http://dx.doi.org/10.5852/eit.2015.144
www. europeani ournaloftaxonomv. eu
2015 • Bello G. & Salman A.
This work is licensed under a Creative Commons Attribution 3.0 License.
Research article
urn:lsid:zoobank.org:pub:llB9BCE3-18F9-429F-8EEA-4E232D9E42E0
Description of a new sepioline species, Sepiola boletzkyi sp. nov.
(Cephalopoda: Sepiolidae), from the Aegean Sea
Giambattista BELLO 1 -* & Alp SALMAN 2
1 Arion, Via Colombo 34, 70042 Mola di Bari, Italy.
2 Ege University, Faculty of Fisheries, Department of Hydrobiology, 35100, Bomova, Izmir, Turkey.
E-mail: [email protected]
* Corresponding author: [email protected]
1 urn: lsid: zoobank. org: author: 31A5 0D6F-5126-48D1-B63 0-FBED A63944D9
2 um:lsid:zoobank.org:author:76C095B6-A975-49D4-BDF0-0802B03E9B4C
Abstract. A new sepioline species, Sepiola boletzkyi sp. nov. (Cephalopoda: Sepiolidae), is described
based on two specimens from the Aegean Sea (eastern Mediterranean). The type specimens are lodged
in the Ege University Faculty of Fisheries Museum of Izmir (Turkey). The new species belongs to the
Sepiola atlantica group sensu Naef, hence it is compared with the species in this group, namely Sepiola
affims, Sepiola atlantica , Sepiola bursadhaesa , Sepiola intermedia , Sepiola robusta, Sepiola rondeletii ,
Sepiola steenstrupiana and Sepiola tridens. The male of S. boletzkyi sp. nov. differs from all the others
in having the combination of homomorphous ventral arm tips, eight enlarged suckers, subdivided into
two groups, in the dorsal row of the distal part of the hectocotylus and a dorsal lobe complementing
the copulatory apparatus. In females of S. boletzkyi sp. nov. the bursa copulatrix is large and, unlike all
other species in the genus, has a large fungiform process towards the mantle wall. The discovery of S.
boletzkyi sp. nov. increases to 12 the number of NE Atlantic-Mediterranean species of Sepiola.
Keywords. Cephalopoda, Sepiolidae, Sepiola boletzkyi sp. nov., Aegean Sea, Mediterranean Sea.
Bello G. & Salman A. 2015. Description of a new sepioline species, Sepiola boletzkyi sp. nov. (Cephalopoda:
Sepiolidae), from the Aegean Sea. European Journal of Taxonomy 144: 1-12. http://dx.doi.org/10.5852/
eit.2015,144
Introduction
The genus Sepiola Leach, 1817 (Cephalopoda: Sepiolida: Sepiolidae) is the most speciose sepioline
genus in the NE Atlantic-Mediterranean region, where it is represented by 11 species (Bello 2013). Eight
of them, Sepiola affinis Naef, 1912, Sepiola atlantica d’Orbigny, 1841, Sepiola bursadhaesa Bello,
2013, Sepiola intermedia Naef, 1912, Sepiola robusta Naef, 1912, Sepiola rondeletii Leach, 1817,
Sepiola steenstrupiana Levy, 1912 and Sepiola tridens de Heij & Goud, 2010, belong to the Sepiola
atlantica group sensu Naef (1923), which is characterized by the peculiar morphology of the male
copulatory apparatus (Fig. 1). Sepiola atlantica and S. tridens are endemic to the Atlantic Ocean, all
other species live in the Mediterranean Sea (Bello 2013). Two more species occur in the Mediterranean,
namely Sepiola aurantiaca Jatta, 1896 and Sepiola ligulata Naef, 1912 (Bello 2003).
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European Journal of Taxonomy 144: 1-12 (2015)
In the paper describing S. bursadhaesa, the most recently defined Sepiola species, Bello (2013: 498)
stated “Because of the [... ] combination of factors that facilitates speciation [in Sepiola ], it is likely that
other, still undescribed species of this genus await discovery.” An additional Mediterranean species, viz.
Sepiola boletzkyi sp. nov., was discovered in the Aegean Sea and is described in the present paper.
Material and methods
The sepiolines were caught by a hand-operated 1 mm mesh size tulle net, during daytime. In the
laboratory they were measured (mantle length) and fixed in 10% formalin in seawater. Afterwards they
were preserved in 70% ethyl alcohol.
The sample consists of two specimens which are housed in the Ege Universitesi Su Uriinleri Fakiiltesi
Miizesi (ESFM) (Ege University Faculty of Fisheries Museum) of Izmir (sample nos. CEP/1994-1 and
CEP/1994-2). Subsequent samplings, carried out in recent times in the same area, did not provide any
additional specimens of this new species.
The photographs of anatomical details were taken with a digital camera (DFC 420, Leica, Cambridge,
U.K.) attached to a stereo microscope.
Measurements were taken on the preserved specimens, except mantle length, which was also taken
before preservation, following Roper & Voss (1983) and Reid (2009). Indices were calculated by
expressing each measure as a percentage of mantle length; their acronyms are defined in Table 1 next to
each measurement designation.
Abbreviations
ML = dorsal mantle length
arms I to IV = first to fourth pair of arms, i.e., dorsal, dorso-lateral, ventro-lateral, ventral arms,
respectively
VML = ventral mantle length
copulatory
apparatus
dorsal row
of suckers
distal
part
ventral row
of suckers
Fig. 1 . Schematic rendering of the
general hectocotylus (left arm I) in the
Sepiola atlantica group sensu Naef
(1923) (after Bello 1995, modified).
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BELLO G. & SALMAN A., Sepiola boletzkyi sp. nov. from the Aegean Sea
Results
Class Cephalopoda Cuvier, 1797
Subclass Coleoidea Bather, 1888
Order SepiolidaLioroni, 1981
Lamily Sepiolidae Leach, 1817
Subfamily Sepiolinae Leach, 1817
Genus Sepiola Leach, 1817
Sepiola Leach, 1817: 137. Type species Sepiola rondeletii Leach, 1817, by monotypy.
Diagnosis
Sepiolines with pair of kidney-shaped light-organ lenses on ventral surface of ink sac; suckers biserial
on arms I to III and at least proximally on arms IV; tentacle club suckers in four to eight oblique rows;
females with ear-shaped bursa copulatrix on left ventral side of mantle cavity; males with left arm I
hectocotylized: suckers and sucker stalks typically arranged in a basal part, a copulatory apparatus com¬
prised of modified suckerless stalks, and a distal part (Bello 2013).
Sepiola boletzkyi sp. nov.
um:lsid:zoobank.org:act:A6A9644D-61E6-4964-9460-A803 703539 ID
Tigs 2-7, Tables 1-2
Diagnosis
Sepiola with biserial suckers on all arms, including tips of arms IV. Tentacle clubs with six oblique rows
of equal-sized suckers. Hectocotylus typical of the Sepiola atlantica group sensu Naef (1923) (Tig. 1):
(1) three regular equal-sized suckers in basal part; (2) copulatory apparatus with roundish dorsal tuber¬
cle, wealdy developed, complemented by broad lobe, dorsomedial to tubercle; (3) sequence of modified
suckers in dorsal row of distal part: (a) two small, (b) six markedly enlarged, (c) one small and (d) two
enlarged suckers; (4) no dorso-lateral groove on medial side of distal part. Bursa copulatrix moderately
large and ear-shaped, anteriorly extended below the left light-organ, with a large fungiform process
towards mantle wall posterior to gill insertion.
Etymology
Lrom the family name of the eminent teuthologist Sigurd von Boletzky, in recognition of his enormous
contribution to teuthology.
Material examined
Holotype
S, ESFM-CEP/1994-1, mature, ML: 20 mm fresh, 16.8 mm preserved.
Paratype
5, ESFM-CEP/1994-2, immature, ML: 19 mm fresh, 17.2 mm preserved. Same locality as holotype.
Type locality
TURKEY: Gulf of Ildir, Aegean Sea, eastern Mediterranean Sea, off Sifne (£e§me District, Izmir
Province), 38°19'32"N 26°24T4"E, on a sandy bottom covered by eel-grass, Zostera sp., at a depth of
2 m, Apr. 1994.
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European Journal of Taxonomy 144: 1-12 (2015)
Description
Medium size Sepiola', maximum recorded ML = 20 mm . male (fresh specimen); maximum recorded
TL, without tentacles = 40.5 mm, male (arms are variously curled up in both specimens). Tentacles
exceed TL by 6-9 mm (tentacle and clubs are variously curled up in both specimens). Measurements
are reported in Table 1.
Body shape corresponds to general shape of NE Atlantic-Mediterranean Sepiola species (Fig. 2).
Mantle sac-shaped, broadly rounded at posterior end, length exceeds width, outline U-shaped. Dorsal
mantle margin fused to head by cutaneous nuchal band 27-31% of ML. Ventral mantle margin very
slightly sinuate, with very low barely visible rounded projection on each side of funnel (Fig. 2). VML
approximately 89-98% of ML. Fins typical for Sepiola , each inserted midway along sides of mantle,
broadly rounded; fin length slightly exceeds half ML, fin width about 70-80% of fin length, insertion
line about 60% of overall fin length. Head width 65-69% of ML in both specimens. Eyes large, bulging
beyond sides of the head, covered by corneal membrane and bordered by secondary eyelid. Funnel long
about 60% of ML, covered posteriorly by ventral margin of mantle, funnel free length about 40% of ML;
distal end does not reach junction of ventral arms.
Arm formula II = III > IV > I in both specimens. Arms slightly longer in male than in female. Suckers
stalked, alternately placed in two series on oral side of all arms, including distal-most part of arms IV.
Fig. 2. Sepiola boletzkyi sp. nov., holotype, S, ESFM-CEP/1994-1. The hectocotylus was removed for
closer examination. A. Dorsal view. B. Ventral view. Scale bar = 1 cm.
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BELLO G. & SALMAN A., Sepiola boletzkyi sp. nov. from the Aegean Sea
Table 1 . Measurements and indices of type specimens of Sepiola boletzkyi sp. nov., MLs in square
brackets are those taken on fresh specimens prior to preservation.
Holotype, $
(mm / %)
Paratype, $
(mm / %)
dorsal mantle length / ML
16.8 [20]
17.2 [19]
ventral mantle length / VMLI
16.4/97.6
15.3/89.0
total length
40.5
38.6
total length, including tentacles
46.8
47.2
arm I length, right and left (hectocotylus) / A1LI
12.9/76.8-8.5/50.6
9.3/54.1
arm II length / A2LI
14.9/88.7
13.0/75.6
arm III length / A3LI
14.8/88.1
13.3/77.3
arm IV length / A4LI
13.3/79.2
10.7/62.2
mantle width / MWI
14.1/83.9
13.5/78.5
head width / HWI
11.0/65.5
11.8/68.6
head length / HLI
7.1/42.3
6.9/40.1
fin length / FLI
9.3/55.4
9.1/52.9
fin width / FWI
6.4/38.1
7.2/41.9
fin insertion / FII
5.7/33.9
5.6/32.6
distance fin anterior insertion - mantle border / Fial
5.6/33.3
5.4/31.4
nuchal commissure width / NCI
4.3/25.6
5.4/31.4
funnel length / FuLI
10.1/60.1
10.6/61.6
free funnel length / FfuLI
6.7/39.9
7.1/41.3
funnel locking cartilage length / FuCartLI
4.0/23.8
4.5/26.2
tentacle club length / C1LI
8.0/47.6
8.0/46.5
gladius length / GLI
-
8.5/49.4
Fig. 3. Right tentacle club of Sepiola boletzkyi sp. nov., holotype, S, ESFM-CEP/1994-1. A. Oral view
(drawing). B. Oral view. C. Dorsal view. Scale bar = 1 mm.
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European Journal of Taxonomy 144: 1-12 (2015)
Table 2. Measurements taken on beaks of the paratype, §, ESFM-CEP/1994-2, of Sepiola boletzkyi
sp. nov., definitions according to Clarke (1986) and A 9 & & Salman (2010).
Upper beak
Lengths (mm)
Rostrum
0.7
Hood
2.6
Total
4.8
Wing
1.6
Lower beak
Lengths (mm)
Rostrum
0.8
Hood
1.2
Wing
1.1
Crest
2.6
Brachial crown of mature males displays sexual dimorphic features (see below for detailed description).
Arm web reduced; encloses base of tentacles between arms III and IV.
Tentacles thin and delicate. Tentacular club longitudinally lined by protective membranes more
developed and ventrally trabeculate (width 1.5-2 mm); distal third of club abruptly tapered, elongate;
club suckers equal sized in six oblique rows (Fig. 3).
Overall skin colouration pale violet brown; most chromatophores not expanded. (Actual colouration
undoubtedly altered by several year storage in ethyl alcohol.) No particular chromatophore distribution
pattern observed.
Mantle-locking apparatus straight, as typical for the genus, long about % ML.
Gladius positioned sagittally on inner face of dorsal mantle, encased in shell sac (see Bizikov 2008 for
general description of sepiolid gladius). Gladius reduced, extends from anterior mantle margin to about
45% of ML, posterior tip does not reach posterior end of mantle (as is typical for the genus). Gladius
shape similar to those reported for S. bursadhaesa (Bello 2013) and S. aurantiaca (Jatta 1896) (Fig. 4).
Chitinous rachis light tan-coloured, widely open V-shaped in cross section. Gladius bluntly pointed at
anterior extremity then gradually enlarges to reach width of about 4% of GL (0.32 mm in examined
specimen); posteriorly, two sides of rachis run almost parallel to about 55% of GL and subsequently
converge to a point short of posterior end. Point where rachis starts to narrow corresponds to anterior
insertion of thin and transparent lateral plates. Posterior part of gladius spear-head shaped where rachis
is the shaft and lateral plates the two lateral blades that taper posteriorly to a narrow point exceeding the
convergence point of the rachis by about 2.5% of GL.
Beaks indistinguishable from those of other NE Atlantic-Mediterranean Sepiola species ( cf Ayik &
Salman 2010) (Fig. 5). Female beak measurements are shown in Table 2. Upper beak rostrum unifor ml y
curved; jaw angle obtuse; wing long as anterior lateral wall margin, shoulder (cutting edge) straight,
its margin neatly separated at an obtuse angle from rostrum; no grooves at hood sides; no lateral wall
Fig. 4. Diagram of the gladius of Sepiola boletzkyi sp. nov., paratype, §, ESFM-CEP/1994-2 (anterior
extremity on the left). Abbreviations: R = rachis, LP = lateral plate. Scale bar = 1 mm.
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BELLO G. & SALMAN A., Sepiola boletzkyi sp. nov. from the Aegean Sea
fold; most of lateral wall, posterior part of hood and inferior part of wing transparent. Lower beak
comparatively wide; rostrum with blunt tip; jaw edge profile convex, jaw angle indistinct; blunt tooth
on shoulder; no notch in hood; lateral wall without fold or ridge, roughly rectangular with lower edge
slightly convex, corner faintly produced; wall and wings transparent.
Fig. 5. Beaks of Sepiola boletzkyi sp. nov., paratype, ESFM-CEP/1994-2. A. Upper beak, lateral
view. B. Lower beak, lateral-oblique view. C. Lower beak, upper (= ventral) view. Scale bar = 1 mm.
Male
Mature. Spermatophores 6.5 mm long. Mantle cavity does not show any unique traits that allows it to
be distinguished from other Sepiola species. Brachial crown dimorphic. Left arm I hectocotylized (see
below), arms II bear 6 pairs of enlarged suckers in middle third; arms III not evidently more robust
than remaining arms and not bent inward (contrary to several Sepiola species); arms IV bear 3 pairs of
enlarged suckers in middle third.
Hectocotylus (left arm I) shorter than right arm I. Basal part (refer to diagram in Fig. 1) bears three
equal-sized, not markedly enlarged, suckers, two in ventral and one in dorsal row (the latter missing
in holotype as result of damage) (Fig. 6A). Copulatory apparatus formed by fusion of four suckerless
stalks, elongated and curved medially, projecting away from arm; ventral-most ones correspond to third
and fourth stalks of ventral row; dorsal-most ones derive from second and third stalks of dorsal row,
modified into roundish tubercle (Fig. 6A-B). Copulatory apparatus complemented by broad smooth-
pointed triangular lobe, dorsomedial to tubercle, formed from more than one dorsal suckerless stalks
fused with each other (i.e., likely fourth, fifth and sixth stalks of dorsal sucker row); lobe forms groove
with tubercle (Fig. 6B-C).
Distal part of hectocotylus somewhat bent, narrow (i.e., two rows of suckers not spread apart to form
a spoon-like widening) (Fig. 6A). Arm tip blunt. No groove on hectocotylus lateral side facing right
arm I (Fig. 6B). First two suckers of dorsal row (Fig. 6A, suckers 1-2) in distal part smaller than facing
suckers of ventral row; followed by six markedly enlarged (Fig. 6A, suckers 3-8), one small (smaller
than facing ventral row suckers) (Fig. 6A, sucker 9), two enlarged suckers (Fig. 6A, suckers 10-11). All
enlarged suckers about the same size. Distal to enlarged suckers, remaining eight suckers gradually taper
to arm tip. Sucker stalks of enlarged suckers proportionally larger and elongated. Suckers of ventral row
regularly sized.
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European Journal of Taxonomy 144: 1-12 (2015)
Fig. 6. Hectocotylus (left arm I) of Sepiola boletzkyi sp. nov., holotype, ESFM-CEP/1994-1.
Abbreviations: a-d = suckerless stalks forming the copulatory apparatus proper (a = ventral-most, d =
dorsal-most one, i.e. tubercle); * = lobe in the dorsal row of suckers, 1-11 = sequence of heteromorphous
suckers in the dorsal row of distal part. A. Oral view; the arm is slightly tilted, with the tip towards the
observer, in order to show details (lobe and small suckers 1-2), otherwise hidden by the four suckerless
stalks of the copulatory apparatus. Basal part: only the two ventral suckers are visible, the dorsal one
is missing as a result of damage. Copulatory apparatus: the tubercle (d) partially hides the lobe in the
dorsal row (*). Distal part: the heteromorphous suckers in the dorsal row are 1-2 small, 3-8 enlarged,
9 small, 10-11 enlarged; the remaining suckers are regular; note that some suckers (2, 3 and 7) are
displaced towards the centre of the arm, i.e., not aligned with the others of their same row; it is not
known whether this is natural or caused by preservation. B. Inner-lateral view of copulatory apparatus;
the lobe (*) forms a groove with the tubercle (d). C. Oblique inner-lateral view of copulatory apparatus;
two barely visible longitudinal lines on the lobe (*) are likely fusion lines between adjacent stalks
forming it. Scale bars = 1 mm.
Female
Immature. Bursa copulatrix large, roughly ear-shaped, longer than wide; anteriorly extends below left
light-organ lens, posteriorly reaches posterior end of mantle cavity (most probably this is not so in
mature females with enlarged ovary), medially reaches sagittal mantle cavity septum (Fig. 7A). Folds
converge towards centre of bursa where they deepen into a longitudinal opening. Posterior half of bursa
bordered by elevated rim; contour elliptical, slightly pointed (like a barge prow). Farge fungiform
process, towards mantle wall rear of gill insertion, sticking out of bursa laterally from converging folds;
stalk of process longitudinally folded; distal end larger than stalk, flat, roughly triangular with rounded
corners, might be fused with mantle wall (slight possible scar on it) (Fig. 7B).
8
BELLO G. & SALMAN A., Sepiola boletzkyi sp. nov. from the Aegean Sea
Fig. 7. Bursa copulatrix of Sepiola boletzkyi sp. nov., paratype, ESFM-CEP/1994-2. A. Ventral view;
the dashed line encloses the distal surface of the large fungiform process. B. Oblique ventral view; the
fungiform process sticks out of the bursa, laterally to the folds convergence; an arrow points to the
longitudinally folded stalk of the process. Scale bar for both figures = 1 mm.
Remarks
Sepiola boletzkyi sp. nov. can be distinguished from other species in the genus based on the hectocotylus
and the bursa copulatrix. As discovered by Naef (1912a, b, 1923) these copulatory organs are species-
specific, hence they discriminate the Sepiola species from each other (see also Bello 1995, 2013).
In particular S. boletzkyi sp. nov. belongs to the Sepiola atlantica group as defined by Naef (1923),
characterized by the peculiar morphology of the male copulatory apparatus (Fig. 1).
The male S. boletzkyi sp. nov. differs from all other species in the S. atlantica group because it is unique
in exhibiting simultaneously the characters (1) homomorphous tip of ventral arms and (2) enlarged
suckers of the dorsal row of the hectocotylus distal part that are subdivided into two groups. Character 1
is shared by S. affinis, S. bursadhaesa, S. intermedia, S. robusta and S. rondeletii as opposed to S.
atlantica, S. steenstrupiana and S. tridens ; character 2 is displayed only by S. atlantica and S. tridens as
opposed to all other species.
It is not clear to which Sepiola species S. boletzkyi sp. nov. is closer because this new sepioline differs
from all four subgroups delineated by Bello (2013) within the atlantica group. Although it shares with
S. atlantica + S. tridens the presence of several enlarged suckers of the dorsal row of the hectocotylus
distal part (character 2), it neatly differs from them because of the homomorphous tip of ventral arms
(character 1). Moreover the grouping of the enlarged suckers in S. boletzkyi sp. nov. diverges from that
of the other two species for the number of suckers in both the proximal and midway groups as well as.
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European Journal of Taxonomy 144: 1-12 (2015)
Table 3. Sequence, from proximal to distal, of sucker-sizes in the dorsal row of the part of the hectocotylus
distal to copulatory apparatus in Sepiola atlantica (after Guerra 1986), Sepiola tridens (after de Heij &
Goud 2010) and Sepiola boletzkyi sp. nov. Peculiarities of each species are highlighted in bold; the
enlarged suckers are followed by regularly tapering ones to the arm tip (Guerra 1986: fig. 3; de Heij &
Goud 2010: fig. 5; present work: Fig. 6A).
Sepiola atlantica 3-4 enlarged, 1-5 small, 3-4 enlarged
Sepiola tridens 3-4 enlarged, 2-3 small, 5-8 enlarged
Sepiola boletzkyi sp. nov, _ 2 small, 6 enlarged, 1 small, 2 enlarged
distinctively, because the first two suckers of the dorsal row are markedly small. Table 3 reports the
sequence of enlarged suckers in the three species with two groups of sucker-sizes.
With respect to (S. affinis + S. bursadhaesa + S. intermedia ), which have just one group of enlarged
suckers in the dorsal row of the distal part of the hectocotylus, S. boletzkyi sp. nov. lacks the groove on
the medial side of the same distal part.
Sepiola boletzkyi sp. nov. is easily distinguished from S. rondeletii and S. robusta based on the fact
they bear no enlarged suckers on the distal part of the hectocotylus and have 8 rows of suckers on the
tentacle club, i.e., 2 more than in S. boletzkyi sp. nov. Lastly S. boletzkyi sp. nov. differs mostly from
S. steenstrupiana because of the homomorphous tips of the ventral arms, the structure of the copulatory
apparatus, the presence of enlarged suckers on the distal part of the hectocotylus, and the occurrence of
6 rows of suckers instead of 4-5 on the tentacle club.
In addition to the abovementioned differences, the hectocotylus of S. boletzkyi sp. nov. is the only one
among the members of the S. atlantica group to have a quite large lobe in the dorsal sucker row next to
the copulatory apparatus tubercle.
The female S. boletzkyi sp. nov. may be discriminated from all other Sepiola species of the S. atlantica
group by its unique bursa copulatrix. It differs from all members of this group, except S. affinis and
S. rondeletii , because of the process projecting towards the mantle wall near the gill insertion; the
process in S. boletzkyi sp. nov., however, is fungiform and larger than in both other species. Furthermore
the bursa copulatrix of S. boletzkyi sp. nov. extends more, both anteriorly and posteriorly, than that of
S. affinis and, in comparison with S. rondeletii , has no caecum (cf. Naef 1923: figs 364 and 368).
The male and female specimens herein described as holotype and paratype, respectively, of S. boletzkyi
sp. nov. were assigned to the same specific entity thanks to the similarity with each other in characters
other than sexual, primarily the tentacle club, in addition to having been collected in the same place.
In the only available female (the paratype), an immature specimen, the bursa copulatrix reaches the
posterior end of the mantle cavity but probably it does not do so in the sexually mature female whose
ovary enlarges as oocytes mature.
Discussion
Sepiola boletzkyi sp. nov. is the ninth species of the genus to be discovered in the Mediterranean Sea. It
also adds to the six Sepiola species already recorded in the eastern Mediterranean Sea by Salman (2009).
As stressed in the section “Remarks”, the new species is assigned, because of its hectocotylus, to the
Sepiola atlantica group as defined by Naef (1923), who notably did not grant any genus-group rank to it.
We adhere to Naef’s concept and do not endorse any generic status for the atlantica group. The species in
10
BELLO G. & SALMAN A., Sepiola boletzkyi sp. nov. from the Aegean Sea
the group share, in addition to the general body bauplan , the synapomorphy of the typical hectocotylus
and particularly the peculiar copulatory apparatus. According to Naef (1923: 582), in these species
“the base of the hectocotylus bears 3 normal suckers, followed by a typical copulatory apparatus which
develops from the stalks of 4 suckers, the 2nd and 3rd sucker of the median [i.e. dorsal] row and the 3rd
and 4th of the lateral [i.e. ventral] row.” These four suckerless stalks are enlarged and fused together
to form a typical organ followed by the distal part of the arm that may bear some enlarged suckers in
the dorsal row (Tigs 1, 6A). At this moment the Sepiola atlantica group includes S. affinis, S. atlantica,
S. boletzkyi sp. nov., S. bursadhaesa, S. intermedia, S. robusta, S. rondeletii, S. steenstrupiana and S.
tr idens.
It is remarkable that three new Sepiola species, all belonging to the atlantica group - namely S. tridens ,
S. bursadhaesa and S. boletzkyi sp. nov. - have been discovered in the Atlantic-Mediterranean area in
the last few years, after a century-long lapse of time since Naef (1912a, 1912b, 1916, 1923) described
several new Mediterranean sepioline species and, most importantly, disentangled and re-shaped the
muddled systematics of the Sepiolinae.
The genus Sepiola displays a great diversity in the Atlantic-Mediterranean area, where it is by far the
most speciose cephalopod genus thanks to its 12 species. The reasons underlying this phenomenon
were discussed in Bello (2013). The fairly diverse copulatory organs in males (hectocotylus) and, less
conspicuously, in females (bursa copulatrix), supposedly give rise to sexual segregation and, therefore,
most probably play an important role in sepioline speciation. The discovery of S. boletzkyi sp. nov.
corroborates the hypothesis that the NE Atlantic-Mediterranean species of Sepiola are a suite of species
that largely arose from in situ speciation on either side of the Strait of Gibraltar after the vicariant
event(s) of (recurring) connection to the Atlantic and isolation of the Mediterranean (Bello 2013).
Sepiola boletzkyi sp. nov. is seemingly endemic to the eastern Mediterranean. Its discovery reinforces
the statement by Mangold & Boletzlcy (1988: 326) that “It is interesting that the only cephalopod species
exclusively known from the Mediterranean are among the Sepiolinae”. Indeed, as shown by Bello
(2013), all or almost all Mediterranean species in the genus Sepiola are endemic or quasi-endemic; on
the other side of the Strait, the NE Atlantic Sepiola species are also endemic to that area.
The description of Sepiola boletzkyi sp. nov. is based on two specimens only, a male and a female. The
collection of additional specimens is necessary to define both the variability of the markedly enlarged
suckers in the dorsal row of the hectocotylus distal part, in males, and the shape of the bursa copulatrix
in mated females. Moreover, new supplementary material would allow tissue sampling for genetic
analyses and DNA barcoding.
Acknowledgements
We thank Bahadir Onsoy (Sitki Ko 9 man University, Turkey) for producing the illustration of the
tentacle club and for improving the beak photographs, and Aldo Corriero and Rosa Zupa (Department
of Emergency and Organ Transplant, University of Bari, Italy) for their hospitality and assistance to the
first author with the stereo microscope photographs. We are most grateful to Chung-Cheng Lu (Museum
of Victoria, Melbourne, Australia and National Chung Hsing University, Taichung, Taiwan) for critically
reading and thoroughly reviewing our paper. We also acknowledge our debt to an anonymous reviewer
for her/his constructive suggestions.
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Manuscript received: 4 May 2015
Manuscript accepted: 12 August 2015
Published on: 12 October 2015
Topic editor: Rudy Jocque
Desk editor: Kristiaan Hoedemakers
Printed versions of all papers are also deposited in the libraries of the institutes that are members of
the EJT consortium: Museum National d’Histoire Naturelle, Paris, France; Botanic Garden Meise,
Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London,
United Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History
Museum of Denmark, Copenhagen, Denmark.
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