).
Notopodial postchaetal lamellae broad, rounded,
completely fused with branchiae through anterior
and middle chaetigers (Figure 3B), folded along
outer margin (Figure 2C), not reaching the tips of
branchiae from around chaetigers 19–34 or comple­
tely fused to branchiae to posteriormost chaetigers;
in far posterior chaetigers notopodial postchaetal
lamellae relatively large, foliaceous (Figure 2E).
Notopodial prechaetal lamellae low, rounded to
conical, best developed in chaetigers 2–8, then gra­
dually reduced in size (Figure 2B, C).
Neuropodial postchaetal lamellae rounded, auri­
culate anteriorly (Figure 2A, B), more elongate,
lower and rectangular in subsequent chaetigers
(Figure 2C), with an elongated superior lobe by
chaetigers 12–14, situated dorsal to chaetae by chae­
tiger 16–21, triangular at first (Figure 2D), becom­
ing more rounded in posterior chaetigers
(Figure 2E).
Anterior chaetae all narrowly winged capillaries
arranged in two rows. Notopodial capillaries
longer, longitudinally striated, those of anterior
row shorter (Figure 1C); capillaries of posterior
notopodia arranged in indistinct double rows; addi­
tional dorsal superior fascicle of 3–5 long, thin
capillaries. Neuropodial capillaries in distinct dou­
ble rows: up to eight capillaries in anterior row,
these short, broad, uniformly granulated; chaetae
of posterior row longer, thinner, lacking granula­
tions and an inferior bundle of 2–5 thin capillaries.
Posterior capillaries slender, alimbate
(Figure 1E, F).
Neuropodial hooded hooks first present from
chaetigers 15–17 in posterior row, numbering up
to 16 per fascicle, accompanied by an anterior row
of 0–7 slender alternating capillaries (in dorsalmost
position) (Figure 1F) and 2–4 ventral inferior slen­
der capillaries (Figures 1E, 2D); number of alter­
nating capillaries gradually diminishing towards
posterior region. Hooded hooks with very long,
sharp main fang bent at an almost right angle to
shaft, surmounted by 2–3 small, closely applied api­
cal teeth (Figures 1D, 4B–D); shaft gently curved
(Figure 1D). Hoods short, 1.7–2.8 times as long as
main fang (Figures 1D, 4A). Ventral sabre chaetae
absent. Notopodial hooded hooks absent.
Pygidium cushion-like, with slightly bilobed ven­
tral margin and dorsal anus (Figure 3A).
Staining pattern
Prostomium, peristomium, caruncle, occipital tentacle,
branchial tips, margins of notopodial pre- and post­
chaetal lamellae, neuropodial postchaetal lamellae and
pygidium more intensely stained (Figure 3A, B). Each
segment has a broad band of stained speckles on the
ventrum, more intense on anterior margin, leaving two
whitish (without speckles) paramedian lines.
Habitat and ecology
The species inhabits the upper sublittoral and is
found in Laminaria roots or in dead Posidonia.
 A new species of Scolelepis from the Black Sea 589

a b

d e

Figure 2. Scolelepis tridentata (Southern, 1914), syntypes (A–D – NMINH 1914.325.7; E – NMINH 1910.42.4): A. Parapodium of
chaetiger 1, anterior view. B. Parapodium of chaetiger 2, anterior view. C. Parapodium of chaetiger 8, anterior view. D. Parapodium of
chaetiger 21, anterior view. E. Parapodium of posterior chaetiger, anterior view. Scale bars: A–E = 0.2 mm.

Biology
description and illustrations for his new species.
Ovoid (180 × 279 µm) oocytes with honeycombed Unfortunately, the designated holotype is a specimen
envelope are present in the coelom from chaeti­ that seems to have regenerating the prostomium, peri­
ger 17. stomium and the first four chaetigers (judging from
the smaller size of segments, branchiae and neuro- and
Taxonomic discussion
notopodial lamellae). Palps were missing in all exam­
Southern (1914) provided an exceptionally good ined specimens. However, Southern (1914) reported
590 V. Surugiu

Figure 3. Scolelepis tridentata (Southern, 1914), syntype (NMINH 1914.325.7), LM micrographs, stained with MB: A. Pygidium, postero-
latero-dorsal view; B, chaetigers 22–27, dorsal view. Scale bars: A = 0.25 mm, B = 0.2 mm.

neuropodial lamellae in posterior chaetigers and mul­

that “the tentacles [=palps] are short, thick and firmly
tidentate hooded hooks. However, S. tridentata can
adherent, of a deep chocolate colour”. To my knowl­
be distinguished from the other species by having the
edge, there are no subsequent descriptions of the palps
shortest lengths of the hoods as compared to the
of S. tridentata, but papillae at the bases of the palps are
height of the main fang. Thus, this ratio is 1.9–2.8
expected to be present. According to Southern (1914),
in S. tridentata (Southern 1914: fig. 23j; Light 1977:
the notopodial lamella of chaetiger 1 is composed of
fig. 3d; Parapar et al. 1992: fig. 2c; present study) vs.
two rounded lobes (see his Fig. 23C). However, it
3.1–4.6 in S. papillosa (Okuda 1937: fig. 2 f; Lee et al.
is likely that he misinterpreted the prechaetal lamella
2021: fig. 4b), 3.2–6.5 in S. quinquedentata
of chaetiger 2 as the second upper lobe. This was
(Hartmann-Schröder 1965: fig. 190; Carrasco 1974:
corrected by McIntosh (1922), who showed that in
fig. 14), 3.4–4.3 in S. texana (Foster 1971: fig. 139;
chaetiger 1 there is “a small conical papilla or cirrus”
Maciolek 1987: fig. 10c), 3.3 in S. towra (Blake &
[=notopodial postchaetal lamella]. Light (1977) and
Kudenov 1978: fig. 5e), and 3.0–6.2 in S. korsuni
Maciolek (1987) re-examined some of the type speci­
(Sikorski 1994: fig. 1 g; present study). In
mens, but incorrectly reported notopodial capillaries
S. tridentata notopodial postchaetal lamellae are not
as being present in chaetiger 1 and hooded hooks as
distinctly separated from branchiae, but rather do not
having 3–5 (usually 4 or 5, re: Light 1977) apical teeth.
reach the tips of the latter in middle and posterior
Careful examination of the type material revealed that
chaetigers. The same is true also for S. papillosa (Lee
the original description was correct and that the noto­
et al. 2021).
chaetae are absent in chaetiger 1 in all examined speci­
Apart from the ratio of lengths of hood to main
mens. Also, close examination of hooded hooks under
fang, S. tridentata differs from S. papillosa only by
SEM revealed that these have only two or three teeth
having fewer hooded hooks per fascicle (10–16 in
above the main fang (Figure 4B–D). Specimens of
the former vs. 18–21 in the latter).
Scolelepis cf. tridentata reported from Northern
California (Light 1977; Maciolek 1987; Blake 1996)
Occurrence and distribution
do have notochaetae on chaetiger 1 and thus most
likely belong to another, yet undescribed species. Atlantic coasts of Ireland (Southern 1914; Maciolek
Fixing these errors will enable further correct identifi­ 1987), France (Fauvel 1927), and Spain (Parapar
cation of this species. et al. 1992) and the Western Mediterranean Sea
Scolelepis tridentata is closely related to S. papillosa (present study).
(Okuda, 1937), S. quinquedentata (Hartmann-
Schröder, 1965), S. texana Foster, 1971, S. towra Scolelepis bellani sp. nov.
Blake and Kudenov, 1978, and S. korsuni Sikorski,
1994. All these species lack notochaetae in chaetiger (Figures 5–7)
1, have notopodial postchaetal lamellae completely Nerinides tridentata.—Vinogradov 1931: 7–9, fig. 2.
fused with branchiae in anterior chaetigers, undivided Not Southern 1914
 A new species of Scolelepis from the Black Sea 591

Figure 4. Scolelepis tridentata (Southern, 1914), syntype (NMINH 1914.325.7), SEM micrographs: A, Neuropodial hooks with hoods,
lateral view; B, Neuropodial hook with three apical teeth, latero-apical view, hood removed; C, Neuropodial hooks with two and three
apical teeth, latero-apical view, hoods removed; D, Neuropodial hook with two apical teeth, apical view, hood removed. Scale bars:
A = 20 μm; B–D = 10 μm.

Nerinides cantabra.—Dumitrescu 1963: 186 (partim).
Not Rioja 1918
Pseudomalacoceros tridentata.—Kisseleva 2004: 251–
252, fig. 99. Not Southern 1914
Material examined.
Type material
Holotype: BLACK SEA: UKRAINE,
Grigorievsky Liman, Sta. 18, 46.601745°N,
31.020987°E, depth 0.3 m, temperature 22°C, sali­
nity 15.54, dissolved oxygen 7.4 mg L–1, coll. I.A.
Sinegub, 23 Jul. 2015, MGAB PLY0166.
Paratypes: Same data as for holotype, 1 cs + 4
af, MGAB PLY0167; Grigorievsky Liman,
46.6096°N, 31.0137°E, depth 0.3 m, fine compact
sand with mud, coll. I.A. Sinegub, 12 and 29
Jul. 2013, det. O. Bondarenko as Nerinides
tridentata, 3 af of which one as SEM stub, MGAB
PLY0168; Gulf of Odessa, Cape Malyi Fontan,
Sta. D90, 46.441667°N, 30.775000°E, depth
6.4 m, sand, pooled with Sta. 24, 46.3250°N,
30.7000°E, depth 11 m, temperature 17.6°C, dis­
solved oxygen 10.01 mg L–1, coll. A.P. Kurakin,
Jul. 2012, 4 af + 1 mf + 1 pf, MGAB PLY0169;
same data as for preceding, 3 af + 2 mf + 1 pf,
NMINH 2023.3.1.
Additional material
BLACK SEA: ROMANIA, original label “Nerinides
cantabra; Rioja, 1918, Marea Neagră”, 8 af, poorly
preserved, MGAB PLY0170; UKRAINE,
Dzharylhach Island, marine side, 46.013833°N,
32.904056°E, midlittoral zone in the excavated pit
(higher than edge of the water), medium sand mixed
with broken shells, coll. M.O. Son, 21 Aug. 2015, 3
592 V. Surugiu

Figure 5. Scolelepis bellani n. sp., paratype (A–C, F – MGAB PLY0169), holotype (D – MGAB PLY0166), and voucher specimen (E –
VS), LM micrographs, stained with MB: A. Anterior end, dorsal view, palps missing. B. Anterior end, lateral view, palps missing.
C. Anterior end, frontal-dorsal view, palps missing. D. Middle body chaetigers (chaetigers 36–41), dorsal view, showing notopodial
postchaetal lamellae divided into a superior flag-like process and inferior narrow part. E. Anterior end, lateral view, showing palps with
papillate basal sheaths. F. Pygidium, dorsal view. Scale bars: A–D, F = 0.5 mm; E = 0.25 mm.

af, MGAB PLY0171; CRIMEA, Karkinit Bay,
45.86469°N, 33.51307°E, depth 0.6–0.8 m, coll.
N. Boltachova, 24 Aug. 2008, identified as
Scolelepis tridentata, 3 af, MGAB PLY0172;
Sevastopol Bay, 44.61977°N, 33.54409°E, depth
17 m, coll. N. Boltachova, 10 Nov. 2006, identified
as Scolelepis tridentata, 3 af + 1 mf, MGAB
PLY0173; SEA OF AZOV, RUSSIA, Taman
Bay, Sta. 33–2, 45.29050°N, 36.806033°E, depth
4.5 m, coll. & det. V. Syomin as Scolelepis sp., 10
Jul. 2013, det. as Scolelepis korsuni by
V. Radashevsky in 2016, 3 very small af of which
one was lost during the study, MGAB PLY0174;
Taman Bay, Sta. 43–2, 45.338252°N,
36.781862°E, coll. & det. V. Syomin as Scolelepis
cf. cantabra, 9 Aug. 2012, 1 af, MGAB PLY0175;
Taman Bay, Sta. 3–2, 45.27933°N, 36.95785°E,
depth 1.8 m, coll. & det. V. Syomin as Scolelepis cf.
cantabra, 3 Jul. 2013, 1 af + 2 mf, MGAB PLY0176;
Taman Bay, Sta. 3–3, same data as for preceding,
1 af with palps, VS personal collection; Taman
Bay, Sta. 3–4, same data as for preceding, 2 af,
MGAB PLY0177.
TYRRHENIAN SEA: ITALY: Campania, N. of
Napoli, Gulf of Gaeta, Licola, 1966–1968, leg.
J. Dörjes, 1 af, SMF 12914.

Comparative material
Scolelepis korsuni. Holotype: NORTH SEA:
NORWAY: Lille Frigg II, Sta. 4–1, 59.96167°N,
2.395556°E, depth 108 m, Van-Veen grab (0.1 m2),
muddy sand, coll. Akvaplan-niva, 11 May 1992, det.
A. Sikorski, 1 Dec. 1992, NHMD 108731 (POL-
000948); non-type material: NOWEGIAN SEA:
NORWAY: Gullfaks oil field, station 4–2,
61.19725°N, 2.302004°E, depth 211 m, coll.
A. Sikorski, 5 Jun. 2002, 1 cs + 1 af + 1 mf + 2
palps, MGAB PLY0178; Mosjøen, Vefsnfjorden, sta­
tion 4, 65.85503°N, 13.17673°E, depth 107 m, coll.
A. Sikorski, 30 Jun. 2006, 2 af, MGAB PLY0179;
Breisundet 17, Ref. 2–1, 67.419967°N, 13.884983°
E, depth 42 m, coll. A. Sikorski, 24 Jan. 2017, 1 cs + 1
af, MGAB PLY0180.
Scolelepis quinquedentata. Holotype: CHILE, Isla
Mocha, Sta. 75, black fine sand with small amount
of detritus, depth 26 m, 10.2°C and 1.84 ml L–1 O2,
11 Mar. 1960, 1 af, ZMH P-14928. Paratypes: same
data as for holotype, 4 af, poorly preserved, ZMH
P-14929.
Scolelepis cantabra. NE ATLANTIC: PORTUGAL,
continental shelf off Cascais, Guia, Sta. G6,
38.678333°N, 9.471667°W, 40 m depth, Jun. 1998,
Smith-McIntyre grab, RV Andrómeda/Auriga, cruise
GUIA98, 1 af, DBUA 0170.02; shelf off Cascais,
Guia, Sta. G18, 38.666667°N, 9.436667°W, 40 m
depth, Jun. 1998, Smith-McIntyre grab, RV
Andrómeda/Auriga, cruise GUIA98, 1 af, DBUA
0190.01; Sado estuary, Sta. S4, 38.472933°N,
8.780000°W, 8 m depth, Feb. 1997, Smith-McIntyre
grab, RV Mestre Costeiro, cruise TOXSADO97, 1 af
and 1 mf, DBUA 0191.01; continental shelf off
Aveiro, Sta. S16, 40.636550°N, 8.773150°W, 12 m,
Dec. 2002, Smith-McIntyre grab, RV Andrómeda/
Auriga, cruise SIMRIA02, 1 af, DBUA 2022.01;
IRELAND, Blacksod Bay, Fishery Station W166,
sand, Clare Islands Survey, coll. R. Southern, 16–23
Sep. 1910, det. as Nerine longirostris, 1 af, poorly pre­
served, NMINH 910.448.5; FRANCE, Dinard,
1893, coll. de M. le Baron de St. Joseph, No. 20–
1911, det. as Nerine longirostris Qfg., re-det. as
Nerinides cantabra by N. Maciolek (?), 1 af + 1 mf,
MNHN-IA-PNT 143; Dinard, St. Vaast, 1882, coll.
de M. le Baron de St. Joseph, No. 20–1911, det. as
Nerine longirostris Qfg., 1 af + 3 mf, MNHN-IA-PNT
144; Dinard, 1893, coll. de M. le Baron de St. Joseph,
No. 20–1911, det. as Nerine longirostris Qfg., 68c, 1 af,
MNHN-IA-PNT 145; Tatihou, May 1895, leg. & det.
as. Nerine foliosa by P. Fauvel, 1 af + 1 mf, MNHN-IA-
PNT 146; BLACK SEA: ROMANIA, original label
A new species of Scolelepis from the Black Sea 593
“Nerinides cantabra Rioja, 1918, Marea Neagră”, 3 af,
poorly preserved, MGAB 30.288.
Description
Holotype incomplete specimen, ca. 9.0 mm long,
2.2 mm wide for 41 chaetigers. Longest complete
specimen 8.5 mm long, 1.5 mm wide for 60 chae­
tigers; longest incomplete specimen 22 mm for
about 61 chaetigers; largest specimen 3.1 mm
wide (at chaetiger 17) for 64 chaetigers. Colour in
alcohol opaque off-white, without pigmentation.
Body very fragile, broad, flattened anteriorly, nar­
row and almost cylindrical in cross section in mid-
body and posterior segments. Anterior segments
short, mid-body segments longer.
Prostomium pointed anteriorly, terminating pos­
teriorly in narrow caruncle reaching anterior margin
of chaetiger 2, posterior part of caruncle pointed and
raised as short, stubby, anteriorly directed occipital
antenna (Figures 5A, C, 8A). Two pairs of small,
indistinct, equal, dark-brown oval eyes, arranged in
a wide trapezium or in a nearly straight transverse
row just in front of palp bases, with anterior eyes set
further apart; sometimes eyes absent. Palps (absent
in holotype) very caducous, reaching at most chae­
tiger 20. Edges of basal sheaths papillated; papillae
triangular, pointed, 3–9 in number on lateral and
posterior sides of palp bases, unequal in size, the
largest in posterior most position (Figure 5E).
Peristomium distinct from chaetiger 1, inflated,
without lateral wings (Figures 5B, C, E, 8A).
Metameric dorsal ciliated organs as primary
TCBs, continuous with ciliation along the inner
edge of branchiae; anterior 14–30 chaetigers (22 in
holotype) with additional secondary TCBs located
at the junction between chaetigers (Figure 5A). Low
dorsal transverse folds present between the bran­
chiae on middle chaetigers (Figure 5D).
Chaetiger 1 with short, tapering, cirriform noto­
podial lamellae and larger, auricular neuropodial
lamellae; capillary chaetae present in neuropodia
only (Figures 5C, 6A).
Branchiae from chaetiger 2 (Figures 5C, 6B), best
developed by chaetiger 10–20, then gradually
decreasing in size, continuing until last five segments
(Figures 5F, 7A, B).
Notopodial postchaetal lamellae of anterior chae­
tigers attain full size by chaetigers 5–6, very broad,
with entire folded outer margin, with glandular cells,
fused to branchiae along entire length in first 15–30
chaetigers (Figures 5A, 6C, D). At around chaetiger
15–31 (27 in holotype) a notch divides notopodial
lamella into superior rounded lobe and broad,
594 V. Surugiu

c
b

d
e

Figure 6. Scolelepis bellani n. sp., paratype (MGAB PLY0169): A. Parapodium of chaetiger 1, anterior view. B. Parapodium of chaetiger 2,
anterior view. C. Parapodium of chaetiger 3, anterior view. D. Parapodium of chaetiger 14, anterior view. E. Parapodium of chaetiger 30,
anterior view. Scale bars: A–E = 0.2 mm.

inferior elongated lobe (Figure 6E). In subsequent
chaetigers notch deepens and enlarges, separating
both lobes completely. Superior lobe gradually
diminishes in size to an apical, rounded, flag-like
process; inferior lobe becomes elongate and narrow
(Figures 5D, 7A). In posterior chaetigers notopodial
postchaetal lamellae foliaceous (Figures 5F, 7B).
Notopodial prechaetal lamellae low and rounded,
best developed in chaetigers 5–17 (Figure 6D).
Neuropodial postchaetal lamellae wide, auricular
in anteriormost chaetigers (Figure 6A–D). In follow­
ing chaetigers lamellae become narrower and more
elongate, not notched, with superior lobe projecting
upward (Figures 6E, 7A).
 A new species of Scolelepis from the Black Sea 595

c d

Figure 7. Scolelepis bellani n. sp., paratype (MGAB PLY0169): A. Parapodium of chaetiger 44, anterior view. B. Parapodium of posterior
chaetiger, anterior view. C. Mid-part of a notopodial limbate capillary chaeta, chaetiger 30. D. Neuropodial hook of chaetiger 30. Scale
bars: A, B = 0.2 mm; C, D = 20 μm.

Anterior parapodia with only capillary chaetae in
both rami, all broad, narrowly limbate, arranged in
double rows (Figure 8B); capillaries of anterior row
shorter, broader, with moderately and uniformly
granulated cores in direct light and fibrous in
reflected light; capillaries of posterior row longer,
thinner, lacking granulations (Figure 7C); notopo­
dial capillaries 1.5 times longer than neuropodial
ones; additional superior and inferior fascicle of
long, thin capillaries without granulations.
Notopodial capillaries very long in posterior chae­
tigers (Figure 7B).
596 V. Surugiu

Figure 8. Scolelepis bellani n. sp., paratype (MGAB PLY0168), SEM micrographs: A. Anterior end, dorsal view, palps missing.
B. Neuropodial capillary chaetae of chaetiger 4. C. Neuropodial hooded hooks of chaetiger 28, left latero-ventral view (dorsal side is to
the left and the anterior end is down). D, Neuropodial hook with three apical teeth, latero-apical view, hood removed. E, Neuropodial
hook with four apical teeth, frontal view, hood removed. F, Neuropodial hook with four apical teeth, latero-apical view, hood removed.
Scale bars: A = 0.5 mm; B, C = 50 μm; D–F = 5 μm.

Neuropodial hooded hooks present from chaeti­
gers 15–28 (24 in holotype) in posterior row, num­
bering up to 23 per row (up to 15 in holotype),
accompanied by 1–3 slender alimbate capillaries in
dorsal part (Figures 6E, 7B, 8C). Hooded hooks
with bluntly tipped main fang, forming a wide
angle (~110°) to shaft, surmounted by three
(Figure 8D) or four slender apical teeth
(Figure 8E, F); shaft long, gently curved, without
constriction. Hood elongated, 4–5 times the length
of main fang (Figure 7D). Ventral sabre chaetae
absent. Notopodial hooded hooks absent.
Pygidium a slightly bilobed ventral pad; anus as
dorsal vertical slit. One or two prepygidial achaetous
segments (Figure 5F).
Staining pattern
Occipital tentacle, caruncle, notopodial lamella of
chaetiger 1 and tips of palpal papillae most intensely

stained (Figure 5A, C, E). Blue pigmentation on
margins of both notopodial and neuropodial post­
chaetal lamellae, intense blue elongate spots at pos­
terior ventral base of each neuropodial postchaetal
lamella (Figure 5B). Prostomium and peristomium
with uniformly dispersed speckles (Figure 5A–C, E).
Ventrally peristomium with longitudinal rows of
blue speckles. Each segment presenting ventrally
two speckled transverse strips, interrupted mid-
ventrally, with anterior strip longer and more intense
than the posterior one (Figure 5B). Palpal sheaths
with dispersed speckles (Figure 5E).
Etymology
This species is named in honour of Dr Gérard Bellan,
Centre d’Océanologie de Marseille, France, in recogni­
tion of his extensive studies of the polychaete fauna of
the Mediterranean Sea. I dedicate this species to him
also as a moral amend for mistakenly synonymyzing his
species Scolelepis mesnili (Bellan & Lagardère, 1971)
with S. squamata (Surugiu 2016), which subsequently
was confirmed as a distinct species based on molecular
markers (Surugiu et al. 2022).
Gender: Masculine
Type locality
Black Sea, Ukraine, Grigorievsky Liman,
46.601745°N, 31.020987°E, depth 0.3 m.
Habitat and ecology
Species inhabits coarse sand between algae, gravely
sand, sand with shell debris, fine sand and muddy
sand at 0–30 m depth (Vinogradov 1949; Caspers
1951; Dumitrescu 1957; 1963; present study).
Biology
Scolelepis bellani n. sp. is gonochoristic. Ovoid
(123 × 197 µm) oocytes with reticulated envelope
from chaetiger 26–38.
Morphometric analysis
The non-metric Multi-Dimensional Scaling
(nMDS) plot (Figure 9A) provided a fairly good
representation of differences between species (2D
Stress value = 0.16). There appears to be a good
segregation of S. cantabra, S. papillosa,
S. quinquedentata, and S. tridentata. However,
S. bellani n. sp. and S. korsuni are morphologically
poorly separated.
The Principal Component Analysis (PCA), whose
first principal component (PC1) accounted for
A new species of Scolelepis from the Black Sea 597
75.0% of the total variability in the data set,
the second principal component (PC2) for 15.5%
and the third principal component (PC3) for 8.4%,
gave similar results (Figure 9B). The PCA plot of
the first two principal components showed that
S. quinquedentata forms a distinct group from
remaining species and has the highest PC1 scores,
which generally correspond to the fact that all bran­
chia are not completely fused with the notopodial
postchaetal lamellae. In contrast, S. tridentata and
S. papillosa, which have branchiae completely fused
with the notopodial lamellae to the last segments,
have the lowest PC1 scores. The remaining species,
in which notopodial lamellae separate from the
branchiae at a certain body segment, are placed in
the middle position of the PCA plot.
The ANOSIM analysis resulted in a global
R-value of 0.751 (p < 0.001), indicating that there
are statistically significant differences between
groups of a priori defined morphospecies.
Subsequent pairwise tests established that there are
strong and highly significant differences (p < 0.001)
between some species (e.g., between S. bellani n. sp.
and S. cantabra, between S. tridentata and
S. cantabra, between S. bellani n. sp. and
S. tridentata, and between S. tridentata and
S. korsuni) (Table I). However, the morphometric
differences between S. bellani n. sp. and S. korsuni
are weakest, but still significant (R = 0.349, p =
0.003).
The Spearman Rank Correlation Coefficient
between the Principal Component’s scores and the
measured character values of specimens revealed that
the segment on which notopodial lamellae starts
separation from branchiae (Br), the maximum num­
ber of hooks per row (MH), and the size of worms
expressed as the width of anterior chaetigers (W) are
the most important characters in discriminating
between analysed species (rS > 0.50 at p < 0.01).
Also, fairly good discriminating characters in distin­
guishing between analysed species (along PC2) are
the first hook-bearing segment (VH) and the ratio of
hood length to main fang height (H/F) (rS > 0.55 at
p < 0.01) (Table S2).
Therefore, to discriminate between S. bellani
n. sp. and S. korsuni scatter plots of the body
width (W) against segment on which notopodial
lamellae start splitting from branchiae (Br), and the
first hook-bearing segment (VH) were constructed.
The maximum number of hooks per row (MH) was
not taken into account because it gradually increases
towards the posterior end and most of the analysed
specimens lacked posterior parts. The scatter plots
show that both the splitting between branchiae and
598 V. Surugiu

Figure 9. Results of the morphometric multivariate analysis of individuals belonging to six species of Scolelepis: A. Non-metric multi­
dimensional scaling (nMDS) plot. B. Principal Component Analysis (PCA) plot. S. tridentata (green upward-pointing triangles),
S. papillosa (blue circles), S. korsuni (blue downward-pointing triangles), S. quinquedentata (pink squares), S. bellani n. sp. (red diamonds),
and S. cantabra (grey asterisks). Holotypes represented by hollow symbols.

notopodial lamellae (Br) and the neuropodial hooks

(VH) appear earlier in S. korsuni than in S. bellani
(Figure 10).
Taxonomic discussion
S. bellani n. sp. is most closely related to
S. tridentata from Ireland, S. papillosa from Korea,
S. quinquedentata from Chile, and S. korsuni from
the North Sea. All these species are characterized by
a pointed prostomium, presence of an occipital
antenna, palps with papillated basal sheaths, the
absence of notochaetae in chaetiger 1, posterior
neuropodia without ventral papilla, and a bilobed
pygidium.
However, S. bellani n. sp. can be distinguished from
S. tridentata in that it has a superior flag-like process on
mid-body notopodial postchaetal lamellae (absent in
S. tridentata) and branchiae are separated from notopo­
dial lamellae from chaetigers 25–30 (completely fused
or only weakly separated to end of body in S. tridentata).
Also, in S. bellani n. sp., the angle between the main
fang and shaft of the hooks is larger than 100°, while in
S. tridentata it is around 90–100°. Finally, in S. tridentata
neuropodial hooded hooks have 2–3 apical teeth (rather
than the 3–4 in S. bellani n. sp.), and an inferior fascicle
 A new species of Scolelepis from the Black Sea 599

Table I. Results of ANOSIM pairwise comparisons between analysed species of Scolelepis,

showing R statistics and p values.

Species R statistic p value

Scolelepis tridentata vs Scolelepis papillosa 0.628 0.001**

Scolelepis tridentata vs Scolelepis korsuni 0.762 0.001**
Scolelepis tridentata vs Scolelepis quinquedentata 0.999 0.006**
Scolelepis tridentata vs Scolelepis bellani n. sp. 0.823 0.001**
Scolelepis tridentata vs Scolelepis cantabra 0.975 0.001**
Scolelepis papillosa vs Scolelepis korsuni 0.733 0.001**
Scolelepis papillosa vs Scolelepis quinquedentata 1 0.048*
Scolelepis papillosa vs Scolelepis bellani n. sp. 0.822 0.001**
Scolelepis papillosa vs Scolelepis cantabra 0.963 0.002**
Scolelepis korsuni vs Scolelepis quinquedentata 0.864 0.028*
Scolelepis korsuni vs Scolelepis bellani n. sp. 0.349 0.003**
Scolelepis korsuni vs Scolelepis cantabra 0.921 0.001**
Scolelepis quinquedentata vs Scolelepis bellani n. sp. 0.562 0.011*
Scolelepis quinquedentata vs Scolelepis cantabra 0.865 0.036*
Scolelepis bellani n. sp. vs Scolelepis cantabra 0.933 0.001**

*Significant at p < 0.05; **significant at p < 0.01.

of neurochaetae is present (rather than absent as in
S. bellani n. sp.). The main fang of hooks is thinner,
with a sharply pointed tip in S. tridentata (Figure 4B–D),
while in S. bellani n. sp. it is shorter and has a blunt tip
(Figure 7D–F). Unfortunately, this latter feature can
only be seen under SEM.
Differences between S. bellani n. sp. and
S. papillosa are, in general, similar to those between
S. bellani n. sp. and S. tridentata. However, in
S. papillosa the ratio of the length of the hood to
the height of the main fang is closer to that of
S. bellani n. sp. (3–4.6 times in the former vs. 4–5
times in the latter).
S. quinquedentata can be separated from S. bellani
n. sp. by its hooks having very long hoods, ca. 5–6
times the length of the main fang (see Taxonomic
discussion for S. tridentata). Also, all branchae in
S. quinquedentata are distally free from the notopo­
dial postchaetal lamellae.
S. bellani n. sp. is very similar morphologically to
S. korsuni (Figure 9). Nevertheless, the two species
differ in the following characters: (1) eyes are usually
absent in S. korsuni (eyes are usually present in
S. bellani n. sp.); (2) neuropodial hooded hooks
have the median tooth situated below the lateral
ones in S. korsuni (the median unpaired tooth is
situated on the top and the two side-by-side below
in S. bellani n. sp.); (3) neuropodial hooks begin on
chaetigers 14–18 in S. korsuni (hooks begin on chae­
tigers 15–28 in S. bellani n. sp.); and (4) branchial
separation starts on chaetigers 12–19 in S. korsuni
(versus chaetigers 15–29 in S. bellani n. sp.).
However, small individuals of S. bellani n. sp.
could not be differentiated from S. korsuni
(Figure 10A, B). There are also differences in the
ecology of the two species. S. korsuni lives in full
strength marine waters at 95–450 m depth and tem­
peratures below 7°C, while S. bellani n. sp. lives in
brackish waters at 0–30 m depth, where tempera­
tures may raise to more than 22°C.
In the Black Sea, this species was also confused
with Scolelepis cantabra. Fauvel (1927) even consid­
ered Nerinides tridentata as juveniles of Nerinides can­
tabra. Like S. bellani n. sp., S. cantabra also has an
occipital tentacle, chaetiger 1 with only neurochae­
tae, and mid-body branchiae with a flag-like end.
Differences between S. bellani n. sp. and
S. cantabra consist in the shape of hooks, the num­
ber of apical teeth on the hooks, and the shape of the
prostomium.
Occurrence and distribution
Black Sea: Sevastopol – Uchkuyevka (Kisseleva
1981, as Nerinides tridentata; Boltachova et al.
2006, as Pseudomalacoceros tridentata); southern
Crimea (Kisseleva & Slavina 1963, as Nerinides tri­
dentata) – Lis’ya Bay (Kisseleva 1992, as Nerinides
tridentata), Karadag region (Vinogradov 1931; 1949,
as Nerinides tridentata), Caucasian coast (Kisseleva &
Slavina 1966, as Nerinides tridentata), Dzharylhach
Bay and the brackish water lagoons Sasyk, Shagani,
Alibey, Burnas, and Shabolat (Vinogradov &
Losovskaya 1963, as Nerinides tridentata),
Romanian coast (Dumitrescu 1957; 1963; Băcescu
et al. 1965, as Nerinides cantabra, in part), Bulgarian
coast – Varna Bay (Caspers 1951; Marinov 1957, as
Nerinides tridentata; Vorobyova & Bondarenko 2009,
as Pseudomalacoceros tridentata); Sea of Azov
(Kisseleva 1987, as Nerinides tridentata; Syomin
600 V. Surugiu

Figure 10. Scatter plots of relationships between: A. The body width (W) and the number of the first chaetiger bearing neuropodial
hooded hooks (VH). B. The body width (W) and the number of the first chaetiger on which notopodial postchaetal lamella splits from
branchia (Br) (red diamonds – Scolelepis bellani n. sp.; blue downward-pointing triangles – Scolelepis korsuni). Holotypes represented by
hollow symbols.

2011, as Pseudomalacoceros cantabra); and
Tyrrhenian Sea – Gulf of Gaeta (present study).
Most probably reports of S. tridentata from the Sea
of Marmara (Gillet & Ünsal 2000; Çinar et al. 2011)
belong to this species.
Concluding remarks
The Black Sea is characterized by a very low species
richness because of the reduced surface water salinity
(on average 17–18 psu) and the presence of hydrogen
sulphide in deep waters (about 87% of its volume is
anoxic). The number of species living in the Black Sea
is estimated to represent only 20–25% of those known
in the Mediterranean Sea (see references in Marin &
Antokhina 2022). Most of the Black Sea species, how­
ever, have immigrated from the neighbouring
Mediterranean Sea during the last 7–9 thousands of
years, when the last connection between these seas has
been established through the Bosphorus and the
Dardanelles straits. Therefore, most of the Black Sea
species are of Mediterranean origin (Surugiu 2008).
Despite being one of the most studied seas, recent
descriptions of new species in the Black Sea (Surugiu
& San Martín 2017; Marin & Antokhina 2022;
Mureșan et al. 2022; Agannemone & Micali 2023;
this study) indicate a yet uncovered cryptic diversity.
With the advent of molecular techniques, the rate of the
descriptions of new species in the Black Sea is pre­
sumed to increase rapidly. However, it is very likely
that species originally described from the Black Sea
are also present in the Mediterranean Sea as in the
Black Sea there are virtually no endemic species
(Surugiu 2008).
Scolelepis cantabra is an enigmatic species.
Though this study confirms its presence in the

Black Sea, it is still unclear whether the species is
still present in the Black Sea or not, because there
are no other reports since 1960s. Scolelepis cantabra
seems to be is also very rare in other European seas,
the only most recent confirmed findings being those
from the coasts of Portugal. Its large size and occur­
rence in shallow subtidal might be conducted to its
overcollection in some areas by recreational fisher­
men to be used as bait (Tarik Meziane, pers.
comm.). Accordingly, its status should be evaluated
through the IUCN Red List categories and criteria
(IUCN 2012).
Acknowledgements
I am much indebted to Nancy Maciolek, Danny Eibye-
Jacobsen and to two anonymous reviewers for critical
appraisal of the earlier version of the manuscript and
whose valuable comments and suggestions greatly
improved the quality of the paper. I would like to
thank Melanya Stan (MGAB), Dieter Fiege (SMF),
Adrian Glover (NHMUK), Emma Sherlock
(NHMUK), Nigel T. Monaghan (NMINH), Paolo
Viscardi (NMINH), Amy Geraghty (NMINH), and
Danny Eibye-Jacobsen (NHMD) for arranging the
loans of the material used in this study. Elena
Lisitskaya and Natalia Boltachova (Kovalevsky
Institute of Marine Biological Research RAS,
Sevastopol, Crimea), Vitaly Syomin (Shirshov
Institute of Oceanology of Russian Academy of
Sciences, Gelendjik, Russia), Olena Bondarenko and
Mikhail Son (Institute of Marine Biology of the NAS of
Ukraine, Odessa, Ukraine), and Andrey Sikorski
(Akvaplan-niva AS, Tromsø, Norway) are kindly
acknowledged for providing specimens for the present
study. I thank Geon Hyeok Lee (Inha University,
Korea) for providing measurements of specimens of
Scolelepis papillosa from Korea. I am also very grateful
to Ascensão Ravara (DBUA), Martin Schwentner
(ZMH), and Tarik Meziane (MNHN) for facilitating
the study of their museum’s polychaete collection.
Many thanks to Irina Gostin, Maria-Magdalena Fusu
and Ștefan Mihăiță Olaru for their assistance with SEM
and LM studies. This research was partially supported
by the German Academic Exchange Service personal
research fellowship (DAAD No. 57378441) and by the
European Commission SYNTHESYS+ Transnational
Access grant (FR-TAF_Call3_042). Financial support
was also provided by the Romanian Ministry of
Research, Innovation and Digitization, within
Program 1 – Development of the national RD system,
Subprogram 1.2 – Institutional Performance – RDI
excellence funding projects, Contract no. 11PFE/
30.12.2021.
A new species of Scolelepis from the Black Sea 601
Funding
This work was supported by the Deutscher
Akademischer Austauschdienst [57378441];
European Commission [FR-TAF_Call3_042].
Disclosure statement
No potential conflict of interest was reported by the
author(s).
Supplementary material
Supplemental data for this article can be accessed
online at https://doi.org/10.1080/24750263.2023.
2229855.
ORCID
Victor Surugiu http://orcid.org/0000-0003-4933-7678
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