ISSN 1867-1616, Volume 40, Number 3
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Mar Biodiv (2010) 40:161–180
DOI 10.1007/s12526-010-0040-5
Author's personal copy
ORIGINAL PAPER
A contribution to the taxonomy of Spio (Spionidae,
Polychaeta, Annelida) occurring in the North and Baltic
Seas, with a key to species recorded in this area
Andreas Bick & Katrin Otte & Karin Meißner
Received: 28 October 2009 / Revised: 12 February 2010 / Accepted: 23 February 2010 / Published online: 9 April 2010
# Senckenberg, Gesellschaft für Naturforschung and Springer 2010
Abstract In order to resolve taxonomic problems known
from literature and diagnostic practices, Spio species
currently recognised in the North Sea and the Baltic Sea
have been re-examined. Spio species recorded in the area of
interest are S. decoratus, Spio cf. filicornis, S. goniocephala,
S. martinensis, and S. armata. Spio decoratus is restricted to
the North Sea, whereas S. armata occurs only in the Baltic
Sea. Revised descriptions with illustrations and notes on the
distributions, as well as remarks on size-dependent differences of characters, are presented for all species. Specimens
of Spio from the North and Baltic Seas, which are assigned
to S. filicornis based on current taxonomic literature, exhibit
significant morphological differences, and are here regarded
as two different species. A definite assignment of specimens
from the two morphological distinct groups to one of the
known Spio species was impossible based on the material
available. The significance of diagnostic characters used for
the identification of Spio spp. is discussed. An identification
key to Spio species found in the investigation area is
provided.
A. Bick (*) : K. Otte
Institut für Biowissenschaften,
Allgemeine und Spezielle Zoologie,
Universität Rostock,
Universitätsplatz 2,
18055 Rostock, Germany
e-mail: andreas.bick@uni-rostock.de
K. Meißner
Senckenberg Forschungsinstitute und Naturmuseen,
Deutsches Zentrum für Marine Biodiversitätsforschung,
Biozentrum Grindel, Martin-Luther-King-Platz 3,
20146 Hamburg, Germany
Keywords Spio . Taxonomy . Key to species .
Size-dependent characters . Northeast Atlantic Ocean .
Distribution
Introduction
Within the Spionidae, about 28 species of the genus Spio
Fabricius, 1785 are currently considered valid (Maciolek
1990; Blake 1996; Sikorski 2001). However, Spio species
show great morphological similarity, and usually only a few
characters have been used to separate them (Maciolek
1990). Such characters include the shape of the anterior
margin of the prostomium, the length of the branchiae on the
first chaetiger, the first appearance as well as the number of
neuropodial hooded hooks, and the number of apical teeth
on these hooks. Because many taxonomic character states
are shared by different species, unambiguous identification
of Spio species is often very difficult and several authors
have indicated the need for a generic revision (Holmquist
1967; Foster 1971; Blake and Kudenov 1978; Maciolek
1990; Worsaae 1999).
The present paper deals with Spio species from the North
and Baltic Seas. According to Bergfeld and Kröncke (2003)
six species occur in this region: Spio filicornis (Müller,
1776), S. goniocephala Thulin, 1957, S. armata (Thulin,
1957), S. decoratus Bobretzky, 1870, S. martinensis Mesnil,
1896, and S. multioculata Rioja, 1918. Several taxonomic
problems have been discussed in relation to these species.
For example, both S. martinensis and S. goniocephala
have been considered junior synonyms of S. filicornis
[Söderström 1920 and Hartmann-Schröder (cited in: Bick
and Gosselck 1985)]. Spio armata was considered as
incertae sedis by Maciolek (1990) and later synonymised
with S. goniocephala by Hartmann-Schröder (1996).
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Böggemann (1997) doubted records of S. decoratus from
northern waters. Spio multioculata has been recorded only
once in the investigation area (as larvae from the Gullmar
Fjord, Kattegat, Sweden) by Hannerz (1956), but he
himself had doubts about his findings.
The study presents revised species descriptions of Spio
found in the North and Baltic Seas including new illustrations and notes on their distribution. Moreover, we document
that certain morphological characters are size-dependent, and
their usefulness in diagnoses is discussed. An identification
key to all Spio known from the investigation area is
provided, and a major step towards a revision of the genus
is taken.
Mar Biodiv (2010) 40:161–180
Zoologische Sammlung der Universität Rostock, Germany
(ZSRO). Type material was borrowed from Zoologiska
Museet, Lunds Universitet, Sweden (L).
Results
Spionidae, Grube, 1850
Genus Spio Fabricius, 1785
Type species: Nereis filicornis Müller, 1776
Type locality: Iluilârssuk, near Frederikshaab (Paamiut),
Westgreenland
Synonym: Paraspio Czerniavsky, 1881; type species: Spio
decoratus Bobretzky, 1870, by monotypy. Euspio McIntosh,
1915; type species: Euspio mesnili McIntosh, 1915.
Material and methods
The material examined in this study comes predominately
from several regions in the North Sea and in the Baltic Sea.
Also, Spio specimens from West Greenland were examined.
In addition, the types of S. armata and S. goniocephala
were studied.
Polychaetes were investigated using light and scanning
electron microscopy (SEM). Shirlastain A was applied to
improve the visualisation of some characters, e.g. tiny or
papillate branchiae on chaetiger 1, and the length and shape
of nuchal and metameric dorsal ciliated organs. Measurements of width refer to the distance between the distal-most
structures on the widest chaetiger seen in dorsal view
(without parapodia and chaetae). Drawings were made
using a camera lucida attached to a light microscope; the
anterior ends were drawn without camera lucida. Since the
capillaries of the species under consideration are similar,
capillaries were illustrated only for S. goniocephala.
Because of taxonomic problems mentioned in the
introduction, information on distribution given in this paper
is only based on material investigated here. For the same
reason, it was abstained from the presentation of synonymy
lists for the two species listed as Spio cf. filicornis (either
from the North Sea or from the Baltic Sea) in this paper.
Additional information included in the diagnoses from the
literature is indicated as such.
For SEM studies specimens were dehydrated in a graded
ethanol series, critical-point dried, attached to aluminium
stubs, sputter coated with gold palladium and examined
with a Zeiss DSM 960 A scanning electron microscope.
The material examined was borrowed from the following
museums and institutions: Landesamt für Natur und Umwelt
des Landes Schleswig-Holstein, Flintbek, Germany
(LANU); MariLim, Kiel, Germany (ML); Institut für
Angewandte Ökologie, Broderstorf, Germany (SBRO);
Senckenberg Museum Frankfurt, Germany (SMF); Zoologisk
Museum, Københavns Universitet, Denmark (ZMUC);
Description Prostomium anteriorly rounded or slightly
incised, lacking frontal or lateral horns; eyes present or
absent; digitiform occipital tentacle absent, but posterior
portion of prostomium may be raised or inflated. Paired
recurved nuchal organs extending onto chaetiger 2 or 3.
Metameric dorsal ciliated organs present. Branchiae
present from chaetiger 1, continuing throughout body,
branchiae completely separate from or basally fused with
notopodial lamellae, branchiae often reduced in size on
chaetiger 1. Notochaetae all capillaries; neurochaetae
capillaries anteriorly, capillaries, hooded hooks and
inferior sabre chaetae on middle and posterior chaetigers.
Pygidium with four anal cirri.
Spio armata (Thulin, 1957)
Figures 1, 9f, 11, 12
Microspio armata Thulin, 1971: 57–61, Fig. 4.
Spio armata—Hartmann-Schröder 1971: 302–304,
Fig. 102.—Sikorski 2001: 319–321, Figs. 1–11.
Type material. Holotype: Øresund, between Landskrona
and Ven, Sweden, several middle fragments (L 916/3745).
Non-type material Baltic Sea: 54°21.99′N, 11°15.63′E, 26
May 1997, sand, 17 m, two specimens (ZSRO-P 390); 54°
34.32′N, 11°20.14′E, 25 June 1997, sand/mud, 27 m, two
specimens (ZSRO-P 483); 54°35.191′N, 10°57.231′E, 20
August 2004, coarse sand, 17 m, one specimen (ZSRO-P
1667); 54°36.860′N, 11°00.753′E, 21 August 2004, one
specimen.
Description Only fragments available; maximum number
of chaetigers 59 (width 0.53 mm), maximum width
0.63 mm (anterior fragment with 42 chaetigers).
Prostomium anteriorly bluntly rounded or slightly
convex, anterior part distinctly extended; posterior end
short, extending to chaetiger 1, tapered (Fig. 1a); two pairs
of black eyes, arranged trapezoidally, anterior pair crescent-
Mar Biodiv (2010) 40:161–180
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Fig. 1 Spio armata (Thulin, 1957). a Anterior end, dorsal view
(pigment typical for the species (see diagnosis) partially lacking in
illustrated specimen); b ventral sabre chaeta from chaetiger 34; c
neuropodial hooded hook from chaetiger 34; d chaetiger 1 anterior
view; e chaetiger 24, anterior view; f chaetiger 34, anterior view; g
chaetiger 48, anterior view. (Scale bar: a 125 µm; b, c 64 µm; d
320 µm; e–g 640 µm)
shaped or rounded, posterior pair oval; prostomium
distinctly separated from peristomium by a furrow (Fig. 1a).
Nuchal organs with short median and long lateral ciliary
bands, lateral ciliary bands posteriorly curved and go nearly
completely back to the transverse ciliary band of chaetiger
2 (Fig. 9f). Metameric dorsal ciliated organs double-paired,
present from between branchiae 3 and 4, posterior
extension not known (Fig. 9f).
Branchiae present from chaetiger 1 until almost to the
end of the body, only the last eight to ten chaetigers without
branchiae; branchiae on chaetiger 1 small, slightly longer or
as long as notopodial postchaetal lamellae; branchiae with
narrow base, tapered distally (Fig. 1a), becoming thinner
and shorter in posterior chaetigers; on anterior chaetigers,
branchiae fused basally with notopodial postchaetal lamellae,
increasingly separated from lamellae in subsequent chaetigers
(Fig. 1d–g); branchiae not touching mid-dorsally on anterior
chaetigers; length of branchiae decreasing posteriorly.
First notopodium shifted dorsally. Notopodial postchaetal
lamellae in anterior chaetigers oval, becoming shorter and
rounded in subsequent chaetigers, on posterior chaetigers
lamellae large, rounded (Fig. 1d–g). Neuropodial postchaetal
lamellae rounded in chaetiger 1 (Fig. 1d), oval in other
anterior chaetigers, becoming smaller and shorter in posterior
chaetigers (Fig. 1e–g).
Alcohol-preserved specimens with dark brown pigment
on anterior part of body: pigmented patches between
peristomium and first chaetiger; scattered pigment dorsolaterally before and/or behind transverse ciliary bands and
around metameric dorsal ciliated organs; longitudinal
pigmented stripes on peristomium indistinct; median pigmented patch laterally between parapodia and on edges of
notopodial postchaetal lamellae, close to branchiae.
Notopodial chaetae all limbate capillaries; capillaries
of anterior chaetigers arranged in two rows: capillaries
of anterior row short, broad, uniformly granulated (as
in S. goniocephala, Fig. 5f), capillaries of posterior row
longer, thinner, lacking granulations (as in S. goniocephala,
Fig. 5g); additional superior fascicle of very long and thin
capillaries without granulations; capillaries of posterior
chaetigers arranged in irregular rows, in last three to five
posterior-most chaetigers row not clearly discernable. Neuro-
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podia with rows of capillaries and hooded hooks as well as
an inferior fascicle of capillaries; capillaries of anterior
neuropodia arranged in two rows similar to notopodial
chaetae; from chaetiger 17 posterior row replaced by single
row of six to eight hooded hooks, accompanied by
alternating short, thin, alimbate non-granulated capillaries
(as in S. goniocephala, Fig. 5e); hooks bidentate, strongly
curved, slightly narrowed subdistally, with long hood and
conspicuous large main fang at nearly right angle to shaft
(Fig. 1c); inferior fascicles with long, thin, limbate capillaries,
without granulations, replaced from about chaetiger 16/17 by
two to four sabre chaetae, each uniformly granulated
(Fig. 1b).
Pygidium unknown.
Distribution and ecology Western Baltic Sea, Øresund
(from literature: North Sea, Norwegian Sea, Arctic Ocean);
sublittoral; clay, sand.
Remarks Spio armata is readily identified by the following
combination of characters: short branchiae on chaetiger 1,
strongly curved hooded hooks with a conspicuous large main
fang present in neuropodia from chaetiger 17, co-appearance
of ventral sabre chaetae and hooded hooks from chaetiger 17,
and rounded notopodial postchaetal lamellae in posterior
chaetigers, rather than elongate lamellae as found in other
species in the North and Baltic Seas.
In S. armata, the first appearance of hooded hooks might
vary with the size of the specimen. In Arctic specimens,
hooded hooks are reported to be present from chaetiger 8 to
20, i.e., from chaetiger 8 to 11 in small specimens and from
chaetiger 18 to 20 in large specimens (Sikorski 2001).
Thulin (1957) observed hooded hooks to start on chaetiger
14 and 16. For this study only six anterior fragments of
comparable length were available for investigation. Therefore
it was not possible to demonstrate a correlation between size
and the start of hooded hooks for the Baltic Sea specimens
(Fig. 11), although it might exist.
The type material of S. armata consists of some middle
fragments only, all without prostomium and pygidium.
These fragments were selected among S. filicornis specimens and assigned to S. armata by Anders Eliason after the
death of Thulin. The shape of the hooded hooks differs
considerably between these two species. Now these
fragments are actually labelled as holotype of S. armata
and are kept in the Museum of Zoology in Lund, Sweden.
Because S. armata was originally described from the
Baltic Sea and no other species in North and Baltic Seas
exhibits these hooks with long hood and conspicuous large
main fang, the type material and original description are
considered adequate to recognize the species. However,
Maciolek (1990) considered this species to be incertae sedis.
For this reason, additional specimens from the type-locality
Mar Biodiv (2010) 40:161–180
(Øresund) could be necessary to allow a doubtless determination of this species in the future.
Spio decoratus Bobretzky, 1870
Figures 2, 9b, 11, 12
Spio decoratus Bobretzky, 1870: 256–257, Fig. 74–77.—
Giordanella 1969: 347, Fig. 1.—Guerin 1972: 330–337,
Figs. 1, 2 and 3.—Dauvin 1989: 169–170, Fig. 1.—
Giangrande 1992: 87–89, Figs. 2, 3 and 4.—Böggemann
1997: 120, Fig. 94.
Spio filicornis—Fauvel 1927: 43–44, Fig. 15A–G (partim).
Non-type material North Sea: 54°43.38′N, 8°03.50′E, 12
August 1997, sand, 13 m, ten specimens (ZSRO-P 442);
54°25.86′N, 8°10.13′E, 11 August 1997, sand, 18 m, six
specimens (ZSRO-P 484); 60°36.99′N, 2°46.07′E, 6 May
1998, mud, 110 m, one specimen (ZSRO-P 720); 54°43.93′
N, 8°03.48′E, 18 August 1998, sand, 13 m, three specimens
(ZSRO-P 905); 55°17.665′N, 6°47.913′E, 13/16 May 2000,
sand, 30 m, one specimen (ZSRO-P 1555); 55°08.403′N,
6°40.302′E, May 2002, muddy sand, 39 m, four specimens
(SBRO-P 67); 54°02.207′N, 6°55.016′E, March 2002, fine
sand, 32 m, two specimens (SBRO-P 225); 54°01.709′N,
6°50.454′E, May 2003, muddy sand, 30.5 m, four specimens
(SBRO-P 1292).
Description Specimens with up to 50 chaetigers (width
0.5 mm), maximum width 0.63 mm (anterior fragment with
30 chaetigers).
Prostomium anteriorly bluntly rounded, anterior part
slightly to distinctly extended; posteriorly extending to
chaetiger 1, tapered (Fig. 2a); two pairs of black eyes,
arranged in trapezoid or rectangle, anterior pair crescentshaped or rounded, posterior pair oval; prostomium
distinctly separated from peristomium by a furrow (Fig. 2a).
Nuchal organs with short median and long lateral ciliary
bands, lateral ciliary bands posteriorly curved and go only
partly back to the transverse ciliary band of chaetiger 2
(Fig. 9b). Metameric dorsal ciliated organs double-paired,
present from between branchiae 5 and 6 (Fig. 9b); middle
chaetigers with additional short transverse ciliary band
between the long transverse ciliary bands connecting the
branchiae (Fig. 9b).
Branchiae from chaetiger 1, continuing to near end of
body; branchiae on chaetiger 1 long, almost as long as
those on following chaetigers; branchiae with broad base,
tapered distally (Fig. 2a), becoming thinner and shorter
with narrow base in posterior chaetigers; branchiae on
anterior chaetigers fused basally with notopodial postchaetal
lamellae, separated from lamellae in posterior chaetigers
(Fig. 2d–g); branchiae not reaching dorsal midline on
anterior chaetigers, length of branchiae decreasing in
subsequent chaetigers.
Mar Biodiv (2010) 40:161–180
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165
Fig. 2 Spio decoratus Bobretzky, 1870. a Anterior end, dorsal view;
b neuropodial hooded hooks from chaetiger 34; c ventral sabre
chaeta from chaetiger 34; d chaetiger 11, anterior view; e chaetiger
24, anterior view; f chaetiger 40, anterior view; g chaetiger 1,
anterior view; h pygidium, dorsal view. (Scale bar: a 0.125 mm; b, c
64 µm; d-f 640 μm; g 320 μm; h 160 µm)
First notopodium shifted dorsally. Notopodial postchaetal lamella slightly pointed on chaetiger 1, oval in other
anterior chaetigers, becoming shorter and rounded in
subsequent chaetigers, elongated in posterior chaetigers
(Fig. 2d–g). Neuropodial postchaetal lamellae rounded in
chaetigers 1 and 2, oval in other anterior chaetigers,
becoming smaller and shorter in middle chaetigers, rounded
in posterior chaetigers (Fig. 2d–g).
Alcohol-preserved specimens with dark-brown pigment
on anterior part of body: prostomium and peristomium of
juveniles completely pigmented, adults dorsally with broad
transverse pigmented stripe on prostomium and peristomium, longitudinal pigmented stripes between prostomium
and peristomium stable in alcohol and formalin-preserved
specimens; scattered pigment dorso-laterally on anterior
chaetigers before and/or behind transverse ciliary bands and
at the end of nuchal organs (Fig. 2a); scattered pigment
ventrally on peristomium; short median pigmented stripe
ventrally between anterior chaetigers; pigmented patches
ventro-laterally on base of neuropodia; a median pigmented
patch between parapodia of subsequent chaetigers; isolated
pigmented stripes laterally before and/or behind noto- and
neuropodia; one to two pigmented patches on branchiae
near base.
Notopodial chaetae limbate capillaries with those of anterior
chaetigers arranged in two rows: capillaries of anterior row
short, broad, uniformly granulated (as in S. goniocephala,
Fig. 5f); chaetae of posterior row longer, thinner, lacking
granulations (as in S. goniocephala, Fig. 5g); additional
superior fascicle of long, thin capillaries without granulations;
capillaries of posterior notopodia arranged in irregular rows,
thin, non-granulated, of different lengths. Neuropodia with
rows of capillaries and hooded hooks as well as an inferior
fascicle of capillaries; capillaries of anterior chaetigers
arranged in two rows, similar to notopodial chaetae; posterior
row replaced by a single row of nine to 11 hooded hooks from
chaetiger 11 (Fig. 11), accompanied by alternating short, thin,
alimbate, non-granulated capillaries (as in S. goniocephala,
Fig. 5e); hooks tridentate, with long hood, slightly
narrowed subdistally; third tooth distinct, rarely surmounted by fourth apical tooth (Fig. 2b); inferior fascicle
with one to three long, thin, limbate capillaries, without
granulations, replaced from about chaetiger 20 by two to
three sabre chaetae, each uniformly granulated (Fig. 2c).
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Pygidium with four anal cirri; dorsal pair longer than
ventral pair or of same length; dorsal pair slender, cirriform,
widely spaced; ventral pair broad and oval, closely spaced,
touching each other at base (Fig. 2h).
Distribution and ecology North Sea, [from literature: Black
Sea, Mediterranean Sea, Northeast Atlantic, Arctic Ocean
(?)]; sublittoral; coarse and fine sand.
Remarks Spio decoratus differs from other Spio species in
the North and Baltic Seas by its comparatively small size.
Furthermore, it possesses a characteristic broad transverse
pigmented band on the prostomium and peristomium and
tridentate hooded hooks with a clearly discernible third
tooth.
Juvenile specimens of Spio cf. filicornis (North Sea), in
particular, are morphologically very similar to S. decoratus.
Both species are conspicuously pigmented and possess
tridentate hooded hooks from chaetiger 11 (Fig. 11). A
character that can be used to separate these species is the
furrow between the prostomium and peristomium which is
present in S. decoratus and absent in Spio cf. filicornis
(North Sea). Spio decoratus also differs from Spio cf.
filicornis (North Sea) in having a broad transverse
pigmented band on prostomium and peristomium rather
than randomly distributed pigmented patches, metameric
dorsal ciliated organs beginning between branchiae 5 and 6
rather than branchiae 4 and 5, and the lateral ciliary bands
of the nuchal organs go only partly back to the transverse
ciliary band of chaetiger 2 and not completely as in S. cf.
filicornis (North Sea). In S. decoratus the third apical tooth
of the hooded hooks is clearly visible, but is hardly
discernable in Spio cf. filicornis (North Sea).
There is still uncertainty whether the specimens from the
North Sea actually are S. decoratus sensu Bobretzky 1870
since neither type material nor material from the type
locality (Black Sea) was available. The morphological
characteristics of the specimens from the North Sea are in
good agreement with the descriptions of S. decoratus from
the English Channel (Dauvin 1989). However, S. decoratus
from the Black Sea possesses bidentate hooded hooks, and
the transverse pigmented band on the peristomium and
peristomium is absent (Bobretzky 1870). Later Giordanella
(1969) examined some material from the Black Sea and
described the hooks as tridentate, modifying Bobretzky’s
description in this aspect.
Spio cf. filicornis (North Sea)
Figures 3, 9d, 10a–d, 11, 12
Non-type material North Sea: 55°17.665′N, 6°47.913′E, 15
October 2003, sand, 28 m, 14 specimens (ZSRO-P 1554);
54°59.70′N, 7°06.40′E, 15 October 2003, sand, three
Mar Biodiv (2010) 40:161–180
specimens (ZSRO-P 1861); 55°08.403′N, 6°40.302′E,
May 2002, fine sand, 30 m, four specimens (SBRO-P 63);
55°08.403′N, 6°40.302′E, September 2002, sand, 37 m,
four specimens (SBRO-P 64); 55°13.060′N, 6°51.297′E,
September 2002, sand, 29 m, four specimens (SBRO-P 65);
54°35.571′N, 6°21.456′E, Autumn 2002, muddy sand,
40 m, two specimens (SBRO-P 445); 54°03.686′N, 7°
15.390′E, August 2003, muddy sand, 25.5 m, three specimens (SBRO-P 845); 54°02.207′N, 6°55.016′E, September
2003, muddy sand, 33 m, three specimens (SBRO-P 1313).
Description Only fragments available, maximum number
of chaetigers 58 (width 0.75 mm), maximum width 1.4 mm
(anterior fragment with 45 chaetigers).
Prostomium anteriorly rounded, antero-lateral part
slightly expanded; posterior part of prostomium pointed,
extending posteriorly to chaetiger 1 (Fig. 3a); two pairs of
black eyes present, with anterior pair crescent-shaped and
widely spaced, posterior pair oval, closely spaced; prostomium and peristomium not distinctly separated by a furrow
(Fig. 3a).
Nuchal organs with short median and long lateral ciliary
bands, lateral ciliary bands outward curved in the range of
the first transverse ciliary band, posteriorly curved and go
nearly completely back to the transverse ciliary band of
chaetiger 2 (Figs. 9d, 10a). Metameric dorsal ciliated
organs double-paired, present from between branchiae 4
and 5 continuing to branchiae 32 and 33 (Figs. 9d, 10c–d);
middle chaetigers with short transverse ciliary band
between metameric dorsal ciliated organs and long transverse
ciliary bands connecting branchiae (Figs. 9b, 10b).
Branchiae present from chaetiger 1, continuing nearly to
end of body; branchiae on chaetiger 1 long, nearly as long
as those on following chaetigers; branchiae with narrow
base, tapered distally (Fig. 3a), becoming thinner and
shorter in posterior chaetigers; branchiae on anterior
chaetigers fused basally with notopodial postchaetal
lamellae; branchiae separated from lamellae in posterior
chaetigers (Fig. 3b–e); branchiae not reaching dorsal
midline, length of branchiae decreasing posteriorly.
First notopodium shifted dorsally. Notopodial postchaetal
lamella on chaetiger 1 elongate, narrow and slightly pointed
apically (Fig. 3b), in other anterior chaetigers short and wide
(Fig. 3c), becoming smaller and rounded in middle (Fig. 3d)
and again long and narrow in posterior chaetigers (Fig. 3e).
Neuropodial postchaetal lamellae elongate and rounded on
chaetigers 1 and 2 (Fig. 3b), short and wide in other anterior
and middle chaetigers (Fig. 3c, d) and rounded in posterior
chaetigers (Fig. 3e).
Alcohol-preserved specimens with dark-brown pigment
on anterior part of body: pigmented stripes and patches on
dorsal part of prostomium and peristomium; anterior
chaetigers completely pigmented dorsally, pigment absent
Mar Biodiv (2010) 40:161–180
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167
Fig. 3 Spio cf. filicornis (Müller, 1776) from the North Sea. a Anterior
end, dorsal view; b chaetiger 1, anterior view; c chaetiger 11,
anterior view; d chaetiger 34, anterior view; e chaetiger 56, anterior
view; f neuropodial hooded hooks from chaetiger 34; g pygidium,
dorsal view. (Scale bar: a 125 µm; b 320 µm; c–e 640 µm; f 64 µm;
g 320 µm)
only where nuchal organs, metameric dorsal ciliated organs
and transverse ciliary bands occur; pigmented patches
present dorso-laterally anterior to and/or posterior to
transverse ciliary bands and at end of nuchal organs
(Fig. 3a); indistinct longitudinal pigmented stripes present
on ventral side of peristomium; pigment bordering anterior
chaetigers ventrally, with short median disrupted pigmented
stripe, not fading in alcohol; median pigmented patch
between neuro- and notopodia present; pigmented stripes
present laterally before and/or behind notopodia; base of
branchiae with one to two short longitudinal pigmented
stripes, also visible in alcohol preserved specimens
(Fig. 3a); pigmented patches on palps and large pigmented
patch on base.
Notopodial chaetae all limbate capillaries; notochaetae of
anterior chaetigers arranged in two rows: chaetae of anterior
row short, broad, uniformly granulated (as in S. goniocephala,
Fig. 5f); chaetae of posterior row longer, thinner, lacking
granulations (as in S. goniocephala, Fig. 5g); additional
superior fascicle of long, thin capillaries without granulations
present; chaetae of posterior chaetigers arranged in irregular
rows, with capillaries thin, non-granulated, with variable
length. Neuropodia with rows of capillaries and hooded
hooks as well as an inferior fascicle of capillaries; capillaries
of anterior neuropodia arranged in two rows, similar to
notochaetae; posterior row replaced by single row of 10–12
hooded hooks from chaetiger 11 (rarely from chaetiger 10,
Fig. 11), accompanied by alternating short, thin, alimbate,
non-granulated capillaries (as in S. goniocephala, Fig. 5e);
hooks with long hood, slightly narrowed subdistally, usually
bidentate, rarely (and only in juveniles) tridentate, third tooth
barely visible (Fig. 3f); inferior fascicle with two to four
long, thin, limbate capillaries without granulations, replaced
by two to three sabre chaetae from about chaetiger 20–24,
each uniformly granulated (Fig. 5d).
Pygidium with four anal cirri of similar length; dorsal
pair slender, cirriform, widely spaced; ventral pair broad,
and cone-shaped, closely spaced (Fig. 3g).
Distribution and ecology North Sea; sublitoral; fine sand,
coarse sand, gravel.
Remarks We have abstained from naming the species, as
definite assignment was impossible on the basis of the
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available material. Based on previously used diagnostic
characters, the species could have been identified as Spio
filicornis (Müller, 1776). But the same applies to specimens
from the Baltic Sea in this paper referred to as Spio cf.
filicornis (Baltic Sea). However, specimens of Spio cf.
filicornis from the North and Baltic Seas exhibit significant
morphological differences and we regard them as two
different species. Hence, it has to be decided whether one
of the two species indeed represents Spio filicornis. The
type material of S. filicornis no longer exists [see also
Remarks on Spio cf. filicornis (Baltic Sea)]. An assignment
of Spio cf. filicornis (North Sea) to a valid species or a
description of a new species will not be possible until
specimens from the locus typicus of S. filicornis near
Paamiut, Greenland (see Worsaae 1999) are obtained and a
neotype is designated. At this point, it should be mentioned
that there is a mix-up in regard to the type-locality of S.
filicornis in the literature. The type locality of S. filicornis
was stated to be Denmark instead of West Greenland by
Hartman (1959) and Maciolek (1990). Though Greenland
in former times was part of Denmark and even today still is
an autonomous country within the Kingdom of Denmark
the type locality should certainly rather be stated to be
Greenland. Spio filicornis was described by Otto Friderich
Müller (1776) in his Zoologiae Danicae Prodomus. After a
very short description, Müller (1776) noted that he had
received the specimens from Greenland by Otto Fabricius.
Fabricius was a missionary in the Frederikshaab colony in
Greenland from 1768 to 1773, where he collected various
animals. These animals have been described in more detail
in his comprehensive work Fauna Groenlandica (1780).
First illustrations of this species, and some remarks about
the habitat in Greenland and the biology were given also by
Fabricius (1785).
Spio cf. filicornis (North Sea) differs from other Spio
species in the North and Baltic Seas by the characteristic
fusion of prostomium and peristomium. Whereas in
juveniles three conspicuous teeth on hooded hooks are
present, the third tooth is minute or completely absent in
adults. Branchiae on chaetiger 1 are nearly as large as those
on subsequent chaetigers in adults, but they are small and
barely as large as the notopodial postchaetal lamella in
juveniles. A comparable delayed development of the first
branchiae was described for Spio setosa Verrill, 1873:
newly settled juveniles (<30 chaetigers) of this species lack
branchiae on chaetigers 1 and 2 (Simon 1967, 1968).
Juveniles of Spio cf. filicornis (North Sea) are very similar
to S. decoratus (for differentiation see Remarks under S.
decoratus).
Morphological differences between specimens of Spio
cf. filicornis from the North and Baltic Seas are significant.
The prostomium and peristomium are fused and pigmented
patches on the peristomium are randomly distributed in
Mar Biodiv (2010) 40:161–180
North Sea specimens, whereas in specimens from the Baltic
Sea the prostomium and peristomium are distinctly separated
and the peristomium is dark-brown pigmented. Although
specimens from the North and Baltic Seas have large
branchiae on chaetiger 1 and hooded hooks first appear on
chaetiger 11, there are differences in the shape of the
branchiae on the anterior chaetigers and hooded hooks. In
North Sea specimens, the branchiae of anterior chaetigers are
large but do not touch dorsally whereas in Baltic Sea
specimens they touch each other. Furthermore, the hooded
hooks of North Sea specimens are bi- (adults) or tridentate
(juveniles), slightly narrowed subdistally and exhibit a long
hood (Fig. 3f), whereas the hooks of Baltic Sea specimens
are always bidentate, not narrowed subdistally and possess a
short hood (Fig. 4d). Hence, Spio cf. filicornis (North Sea)
and Spio cf. filicornis (Baltic Sea) are here considered to be
two different species.
Spio cf. filicornis (Baltic Sea)
Figures 4, 9e, 11, 12
Non-type material Baltic Sea: 54°18.405′N, 11°12.030′E,
15 October 1997, sand, 16 m, four specimens (ZSRO-P 409);
54°20.10′N, 12°04.58′E, 28 August 1997, mud, 19 m, four
specimens (ZSRO-P 410); 54°10′N, 11°10′E, 29 July 1980,
22 m, three specimens (ZSRO-P 1363); 54°23.18′N, 11°
57.48′E, 27 October 1995, 17 m, one specimen (ZSRO-P
1748); 54°04.490′N, 11°24.350′E, September 2003, mud,
16 m, one specimen (SBRO-P 2557); 54°10.550′N, 11°
45.000′E, August 2002, mud, 17 m, two specimens (SBRO-P
2568); 54°10.100′N, 11°45.000′E, September 2004, coarse
sand, 5 m, one specimen (SBRO-P 3493); 54°28.800′N, 12°
15.800′E, September 2004, sand, 24 m, two specimens
(SBRO-P 3523); 54°10.550′N, 11°45.000′E, February 2004,
sand mud, 19.5 m, one specimen (SBRO-P 3541); 54°31.012′
N, 12°18.584′E, March 2004, sand, 22 m, one specimen
(SBRO-P 3542); 54°27.93′N, 12°17.750′E, February 2004,
fine sand, 18 m, one specimen (SBRO-P 3543).
Additional material Greenland (Nipisat, Disko Island): 69°
26.23′N, 54°13.70′W, sandy mud, 1.5 m, 13 specimens
(ZMUC-POL 1769–1777).
Description Specimens with up to 38 chaetigers (width
0.65 mm), maximum width 1.1 mm (anterior fragment with
22 chaetigers).
Prostomium elongated, bluntly rounded, anterior part
slightly expanded, often weakly incised; posterior part of
prostomium with high, narrow, keel-shaped elevation,
beginning at the level of the eyes and terminating on
chaetiger 1 (not shown in Fig. 4a); three pairs of black eyes
arranged trapeziformly, anterior pair crescent-shaped or
oval, widely spaced, middle pair oval or irregularly shaped,
Mar Biodiv (2010) 40:161–180
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169
Fig. 4 Spio cf. filicornis (Müller, 1776) from the Baltic Sea. a Anterior
end, dorsal view; b chaetiger 1, anterior view; c chaetiger 12, anterior
view; d neuropodial hooded hook from chaetiger 24; e ventral sabre
chaeta from chaetiger 24; f pygidium, ventral view. (Scale bar: a
125 µm; b, c 640 µm; d, e 64 µm; f 160 µm)
posterior pair oval or round, closely spaced; prostomium
distinctly separated from peristomium by furrow (Fig. 4a).
Nuchal organs with short median and long lateral ciliary
bands, lateral ciliary bands outward curved in the range of
the first transverse ciliary band, posteriorly curved and go
nearly completely back to the transverse ciliary band of
chaetiger 2 (Fig. 9e). Metameric dorsal ciliated organs
double-paired extending, anterior to posterior, from between
branchiae 3 and 4, and continuing on subsequent segments;
posterior extent not defined (Fig. 9e).
Branchiae from chaetiger 1, continuing to near to last
chaetiger; branchiae on first chaetiger nearly as long as
those on the following chaetigers; branchiae with narrow
base, tapering distally (Fig. 4a), becoming thinner and
shorter posteriorly; branchiae on anterior chaetigers basally
fused with notopodial postchaetal lamellae, separated from
lamellae in posterior chaetigers; branchiae reaching midline
dorsally and touching on anterior chaetigers.
First notopodium shifted dorsally. Notopodial postchaetal
lamellae on anterior chaetigers oval, becoming smaller and
rounded on subsequent chaetigers and appearing elongated
in posterior chaetigers. Neuropodial postchaetal lamellae in
anterior chaetigers oval, becoming smaller and shorter on
subsequent chaetigers, rounded in posterior chaetigers.
Alcohol-preserved specimens with dark-brown pigment
on anterior part of the body distributed as follows:
peristomium heavily pigmented (Fig. 4a); pigmented
patches dorsolaterally on anterior chaetigers before and/or
behind transverse ciliary bands, around nuchal organs and
metameric dorsal ciliated organs; indistinct longitudinal
pigmented stripes ventrally on peristomium; pigmented
border on anterior chaetigers ventrally; short median
disrupted pigmented stripe and one to two pigmented
patches ventro-laterally at base of neuropodia; median
pigmented patch and several pigmented stripes between
neuro- and notopodia; particularly conspicuous on the first
two chaetigers; base of branchiae with one to two short
longitudinal pigmented stripes (pattern of pigmentation
not depicted in Fig. 4a); two weak longitudinal pigmented
stripes on palps and a large pigmented patch on the base of
palps.
Notopodial chaetae all limbate capillaries; capillaries of
anterior chaetigers arranged in two rows: chaetae of anterior
row short, broad, uniformly granulated (as in S. gonioce-
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Mar Biodiv (2010) 40:161–180
Distribution and ecology Baltic Sea, North Atlantic: West
Greenland; sublitoral; mud, fine sand, coarse sand.
phala, Fig. 5f); chaetae of posterior row longer, thinner,
lacking granulations (as in S. goniocephala, Fig. 5g);
additionally, superior fascicle of very long, thin capillaries
without granulations; capillaries of posterior chaetigers
arranged in irregular rows, thin, non-granulated, of different
length. Neuropodia with rows of capillaries and hooded
hooks as well as an inferior fascicle of capillaries;
capillaries of anterior neuropodia arranged in two rows,
similar to notochaetae; posterior row replaced by a single
row of six to eight hooded hooks from chaetiger 11
(exceptionally from chaetiger 10, Fig. 11), accompanied
by alternating short, thin, alimbate, non-granulated capillaries (as in S. goniocephala, Fig. 5e); hooks not narrowed
subdistally, with a short hood, bidentate, main fang and
apical tooth well developed (Fig. 4d); inferior fascicle with
two to three long, thin, limbate capillaries without granulations, replaced by two sabre chaetae from about
chaetiger 20–24, each distally granulated (Fig. 4e).
Pygidium with four anal cirri; dorsal pair longer than
ventral pair or of same size; dorsal pair more widely spaced
than ventral pair (Fig. 4f).
Remarks Baltic Sea specimens of Spio cf. filicornis differ
significantly from other Spio species in North Sea and
Baltic Sea in having a heavily dark brown pigmented
peristomium. The slight medial incision on the anterior
margin of the prostomium, and the high, narrow, keelshaped elevation on the posterior part of the prostomium
are also useful diagnostic characters.
Spio cf. filicornis (Baltic Sea) is similar to S. armata.
Both species occur in the Baltic Sea and the adults are large
and robust (for differentiation see Remarks on S. armata).
Specimens of Spio cf. filicornis from the North and Baltic
Seas exhibit significant differences and we here regard them
to be different species [see Remarks on Spio cf. filicornis
(North Sea)]. Spio cf. filicornis (Baltic Sea) is similar to S.
filicornis from Øresund (Thulin 1957), the western North
Atlantic (Maciolek 1990) and West Greenland (Worsaae
1999). Our study of additional material from Greenland
(Nipisat, Disko Island) supports this conclusion. We have
Fig. 5 Spio goniocephala Thulin, 1957. a Anterior end of a North
Sea specimen, dorsal view; b anterior end of a Baltic Sea specimen,
dorsal view; c neuropodial hooded hook from chaetiger 48; d ventral
sabre chaeta from chaetiger 34; e short capillary from chaetiger 56; f
capillary from anterior row of chaetae in neuropodium of chaetiger 20;
g neuropodial capillary from posterior row of chaetae of chaetiger 20;
h pygidium, dorsal view. (Scale bar: a, b 125 µm; c–g 64 µm;
h 320 µm)
Mar Biodiv (2010) 40:161–180
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abstained from naming this species for the same reasons
already mentioned for Spio cf. filicornis (North Sea) (see
Remarks there).
Spio cf. filicornis (Baltic Sea) was usually only found in
small numbers (one to three specimens per sample) when
collected.
Spio goniocephala Thulin, 1957
Figures 5, 6, 8a, b, 9a, 11, 12
Spio goniocephala Thulin, 1957: 53–57, Figs. 2 and 3.—
Hartmann-Schröder 1971: 301–302, Fig. 101.—HartmannSchröder 1996: 339, Fig. 155.—Böggemann 1997: 120,
Fig. 96.—Sikorski 2001: 321, Figs. 1–5.
Spio filicornis—Eliason 1920: 40–42, Fig. 7 (partim).
Type material. Holotype: Øresund, south of Helsingør
(L 916/3701); Paratype: Øresund, south of Helsingør, three
specimens (L916/3702–3704).
Fig. 6 Spio goniocephala
Thulin, 1957. a Chaetiger 1,
anterior view; b chaetiger
2, anterior view; c chaetiger 20,
anterior view; d chaetiger 48,
anterior view; e chaetiger
56, anterior view; f chaetiger 62,
anterior view. (Scale bar:
a–f 320 µm)
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Non-type material North Sea: Weser estuary May 1997,
muddy sand, 19,5 m, two specimens (ZSRO-P 412); 55°
17.665′N, 6°47.913′E, 13/16 May 2002, sand, 28 m, seven
specimens (ZSRO-P 1556); 53°41.513′N, 6°28.806′E,
May 2003, fine sand, 20 m, one specimen (SBRO-P
501); 53°43.342′N, 6°28.683′E, May 2003, sand, 24 m,
four specimens (SBRO-P 502); 53°41.513′N, 6°28.806′E,
May 2003, fine sand, 20 m, ten specimens (SBRO-P 507);
53°43,.342′N, 6°28.683′E, May 2003, sand, 25 m, five
specimens (SBRO-P 508); 53°43.342′N, 6°28.683′E,
September 2002, sand, 22 m, 18 specimens (SBRO-P
572); 54°30.507′N, 7°51.701′E, November 2002, fine
sand, 20.5 m, one specimen (SBRO-P 688). Baltic Sea:
54°10.1′N, 11°21.4′E, 25 June 1981, 23 m, one specimen
(ZSRO-P 179); Mecklenburg Bight, October 1997, sand,
17 m, 29 specimens (ZSRO-P 411); 54°02.440′N, 11°
05.800′E, 20 m, ten specimens (ZSRO-P 1127).
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Description Specimens up to 77 (width 0.63 mm, North
Sea) and up to 53 chaetigers (width 0.6 mm, Baltic Sea),
maximum width 0.85 mm (North Sea, anterior fragment
with 52 chaetigers) and 0.75 mm (Baltic Sea, anterior
fragment with about 30 chaetigers).
Prostomium elongated, anteriorly tapered, anterior edges
rounded, antero-lateral part with rounded projections
(Fig. 8a); posterior part of prostomium rounded and not
conspicuously elevated, extending posteriorly to chaetiger 1
(Fig. 5a, b); two pairs of black eyes, arranged trapeziformly
or rectangular; anterior pair often irregularly shaped,
crescent-shaped or rounded, posterior pair oval; prostomium
not or little broadened in range of eyes; prostomium and
peristomium distinctly separated by furrow (Fig. 5a, b).
Nuchal organs with short median and long lateral
ciliary bands, lateral ciliary bands slightly outwardly
curved in the range of the first transverse ciliary band
(Figs. 8a, 9a). Paired metameric dorsal ciliated organs
slightly outward curved, from between branchiae 3 and 4
(Figs. 8b and 9a).
Branchiae present from chaetiger 1, continuing nearly to
last chaetiger; branchiae on chaetiger 1 small, about as half
as long and wide as those on the following chaetigers;
branchiae with broad base, tapered distally, becoming
thinner and shorter in subsequent chaetigers, with narrow
base in posterior chaetigers (Fig. 5a, b); branchiae on
anterior chaetigers fused basally with notopodial postchaetal lamellae, separated from lamellae in posterior
chaetigers (Fig. 6a–f); branchiae on anterior chaetigers
almost reaching dorsal midline.
First notopodium shifted dorsally. Notopodial postchaetal
lamellae on chaetiger 1 slightly pointed apically, oval in
other anterior chaetigers, becoming shorter and rounded in
middle chaetigers and then elongated in posterior chaetigers
(Fig. 6a–f). Neuropodial postchaetal lamellae rounded on
chaetigers 1 and 2, oval in other anterior chaetigers,
becoming smaller and shorter subsequently, rounded in
posterior chaetigers (Fig. 6a–f).
Alcohol-preserved specimens with dark-brown pigment
on anterior part of body: prostomium of juveniles completely pigmented dorsally, adults with a dorsal median
longitudinal pigmented stripe and with scattered pigment
near the anterior margin, around eyes and on posterior part
of prostomium (Fig. 5a, b); anterior chaetigers completely
pigmented dorsally, except for nuchal organs, metameric
dorsal ciliated organs and transverse ciliary bands; pigmented
patches dorsally on anterior chaetigers, stable in alcoholpreserved specimens (Fig. 5a, b); two indistinct longitudinal
pigmented stripes on palps; anal cirri also pigmented: ventral
pair more strongly pigmented than dorsal pair; dorsal anal
cirri with pigmented patch subdistally.
Notopodial chaetae all limbate capillaries; capillaries of
anterior chaetigers arranged in two rows: chaetae of
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anterior row short, broad, uniformly granulated (Fig. 5f);
chaetae of posterior row longer, thinner, lacking granulations (Fig. 5g); superior fascicle of long, thin capillaries
without granulations also present; capillaries of posterior
chaetigers arranged in irregular rows, thin, non-granulated,
of different lengths. Neuropodia with rows of capillaries
and hooded hooks as well as an inferior fascicle of
capillaries; capillaries of anterior neuropodia arranged in
two rows, similar to notochaete; from chaetiger 12 in small
specimens (North Sea and Baltic Sea) and from chaetiger 23
or 20 in large specimens (North Sea or Baltic Sea) one to
two hooded hooks present in posterior row, subsequently
capillaries replaced by a single row of seven to ten hooks
(Fig. 11), accompanied by alternating short, thin, alimbate,
non-granulated capillaries (Fig. 5e); hooks bidentate, uniformly curved, slightly narrowed subdistally, with long
hood, main fang and apical tooth weakly developed
(Fig. 5c); inferior fascicle with three to seven (North Sea)
or two to five (Baltic Sea) long, thin, limbate capillaries
without granulations, replaced by three to four sabre chaetae
from about chaetiger 20–22, each uniformly granulated
(Fig. 5d).
Pygidium with four anal cirri of about same size; dorsal
pair slender, cirriform and widely spaced; ventral pair
broadened oval, touching each other at base (Fig. 5h).
Distribution and ecology Baltic Sea, North Sea, [from
literature: Baffin Island (?)]; sublittoral; mud, fine sand,
coarse sand.
Remarks Spio goniocephala is characterised by the tapered
anterior part of its prostomium, the inconspicously elevated,
posterior part of the prostomium, and the median (not
dorsolateral) pigmented patches dorsally on anterior
chaetigers. Branchiae on chaetiger 1 are small, about half
as long and wide as those on the following chaetigers.
Furthermore, neuropodial capillaries are replaced first by
one to two hooded hooks and subsequently by a single
row of seven to ten hooded hooks.
As in S. armata, the first appearance of hooded hooks is
size-dependent (Fig. 11). Several authors have found
neuropodial hooded hooks present from chaetiger 16–21
(Hartmann-Schröder 1996; Thulin 1957) or 16–19 (Bick
and Gosselck 1985). This range has been expanded to
chaetiger 12–23, and it is likely that in very small
specimens hooded hooks may appear before chaetiger 12
and in larger specimens after chaetiger 23. In S. goniocephala
the branchiae on chaetiger 1 are generally small, but in small
specimens (0.25–0.38 mm width, max. 35 chaetigers)
branchiae are exceptionally diminutive or papilliform and
may be easily overlooked. A comparable delayed development of the first branchiae was also described for S. setosa
Verrill, 1873 (Simon 1967, 1968), and it is also assumed to
Mar Biodiv (2010) 40:161–180
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occur in Spio cf. filicornis (North Sea) [see Remarks on Spio
cf. filicornis (North Sea)].
The type material of S. goniocephala (Zoologiska
Museet, Lund University; L916/ 3701–3704) was examined.
Although the material was dried out, all diagnostic characters
could still be observed and are consistent with the material
from the North and Baltic Seas.
Spio martinensis Mesnil, 1896
Figures 7, 9c, 11, 12
Spio martinensis Mesnil, 1896: 122–128, VII, 1–20.—
Hannerz 1956: 74–85, Figs. 24–28.—Giordanella 1969: 327,
342–343.—Fig. 6.—Rasmussen 1973: 96.—Dauvin 1989:
170–172.—Hartmann-Schröder 1996: 340.—Böggemann
1997: 121.—Bergfeld and Kröncke 2003: 27–29.—Not:
Southern 1914: 95–96.
Spio filicornis—Fauvel 1927: 43–44, Fig. 15A–G (partim).
Non-type material North Sea: 54°18.00′N, 8°17.97′E, 15
December 1987, sand, 14 m, one specimen (ZSRO-P 391);
54°28.36′N, 8°48.00′E, 26 August 1997, sand, eulittoral, one
specimen (ZSRO-P 485); 54°43.93′N, 8°03.48′E, 18 August
Fig. 7 Spio martinensis Mesnil, 1896. a Anterior end of a North Sea
specimen, dorsal view; b anterior end of a Baltic Sea specimen,
dorsal view; c chaetiger 1, anterior view; d chaetiger 14, anterior
173
1998, 13 m, six specimens (ZSRO-P 906); 54°18.00′N, 8°
18.00′E, 15 August 2005, sand, 15 m, four specimens
(ZSRO-P 1853); 54°43.50′N, 8°03.05′E, 17 August 2005,
sand, 14 m, eight specimens (ZSRO-P 1855); 54°35.00′N, 8°
08.00′E, 16 August 2005, sand 14 m, four specimens
(ZSRO-P 1856); 54°26.00′N, 8°10.00′E, 16 August 2005,
sand, 19 m, four specimens (ZSRO-P 1857); 54°43.38′N, 8°
41.82′E, 6 September 2005, sand eulitoral, two specimens
(ZSRO-P 1858). Baltic Sea: 54°36,404′N, 12°39,448′E,
March 2002, four specimens (SBRO-P 793); Wismarbucht,
Februar 2003, fine to medium sand, 10 m, nine specimens
(SBRO-P 809); Graal-Müritz (to the east of Rostock),
Autumn 2004, fine sand, 10 m, two specimens (SBRO-P
3283); Graal-Müritz (to the east of Rostock), Autumn 2004,
sand, 11 m, five specimens (SBRO-P 3295).
Description Maximum number of chaetigers 50 (width
0.5 mm), maximum width 0.63 mm (anterior fragment
with 42 chaetigers).
Prostomium narrow, anteriorly rounded, anterior part
sometimes slightly expanded; posterior part of prostomium
view; e chaetiger 34, anterior view; f neuropodial hooded hook from
chaetiger 34; g pygidium, dorsal view. (Scale bar: a, b 125 µm; c–e
320 µm; f 64 µm; g 160 µm)
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rounded or tapered, posteriorly extending to chaetiger 1
(Fig. 7a, b); two pairs of black eyes arranged trapezoidally,
anterior pair crescent-shaped, rarely rounded, posterior pair
oval; prostomium and peristomium of North Sea specimens
completely separated by a depression, on Baltic Sea specimens a deep groove between prostomium and peristomium only exists anteriorly (Fig. 7a, b).
Nuchal organs with short median and long lateral ciliary
bands, lateral ciliary bands posteriorly curved and go only
partly back to the transverse ciliary band of chaetiger 2
(Fig. 9c); Metameric dorsal ciliated organs double-paired,
indistinct from between branchiae 4 and 5 to between
branchiae 5 and 6 but distinct from between branchiae 6
and 7 or 7 and 8 (Fig. 9c); its posterior extension has not
been defined.
Branchiae from chaetiger 1, continuing to near end of
body; branchiae on chaetiger 1 almost as long as those on
following chaetigers (Fig. 7a, b); branchiae on anterior
chaetigers basally fused with notopodial postchaetal lamellae, becoming separated posteriorly; minor differences in
length and shape of branchiae of North Sea and Baltic Sea
with North Sea specimens branchiae slender, cirriform,
without wide base, slightly tapered distally, becoming
thinner and shorter in posterior chaetigers (Fig. 7a);
branchiae on anterior chaetigers reach the midline dorsally,
touch each other; branchiae of Baltic Sea specimens with
broad base anteriorly, slightly tapered distally, becoming
thinner and shorter posteriorly, without broad base in
posterior chaetigers (Fig. 7b); branchiae on anterior
chaetigers almost reaching dorsal midline, not touching
each other.
First notopodium shifted dorsally. Notopodial postchaetal lamellae on chaetiger 1 slightly pointed, in other anterior
chaetigers oval, becoming shorter and rounded thereafter,
notopodial lamellae elongated in posterior chaetigers
(Fig. 7c–e). Neuropodial postchaetal lamellae rounded on
chaetigers 1 and 2, in other anterior chaetigers oval,
becoming smaller and shorter thereafter, rounded in
posterior chaetigers (Fig. 7c–e).
Alcohol preserved specimens with dark brown pigment
on anterior part of body: pigmented patch dorsally on
anterior margin of prostomium; pigmented patches dorsolaterally on anterior chaetigers before and/or behind
transverse ciliary bands; pigment indicates end of nuchal
organs (Fig. 7a, b); median pigmented patch between
neuro- and notopodia; row of pigmented patches on palps;
anal cirri also pigmented, ventral pair more intensive than
dorsal pair; on specimens from North Sea additional
pigment: a median longitudinal pigmented stripe dorsally
on prostomium; indistinct pigment scattered dorsally on
peristomium and distinct pigmented patch at about level of
eyes (Fig. 7a); pigment between peristomium and first
chaetiger; a short median pigmented stripe ventrally
Mar Biodiv (2010) 40:161–180
between anterior chaetigers; pigmented stripes or patches
ventro-laterally on base of neuropodia; median transverse
pigmented stripe ventrally; several pigmented stripes
between noto- and neuropodia; small pigmented patch at
the base of palps; pigment at margin of notopodial lamellae,
close to branchiae.
Notopodial chaetae all limbate capillaries; chaetae of
anterior chaetigers arranged in two rows: capillaries of
anterior row short, broad, uniformly granulated (Fig. 5f);
chaetae of posterior row longer, thinner, lacking granulations (Fig. 5g); additional superior fascicle of very long,
thin capillaries without granulations; capillaries of posterior
chaetigers arranged in irregular rows, thin, non-granulated,
varying length of capillaries. Neuropodia with rows of
capillaries and hooded hooks as well as an inferior fascicle
of capillaries; chaetae of anterior neuropodia arranged in
two rows, similar to notopodial chaetae; posterior row
replaced by a single row of five to eight hooded hooks from
chaetiger 14 (exceptionally from chaetiger 13), accompanied by alternating short, thin, alimbate, non-granulated
capillaries (Fig. 5e); hooks with long hood, slightly
narrowed subdistally, tridentate, third tooth hardly visible
(Fig. 7f); inferior fascicle with two to three long, thin,
limbate capillaries without granulations, replaced by two to
three sabre chaetae from about chaetiger 20–24, each
uniformly granulated (Fig. 5d).
Pygidium with four anal cirri; dorsal pair slender,
cirriform and widely spaced; ventral pair broad, oval or
slightly cone-shaped, closely spaced and touching each
other at the base (Fig. 7g); dorsal pair of North Sea
specimens slightly longer than ventral pair and dorsal pair
of Baltic Sea specimens shorter than ventral pair or of the
same size.
Distribution and ecology Baltic Sea, North Sea, [from
literature: northeast Atlantic, Arctic Ocean (?), northeast
Pacific (?)]; sublitoral; mud, fine sand, coarse sand.
Remarks Spio martinensis is characterised by its hooded
hooks present from chaetiger 14 (rarely from chaetiger 13).
There are differences between the North Sea and Baltic Sea
specimens in the length and shape of branchiae. Whereas
the North Sea specimens possess long and slender
branchiae without a wide base, specimens from the Baltic
Sea have shorter, wide-based branchiae. Furthermore, only
Baltic Sea specimens have the lateral margin of the
prostomium slightly expanded. North Sea specimens
possess a flat depression between prostomium and peristomium, whereas it is deep and limited to the anterior part
in Baltic Sea specimens. The anal cirri are also slightly
different.
Since type material or material from the type locality
could not be tracked by the authors and is supposedly lost,
Mar Biodiv (2010) 40:161–180
Author's personal copy
it was not possible to decide whether the specimens found
in the North Sea and Baltic Sea belong to the same species
or whether the assignment to S. martinensis is reliable.
The characters found in specimens from both the North
and Baltic Seas are generally similar to the original
description of S. martinensis from the English Channel
(Mesnil 1896).
Key to species of Spio from the North and Baltic Seas
1 Branchiae on chaetiger 1 distinctly shorter than branchiae
on chaetiger 2; bidentate neuropodial hooded hooks first
appear on chaetigers 16–23 (12–15 in juveniles, width
<0.4 mm)........................................................................ 2
- Branchiae on chaetiger 1 as long as those on chaetiger
2, or only slightly shorter; bi- or tridentate neuropodial
hooded hooks first appear on chaetigers 11–14 in
juveniles and adults...................................................... 3
2 Branchiae on chaetiger 1 very short, only slightly
longer or as long as notopodial postchaetal lamellae;
anterior part of prostomium broadly rounded, posterior part tapered; lateral ciliary bands posteriorly curved
and go nearly completely back to the transverse
ciliary band of chaetiger 2 (Fig. 9f); metameric dorsal
ciliated organs double-paired; neuropodial hooded
hooks from chaetigers 17 (14–16, Thulin 1957; 8–20,
Sikorski 2001), strongly curved, main fang large, of
nearly right angle to shaft (Fig. 1c); inferior sabre
chaetae and neuropodial hooks conjointly appear
...................................................................... S. armata
- Branchiae on chaetiger 1 short, but distinctly longer
than notopodial postchaetal lamellae; anterior part of
prostomium tapered, posterior part broadly rounded;
lateral ciliary bands slightly outward curved in the
range of the first transverse ciliary band (Figs. 8a and
9a); single-paired metameric dorsal ciliated organs;
neuropodial hooded hooks from chaetigers 16–23
(12–15 in juveniles, width <0.5 mm), uniformly
curved, weakly developed main fang (Fig. 5c); inferior sabre chaetae first present after first appearance of
neuropodial hooded hooks ................ S. goniocephala
3 Neuropodial hooded hooks bi- or tridentate from
chaetiger 11 (rarely 10) .............................................. 4
- Neuropodial hooded hooks tridentate, from chaetiger
14 (rarely 13); posterior part of prostomium tapered;
lateral ciliary bands posteriorly curved and go only
partly back to the transverse ciliary band of chaetiger 2
(Fig. 9c); metameric dorsal ciliated organs doublepaired ..................................................... S. martinensis
4 Neuropodial hooded hooks bidentate; metameric
dorsal ciliated organs present from between branchial pairs 3 and 4; posterior part of prostomium with
distinct long, keel-shaped elevation (visible in lateral
175
view); lateral ciliary bands outward curved in the
range of the first transverse ciliary band, posteriorly
curved and go nearly completely back to the
transverse ciliary band of chaetiger 2 (Fig. 9e);
conspicuous dark-brown pigment on peristomium and
no or very faint pigment on prostomium (Fig. 4a)
.................................... Spio cf. filicornis (Baltic Sea)
- Neuropodial hooded hooks usually tridentate; metameric dorsal ciliated organs present from between
branchial pairs 4 and 5 or 5 and 6; posterior part of
prostomium pointed or tapered; distinct brown pigment
on peristomium and prostomium (Figs. 2a and 3a)
..................................................................................... 5
5 Prostomium not distinctly separated from peristomium
by a furrow (Fig. 3a); brown pigment on prostomium
and peristomium in irregular patches; lateral ciliary
bands outward curved in the range of the first transverse
ciliary band, posteriorly curved and go nearly completely back to the transverse ciliary band of chaetiger 2
(Figs. 9d, 10a); metameric dorsal ciliated organs from
between branchial pairs 4 and 5; the third tooth on the
hooded hooks hardly visible (Fig. 3f, often worn
down?) ........................... Spio cf. filicornis (North Sea)
- Prostomium distinctly separated from peristomium by a
furrow (Fig. 2a); broad transverse pigmented band on
prostomium and peristomium (Fig. 2a); lateral ciliary
bands posteriorly curved and go only partly back to the
transverse ciliary band of chaetiger 2 (Fig. 9b);
metameric dorsal ciliated organs present from between
branchial pairs 5 and 6; distinct third tooth on the
hooded hooks (Fig. 2b) S. decoratus
Discussion
At least six Spio species occur in our study area: S.
decoratus, Spio cf. filicornis (North Sea), S. goniocephala
and S. martinensis in the North Sea, and S. armata, Spio cf.
filicornis (Baltic Sea), S. goniocephala and S. martinensis
in the Baltic Sea. Based on detailed morphological studies
it was possible to find diagnostic characters for all
investigated species and to provide a key for identification.
Of particular importance is the discovery that specimens of
the supposedly widespread S. filicornis belong to two
different species, of which one occurs in the Baltic Sea and
the other one in the North Sea. However, not all taxonomic
problems have been resolved. For example, specimens of S.
martinensis from the North and Baltic Seas differ in some
respects, but are here regarded as con-specific. Taxonomic
problems within Spio can be attributed, on one hand, to the
loss or poor condition of type material, as is the case with
S. filicornis, S. armata and S. goniocephala. On the other
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Mar Biodiv (2010) 40:161–180
Fig. 8 Spio goniocephala
Thulin, 1957, North Sea.
a Anterior end, with nuchal
organs and single-paired
metameric dorsal ciliated
organs, slightly outward curved,
present from between branchial
pairs 3 and 4; b chaetigers
3–8 with single-paired, slightly
outward curved metameric
dorsal ciliated organs
hand, diagnostic characters that have been previously used
to differentiate Spio species are not sufficiently discrimantory. For example, S. filicornis has been characterised by a
rounded, sometimes slightly incised anterior margin of the
prostomium, long branchiae on first chaetigers that are of
the same length as those on the second chaetiger, and
bidentate hooded hooks first appearing from chaetiger 11
(e.g. Maciolek 1990). The inadequacy of this set of
characters as diagnostic is revealed by the following data:
out of 28 Spio species, 16 species possess a rounded
anterior margin of the prostomium, 15 species have long
branchiae on chaetiger 1, and bidentate hooded hooks from
chaetiger 11 are reported for nine species (based on species
descriptions in Maciolek 1990; Blake 1996, and Sikorski
2001). The distribution range of the species includes a great
variety of geographically diverse regions: S. filicornis has
been recorded in western Greenland, southern and northern
Europe, including the North Sea, Baltic Sea, Black Sea and
White Sea, Arctic Sea, Bering Sea, North Pacific, Red Sea,
South Africa, and Ross Sea. It seems likely that the
cosmopolitan species S. filicornis actually represents a
species complex.
The fact that certain morphological characters do not
serve well as diagnostic characters may also be explained
by their size-dependency, or in other words, their change in
the course of development. The recognition of sizedependency of character states is important and has to be
considered in the species diagnosis. Within Spio, that
concerns the shape of the caruncle, the beginning of
branchiae and their length, particularly on the first
chaetiger, the start and the number of hooded hooks, the
number of apical teeth on these hooks, the pigmentation
and the length and shape of the anal cirri (Augener 1914;
Blake 1996; Day 1973; Guerin 1972; Hannerz 1956;
Hartmann-Schröder 1996; Mesnil 1896; Sikorski 2001;
Simon 1967, 1968; Thulin 1957). One of the most
important diagnostic characters is the first appearance of
neuropodial hooded hooks. Three different patterns
concerning that character were found among the investigated
species (Fig. 11): (1) no size-dependency of the character—
hooded hooks always start at the same chaetiger; (2) a minor
shift occurs and the first appearance of hooded hooks is
moved posteriorly; (3) the first appearance of hooks differs
considerably between juveniles (from chaetiger 11) and
adults (from chaetiger 20–23). Pattern 1 was observed for
Spio cf. filicornis (North Sea), Spio cf. filicornis (Baltic Sea),
and S. decoratus. Pattern 2 applies to S. martinensis and
pattern 3 to S. goniocephala. For the sixth species
investigated, S. armata, only six specimens, all of comparable length, were available. Sikorski (2001) reported that for
S. armata from Arctic regions, hooded hooks are present
from chaetigers 8–11 in small and chaetigers 18–20 in large
specimens, and Thulin (1957) found hooks to start on
chaetiger 14 and 16 in this species. A correlation between
size and the beginning of hooded hooks in Baltic Sea
specimens may also exist.
Not only the first appearance but also the number of
hooks is size-dependent: the smaller the specimens, the
fewer the number of neuropodial hooks (Fig. 12). This is
true for all Spio species occurring in both the North and
Baltic Seas and it can also be assumed for all other Spio
species. In certain cases, the number of apical teeth also
varies with body size. The hooded hooks of juvenile Spio
cf. filicornis (North Sea) are tridentate but usually bidentate
in adults. The same observation was made by Hannerz
(1956) and Bergfeld and Kröncke (2003) for S. martinensis.
However, we cannot support this observation based on our
material of S. martinensis.
Another important character is the length of the
branchiae on the first chaetiger. This length varies depending on size of the specimens in Spio cf. filicornis (North
Sea) and in S. goniocephala. The branchiae of S.
goniocephala are very small and papilliform in juveniles
and half as long as the branchiae on the second chaetiger in
Mar Biodiv (2010) 40:161–180
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177
Fig. 9 Schematic representation of nuchal organs, metameric dorsal
ciliated organs and transverse ciliary bands on Spio species from the
North Sea and Baltic Sea. a S. goniocephala Thulin, 1957; b S.
decoratus Bobretzky, 1870; c S. martinensis Mesnil, 1896; d S. cf.
filicornis (Müller, 1776), North Sea; e S. cf. filicornis (Müller, 1776),
Baltic Sea; f S. armata (Thulin, 1957) (lcb lateral ciliary bands of
nuchal organs, mcb median ciliary bands of nuchal organs, stcb short
transversal ciliary band, tcb2 transversal ciliary band of chaetiger 2)
adults. The first branchiae on small specimens of Spio cf.
filicornis (North Sea) are conspicuously shorter than those
on the following chaetigers, but on adults they are usually
as large as those on subsequent chaetigers. In conclusion,
characters previously used as diagnostic for Spio species
are insufficient, partly because their size-dependency has
not been taken into account.
A character we found to be useful also for Spio are the
nuchal organs, together with the metameric dorsal ciliated
organs. They have already been proven extremely useful
diagnostic characters in other spionid genera, e.g. in
Marenzelleria (Bick 2005), Spiophanes (Meißner 2005),
and Laonice (Sikorski 2002). The nuchal organs of Spio
species from the North and Baltic Seas consists of two
median and two lateral bands of cilia (Figs. 8a and 10a).
Median ciliary bands are short, originate directly behind the
prostomium and extend posteriorly to the transverse ciliary
band of chaetiger 2. The lateral ciliary bands are longer,
originate on the palps and extend posteriorly to the middle
of chaetiger 3; they are posteriorly curved and go
completely or nearly completely (e.g. S. armata, S. cf.
filicornis; Figs. 9d–f and 10a), partly (e.g. S. decoratus, S.
martinensis; Fig. 9b, c) or never (e.g. S. goniocephala;
Figs. 8a and 9a) back to the transverse ciliary band of
chaetiger 2. The nuchal organs and the metameric dorsal
ciliated organs are readily visible well-defined structures on
the epithelium in well-preserved adults. In most juveniles,
they comprise lighter coloured areas on the pigmented
epithelium. Shirlastain A was found particularly useful to
visualise nuchal organs and metameric dorsal ciliated
organs in faded or poorly preserved specimens.
Blake and Kudenov (1978) noticed that “dorsal sense
organs ... are important to species concepts” and “any
revision of Spio and Microspio should thus consider dorsal
sense organs at the species level”. This approach was used
by Söderström (1920), who tried to establish the separation of the genera Spio and Microspio by means of the
metameric dorsal ciliated organs, specifying the number of
longitudinal ciliary bands of the metameric dorsal ciliated
organs in Microspio as two and in Spio as four. The
metameric dorsal ciliated organs of Spio spp. consists of
two or four longitudinal ciliary bands arranged in a single
or double row on the dorsum of the first chaetigers
(Figs. 8b and 10c). According to Jelsing (2003), it belongs
to the group IV (paired, metameric dorsal ciliated organs),
occurring within Spionidae. Single dorsal ciliated organs
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Mar Biodiv (2010) 40:161–180
Fig. 10 Spio cf. filicornis
(Müller, 1776), North Sea.
a Anterior end with nuchal
organs, and double-paired
metameric dorsal ciliated organs
present from between branchial
pairs 4 and 5; b additional pointshaped transverse ciliary band
on chaetiger 25; c chaetigers
12–15 with double-paired
metameric dorsal ciliated
organs, long transverse ciliary
bands connecting the branchiae
and additional short transverse
ciliary between dorsal ciliated
organs; d end of double-paired
metameric dorsal ciliated organs
on chaetiger 33
(Fig. 8b) are concave, whereas double-paired dorsal
ciliated organs (Fig. 10c) consist of parallel or slightly
curved ciliary bands. Spio goniocephala (Fig. 9a) is the
only species in this region with single-paired dorsal
ciliated organs; all other species have double-paired dorsal
ciliated organs (Fig. 9b–f). Apart from the shape, differences also exist in the first appearance of the dorsal
ciliated organs. Jelsing (2002) stated that the dorsal
Fig. 11 Relationship between size of specimens (width in millimetres) and first appearance of neuropodial hooded hooks in Spio
species from the North Sea and the Baltic Sea
Fig. 12 Relationship between size of specimens (width in millimetres) and number of neuropodial hooded hooks in Spio species
from the North Sea and the Baltic Sea
Mar Biodiv (2010) 40:161–180
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ciliated organs are intersegmentally located, but in fact it is
difficult to define the accurate position. The branchiae
have, therefore, been used as a surrogate to determine the
beginning of the dorsal ciliated organs. In Spio species
found in North Sea and Baltic Sea, they first appear from
between branchial pairs 3 and 4 [Spio armata, Spio cf.
filicornis (Baltic Sea), S. goniocephala] or between 4 and
5 [Spio cf. filicornis (North Sea), S. martinensis] or six [S.
decoratus]. The ciliary bands of the metameric dorsal
ciliated organs tend to extend in length until they are
nearly as long as the corresponding chaetiger. They
disappear abruptly on a middle chaetiger (Fig. 10d). It is
difficult to decide whether the termination of the dorsal
ciliated organs is species-specific or variable because it
was not always possible to define its posterior extension.
All Spio species possess a transverse ciliary band
connecting the opposite branchiae of the corresponding
chaetiger. This ciliary band is absent on chaetiger 1, short
on chaetiger 2 and fully developed from chaetiger 3
(Fig. 8a). Spio cf. filicornis (North Sea) and S. decoratus
possess an additional shorter transverse ciliary band on
middle chaetigers (Figs. 9b–d and 10b, d). First, it is a
ciliated tuft situated between the longitudinal ciliary bands
of the dorsal ciliated organs (Fig. 10b), then extends onto
the following chaetigers to form finally the short additional
transverse band (Fig. 10c, d).
Summary
The present study provides a detailed morphological study
of Spio species found in the North and Baltic Seas.
Previously used morphological characters were critically
reviewed and new characters were included into the
amended species diagnoses. The following characters were
found to be of particular taxonomic importance: the shape
of the prostomium and the type of extension of its posterior
part, the presence or absence of a groove-like depression
(or furrow) between the prostomium and peristomium, the
shape of the nuchal organs and metameric dorsal ciliated
organs (metameric longitudinal ciliary bands on the dorsum
of anterior and middle chaetigers), the first appearance of
sabre chaetae, and the pigment pattern on the prostomium,
peristomium and anterior chaetigers. A number of taxonomic characters are size-dependent. For the species from
the study region, size-dependent variation may concern the
beginning and the number of neuropodial hooded hooks,
the number of apical teeth on these hooks and the length of
branchiae on the first chaetiger. All these characters are of
taxonomic importance. As a result of the study, it was also
discovered that S. filicornis previously reported from the
North and Baltic Seas include two different species. The
formal assignment of these species to either a known or a
179
new species has to be undertaken as soon as material from
the type locality of S. filicornis becomes available.
Acknowledgements We thank James A. Blake, Nancy J. Maciolek
and three anonymous referees, who reviewed a previous draft of this
manuscript. Their comments greatly improved the quality of the paper.
We would like to thank Volker Schroeren (LANU), Karin Fürhaupter
(MariLim), Michael Zettler (IOW), Regine Bönsch (IfAOE), Katrine
Worsaae and Danny Eibye-Jacobsen (ZMUC) and Dieter Fiege (SMF)
for making their collections available to us. Type material was
supplied by Lars R. Lundqvist from Zoologiska Museet, Lunds
Universitet, Sweden (L). Tatyana Nikitenkova (IfAOE) translated
Russian publications. We are grateful to Prof. Ludwig Jonas, Gerhard
Fulda and Wolfgang Labs from the Electron Microscopic Centre of the
Medical Faculty of the University of Rostock for their support. The
study was funded in part by the Federal Agency for Nature
Conservation, Germany under FKZ 370725201.
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