©
Zoological Institute, St.Petersburg, 2004
Description of two new species of tadpole-gobies
(Teleostei: Gobiidae: Benthophilus)
V.S. Boldyrev & N.G. Bogutskaya
Boldyrev, V.S. & Bogutskaya, N.G. 2004. Description of two new species of tadpolegobies (Teleostei: Gobiidae: Benthophilus). Zoosystematica Rossica, 13(1): 129-135.
Benthophilus durrelli and B. ragimovi, two new species of tadpole-gobies, are described.
Both belong to an assemblage of Benthophilus species with clearly differentiated dermal
ossifications. B. durrelli inhabits estuarine regions, deltas, lower reaches of rivers, and
dam lakes in the Sea of Azov basin and is introduced in the Volga. It was hitherto not
distinguished from the Azov tadpole-goby, B. stellatus, from which it differs in the posterior position of the second dorsal fin (behind vs. in front of the anal-fin origin), comparatively small (vs. considerable) difference in size of tubercles and granules, rectangular
dermal fold with slightly undulated margin (vs. triangular, with clearly undulated margin), narrow (vs. wide) first semiring-like blotch behind the first dorsal fin, and 17-21,
commonly 18-20 transverse rows of neuromasts on the flanks (vs. 20-25). B. ragimovi, a
deepwater species, is only recorded at western coast of middle and south Caspian from
Chechen’ Island to Astara. It differs from the closest congener, B. pinchuki, in the weak
(vs. well-developed) depression on the head, absence of granules in the head depression,
very small, densely-set granules located on whole upper head surface and anterior part of
back, sometimes also on caudal peduncle (vs. enlarged granules located only along the
lateral margins of the head depression), reduced first tubercles of the dorsal row (vs.
anterior dorsal tubercles of the same size as the posterior ones).
V.S. Boldyrev, Volgograd Division, State Institute of Lake and River Fisheries, ul. Pugachevskaya 1, Volgograd 400001, Russia, E-mail: bugs@t-k.ru
N.G. Bogutskaya, Zoological Institute, Russian Academy of Sciences, Universitetskaya
nab. 1, St.Petersburg 199034, Russia. E-mail: office@zin.ru
Introduction
Gobiid fishes of the genus Benthophilus (tadpolegobies) are found in the basins of the Caspian
and Black Sea including Sea of Azov. They are
small size species (maximum total length does
not exceed 150 mm) and may be found in fresh
and brackish waters with salinity up to 20‰ and
slightly higher. They have been never found in
areas with true marine salinity (over 30‰) being
typical inhabitants of deltas, estuaries and coastal waters of the brackish Caspian Sea and Sea of
Azov. Only some Caspian species inhabit deep
waters.
There are over 18 species of Benthophilus.
Most of them are poorly known and may easily
be misidentified. Most our knowledge of these
fishes came from papers by Iljin (1927a, 1927b,
1927c, 1930, 1938, 1949, 1956) and Ragimov
(1969, 1972, 1976, 1978, 1982, 1985a, 1985b,
1998). Quite recently, a summarizing review of
European Benthophilus has appeared (Miller,
2004) mainly based upon unpublished data by
Pinchuk. However, the taxonomic diversity of
Benthophilus appears to be still underestimated,
and taxonomic status of some forms or subspecies is pending wider morphological and genetic
studies (Pinchuk & Miller, 2004a).
As presently understood, the genus is apparently monophyletic (Pinchuk & Miller, 2004a). It is
defined by a number of autapomorphies. Among
them, a distinctively depressed head, absence of
normal scales but modification of dermal ossification into bony plates of different sizes and
shapes, reduction of the first dorsal fin to 1-4 unbranched rays and both dorsal fins widely interspaced, presence of a chin barbel and a dermal
fold just behind the corner of the mouth. The biology of species in this genus is characterized by a
distinct specialization, which we call “ephemery“.
Tadpole-gobies live about a year reaching maturity the following spawning season after a year of
hatching, or some individuals can attain maturity
at an age of 6-7 months (Iljin, 1927b; Ragimov,
1985b; Boldyrev, unpubl. data). All individuals die
after spawning, females earlier than males.
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V.S. Boldyrev & N.G. Bogutskaya: New species of Benthophilus • ZOOSYST. ROSSICA Vol. 13
Species of the genus may be roughly grouped
into two assemblages based upon the structure
of dermal ossifications. In the first group, bony
plates are not differentiated by size and generally small; each plate has a base embedded into
the skin and a portion with more or less developed thorns exposed to outside. In species of the
second group, bony plates are differentiated into
tubercles and granules. Tubercles are relatively
large polygonal bony scutes, which are spiny,
thorny or stellate and located on head and body;
they are usually arranged in longitudinal rows on
the body (dorsal and ventral ones, and 1 or 2 additional lateral rows). Granules are small bony
projections, which resemble dermal ossification
in species of the first group. In mature males,
granules and tubercles are reduced, and in spawning males of most species the body is completely
naked.
We have started a revision of the genus Benthophilus; first results of it revealed two new species described below.
Methods
Standard length (SL) is measured from the tip
of the upper jaw to the end of the hypural complex. Other measurements made point to point
are explained in the table. Last two branched rays
articulated on a single pterigiophore in second
dorsal and anal fin are noted as “1½”. Osteological characters and dermal ossification are examined in cleared-and-stained with alizarin red S
specimens and from radiographs. The number of
tubercles in the body rows does not include those
on the head and the bases of caudal fin rays. Since
a mature male looses tubercles, its head widens
and the fins lengthen, the descriptions are based
only on females and larger immature males.
Abbreviations: ZIN, Zoological Institute, Russian Academy of Sciences, St.Petersburg; ZIAz,
Institute of Zoology, National Academy of Sciences of Azerbaijan, Baku.
Benthophilus durrelli sp. n.
(Fig. 1)
Common name. Don tadpole-goby.
Holotype. M, ZIN 53211, SL 42.9 mm, Russia, Tsimlyansk Reservoir, 25.IV.2003, coll. V. Boldyrev.
Paratypes (all from Russia). 4 specs, ZIN 53212, SL
36.6-54.2 mm, same data as holotype; 7 specs, ZIN 9972,
SL 40-47 mm, Taganrog, 1892, coll. Tarnani; 7 specs,
ZIN 10167, SL 37-51.6 mm, same data; 5 specs, ZIN
44445, SL 47.2-54.5 mm, at Taganrog, VII.1909, coll.
Alferaki; 1 spec., ZIN 44449, SL 35.5, Azov Expedition,
26.VIII.1926, coll. B. Iljin; 4 specs, ZIN 53213, SL 40.248.8 mm, lower Don, VII.2003, coll. V. Boldyrev; 23
specs, ZIN 53214, SL 32.6-40.3 mm, Tsimlyansk Reservoir at Kalach-na-Donu, 10.IV.2004, coll. V. Boldyrev;
25 specs, ZIN 53215, SL 37-66 mm, same locality,
IV.2003, coll. V. Boldyrev.
Description. First dorsal fin with 3 or 4 unbranched rays. Second dorsal fin with 7½ or 8½
(rarely 6½ or 9½) branched rays. Anal fin with
7½ or 8½ (rarely 6½) branched rays. Origin of
second dorsal fin behind vertical through origin
of anal fin. Pectoral fin with (15)16-17 rays. Tubercles small but not reduced in size on head;
26-29, commonly 27-28, tubercles in dorsal row;
20-24 tubercles, commonly 21-23, in ventral row;
19-27 tubercles in upper lateral row markedly
reduced in size on caudal peduncle; 10 tiny tubercles in lower lateral row. One tubercle between
eyes. Dorsal tubercle with two radii of small
thorns on posterior surface. Granules not numerous, sparse on flanks and absent from caudal
peduncle. Chin barbel slightly compressed, with
thick base; its length exceeds half eye diameter.
Dermal fold large (length of its base up to two
eye diameters), clearly rectangular, with slightly
indulated margin. Mouth corner below anterior
eye margin. Number of transverse rows of neuromasts on flanks 17-21, commonly 18-20. Total number of vertebrae 28-29 (9 precaudal and
19-20 caudal).
Morphometric characters given in the Table.
General coloration variably pale. Dark blotches on back often not bright. Dark spot in front of
second dorsal fin present in about 50% of specimens. First semiring blotch not wide behind first
dorsal fin (not reaching base of first ray of second dorsal fin).
Size up to 66 mm SL. Total length up to
81.5 mm in Tsimlyansk Reservoir (our data) and
70 mm in Saratov Reservoir (Kudersky et al.,
1995).
Comparison. Benthophilus durrelli was hitherto not distinguished from the Azov tadpolegoby, B. stellatus (Kudersky et al., 1995; Evlanov
et al., 1998; Pinchuk & Miller, 2004c; etc.). When
compared to B. durrelli (see the diagnosis given
above), B. stellatus clearly differs in the anterior
position of the second dorsal fin (in front of the
anal-fin origin), comparatively great difference
in size between tubercles and granules, triangular dermal fold with a clearly undulated, oblique
free margin, absence of any additional dark spot
in front of the second dorsal fin, wide first semiring-like blotch behind the first dorsal fin (posterior margin of this blotch usually reaches the
base of the first ray of the second dorsal fin), and
20-25 transverse rows of neuromasts on the
flanks. B. stellatus and B. durrelli from the same
locality are also different in the size of adults:
SL 31-98 (41.7) mm and 22-47 (29.9) mm, respectively, in Tsimlyansk Reservoir (April 2002)
and 62-88 (75.1) mm and 22-62 (41) mm in lower Don (May 2003).
B. durrelli is morphologically very close to B.
mahmudbejovi from the North Caspian. Ragimov
ZOOSYST. ROSSICA Vol. 13 • V.S. Boldyrev & N.G. Bogutskaya: New species of Benthophilus
1a
1b
2a
2b
Figs 1, 2. Benthophilus, holotypes, dorsal (a) and lateral (b) views. 1, B. durrelli sp. n.; 2, B. ragimovi sp. n.
131
132
V.S. Boldyrev & N.G. Bogutskaya: New species of Benthophilus • ZOOSYST. ROSSICA Vol. 13
(1976: 1196) identified six specimens from the
Sea of Azov (ZIN 44445 and 44449, now paratypes of B. durrelli sp. n.) as B. mahmudbejovi,
and most subsequent authors followed him in
including the Sea of Azov into the range of B.
mahmudbejovi (Pinchuk & Miller, 2004b). These
two species are similar in having comparatively
small tubercles, narrow first semiring-like blotch
behind the first dorsal fin and the second dorsal
fin starting behind the vertical through the analfin origin. However, B. mahmudbejovi is different in having numerous and densely-set granules,
which cover the whole body (except for the abdominal area) including the caudal peduncle,
where granules are located between the dorsal
and upper lateral rows of tubercles. Besides, in
B. mahmudbejovi sparse granules are present on
the abdomen in front of the pelvics, transverse
rows of neuromasts on the flanks are more numerous (commonly 22-24), and a dorsal tubercle
bears one radius of thorns on its posterior surface instead of two.
Benthophilus species from the Sea of Azov,
Black Sea and North Caspian basins with differentiated dermal ossification are characterized by
a number of common features, among them are
the following ones: presence of tubercles on the
head (though very small in B. abdurahmanovi),
both paired and unpaired ones between the eyes
and in the middle of the posterior head surface;
24-30 tubercles in the dorsal row; commonly 3
neuromasts in the abdominal transverse rows; and
a well-developed chin barbel. A key is given below for identification of species from this group.
1. Dark brown saddles or blotches on back absent or
indistinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
– Three distinct large dark semiring-shaped blotches
or saddles along midline of back and flanks . . . . . . 4
2. Tubercles on head and body well distinguishable from
very small granules . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . B. macrocephalus (Pallas, 1787)
– Tubercles and granules on head very small and of
almost equal size, hardly distinguishable from each
other; only 1 or 2 of them are identifiable at the border between head and back; tubercles of dorsal row
increase in size from head to base of first dorsal; no
tubercle in middle of head posterior surface . . . . . . 3
3. Distance between mouth corners over 4.5 eye diameters; commonly 9½ rays in second dorsal and 8½
branched rays in anal fin . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . B. abdurahmanovi Ragimov, 1978
– Distance between mouth corners less than 4.5 eye
diameters; commonly 10½ rays in second dorsal and
9½ branched rays in anal fin . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . B. magistri Iljin, 1927
4. Groove between eyes along midline of head; tubercles behind eyes large; granules on head behind eyes
large (some of them almost of same size as nearby
tubercles) . . . . . . . . . . B. casachicus Ragimov, 1978
– No groove between eyes along midline of head; granules on head minute . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Two tubercles between eyes; upper jaw considerably
projected over lower jaw . . B. leobergius Berg, 1949
– One tubercle between eyes . . . . . . . . . . . . . . . . . . . 6
6. Origin of second dorsal fin behind vertical through
origin of anal fin . . . . . . . . . . . . . . . . . . . . . . . . . . 7
– Origin of second dorsal fin in front of vertical through
origin of anal fin . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7. Granules present between dorsal and upper lateral
rows of tubercles on whole body including caudal
peduncle; sparse granules present on abdomen in front
of pelvics; commonly 22-24 transverse rows of neuromasts on flanks . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . B. mahmudbejovi Ragimov, 1976
– Granules sparse on flanks and absent from caudal
peduncle; no granules in abdomen area; commonly
18-20 transverse rows of neuromasts . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . B. durrelli sp. n.
8. Head elongate, its width 77-91% of its length; first
semiring-like blotch not wide behind first dorsal fin
and never touches base of first ray of second dorsal
fin; commonly bright dark additional spot in front of
second dorsal fin present . . . . . B. nudus Berg, 1898
– Head widened, its width 94-104% of its length; first
semiring-like blotch wide behind first dorsal fin and
usually reaches base of first ray of second dorsal fin;
no additional spot in front of second dorsal fin . . . . .
. . . . . . . . . . . . . . . . . . . . B. stellatus (Sauvage, 1874)
Distribution. Gulf of Taganrog of the Sea of
Azov. In the River Don, from delta to Tsimlyansk
Reservoir. Probably in tributaries: Severskii Donetz and Manych (Solodovnikov, 1930; Vitkovsky, 2000; etc.). Unintentionally introduced (with
mysids introduction from Tsimlyansk Reservoir)
to Kuibyshev Reservoir on Volga, from which it
widely spread in Volga from Gorky Reservoir to
Volgograd Reservoir (Gavlena, 1973; Kudersky
et al., 1995; Evlanov et al., 1998; our data). B.
durrelli inhabits both fresh and brackish waters.
It is abundant in coastal areas of the Sea of Azov,
especially in estuarine regions at river mouths
with considerably freshened water (less than 1.53‰), lower reaches of rivers, and dam lakes. The
species prefers silty sand with mollusc shells.
Etymology. The species is named after Jerald
Durrell, famous English animal writer.
Benthophilus ragimovi sp. n.
(Fig. 2)
Benthophilus ctenolepidus – Berg, 1949: 1113, 1114
(part.: Figs. 852-855).
Benthophilus ctenolepidus ctenolepidus – Ragimov, 1982:
48 (western coast of Caspian Sea, up to Astara in the
south).
Common name. Ragimov’s tadpole-goby.
Holotype. M, ZIN 53216, SL 67.8 mm, Azerbaijan,
Caspian Sea, western coast, off Yamma-Kilyazi, depth
50 m, 27.VI.1984, coll. D. Ragimov.
Paratypes. Azerbaijan: 1 spec., ZIN 53217, SL 62.5
mm, same data as holotype; 3 specs, ZIN 23126, SL 63.576.5 mm, north of Apsheron Peninsula, Caspian Exped.,
11.II.1915, coll. N.M. Knipowitch; 8 specs, ZIAz 93, SL
66.4-73.5 mm, same data as holotype; 6 specs, ZIAz 467,
SL 63.4-72 mm, middle Caspian Sea, western coast,
X.1984, coll. D. Ragimov; 7 specs, ZIN 53218, SL 6578.3 mm, Andreyeva Bank, depth 150 m, 22.IX.1984,
ZOOSYST. ROSSICA Vol. 13 • V.S. Boldyrev & N.G. Bogutskaya: New species of Benthophilus
coll. D. Ragimov; 4 specs, ZIN 53219, SL 60.2-67.2 mm,
off Yamma-Kilyazi, 1984, coll. D. Ragimov.
Additional material (poorly preserved or not located
currently). Russia: 1 spec., ZIN 33122, SL 28.6 mm, off
Makhachkala, depth 17 m, Caspian Exped., 31.V.1912;
Azerbaijan: 3 specs, ZIAz 100, SL 28.6-58.3 mm, Shakhova Spit, depth 50 m, 30.VII.1985, coll. D. Ragimov;
5 specs, ZIAz 469, SL 66.4-79.6 mm, Andreyeva Bank,
depth 150 m, 27.IX.1984, coll. D. Ragimov.
Description. First dorsal fin with 4 unbranched
rays. Second dorsal fin with 9½ or, commonly,
10½ branched rays. Anal fin with 8½ or 9½ (rarely 10½) branched rays. Origin of second dorsal
fin on vertical through origin of anal fin. Pectoral fin with 15-17 rays. No tubercles on head.
First tubercles of dorsal row (in front of first dor-
133
sal fin) commonly reduced in size; 30-33, usually 31 or 32, tubercles in dorsal row; 24-25 tubercles in ventral row; over 20 tubercles in upper
lateral row markedly reduced in size on caudal
peduncle; 19 to 16 tiny tubercles in lower lateral
row. Granules very small, densely-set on whole
upper head surface and anterior part of back,
sometimes also on caudal peduncle between dorsal and upper lateral rows of tubercles. Chin barbel thin and short, its length less than half eye
diameter. Dermal fold small (its length less than
eye diameter), triangular. Mouth corner below
posterior half of eye. Head depression shallow,
often well-developed only between eyes. Number
of transverse rows of neuromasts on flanks 23-
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V.S. Boldyrev & N.G. Bogutskaya: New species of Benthophilus • ZOOSYST. ROSSICA Vol. 13
26, commonly 23-24. Total number of vertebrae
30-31 (9 precaudal and 21-22 caudal).
Morphometric characters given in the Table.
Coloration pale. No dark spots or blotches.
SL up to 80 mm. Maximum total length recorded is 94 mm (Ragimov, 1982).
Comparison. Benthophilus ragimovi sp. n. belongs to another group of Benthophilus species
with differentiated dermal ossifications. Species
of this group occur only in the middle and south
Caspian never entering fresh waters. In addition
to B. ragimovi sp. n., the group includes B. ctenolepidus Kessler, 1877, B. leptocephalus Kessler,
1877, and B. pinchuki Ragimov, 1982. They share
a number of characters, among them the complete
absence of tubercles on the upper head surface,
usually 4 abdominal transverse rows of neuromasts, and a poorly developed chin barbel.
B. pinchuki is the closest congener to B. ragimovi having a complete dorsal row (incomplete
in B. ctenolepidus and B. leptocephalus) with an
increased number of tubercles (31-33), a small
triangular dermal fold and comparatively numerous vertebrae (30-31 total vertebrae with 21 or
22 caudal ones). However, B. pinchuki differs in
the following characters: well-developed depression on the head between and behind the eyes;
absence of granules in the head depression; presence of granules only along the lateral margins
of the head depression and absence from the body
(if granules present on the back, they are relatively large, very sparse, and only located in front
of the first dorsal fin); tubercles in the dorsal row
comparatively large; anterior dorsal tubercles of
the same size as the posterior ones.
Distribution. Recorded off the western coast
of middle and south Caspian Sea from Chechen’
Island to Astara (Ragimov, 1965, 1982). Deepwater species.
Etymology. The species is named after the late
Dadash Ragimov, well-known ichthyologist, expert in Caspian gobiid fishes.
Acknowledgements
The project is supported by grant 02-04-49993 of the
Russian Foundation for Basic Research. We appreciate
very much the valuable help of the late Dr. D. Ragimov
who provided material from the ZIAz collection. We are
thankful to Dr. Sh. Ibragimov for his assistance during V.
Boldyrev’s stay in Baku, and to Dr. E.D. Vasil’eva (Zoological Museum, Moscow State University) and G. Volkova (ZIN) who helped us during our work in the collections under their care. Thanks also go to A.M. Naseka
(ZIN) for making photos.
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Received 2 November 2004