Two new freshwater fish species of the genus Telestes (Actinopterygii, Cyprinidae) from karst poljes in Eastern Herzegovina and Dubrovnik littoral (Bosnia and Herzegovina and Croatia)

Abstract

Two new species, Telestes dabar and Telestes miloradi, are described on the basis of morphological comparisons of isolated geographical populations of fishes identified earlier as Telestes metohiensis. A lectotype is designated for Telestes metohiensis, whose range is shown to include waters of Gatačko, Cerničko, and Nevesinjsko poljes in Eastern Herzegovina. Telestes dabar from Dabarsko Polje (Eastern Herzegovina) and Telestes miloradi from Konavosko Polje (south Croatia) share with Telestes metohiensis the following combination of characters that distinguish them from the rest of the genus Telestes: pharyngeal teeth in one row, usually 5-4; preoperculo-mandibular canal not communicating with the infraorbital canal; mouth subterminal, the tip of the mouth cleft on or below the level of the ventral margin of the eye; postcleithrum minute or absent; ventral portion of the trunk with a dark stripe on a pale background; and dorsal portion of trunk uniformly dark and bordered ventrally by a dark midlateral stripe. Telestes dabar and Telestes miloradi are distinguishable from Telestes metohiensis in usually having 8½ branched dorsal-fin rays (vs. usually 7½), 9 or 10 gill rakers (vs. 7-10, usually 8), and the dark stripe on the ventral portion of the trunk below the main pigmented area of the back narrow and usually not reaching posteriorly to the caudal peduncle (vs. dark stripe wide and extending posteriorly to the caudal peduncle). Telestes dabar is distinguished from Telestes miloradi by having scales on most of the body situated close to one another and overlapping in a region behind the pectoral girdle and usually on the caudal peduncle (vs. overlapping scales on most of the body); the lateral line usually incomplete and interrupted, with 24-69, usually 54-65, total scales (vs. lateral line usually complete, with 55-67 total scales); scales above and below the lateral line slightly smaller than lateral-line scales (vs. of about equal size); head width 43-52% HL (vs. 48-58% HL); and lower jaw length 10-12% SL or 36-41% HL (vs. 8-10% SL or 33-38% HL). Telestes miloradi, a very local endemic species,is known only by historical samples. Telestes dabar is an abundant fish in Dabarsko Polje, but its range is critically restricted during the dry season by a few permanent sources. Nothing is known about its occurrence in underground karst waters.

Full text

Two new freshwater sh species of the genus Telestes (Actinopterygii, Cyprinidae)... 53
Two new freshwater fish species of the genus Telestes
(Actinopterygii, Cyprinidae) from karst poljes in
Eastern Herzegovina and Dubrovnik littoral
(Bosnia and Herzegovina and Croatia)
Nina G. Bogutskaya1,†, Primož Zupančič2,‡, Ivan Bogut3,§, Alexander M. Naseka1,|
1 Zoological Institute, Russian Academy of Sciences, 1 Universitetskaya Emb., St Petersburg 199034, Russia
2 Dolsko 14, 1262 Slovenia 3 Agronomski fakultet u Sveučilišta u Mostaru, Zavod za ribarstvo, zoologiju i
zaštitu voda, Biskupa Čule 10, 88000 Mostar, Bosnia and Hercegovina
† urn:lsid:zoobank.org:author:F6957EF1-366E-484E-BF40-211E17A93A1B
‡ urn:lsid:zoobank.org:author:0DC4A949-8AF7-4E80-8305-FDF271318A36
§ urn:lsid:zoobank.org:author:248F52CE-3AC6-4AA4-9EA2-FB5BCD95D7EF
| urn:lsid:zoobank.org:pub:D89D42DE-5B28-48B7-9431-2D603C985CAF
Corresponding author: Nina G. Bogutskaya (nbogutskaya@rambler.ru)
Academic editor: C. Baldwin|Received 23 September 2011|Accepted 19 March 2012|Published 5 April 2012
urn:lsid:zoobank.org:pub:D89D42DE-5B28-48B7-9431-2D603C985CAF
Citation: Bogutskaya NG, Zupančič P, Bogut I, Naseka AM (2012) Two new freshwater sh species of the genus Telestes
(Actinopterygii, Cyprinidae) from karst poljes in Eastern Herzegovina and Dubrovnik littoral (Bosnia and Herzegovina
and Croatia). ZooKeys 180: 53–80. doi: 10.3897/zookeys.180.2127
Abstract
Two new species, Telestes dabar and T. miloradi, are described on the basis of morphological comparisons
of isolated geographical populations of shes identied earlier as T. metohiensis. A lectotype is designated
for T. metohiensis, whose range is shown to include waters of Gatačko, Cerničko, and Nevesinjsko poljes
in Eastern Herzegovina. Telestes dabar from Dabarsko Polje (Eastern Herzegovina) and T. miloradi from
Konavosko Polje (south Croatia) share with T. metohiensis the following combination of characters that
distinguish them from the rest of the genus Telestes: pharyngeal teeth in one row, usually 5–4; preoperculo-
mandibular canal not communicating with the infraorbital canal; mouth subterminal, the tip of the mouth
cleft on or below the level of the ventral margin of the eye; postcleithrum minute or absent; ventral por-
tion of the trunk with a dark stripe on a pale background; and dorsal portion of trunk uniformly dark
and bordered ventrally by a dark midlateral stripe. Telestes dabar and T. miloradi are distinguishable from
T. metohiensis in usually having 8½ branched dorsal-n rays (vs. usually 7½), 9 or 10 gill rakers (vs. 7–10,
ZooKeys 180: 53–80 (2012)
doi: 10.3897/zookeys.180.2127
www.zookeys.org
Copyright Nina G. Bogutskaya et al. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0
(CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Nina G. Bogutskaya et al. / ZooKeys 180: 53–80 (2012)
54
usually 8), and the dark stripe on the ventral portion of the trunk below the main pigmented area of the
back narrow and usually not reaching posteriorly to the caudal peduncle (vs. dark stripe wide and extending
posteriorly to the caudal peduncle). Telestes dabar is distinguished from T. miloradi by having scales on most
of the body situated close to one another and overlapping in a region behind the pectoral girdle and usually
on the caudal peduncle (vs. overlapping scales on most of the body); the lateral line usually incomplete
and interrupted, with 24–69, usually 54–65, total scales (vs. lateral line usually complete, with 55–67 total
scales); scales above and below the lateral line slightly smaller than lateral-line scales (vs. of about equal
size); head width 43–52% HL (vs. 48–58% HL); and lower jaw length 10–12% SL or 36–41% HL (vs.
8–10% SL or 33–38% HL). Telestes miloradi, a very local endemic species, is known only by historical sam-
ples. Telestes dabar is an abundant sh in Dabarsko Polje, but its range is critically restricted during the dry
season by a few permanent sources. Nothing is known about its occurrence in underground karst waters.
Keywords
Cypriniformes, West Balkans, Alpha Taxonomy and new taxa
Introduction
Karst basins in Croatia and Bosnia and Herzegovina are populated by many local en-
demic species most of them belonging to the cyprinid subfamily Leuciscinae (Mrakovčić
et al. 2006; Kottelat and Freyhof 2007; Jelić et al. 2008; Zupančič 2008). Among leu-
ciscine cyprinids of the Dinaric karst, most numerous is a group of species that was
formerly assigned to the genus Phoxinellus Heckel, 1843. In a revision of this group
(Zupančič and Bogutskaya 2002; Bogutskaya and Zupančič 2003) ten Phoxinellus spe-
cies were divided into two groups, one with two subgroups. Later (Freyhof et al. 2006),
based on nuclear and mtDNA sequences, Phoxinellus was found to be paraphyletic
because it includes three unrelated monophyletic units. e scientic name Phoxinellus
was therefore restricted to species having plain colouration, a small or absent post-
cleithrum, no genital papilla, and an almost entirely naked body (species included: type
species P. alepidotus Heckel, 1843, P. dalmaticus Zupančič & Bogutskaya, 2000, and P.
pseudalepidotus Bogutskaya & Zupančič, 2003). Species having an irregularly spotted
colour pattern, a large postcleithrum, an increased number of precaudal anal-n ptery-
giophores, and a large female genital papilla were assigned to a new genus, Delminichthys
Freyhof, Lieckfeldt, Bogutskaya, Pitra & Ludwig, 2006 (species included: type species
D. adspersus (Heckel, 1843), D. ghetaldii (Steindachner, 1882), D. jadovensis Zupančič
& Bogutskaya, 2002, and D. krbavensis Zupančič & Bogutskaya, 2002). Finally, species
having a small or absent postcleithrum, no genital papilla, a dark midlateral stripe from
the head to the caudal peduncle, and non-overlapping scales were assigned to Telestes (T.
croaticus (Steindachner, 1866), T. fontinalis (Karaman, 1972), and T. metohiensis (Stein-
dachner, 1901)), bringing the total number of species of Telestes to 11. ese generic
assignments were later supported by a molecular phylogenetic study based on the same
mitochondrial but dierent nuclear markers (Perea et al. 2010).

Two new freshwater sh species of the genus Telestes (Actinopterygii, Cyprinidae)... 55
Zupančič and Bogutskaya (2002) and Bogutskaya and Zupančič (2003) rede-
scribed T. metohiensis based on 251 specimens (including syntypes) from Nevesinjsko
Polje, Gatačko Polje, Cerničko Polje, and Dabarsko Polje karsts in Bosnia and Her-
zegovina, and from Ljuta River in Konavle Region [Konavosko Polje] in southern
Croatia. Bogutskaya and Zupančič (2003: 375) mentioned that the syntypes from
the Ljuta River were distinguished by having 8½ branched anal-n rays (vs. usually
7½ in most other syntypes). Specimens from Dabarsko Polje (Vrijeka and Opačica
rivers) were also only tentatively identied as metohiensis because of having overlap-
ping scales (vs. non-overlapping, narrowly to widely spaced) and usually 8½ branched
anal-n rays (Zupančič and Bogutskaya 2002: 417). Since 2003, we have examined
additional material of the entire T. metohiensis species complex, and two species – one
from Ljuta River in Konavosko Polje and one from Dabarsko Polje – are described
here as new.
Methods
Measurements were made point to point to the nearest 0.1 mm and follow methods
used by Bogutskaya and Zupančič (2010) for Squalius species. In most aspects, they
coincide with the scheme by Kottelat and Freyhof (2007: g. 1), but a few clarica-
tions of the latter are as follows. Standard length (SL) was measured from the anteri-
ormost point of the upper lip (not of the snout) to the end of the hypural complex.
Head length (HL) was measured from the anteriormost point of the upper lip to the
posteriormost point of the opercular membrane. Interorbital width was measured
including the skin fold. e term ‘length of dorsal n’ is used for the length of the
dorsal-n base, and the term ‘depth of dorsal n’ is used for the length of the longest
ray of the dorsal n. Respective terms are used for the measurements of the anal n.
e last two branched rays articulating on a single pterygiophore in the dorsal and
anal ns are noted as “1½”. Total lateral-line scale count includes all pored scales,
from the rst one just behind the posttemporal bone to the posteriormost one located
on the bases of the caudal-n rays. Total number of lateral-row scales includes all
scales, pored and non-pored, from the rst one just behind the posttemporal bone to
the posteriormost one located on the bases of the caudal-n rays. Osteological char-
acters were examined from dissections and from radiographs. Statistical analyses were
performed with Microsoft Excel and Statistica 6.0 packages.
Abbreviations used: NMW, Naturhistorisches Museum, Wien; SMNH, Slovenian
Museum of Natural History; PZC, Collection of P. Zupančič, Dolsko (Slovenia); SMF,
Senckenberg Museum, Frankfurt a. Main; ZISP, Zoological Institute, Russian Acad-
emy of Sciences, St Petersburg; ZMH, Zoologisches Museum und Institut, Universität
Hamburg; (cephalic sensory canals) CIO, infraorbital canal; CPM, preoperculo-man-
dibular canal; CSO, supraorbital canal; and CST, supratemporal canal.

Nina G. Bogutskaya et al. / ZooKeys 180: 53–80 (2012)
56
Results
Telestes dabar sp. n.
urn:lsid:zoobank.org:act:4A50A07B-57BD-472B-9B00-6941D8C6779A
http://species-id.net/wiki/Telestes_dabar
Figs 1a, 2a, 3a
Holotype. NMW 95295, 79.1 mm SL, BOSNIA & HERZEGOVINA: Dabarsko
Polje, Opačica River at Potkom, 43°5.9'N, 18°7.6'E, 15 Sept. 2006, coll. Zupančič.
Paratypes. NMW 95300, 6, 55.2–71.1 mm SL, same data as holotype; PZC 525,
72, 32.5–51.3 mm SL, same data as holotype; PZC 526, 13, 48.0–73.8 mm SL, same
data as holotype; PZC 565, 21, 35.5–56.7 mm SL, same locality, 8 July 2011; ZISP
54995, 15, 38.1–58.9 mm SL, same locality, 8 July 2011; SMNH 444, 35.5–60.8 mm
SL, same locality, 8 July 2011; PZC 279, 12, 44.2–71.9 mm SL, BOSNIA & HER-
ZEGOVINA: Dabarsko Polje, Vrijeka River, 24 May 2001; PZC 521, 13, 54.0–62.6
mm SL, same locality as 279, 15 Sept. 2004; PZC 575, 18, 40.5–69.8 mm SL, same
locality as 279, 15 Sept. 2006.
Diagnosis. Telestes dabar is distinguished from T. metohiensis and T. miloradi by
having the following combination of characters: slightly curved dark stripe (obvious in
live and preserved specimens) present from just behind operculum to vertical just ante-
rior to origin of anal n, this stripe narrow and separated from dark pigmented area on
back along its entire length; scales on most of body not overlapping but situated close
to one another; scales overlapping behind pectoral girdle along lateral line and usually
Figure 1. a Telestes dabar, holotype, female, 79.1 mm SL, NMW 95295, Bosnia & Herzegovina:
Opačica River, Dabarsko Polje b Telestes metohiensis, female, 82.1 mm SL, PZC 293, Bosnia & Herzego-
vina: Zovidolka River (Zalomka River system), Nevesinjsko Polje.
a
b

Two new freshwater sh species of the genus Telestes (Actinopterygii, Cyprinidae)... 57
on caudal peduncle; snout with eshy tip projecting over upper lip; mouth subtermi-
nal with tip of mouth cleft at or below level of ventral margin of eye; lateral line usually
interrupted, with 24–69 total lateral-line scales; branched dorsal-n rays usually 8½;
branched anal-n rays usually 8½; gill rakers 9 or 10; total vertebrae 39–41, mode 40;
abdominal vertebrae 22–24, mode 22; caudal vertebrae 16–18, mode 17; head width
43–52% HL; and lower jaw long, length 10–12% SL.
Description. Morphometric data are summarised in Table 1a, selected counts in
Tables 2–4. General appearance can be seen in Figs 1a and 2a. Body compressed, elon-
Figure 2. a Telestes dabar, male paratype, live specimen, ZISP 54995, 58.9 mm SL, Bosnia & Herzegovina:
Opačica River, Dabarsko Polje b Telestes metohiensis, live specimen, male, 86.2 mm SL, PZC 567, Bosnia &
Herzegovina: spring Ljeskovik in Zalomka River, Nevesinjsko Polje c Telestes metohiensis, live specimen, male,
84.5 mm SL, PZC 566, Bosnia & Herzegovina: Zovidolka River (Zalomka River system), Nevesinjsko Polje.
c
a
b

Nina G. Bogutskaya et al. / ZooKeys 180: 53–80 (2012)
58
gate. Caudal-peduncle depth only slightly less than half maximum body depth; head
length greater than maximum body depth. Eye small, its diameter smaller than snout
length. Snout eshy, slightly to markedly projecting beyond upper lip (similar to a
feature Kottelat and Freyhof [2007: g. 39] called the “rostral cap,” which covers all or
part of upper lip); snout terminating laterally in prominent crease along anterior edge
of rst infraorbital. Mouth subterminal, tip of mouth cleft at level of ventral margin
of eye or, more frequently, below it. Lower jaw-quadrate junction at vertical through
anterior half of eye. Length of lower jaw 10–12% SL or 36–41% HL, or 102–132%
depth of operculum.
Dorsal n with 7½ (9 specimens), 8½ (151) or 9½ (1) branched rays, 8½ in holo-
type. Dorsal-n origin above posterior end of pelvic-n base. Anal n with 8½ (153)
or 9½ (8) branched rays, 8½ in holotype. Outer margin of anal n slightly concave or
almost straight. Caudal n moderately forked, lobes weakly pointed, with 9+8 princi-
pal branched rays.
Total gill rakers (Table 2) 9 (20) or 10 (20), 10 in holotype. Pharyngeal teeth 5–4,
hooked, slightly serrated (examined in 5 specimens).
Scales covering entire body including pre-pectoral area and abdomen, non-over-
lapping on most parts of body but overlapping in triangular-shaped area just behind
pectoral girdle and usually on caudal peduncle at least behind anal n (Fig. 3a); lateral-
line scales always overlapping, sometimes a few posteriormost scales not overlapping.
Scales irregularly set but close to one another. Most ank scales oval, somewhat deeper
than long; scales on caudal peduncle more elongated (longer than deep) having promi-
nent posterior attenuation. Trunk scales smaller than lateral-line scales but not con-
Figure 3. a Telestes dabar, alizarin stained specimen, PZC 575, 63.7 mm SL, Bosnia & Herzegovina:
Vrijeka River, Dabarsko Polje b Telestes metohiensis, alizarin stained specimen, 63.3 mm SL, PZC 312,
Bosnia & Herzegovina: spring Ljeskovik in Zalomka River, Nevesinjsko Polje.
a
b

Two new freshwater sh species of the genus Telestes (Actinopterygii, Cyprinidae)... 59
Table 1a. Morphometric data of T. dabar.
T. dabar
Holo-
type females, n=26 males, n=13
 min max mean SD min max M SD
SL, mm 79.1 60.3 81.8 67.8 55.5 69.0 61.1
Maximum body depth (% SL) 21.6 21.6 26.0 23.6 1.3 21.7 25.0 22.6 0.9
Depth of caudal peduncle (% SL) 9.8 9.8 12.4 11.0 0.7 10.0 12.4 11.2 0.7
Depth of caudal peduncle
(% length of caudal peduncle) 47.9 47.7 59.6 53.6 4.3 44.6 62.2 52.9 5.2
Maximum body width (% SL) 13.4 11.4 16.0 13.7 1.2 12.0 22.0 14.2 2.6
Predorsal length (% SL) 55.3 53.5 58.7 56.9 1.1 53.8 57.3 55.5 1.2
Postdorsal length (% SL) 36.3 32.8 36.6 35.2 1.0 34.4 37.2 35.8 1.0
Prepelvic length (% SL) 51.6 50.4 55.5 52.7 1.4 50.6 54.4 52.2 1.0
Preanal length (% SL) 71.2 68.3 73.2 70.8 1.3 68.6 71.9 69.9 1.0
Pectoral – pelvic-n origin length (% SL) 24.5 24.5 27.7 25.6 0.9 22.3 26.5 24.1 1.2
Pelvic – anal-n origin length (% SL) 20.2 17.2 20.2 19.0 0.6 16.6 20.3 18.4 1.3
Length of caudal peduncle (% SL) 20.6 18.5 21.9 20.5 0.8 19.2 23.5 21.2 1.1
Dorsal-n base length (% SL) 10.3 9.3 12.3 10.9 0.8 9.9 12.2 11.2 0.7
Dorsal n depth (% SL) 18.9 16.6 20.2 18.6 1.4 18.5 21.8 19.7 0.9
Anal-n base length (% SL) 12.1 9.2 12.1 10.6 0.8 10.3 12.4 11.2 0.5
Anal n depth (% SL) 12.3 11.9 14.8 13.2 0.8 13.4 17.2 15.9 1.1
Pectoral n length (% SL) 19.9 18.1 21.2 19.9 0.8 22.5 26.7 24.2 1.3
Pelvic n length (% SL) 14.8 12.9 16.3 14.9 0.9 16.3 17.7 16.9 0.5
Head length (% SL) 27.1 25.6 28.7 27.1 0.9 26.6 29.6 28.1 1.0
Head length (% body depth) 125.1 104.0 125.1 115.3 5.9 110.8 135.1 124.3 7.6
Head depth at nape (% SL) 17.4 17.1 19.3 18.1 0.7 17.0 18.8 18.2 0.5
Head depth at nape (% HL) 64.2 62.4 70.7 66.8 2.3 60.6 69.3 65.0 2.7
Maximum head width (% SL) 13.5 11.6 14.6 13.6 0.7 12.4 14.5 13.6 0.6
Maximum head width (% HL) 50.0 42.8 52.3 50.1 2.4 44.7 52.1 48.3 2.4
Maximum cranial width
(% cranium roof length) 68.9 61.9 72.6 65.8 3.1 59.7 68.8 64.4 2.5
Snout length (% SL) 8.5 7.4 9.2 8.1 0.5 7.8 9.0 8.5 0.4
Snout length (% HL) 31.4 27.7 32.8 29.7 1.3 28.5 31.7 30.3 1.1
Eye horizontal diameter (% SL) 6.6 5.9 7.2 6.6 0.4 6.3 7.8 6.8 0.4
Eye horizontal diameter (% HL) 24.4 22.1 26.8 24.5 1.5 22.1 26.8 24.3 1.4
Eye horizontal diameter
(% interorbital width) 74.5 66.3 81.9 73.3 4.7 69.5 82.0 74.4 4.7
Postorbital distance (% HL) 53.0 48.8 54.0 52.2 1.4 47.1 53.0 50.2 1.9
Interorbital width (% SL) 8.9 8.2 9.6 9.1 0.4 8.5 9.9 9.2 0.4
Interorbital width (% HL) 32.8 31.3 36.0 33.4 1.2 29.0 35.6 32.7 1.9
Length of upper jaw (% HL) 28.6 24.5 30.5 27.8 1.9 26.2 29.5 27.7 1.1
Length of upper jaw (% SL) 7.7 6.6 8.5 7.5 0.6 7.0 8.6 7.8 0.5
Length of lower jaw (% SL) 10.6 9.5 11.2 10.2 0.5 9.7 11.5 10.9 0.5
Length of lower jaw (% HL) 39.2 35.8 40.7 37.6 1.3 35.6 41.1 38.6 1.6
Length of lower jaw (% interorbital width) 119.7 106.4 123.4 112.6 4.4 110.2 129.8 120.2 6.4
Length of lower jaw
(% depth of operculum) 110.2 102.4 120.0 108.2 4.5 105.7 131.7 116.6 8.2

Nina G. Bogutskaya et al. / ZooKeys 180: 53–80 (2012)
60
Table 1b. Morphometric data of T. miloradi.
T. miloradi
Holo-
type females, n=9 males, n=3
 min max mean SD min max Mean SD
SL, mm 66.7 34.1 66.7 57.9 61.8
Maximum body depth (% SL) 22.4 21.7 25.8 23.5 1.5 19.6 24.6 23.0 2.9
Depth of caudal peduncle (% SL) 11.4 10.4 11.6 11.2 0.4 9.6 11.3 10.8 1.0
Depth of caudal peduncle
(% length of caudal peduncle) 53.8 49.8 54.8 52.4 2.1 41.8 52.8 49.1 6.4
Maximum body width (% SL) 14.3 12.9 15.3 14.6 0.7 11.4 16.8 15.0 3.1
Predorsal length (% SL) 55.9 55.9 57.7 57.0 0.7 54.3 54.9 54.5 0.3
Postdorsal length (% SL) 35.2 32.8 35.3 34.3 1.0 35.4 35.6 35.5 0.1
Prepelvic length (% SL) 51.6 51.5 53.3 52.3 0.9 50.0 50.2 50.1 0.1
Preanal length (% SL) 72.8 68.2 72.8 70.3 1.9 68.2 68.4 68.3 0.1
Pectoral – pelvic-n origin length (% SL) 25.1 25.1 27.6 26.2 1.1 23.3 25.0 24.4 1.0
Pelvic – anal-n origin length (% SL) 20.8 16.6 20.8 18.3 1.7 17.6 18.4 18.2 0.5
Length of caudal peduncle (% SL) 21.2 20.5 22.1 21.3 0.7 21.4 23.0 22.0 0.9
Dorsal-n base length (% SL) 11.6 9.9 11.6 10.9 0.7 11.2 12.6 12.2 0.8
Dorsal n depth (% SL) 19.6 19.2 20.9 19.8 0.8 20.9 22.1 21.4 0.6
Anal-n base length (% SL) 12.4 10.7 12.4 11.9 0.6 11.8 11.9 11.8 0.1
Anal n depth (% SL) 13.2 0.0 15.0 12.5 4.7 13.7 15.5 14.3 1.0
Pectoral n length (% SL) 21.7 18.5 21.7 20.8 1.0 23.5 25.4 24.8 1.1
Pelvic n length (% SL) 15.6 13.6 15.6 14.6 0.8 16.9 17.6 17.2 0.4
Head length (% SL) 26.8 25.4 28.0 27.0 0.8 26.5 29.0 28.2 1.4
Head length (% body depth) 119.9 105.0 119.9 115.0 6.0 117.6 135.4 123.5 10.3
Head depth at nape (% SL) 18.3 16.9 18.3 17.6 0.5 16.7 19.0 18.2 1.4
Head depth at nape (% HL) 68.1 62.5 68.1 65.1 2.3 62.9 65.7 64.7 1.6
Maximum head width (% SL) 14.1 12.3 14.5 14.0 0.7 12.6 16.9 15.4 2.5
Maximum head width (% HL) 52.5 48.7 53.6 52.0 1.5 47.5 58.2 54.6 6.2
Maximum cranial width
(% cranium roof length) 62.8 59.6 73.9 66.0 5.0 58.3 65.2 62.9 4.0
Snout length (% SL) 9.0 8.0 9.7 8.9 0.5 8.5 8.6 8.6 0.0
Snout length (% HL) 33.7 31.4 34.7 33.0 1.2 29.5 32.4 30.5 1.7
Eye horizontal diameter (% SL) 6.5 6.3 6.9 6.5 0.2 6.6 6.7 6.7 0.0
Eye horizontal diameter (% HL) 24.4 22.5 26.4 24.2 1.4 22.9 25.2 23.7 1.3
Eye horizontal diameter
(% interorbital width) 70.2 64.7 83.0 71.3 5.9 69.4 83.4 74.1 8.1
Postorbital distance (% HL) 49.1 48.9 51.6 50.2 1.2 50.2 51.0 50.5 0.5
Interorbital width (% SL) 9.3 8.1 9.7 9.2 0.5 8.0 9.6 9.1 0.9
Interorbital width (% HL) 34.8 31.9 34.8 34.1 0.9 30.2 33.1 32.1 1.7
Length of upper jaw (% HL) 27.9 26.0 28.1 27.2 0.9 23.5 25.1 24.0 0.9
Length of upper jaw (% SL) 7.5 6.6 7.9 7.3 0.4 6.7 6.8 6.8 0.1
Length of lower jaw (% SL) 9.3 8.4 10.4 9.5 0.6 8.7 10.4 9.8 1.0
Length of lower jaw (% HL) 34.7 33.1 38.4 35.3 1.8 32.9 35.9 34.9 1.7
Length of lower jaw (% interorbital width) 99.8 99.8 114.3 103.7 6.1 108.5 109.1 108.7 0.4
Length of lower jaw
(% depth of operculum) 104.0 95.6 104.0 101.1 2.9 95.6 107.1 103.3 6.7

Two new freshwater sh species of the genus Telestes (Actinopterygii, Cyprinidae)... 61
Table 1c. Morphometric data of T. metohiensis.
T. metohiensis, Gatačko and Cerničko poljes
lectotype females, n=25 males, n=10
 min max mean SD min max mean SD
SL, mm 87.9 60.26 102.1 74.9 54.4 75.7 63.1
Maximum body depth (% SL) 23.9 19.9 25.1 22.5 1.4 20.7 24.1 22.5 1.0
Depth of caudal peduncle (% SL) 11.3 9.7 11.9 10.7 0.5 10.4 11.8 11.2 0.5
Depth of caudal peduncle
(% length of caudal peduncle) 61.5 46.7 61.5 53.0 4.1 46.7 55.7 51.0 3.1
Maximum body width (% SL) 16.0 11.1 16.4 14.3 1.7 14.1 15.6 14.8 0.6
Predorsal length (% SL) 57.6 49.4 60.1 56.5 2.2 53.7 57.7 55.7 1.3
Postdorsal length (% SL) 33.4 31.0 36.5 33.7 1.6 32.5 38.0 35.2 1.6
Prepelvic length (% SL) 52.0 46.0 55.9 52.7 1.9 50.4 53.8 51.9 1.3
Preanal length (% SL) 69.1 65.6 73.6 70.2 1.9 67.1 69.7 68.5 0.9
Pectoral – pelvic-n origin length
(% SL) 27.5 24.7 29.7 27.0 1.3 24.1 27.2 25.5 1.1
Pelvic – anal-n origin length (% SL) 19.8 16.4 20.1 18.5 1.1 15.6 18.2 17.0 0.8
Length of caudal peduncle (% SL) 18.3 19.1 22.3 20.3 1.2 20.9 22.8 22.0 0.6
Dorsal-n base length (% SL) 11.1 9.5 11.5 10.8 0.6 10.8 13.6 12.0 1.0
Dorsal n depth (% SL) 17.6 14.9 18.3 17.2 0.8 16.3 19.6 18.5 1.1
Anal-n base length (% SL) 11.3 9.1 12.0 10.9 0.7 10.0 13.2 11.9 1.1
Anal n depth (% SL) 12.3 12.4 15.3 13.7 0.8 12.8 15.4 14.3 1.0
Pectoral n length (% SL) 18.4 17.0 19.8 18.4 0.8 22.7 25.3 23.6 0.8
Pelvic n length (% SL) 12.9 12.4 15.3 13.7 0.8 15.4 17.7 16.6 0.7
Head length (% SL) 27.0 23.8 28.2 26.6 1.1 26.1 28.2 27.3 0.6
Head length (% body depth) 112.9 111.0 132.5 120.4 6.6 114.4 131.8 122.8 5.2
Head depth at nape (% SL) 17.4 15.5 18.7 17.3 0.8 15.9 18.1 17.3 0.7
Head depth at nape (% HL) 64.6 62.9 71.8 65.1 2.2 58.5 66.1 63.5 2.7
Maximum head width (% SL) 14.8 12.9 15.7 14.8 0.7 14.5 15.7 14.9 0.4
Maximum head width (% HL) 54.9 50.6 59.3 55.7 2.7 52.8 56.9 54.7 1.6
Maximum cranial width
(% cranium roof length) 63.5 62.9 78.6 68.6 4.4 65.4 71.7 68.2 2.7
Snout length (% SL) 8.1 7.3 8.8 8.2 0.4 8.5 9.4 8.9 0.4
Snout length (% HL) 30.0 27.9 33.1 30.8 1.6 31.2 34.2 32.5 1.1
Eye horizontal diameter (% SL) 6.0 4.6 6.9 5.9 0.5 5.3 6.8 6.0 0.4
Eye horizontal diameter (% HL) 22.4 19.4 25.2 22.1 1.5 19.5 24.8 22.1 1.8
Eye horizontal diameter
(% interorbital width) 70.7 57.0 80.2 68.2 6.9 60.2 71.8 65.1 4.3
Postorbital distance (% HL) 51.9 47.9 55.6 51.9 2.4 46.2 53.1 50.8 2.3
Interorbital width (% SL) 8.5 7.7 9.2 8.7 0.4 8.4 9.8 9.3 0.5
Interorbital width (% HL) 31.6 29.6 34.8 32.6 1.7 30.8 35.8 34.0 1.5
Length of upper jaw (% HL) 27.8 27.0 31.0 29.2 1.0 27.0 30.9 29.0 1.1
Length of upper jaw (% SL) 7.5 6.4 8.3 7.8 0.4 7.5 8.4 7.9 0.3
Length of lower jaw (% SL) 9.9 8.3 11.3 10.3 0.8 9.5 11.0 10.3 0.6
Length of lower jaw (% HL) 36.7 35.0 42.2 38.7 2.2 34.8 41.2 37.8 2.1
Length of lower jaw
(% interorbital width) 116.0 106.5 132.5 120.6 6.4 101.6 121.9 113.7 6.9
Length of lower jaw
(% depth of operculum) 107.2 106.5 117.9 112.5 3.0 109.6 119.2 112.7 3.5

Nina G. Bogutskaya et al. / ZooKeys 180: 53–80 (2012)
62
Table 1d. Morphometric data of T. metohiensis.
T. metohiensis (anis nomen museale), Nevesinjsko Polje
lectotype females, n=33 males, n=14
 min max mean SD min max M sd
SL, mm 87.9 57.8 113.8 79.5 57.7 86.2 70.4
Maximum body depth (% SL) 23.9 19.3 24.1 21.8 1.1 19.8 23.8 21.6 1.4
Depth of caudal peduncle (% SL) 11.3 9.0 12.0 10.7 0.7 9.6 12.3 11.1 0.7
Depth of caudal peduncle
(% length of caudal peduncle) 61.5 42.7 60.0 54.2 4.5 43.6 59.9 51.8 4.2
Maximum body width (% SL) 16.0 11.6 17.6 14.5 1.4 11.0 16.6 14.3 1.7
Predorsal length (% SL) 57.6 54.2 59.4 57.1 1.3 54.4 58.4 56.3 1.0
Postdorsal length (% SL) 33.4 29.9 36.4 34.0 1.3 33.8 36.5 35.2 0.7
Prepelvic length (% SL) 52.0 51.6 55.1 53.3 1.0 50.1 54.6 51.6 1.2
Preanal length (% SL) 69.1 67.8 73.4 70.9 1.3 65.9 70.3 68.8 1.3
Pectoral – pelvic-n origin length (%
SL) 27.5 24.2 30.0 25.9 1.2 21.7 25.4 23.8 1.0
Pelvic – anal-n origin length (% SL) 19.8 16.7 44.1 19.4 4.6 15.6 19.9 17.5 1.3
Length of caudal peduncle (% SL) 18.3 18.0 21.3 19.7 0.9 20.0 23.2 21.4 0.8
Dorsal-n base length (% SL) 11.1 9.4 12.9 11.1 0.8 8.1 12.1 10.4 1.2
Dorsal n depth (% SL) 17.6 15.8 20.4 17.9 1.4 15.9 21.8 18.0 1.5
Anal-n base length (% SL) 11.3 9.4 12.0 10.9 0.6 10.5 13.2 11.4 0.7
Anal n depth (% SL) 12.3 10.6 15.0 13.1 1.1 12.0 16.2 14.0 1.4
Pectoral n length (% SL) 18.4 18.0 21.9 20.2 1.0 21.8 26.4 23.8 1.4
Pelvic n length (% SL) 12.9 12.5 15.6 14.1 0.8 14.2 17.5 15.7 0.9
Head length (% SL) 27.0 26.7 29.4 27.8 0.8 26.3 30.1 28.0 1.1
Head length (% body depth) 112.9 114.5 142.6 127.4 6.6 116.5 142.9 130.0 8.5
Head depth at nape (% SL) 17.4 16.5 18.7 17.6 0.6 15.7 18.8 17.3 0.9
Head depth at nape (% HL) 64.6 60.0 67.9 63.4 1.8 57.1 65.9 62.0 2.9
Maximum head width (% SL) 14.8 13.7 16.6 14.9 0.6 13.3 16.2 14.7 0.8
Maximum head width (% HL) 54.9 50.5 61.2 53.7 2.3 51.0 57.4 53.0 2.4
Maximum cranial width
(% cranium roof length) 63.5 64.9 79.6 72.5 4.3 66.3 75.8 69.1 2.8
Snout length (% SL) 8.1 7.9 9.8 8.7 0.5 7.7 9.3 8.6 0.4
Snout length (% HL) 30.0 29.4 33.4 31.3 1.1 27.2 33.0 30.8 1.6
Eye horizontal diameter (% SL) 6.0 4.7 7.7 6.1 0.7 5.4 8.0 6.3 0.7
Eye horizontal diameter (% HL) 22.4 17.3 27.3 21.9 2.5 20.6 26.4 22.5 2.0
Eye horizontal diameter
(% interorbital width) 70.7 52.0 86.7 67.5 8.3 59.8 92.5 73.1 11.9
Postorbital distance (% HL) 51.9 49.7 57.2 53.3 1.9 48.1 54.4 51.2 1.8
Interorbital width (% SL) 8.5 8.0 10.0 9.0 0.6 7.5 9.9 8.7 0.8
Interorbital width (% HL) 31.6 29.7 35.9 32.5 1.9 26.2 34.8 31.2 2.7
Length of upper jaw (% HL) 27.8 26.7 30.5 28.7 0.9 27.4 29.8 28.5 0.8
Length of upper jaw (% SL) 7.5 7.2 8.7 8.0 0.4 7.3 8.9 8.0 0.5
Length of lower jaw (% SL) 9.9 10.0 12.2 10.7 0.5 10.0 12.5 11.0 0.9
Length of lower jaw (% HL) 36.7 36.3 43.8 38.5 1.5 36.3 44.0 38.9 2.5
Length of lower jaw
(% interorbital width) 116.0 110.0 133.8 121.2 6.0 109.1 138.7 123.0 9.8
Length of lower jaw
(% depth of operculum) 116.5 110.8 133.8 116.1 4.9 111.5 132.3 118.5 7.0

Two new freshwater sh species of the genus Telestes (Actinopterygii, Cyprinidae)... 63
siderably so. All scales well ossied, usually visible without staining. In live specimens,
scales clearly visible because of some silver highlights (Fig. 2a). Lateral line complete (2
specimens), long but incomplete (5) or interrupted (33) as in specimen in Fig. 3a; if
interrupted, gaps typically comprising absence of a few scales in a few places, 24–69 in
total (Table 3), 65 with one interruption in holotype. Lateral line making clear curva-
ture above anal-n origin. Number of scales in total lateral series 62–69 (modal range
65–67), 68 in holotype.
Parietal segment of CSO lacking. CPM not communicating with CIO, terminat-
ing over the upper margin of opercular antedorsal process. CSO complete with 8,
rarely 7 or 9, pores. CIO complete with 14–17 pores and with 4 canal openings on
rst infraorbital. CPM complete or interrupted between the angulo-articular and pre-
operculum and/or between preoperculum and operculum, CPM with 14–17 pores (4,
rarely 5, canal openings on dentary, and 7–9, usually 8, canal openings on preopercu-
lum). CST complete, with 5–7 pores or narrowly interrupted in middle.
Total vertebrae (Table 4) 39 (49), 40 (101) or 41 (11), 40 in holotype; abdominal
vertebrae 22 (101), 23 (58) or 24 (2), 22 in holotype; caudal vertebrae 16 (9), 17 (82)
or 18 (70), 18 in holotype; predorsal vertebrae 13 (24), 14 (126) or 15 (11), 14 in
holotype; intermediate vertebrae 3 (123) or 4 (38), 3 in holotype. Most frequent verte-
bral formulae 22+17 (41), 22+18 (60) and 23+17 (40), 22+18 in holotype.
Colouration. In live specimens, dark back contrasting sharply with pale area
below lateral midline, even in small specimens. Black midlateral stripe extending
from head to caudal peduncle forming ventral border of darkly pigmented area on
Table 2. Gill-raker counts in Telestes dabar, T. miloradi and T. metohiensis.
7 8 9 10 mean SD
T. dabar 20 20 9.5 0.51
T. miloradi 1 11 6 9.3 0.57
T. metohiensis, Gatačko Polje 2 33 15 1 8.3 0.58
T. metohiensis, Nevesinjsko
Polje 6 28 12 1 8.2 0.67
Table 3. Total lateral-line scale and total lateral-series scales counts in Telestes dabar, T. miloradi and
T. metohiensis.
Total lateral-line scales Total lateral-series
scales
0–53 54–57 58–61 62–65 66–69 70–73 mean SD range mean SD
T. dabar 5 17 8 7 3 54.2 9.58 62–69 66.6 2.41
T. miloradi 6 5 5 2 60.3 3.91 58–69 62.8 2.93
T. metohiensis,
Gatačko Polje 11 10 25 5 61.0 3.92 56–71 63.5 3.74
T. metohiensis,
Nevesinjsko Polje 1 9 11 19 5 2 61.1 5.32 60–71 65.0 3.07

Nina G. Bogutskaya et al. / ZooKeys 180: 53–80 (2012)
64
Table 4a. Vertebral counts in Telestes species endemic in Croatia and Bosnia and Herzegovina. Total
vertebrae.
38 39 40 41 42 mean SD
T. dabar 49 101 11 39.8 0.56
T. miloradi 1 10 11 40.5 0.60
T. metohiensis, Gatačko Polje 6 33 12 2 39.2 0.68
T. metohiensis, Nevesinjsko Polje 10 112 35 39.2 0.52
T. croaticus 5 21 38.8 0.40
T. fontinalis 1 7 3 39.2 0.60
T. karsticus 2 2 3 41.1 0.89
T. polylepis 2 9 6 41.2 0.66
T. turskyi 2 9 6 1 40.3 0.77
T. ukliva 8 42.0 0
Table 4b. Vertebral counts in Telestes species endemic in Croatia and Bosnia and Herzegovina. Vertebral
formulae.
21+17 22+16 22+17 22+18 22+19 23+16 23+17 23+18 23+19 24+16 24+17 24+18 24+19
T. dabar 41 60 8 40 10 1 1
T. miloradi 1 6 1 4 10
T. metohiensis,
Gatačko Polje 6 28 2 5 10 2
T. metohiensis,
Nevesinjsko
Polje
7 58 12 33 24
T. croaticus 2 3 16 5
T. fontinalis 1 1 7 3
T. karsticus 1 1 2 1 2
T. polylepis 1 1 8 4 1 2
T. turskyi 14 1561
T. ukliva 3 5
back. Another black lateral stripe occurring more ventrally, on otherwise pale ventral
portion of trunk; this stripe extending from eye or opercle (or just behind oper-
cle) to at least vertical through point halfway between origins of pelvic and anal
ns, sometimes extending as poorly coalesced spots onto caudal peduncle. Dash-like
black marking present along internal procurrent rays of caudal-n dorsal lobe, and
elongate black blotch present at bases of 3rd–7th branched rays of dorsal n. Black
pigment also occurring on rays of dorsal and caudal ns, but its intensity varying
among individuals. Peritoneum black. is general pattern of pigmentation retained
in formaldehyde-xed and ethanol-preserved specimens. Live specimens collected
from May through September, both males and females, exhibiting yellowish-orange
pigment at bases of all ns, especially pectoral and anal ns, and yellowish pigment
on iris and along anterior, dorsal and posterior margins of operculum. Colouration
of specimens in cold season unknown.

Two new freshwater sh species of the genus Telestes (Actinopterygii, Cyprinidae)... 65
Table 4c. Vertebral counts in Telestes species endemic in Croatia and Bosnia and Herzegovina. Abdominal, caudal and predorsal vertebrae in Telestes dabar, T. mi-
loradi and T. metohiensis.
Abdominal vertebrae Caudal vertebrae Predorsal vertebrae
21 22 23 24 Mean SD 15 16 17 18 19 mean SD 12 13 14 15 mean SD
T. dabar 101 58 2 22.4 0.51 9 82 70 17.4 0.59 24 126 11 13.9 0.46
T. miloradi 8 14 22.6 0.49 5 16 1 17.8 0.50 15 7 14.3 0.5
T. metohiensis,
Gatačko Polje 36 17 22.3 0.47 7 42 4 16.9 0.46 3 38 12 14.2 0.50
T. metohiensis,
Nevesinjsko Polje 1 92 64 22.4 0.51 2 43 105 7 16.7 0.56 2 124 31 14.2 0.42
T. croaticus 2 19 5 22.1 0.52 8 18 16.7 0.47 21 5 14.2 0.40
T. fontinalis 2 9 22.8 0.40 7 4 16.4 0.50 2 9 14.8 0.40
T. karsticus 1 4 2 23.1 0.69 1 5 1 18.0 0.58 4 3 13.4 0.53
T. polylepis 1 13 3 23.1 0.49 2 11 4 18.1 0.60 1 16 13.9 0.24
T. turskyi 5 13 22.7 0.46 1 6 10 1 17.6 0.62 16 2 13.1 0.32
T. ukliva 3 5 23.6 0.52 5 3 18.4 0.52 2 5 1 12.9 0.64

Nina G. Bogutskaya et al. / ZooKeys 180: 53–80 (2012)
66
Sexual dimorphism. Genital papilla absent in both males and females. Most mor-
phometric characters not signicantly dierent between males and females (Table 1a)
with ve exceptions. In males, distance between origins of pectoral and pelvic ns long-
er than in females (P<0.0001), dorsal n deeper (P<0.02), anal n deeper (P<0.0001),
pectoral n longer (P<0.0001), pectoral n often reaching pelvic-n origin in males,
and pelvic n longer (P<0.0001), pelvic n often reaching anal-n origin in males.
Figure 4. Map of distribution of Telestes dabar (diamond), T. miloradi (star) and T. metohiensis (square)
in karst elds of Eastern Herzegovina and Dubrovnik littoral; white circles show ponors and white circles
with black dots - springs.

Two new freshwater sh species of the genus Telestes (Actinopterygii, Cyprinidae)... 67
In samples collected in May, ripe males with small but prominent conical breeding
tubercles. Tubercles regularly covering entire body, including dorsal and ventral sur-
faces of caudal peduncle, except for ventralmost surface of head. Single tubercle located
on each scale. On all ns (except for caudal n), tubercles present on both sides along
all rays and on n membrane, being particularly dense along marginal rays. Tubercles
forming rows along outer margins of operculum and pectoral n; tubercles in those
rows larger than others on body. Degree of tubercle development varying between
males with regard to both size of tubercles and their location. Tubercles always present
on head, back, and pectoral n. Males retaining tubercles, though reduced in size and
density, until September.
Distribution. e new species is known from two rivers, Vrijeka and Opačica, in
the Dabarsko Polje of Eastern Herzegovina in Bosnia and Herzegovina (Fig. 4).
Habitat and biology. From May through September T. dabar is found in shal-
low water of those river sections that are adjacent to and lled from underground
Figure 5. a Habitat of Telestes dabar: Opačica River at Potkom, Dabarsko Polje (type locality) b Opačica
River 100 m away from the spring (8 July 2011) c–d habitat of Telestes metohiensis: Nevesinjsko Polje;
c – spring Ljeskovik in Zalomka River (8 July 2011) d Zovidolka River at Udbine (8 July 2011). All in
Bosnia and Herzegovina.
a
c
d
b

Nina G. Bogutskaya et al. / ZooKeys 180: 53–80 (2012)
68
springs. ere is no current, and the water is clean (Fig. 5a). Females with eggs and
just-spent females were caught on 24 May 2001 in Vrijeka River and mature males
and just-spent females on 31 May 2000 in Opačica River. e size of the ripe eggs
was 1.3–1.7 mm in diameter. In all examined samples females predominate. e
smallest spent female was 45.0 mm SL, and the smallest ripe male 43.7 mm SL. No
other shes were caught in Opačica together with T. dabar while D. ghetaldii were
collected in Vrijeka.
Etymology. e specic name, dabar, refers to the type locality, Dabarsko, or Da-
bar Polje; it is a noun in apposition.
Telestes miloradi sp. n.
urn:lsid:zoobank.org:act:9A45C6CE-B7BF-4593-BAF6-0D10C45C78EA
http://species-id.net/wiki/Telestes_miloradi
Fig. 7
Holotype. NMW 95296 (ex 51169), 66.7 mm SL; CROATIA: stream Ljuta at Gruda
[misspelt Grinda in Steindachner (1901: 197)), 1901, coll. Kolombatovič.
Paratypes. NMW 51169, 13 (? syntypes [now paralectotypes] of Paraphoxinus
metohiensis), 31.4–62.6 mm SL, same data as holotype; NMW 51170, 4 (syntypes
[now paralectotypes] of Paraphoxinus metohiensis), 57.9–66.4 mm SL, same data as
holotype; NMW 51171, 3 syntypes [now paralectotypes] of Paraphoxinus metohiensis,
74.6–83.1 mm SL, same data as holotype; NMW 51173, 1 syntype [paralectotype] of
Paraphoxinus metohiensis, 119.3 mm SL, same data as holotype.
Diagnosis. Telestes miloradi is distinguished from T. metohiensis and T. dabar
by having the following combination of characters: slightly curved, relatively nar-
row dark stripe (obvious in most preserved specimens) present on ventral portion of
trunk from just behind operculum to vertical at or anterior to origin of anal n, this
stripe separated from dark pigmented area on back along its entire length; scales on
most of body not overlapping; mouth subterminal with tip of mouth cleft at or be-
low level of ventral margin of eye; snout not eshy; lateral line complete with 55–67
total scales; branched dorsal-n rays 8½; branched anal-n rays 8½; gill rakers usu-
ally 8–10, mode 9; total vertebrae usually 40 or 41; abdominal vertebrae 22–23,
mode 23; caudal vertebrae 16–18, mode 18; head width 48–58% HL, and lower jaw
short, length 8–10% SL.
Description. Morphometric data are summarised in Table 1b, selected counts
in Tables 2–4. General appearance can be seen in Figs 7a and 7b. Body compressed,
elongate. Caudal peduncle depth equal to or only slightly less than half maximum
body depth; head length greater than maximum body depth. Eye small, its diam-
eter smaller than snout length. Snout not eshy, rostral cap covering only part of
upper lip, at least in preserved specimens. Mouth subterminal, tip of mouth cleft

Two new freshwater sh species of the genus Telestes (Actinopterygii, Cyprinidae)... 69
at level of ventral margin of eye or, more frequently, below it. Lower jaw-quadrate
junction at vertical through anterior half of eye. Length of lower jaw 8–10% SL
or 33–38% HL, or 96–107% depth of operculum (equal to depth of operculum
on average).
Dorsal n with 8½ branched rays. Dorsal-n origin above posterior end of pelvic-
n base. Anal n with 8½ branched rays. Outer margin of anal n slightly concave.
Caudal n moderately forked, lobes weakly pointed, with 9+8 principal branched rays.
Total gill rakers (Table 2) 8 (1 specimen), 9 (11) or 10 (6), 10 in holotype. Pharyngeal
teeth 5–4, hooked, slightly serrated (examined in 5 specimens).
Scales covering entire body including pre-pectoral area and abdomen, overlap-
ping on most parts of body. Scales regularly set; lateral-line scales and scales above and
below it of about equal size. Lateral line complete (Table 3), 55–67 scales in total, 61
in holotype. Lateral line not curving above anal-n origin. Number of scales in total
lateral series 58–67 (modal range 62–64), 63 in holotype.
Parietal segment of CSO lacking. CPM not communicating with CIO, terminat-
ing over upper margin of opercular antedorsal process or communicating with CIO
(on one side in 3 specimens). CSO complete with 8, rarely 7 or 9, pores. CIO com-
plete with 14–16 pores and with 4 canal openings on rst infraorbital. CPM complete,
with 14–16 pores (4 canal openings on dentary, and 7–9, usually 8, on preoperculum).
CST complete, with 5 or 7 pores.
Total vertebrae (Table 4) 39 (1), 40 (10) or 41 (11), 41 in holotype; abdominal
vertebrae 22 (8) or 23 (14), 23 in holotype; caudal vertebrae 17 (5), 18 (16) or 19 (1),
18 in holotype; predorsal vertebrae 13 (15) or 14 (7), 14 in holotype; intermediate ver-
tebrae 3 (12) or 4 (9), 3 in holotype. Vertebral formulae 22+17 (1), 22+18 (7), 23+17
(5) and 23+18 (10), 23+18 in holotype.
Colouration. In preserved specimens, dark back contrasting sharply with pale area
below lateral midline. Dark midlateral stripe extending from head to caudal peduncle
forming ventral border of darkly pigmented region on back(faded in some specimens).
Another, more conspicuous, dark lateral stripe, occurring on ventral portion of trunk, nar-
row and not extending posterior to vertical through origin of anal n. Peritoneum dark.
Sexual dimorphism. Genital papilla absent in both males and females. Most mor-
phometric characters not signicantly dierent between males and females (Table 1).
In the three male specimens examined, dorsal n deeper than in females (P=0.0105);
pectoral n longer (P<0.001), pectoral n nearly reaching pelvic-n origin in males
and well short of pelvic n in females; and pelvic n longer (P<0.0001), pelvic n
almost reaching anal-n origin in males, well short of anal n in females.
Distribution. e new species is known from Ljuta River in Konavosko Polje, also
called Konavoska Ljuta, of Dubrovnik littoral (Fig. 4). Only historical NMW samples
are known to us.
Etymology. e species is named for Milorad Mrakovčić, Zagreb, in recognition
of his many contributions to the study of freshwater shes in the Adriatic basin.

Nina G. Bogutskaya et al. / ZooKeys 180: 53–80 (2012)
70
Comparative remarks
Telestes dabar, T. miloradi, T. metohiensis, T. croaticus, and T. fontinalis are distinguished
from all congeners by having the pharyngeal teeth in one row, 5–4 or 5–5 (vs. usually
2.5–5.2 or 2.5–4.2), having the preoperculo-mandibular canal terminating in a free
pore at the upper margin of the opercular antedorsal process and not communicating
with the infraorbital canal (vs. communicating), and in lacking a postcleithrum (vs.
postcleithrum present). All ve species possess a dark midlateral stripe from the head
to the caudal peduncle that forms the ventral border of the darkly pigmented area on
the back. is feature is similar to that in some other Telestes species (Kottelat and
Freyhof 2007: 282–289). Within this group, T. dabar, T. miloradi, and T. metohiensis
dier from T. croaticus and T. fontinalis in having an additional black lateral stripe
occurring on the otherwise pale ventral portion of the trunk. is character was con-
sidered unique for T. metohiensis (Kottelat and Freyhof 2007: 284). In T. dabar and T.
miloradi this ventral stripe (Fig. 1a, 2a, 7) is narrow and extends from just behind the
operculum maximally to a vertical through the origin of the anal n. In T. metohiensis,
the stripe (Fig. 1b, 2b–c) is wide and usually extends posteriorly to the caudal peduncle
where it merges with the main pigmented area. e pale area between the dark area on
the back and the ventral stripe varies in length and depth, being the smallest in females
(Fig. 1b, 2b–c).
Besides the presence of the ventral stripe, Telestes dabar and T. miloradi are fur-
ther distinguishable from T. croaticus and T. fontinalis by usually having 8½ branched
dorsal-n rays (vs. usually 7½). Telestes dabar diers from T. croaticus by usually having
40 total vertebrae (vs. usually 39) (Table 4); a maximum head width of 42–52% HL
(averaging 50% HL in females and 48% HL in males), which is considerably smaller
than the head depth at nape, 61–71% HL (averaging 67% HL in females and 65% HL
in males) (vs. the maximum head width only slightly smaller than the head depth or
about equal to it); and a smaller size, up to 82 mm SL (vs. up to 160 mm). Telestes da-
bar can be further distinguished from T. fontinalis by having 5–4 pharyngeal teeth (vs.
5–5); a usually long, though slightly incomplete and narrowly interrupted, lateral line
that reaches the posterior half of the caudal peduncle and has 24–69, usually 54–65,
total scales (vs. a short, incomplete, and widely interrupted lateral line terminating in
the area between the pectoral and anal ns with usually 23–37 total scales); usually 17
or 18 caudal vertebrae (vs. 16); usually 13 or 14 predorsal vertebrae (vs. 15) (Table 4);
3 or 4 intermediate vertebrae (vs. 5); and a moderately compressed body without any
ventral keel (vs. a markedly laterally compressed body and a scaled ventral keel in front
of the pelvic ns).
Telestes miloradi further diers from T. croaticus and T. fontinalis in usually having
a long, complete lateral line with 55–67 scales vs. an often incomplete and interrupted
lateral line with (18–45)51–70 and (17)23–37(56) scales, respectively. e new spe-
cies further diers from T. croaticus in having 8–10, most frequently 9, gill rakers, (vs.
8–9, most frequently 8); usually 40 or 41 total vertebrae (vs. 38 or 39); 22–23, mode
23, abdominal vertebrae (vs. 21–23, mode 22); and 17–19, mode 18, caudal vertebrae

Two new freshwater sh species of the genus Telestes (Actinopterygii, Cyprinidae)... 71
(vs. 16–17, mode 17) (Table 4). Telestes miloradi can be further distinguished from T.
fontinalis by having 5–4 pharyngeal teeth (vs. 5–5); usually 40 or 41 total vertebrae (vs.
38–40, usually 39); 17–19, usually 18, caudal vertebrae (vs. 16–17, often 16) (Table
4); 3 or 4 intermediate vertebrae (vs. 5); and a moderately compressed body without a
ventral keel (vs. a markedly laterally compressed body and a scaled ventral keel in front
of the pelvic ns).
Telestes miloradi diers from T. croaticus, T. fontinalis, T. metohiensis, and T. dabar
in having comparatively well-overlapped scales, especially on the anterior part of the
ank and on the caudal peduncle; and scales of about the same size in the lateral line
and above and below it (vs. scales usually non-overlapping on most of the body, and
scales above and below the lateral line smaller than the lateral-line scales). e scale
pattern in T. miloradi is very similar to the pattern found in most leuciscine shes, e.g.
T. karsticus Marčić & Mrakovčić, 2011 (Fig. 8), T. turskyi (Heckel, 1843), T. ukliva
(Heckel, 1843). e presence of overlapping scales is a plesiomorphic feature for the
Leuciscinae (Bogutskaya 1990, 1991), as is an interconnection of CPM and CIO (e.g.,
T. karsticus). In addition to overlapping scales, a few specimens of T. miloradi exhibit
interconnected CPM and CIO.
In this study, we compared specimens of T. metohiensis from Gatačko, Cerničko,
and Nevesinjsko poljes and found that specimens from the Nevesinjsko Polje (labeled
as anis by Steindachner although the name anis was never published), though
somewhat dierent in usually having smaller and more scattered scales, are similar
to T. metohiensis from the Gatačko and Cerničko poljes in all other aspects examined
by us (Tables 1c–d, 2–4). We suppose that a reason for Steindachner’s opinion could
be the dierences evident between the NMW specimens of T. metohiensis from the
Gatačko Polje (Mušnica River, Gacko) and those from the Nevesinjsko Polje (Zalomka
River) (Fig. 6). Additional material examined by us revealed that the dierences noted
above in scales may be size-dependent: larger specimens of T. metohiensis in both poljes
usually have more densely set scales. A specimen of T. metohiensis from Gatačko Polje,
NMW 51176:3 (87.9 mm SL) is designated here as lectotype to ensure taxonomic
stability in the event that T. metohiensis from the Nevesinjsko Polje is recognised as
taxonomically distinct in the future.
Both T. dabar and T. miloradi are distinguishable from T. metohiensis by usually
having 8½ branched dorsal-n rays (vs. usually 7½: 7½ found in 192 specimens and
8½ in 22); 39–41, modes 40 and 41, total vertebrae (vs. 38–40, rarely 41, mode 39);
16–19, modes 17 and 18, caudal vertebrae (vs. 15–17, usually 16 or 17) (vs. 15–17,
usually 16 or 17) (Table 4; dierences are statistically signicant at P<0.0001); and
more numerous gill rakers, 9 or 10 (vs. (7)8–9(10), most frequently 8) (Table 2). Te l -
estes dabar further diers from T. metohiensis in usually having an interrupted lateral
line with 24–69, usually 54–65, total scales (vs. usually complete with 58–65 scales)
(Table 3; dierence is statistically signicant at P<0.0001). e scale pattern also dis-
tinguishes T. dabar from T. metohiensis. In T. dabar, the scales (Fig. 3a) are densely set,
but they do not overlap on most parts of the body except behind the pectoral girdle
and on the caudal peduncle; the scales above and below the lateral line are only slightly

Nina G. Bogutskaya et al. / ZooKeys 180: 53–80 (2012)
72
smaller than the lateral-line scales; and the scales are oval, somewhat deeper than long,
on the anks and elongated, longer than deep with a prominent posterior attenuation,
on the caudal peduncle. In T. metohiensis the scales are more or less widely spaced and
do not overlap on the entire body except for the lateral line; this feature can be seen in
live specimens, Fig. 2b); the scales (Fig. 3b) above and below the lateral line usually are
considerably smaller than the lateral-line scales; and the scales are almost circular on
both the anks and the caudal peduncle. Telestes dabar is further distinguished from
T. metohiensis by having a black-and-white general colouration except for yellowish-
orange pigmentation of the n bases and yellow pigmentation in the iris and along the
operculum in adults (vs. yellowish-green or greenish-bronze pigmentation of the whole
body and ns in both young and adults, Fig. 2b–c). With regard to the morphometric
features, T. dabar is rather similar to T. metohiensis, diering only in having a narrower
head, maximum head width 12–15% SL or 43–52% HL and maximum cranial width
60–73% cranium roof length (vs. 13–17% SL or 51–59% HL, and 64–80% cranium
roof length) (Table 1; dierences are statistically signicant at P<0.0001).
Besides the characters mentioned above, Telestes dabar diers from T. miloradi in
having the lateral-line scales larger than the scales above and below it (vs. of about
equal size); usually an interrupted lateral line making a sharp curvature upward above
the anal-n base (vs. complete and making no sharp curvature); the length of lower jaw
10–12% SL, 36–41% HL, 102–132% depth of operculum (vs. 8–10% SL, 33–38%
HL, 96–107% depth of operculum) (Table 1; dierences are statistically signicant at
P<0.0001).
Figure 6. Telestes metohiensis. a NMW 51176:3, lectotype, 87.9 mm SL, ‘Mušica’ [Mušnica] River b
NMW 51090 (labelled as anis nomen museale), 57.9 mm SL, Zalomska [Zalomka River].
a
b

Two new freshwater sh species of the genus Telestes (Actinopterygii, Cyprinidae)... 73
Comments on the distribution and conservation of T. dabar, T. miloradi and T.
metohiensis
e three Telestes species are known from four of the 13 main karst poljes located in
Eastern Herzegovina (Bosnia and Herzegovina) and in Dubrovnik littoral area (Croa-
tia) (Fig. 4). ese karst poljes are part of a high-karst geotectonic unit known as Di-
naric Karst, which consists of Mesozoic carbonate formations. e depth of soluble and
highly karstied rocks here exceeds 3000 m (Milanović 1981, 2006). A polje (means
“eld” in many Slavic languages) is a large closed depression draining underground
with a at oor. Its streams may be permanent, intermittent and perennial, and, in
natural conditions, a polje is subject to periodic ooding and becomes a lake. In East-
ern Herzegovina and in the Dubrovnik littoral area stepwise poljes are distributed from
a
b
Figure 7. Telestes miloradi, Croatia: Ljuta River at Gruda, Konavosko Polje a Holotype, male, 66.7 mm
SL, NMW 95296 b paratype, female, 119.3 mm SL, NMW 51173.
Figure 8. Telestes karsticus, PZC 504, 62.6 mm SL, Croatia: Sušik River, Danube drainage.

Nina G. Bogutskaya et al. / ZooKeys 180: 53–80 (2012)
74
60 m up to 1,080 m above sea level (Fig. 4). Streams and rivers appear from temporary
or permanent springs and sink underground through swallow holes called ponors. In
general, hydro-systems of all poljes under consideration (Fig. 4), except for the Kona-
vosko Polje with direct connection to the Adriatic Sea and the Polje Gradac with no
springs or surface ows, belong to the Neretva drainage area and form a complex net of
underground ows. Within the Neretva drainage, the poljes belong to catchments of
the Buna River (Slato and Nevesinjsko), the Bregava River (Lukavačko and Dabarsko),
and the Trebišnijca River (Gatačko, Fatničko and others). At present, no one polje has
a direct groundwater (surface) ow connection with Neretva or its tributaries. His-
torically, interconnections between variable surface streams and between them and the
main Neretva course occurred during dierent geological epochs. Fish distributions
can provide some evidence of this. Conversely, very local distribution of some shes
may indicate the isolation of some surface drainage systems for a long time.
Slato [Zlato] Polje (1,020‒1,080 m above sea level) is situated at the highest el-
evation of all Eastern Herzegovina poljes. Ćurčić (1915a) reported nding no shes
at Slato Polje, and no shes are known from later literature or from museum collec-
tions. e Slato Polje is connected with Nevesinjsko Polje through a narrow valley
that is now dry. e Nevesinjsko Polje, the largest polje in Eastern Herzegovina, has a
surface area of 170 km2 and is located at an elevation from 870 m to 800 m above sea
level. e lowest point is Biograd Ponor, which is the terminus of the Zalomka River
that starts at Raščelica near the Gatačko Polje. is river has a permanent ow only
between Fojnica and Črni Kuk. Along the river bed downstream from Črni Kuk there
are a lot of ponors. e most prominent leakage zone is in the Rilja section where the
Zalomka is active only 213 days per year, on average (Milanović 1981, 2006). In the
warm season almost the entire river bed of the Zalomka within the Nevesinjsko Polje
is dry, and shes are found in its upper section only. Only four species are known from
there: Salmo sp., Squalius cf. squalus, Squalius svallize Heckel & Kner, 1858, Phoxinus
sp. (PZ personal observations). Ćurčić (1915a) reported that Paraphoxinus metohienis
was the most numerous species around Fojnica, but at present only Phoxinus sp. was
found there by PZ. Further downstream, T. metohiensis occurs in those very short river
sections that are adjacent to permanent springs such as Ljeskovik. is species also oc-
curs in upper reaches of the Zavidolka River that temporarily ows to the Zalomka in
the east from the Biograd Ponor (our data). Delminichthys ghetaldii is absent from the
Nevesinjsko Polje and the Zalomka system.
Dabarsko Polje, about 20 km long and 1 to 3 km wide, is located close to the
Nevesinjsko Polje but isolated from it. e Dabarsko Polje lies more than 400 m of
elevation below the Nevesinjsko Polje. At present, the Dabarsko Polje is a closed basin
without a possibility for surface runo. All waters of the Dabarsko Polje catchment
ow through underground karst conduits toward the springs of the Bregava River,
though historically the Polje drained to the Bregava River canyon that is now dry. e
lowest point is the Ponikva Ponor (471 m above sea level), the terminus of a single
permanent stream in the polje, the Vrijeka River, which is only 2.5 km long (Milanović
1981, 2006). e Opačica River located in the northwestern part of the polje is longer

Two new freshwater sh species of the genus Telestes (Actinopterygii, Cyprinidae)... 75
but intermittent. Only two native species occur in Dabarsko Polje: D. ghetaldii and
T. dabar (recorded earlier under the name Phoxinellus metohiensis) in Vrijeka and only
T. dabar in Opačica (our data). Delminichthys (as Phoxinellus) ghetaldii was rst re-
corded in Sušica and Ljelješnica cave springs by Ćurčić (1915a) and then conrmed by
PZ’s ndings in a stream owing from the Ljelješnica Cave (Zupančič and Bogutskaya
2002) and in the Vrijeka. It is relatively less abundant in Vrijeka than T. dabar. A trout
has been introduced into Vrijeka.
e small Lukavačko Polje (2.5 km2) is located at an elevation of 852–880 m east
from the Dabarsko Polje. e two poljes are divided by about 400 m of elevation.
Fatničko Polje is also a small closed basin (5.6 km2) located southeast from the Dabar-
sko Polje at a much lower elevation, 462–470 m above sea level. e Fatničko Polje is
divided from the Dabarsko Polje by an extremely karstied limestone ridge which is
about 2 km wide. e most prominent karst features of the Fatničko Polje are the Obod
temporary spring and the Pasmica Ponor. About 85% to 90% of the Fatnicko Polje wa-
ter ows to the Trebišnjica springs and 10% to 15% to the Bregava springs (Milanović
1981, 2006). Ćurčić (1915a, 1915b), Taler (1953a, 1953b), Sabioncello (1967) and
Vuković (1977) reported P. metohiensis from the Lukavačko and Fatničko poljes. How-
ever, no extant samples conrm these reports. In 1998–2001, PZ found only D. ghet-
aldii in Fatničko Polje and no D. ghetaldii or T. metohiensis in the Lukavačko Polje
(Zupančič and Bogutskaya 2002). No other data have been received since then.
Gatačko Polje (37.6 km2) consists of two geomorphologically and hydrogeologi-
cally interconnected units: Gatačko Polje itself and Small Gatačko Polje. e largest
ponor zone is situated in Small Gatačko Polje along the 8 km long tectonic contact
between ysch sediments and karstied limestone, from Srdevići to the Šabanov Ponor
(936 m above sea level). e entire Gatačko Polje belongs to the catchment area of the
Trebisnjica River, except a very small eastern part. e longest underground ow (35
km) in Eastern Herzegovina is between the Srdevići Ponor and the Trebisnjica Springs.
e main ow in the Gatačko Polje is the Mušnica River with its tributary Gračanica.
e Mušnica is formed by three streams, Vrba, Ulinjski Potok and Jasenički Potok.
ey ow from the Cemern and Lebršnik mountains. e Mušnica goes from the east-
ern to the western border of the polje and along its western border southwards before
it completely sinks in the Šabanov Ponor. e ow of the Mušnica River re-appears in
the Cerničko Polje where it is named the Ključka River. It originates in a large cave,
Vilina Pećina and terminates about 300 m downstream in a ponor that has a recharge
capacity of approximately 20 m3 s−l (Milanović 2006). Telestes metohiensis (in Paraph-
oxinus or Phoxinellus by earlier authors), Phoxinus sp. and Alburnus neretvae Buj, Sanda
et Perea 2010 are known to occur in Mušnica. Phoxinus sp. and, probably, Salmo sp.
occur in Gračanica. Many literature sources (see in Zupančič and Bogutskaya 2002)
reported Paraphoxinus metohiensis from the Gračanica River at Gacko, but we know
of no extant samples. Only T. metohiensis was found (PZC) in short karstic streams of
the Cerničko Polje.
Telestes metohiensis or close species are absent from other poljes except for the Ko-
navosko Polje. is polje is located rather far in the south from poljes inhabited by T.

Nina G. Bogutskaya et al. / ZooKeys 180: 53–80 (2012)
76
metohiensis and T. dabar. e Konavosko Polje or Polje Konavli is the lowermost polje
in the region (elevation 60 m above sea level) with surface of 48 km2. It is developed
along the most important overthrust of the entire dinaric karst region (“High Karst
Overthrust”). e largest spring Konavoska Ljuta is located at this contact. In natural
conditions it was a temporary ooded polje. Main ows are Ljuta and its tributaries
Konavočica and Opačica. At present, a tunnel between the polje and the sea coast
drains the polje (Milanović 2006). No recent records exist of a Telestes species in the
Ljuta or other springs of the Konavosko Polje. However, we think that eorts are
worth trying to nd T. miloradi, a new species, described here from the Konavoska
Ljuta before it is nally considered extinct. It is known that many species from Phox-
inellus, Telestes and Delminichthys genera are able to enter karstic underground waters
during droughts and dramatic water table level uctuation (Milanović 2006; Jelić,
2008; Marčić et al. 2011). However, this phenomenon probably depends considerably
on the degree of development, size and depth of caverns, siphonic lakes and pools of
estavelles.
e ranges of T. metohiensis and T. dabar are extremely small at least during the
dry season and in surface water bodies. e recognition of T. dabar as a distinct species
will require its Red List evaluation and a re-evaluation of T. metohiensis (now consid-
ered as Vulnerable) because of the loss of a part of its former presumed range. Telestes
dabar would probably deserve the Critically Endangered status because of its very re-
stricted distribution (IUCN 2010). Both species are threatened by habitat destruction,
in particular by construction of tunnels for the draining of poljes and controlling their
inundations, lining of river beds to obstruct water ow into ponors and reversible com-
munication with estavelles.
Key to Telestes species of isolated karst river systems of the Adriatic basin in Bos-
nia and Herzegovina and Croatia, including Krbavsko Polje
1a Pharyngeal teeth in two rows, usually 2.5–5.2 or 2.5–4.2. Preoperculo-man-
dibular canal communicating with infraorbital canal. Postcleithrum present,
of moderate size .......................................................................................... 2
1b Pharyngeal teeth in one row, usually 5–4. Preoperculo-mandibular canal not
communicating with infraorbital canal. Postcleithrum minute or absent .....3
2a Branched dorsal-n rays 8½ and branched anal-n rays 9½. Total lateral-line
scales 60–68 ................................................................................... T. ukliva
2b Branched dorsal-n rays 7½ and branched anal-n rays 8½. Total lateral-line
scales 72–79 ...................................................................................T. turskyi
3a Black stripe on ventral portion of trunk narrow and not reaching caudal pe-
duncle. Branched dorsal-n rays usually 8½ ...............................................4
3b Black stripe on ventral portion of trunk absent or present; if present, stripe
wide anteriorly and extending posteriorly to caudal peduncle. Branched dor-
sal-n rays usually 7½ ................................................................................. 5

Two new freshwater sh species of the genus Telestes (Actinopterygii, Cyprinidae)... 77
4a Scales overlapping on most of body, scales approximately same size in, above
and below lateral line. Lateral line not curving above anal-n origin. Lower-
jaw length 8–10% SL .................................................................T. miloradi
4b Scales non-overlapping on most of body, though densely set. Lateral-line
scales larger than scales above and below lateral line. Lateral line making clear
curvature above anal-n origin. Lower-jaw length 10–12% SL .......T. dabar
5a Body strongly compressed, abdomen with weakly scaled ventral keel in front
of pelvics. Lateral line short, incomplete, terminating between pectoral and
anal ns, and usually widely interrupted, with 17–56, usually 23–37, total
scales. Branched anal-n rays 7½ .............................................. T. fontinalis
5b Body slightly compressed, abdomen rounded. Lateral line long, slightly in-
complete or complete, usually terminating on caudal peduncle, and narrowly
interrupted, with 24–70, usually 54–68, total scales. Branched anal-n rays
usually 8½ .................................................................................................. 6
6a Black stripe on ventral portion of trunk present ..................... T. metohiensis
6b Black stripe on vental portion of trunk absent ........................... T. croaticus
Comparative material
Telestes karsticus. PZC 577, 7, 547–750 mm SL, CROATIA: Sušik River at Tomići,
Dobra River system (Danube drainage), 24 June 2005, coll. Zupančič.
Telestes metohiensis. All from BOSNIA AND HERZEGOVINA. Gatačko Polje:
NMW 51176:3, lectotype, mm SL, 87.9 mm SL, ‘Mušica bei Imotski’ [misspelling,
should be Mušnica River, Gatačko Polje; Imotski is probably an error because Steindach-
ner (1901: 198) clearly indicated that the specimens had been received from karst waters
and streams near Gacko, a town in the north of Gatačko Polje (43°09.4'N, 18°31.8'E);
see also Zupančič & Bogutskaya 2002], 1899, coll. Redel, Sturani [Sturany]; NMW
51176:1–2, 72.2–82.5 mm SL, same data as holotype; NMW 12972, 1 paralectotype,
92.2 mm SL, same data as holotype; NMW 12973–75 (in one jar with 12972), 3 para-
lectotypes, 59.2–95.3 mm SL, same data as holotype; NMW 51172, 2 paralectotypes,
92.0–97.8 mm SL, same data as holotype; NMW 51174, 2 paralectotypes, 76.4–88.5
mm SL, same data as holotype; NMW 51175, 3 paralectotypes, 66.8–75.9 mm SL,
same data as holotype; SMF 805, 2, 53.2–65.6 mm SL, Gacko; ZMH 15137, 7, 51.2–
64.4 mm SL, Gacko, Herzegovina; Cerničko Polje: PZC 223, 9, 45.0–90.6 mm SL,
source of Ključka River in Vilina Pečina (cave), 43°5.6'N 18°29'E, 16 May 2003, coll.
Zupančič; PZC 330, 9, 46.0–88.7 mm SL, same locality and collector, 23 May 2001;
PZC 337, 4, 73.7–85.4 mm SL, same locality and collector, 20 Aug. 2000; PZC 368, 4,
49.0–62.2 mm SL, spring at Ključ, 43°5.6'N 18°29.6'E, 23 May 2000, coll. Zupančič;
PZC 503, 7, same locality and collector, 3 Aug. 2000. Nevesinjsko Polje: (NMW syn-
types [now paralectotypes] of P. metohiensis labeled as Paraphoxinus anis [handwritten
by Steindachner; never published], nomen museale) NMW 9368–9372, 5, Zalomska
[Zalomka River], 1896, coll. Hawelka; NMW 51088, 10, 47.6–57.9 mm SL, same

Nina G. Bogutskaya et al. / ZooKeys 180: 53–80 (2012)
78
data as above; NMW 51089, 10, 49.6–58.1 mm SL, same data as above; NMW 51092,
5, 51.9–65.5 mm SL, same data as above; NMW 51093, 8, 47.4–58.7 mm SL, same
data as above; NMW 51094, 10, same data as above; NMW 51090, 5 (not labeled as
syntypes of P. metohiensis though exclusion of these specimens from the type series is not
evident from the original description), 47.6–57.9 mm SL, same data as above; NMW
51091, 4 (not labeled as syntypes, as above), 63.5–87.1 mm SL, same data as above;
PZC 206, 3, 50.4–73.6 mm SL, Batuša River (Zalomka system), 43°19.3'N 18°7.1'E,
9 May 2000, coll. Zupančič; PZC 355, 17, 39.6–63.8 mm SL, same locality and collec-
tor, 21 May 2001; PZC 356, 6, 53.2–66.1 mm SL, same locality and collector, 5 May
2000; PZC 312, 19, 50.1–79.8 mm SL, spring Ljeskovik in Zalomka River near Rast
and Odžak, 43°12.1'N 18°12.4'E, 21 May 2001, coll. Zupančič; PZC 313, 14, 49.1–
78.7 mm SL, same locality and collector, 1 July 2002; PZC 567, 7, 45.1–86.2 mm
SL, same locality and collector, 8 July 2011; PZC 358, 2, 38.2–42.0 mm SL, spring
in Zalomka River at Budisavlje, 43°13.3'N 18°13.1'E, 22 May 2011, coll. Zupančič;
PZC 523, 15, 39.5–90.9 mm SL, Zovidolka [Zavodoka, Zovidolska] River at Ud-
bine, 43°8.6'N 18°14.7'E, 15 Sept. 2006, coll. Zupančič; PZC 293, 12, 49.1–99.2 mm
SL, Zovidolka [Zavodoka, Zovidolska] River at Udbine, 43°8.6'N 18°14.7'E, 29 Aug.
2006, coll. Zupančič; PZC 315, 17, 57.8–81.4 mm SL, same locality and collector, 16
July 2002; PZC 523, 15, 39.5–90.9 mm SL, same locality and collector, 15 Sept. 2006;
PZC 524, 13, 39.5–90.9 mm SL, same locality and collector, 15 Sept. 2006; PZC 566,
45, same locality and collector, 8 July 2011.
Telestes polylepis. NMW 18931–41, 11, paralectotypes, 93.6–107.2 mm SL, CRO-
ATIA: Josefsthal [Josipdol], 1866; NMW 49713, 3, syntypes, 86.7–90.7 mm SL,
CROATIA: Dobra River, 1866; NMW 49714–1, lectotype, 85.9 mm SL, CROATIA:
Mresnitza [Mrežnica] River, 1866; NMW 49715, 2, paralectotypes, 80.8–84.1 mm
SL, same data as lectotype.
Telestes turskyi. NMW 49629-1, lectotype, 129.5 mm SL, CROATIA: Čicola River
[tributary of Krka], Dernis [Drniš] “Heckels Reise, 1840”; 17, paralectotypes, 55.3–
102.3 mm SL.
Telestes ukliva. NMW 49639-1, lectotype, 85.0 mm SL, CROATIA: Sign “[aus der
Cettina]”, “Heckels Reise, 1840”; NMW 49639-2 and 3, 2, 83.3–84.8 mm SL, same
data as lectotype; NMW 49635, 3, paralectotypes, 58.0–75.8 mm SL, same data as
lectotype; ZMH 15095, 1, CROATIA: Cetina.
Other extensive comparative material is listed in Zupančič and Bogutskaya (2002)
and Bogutskaya and Zupančič (2003).
Acknowledgements
We are very thankful to N. Ančić for his assistance during eld trips. Sincere appre-
ciation goes to B. Herzig, E. Mikschi, C. Prenner and H. Wellendorf (NMW) for
providing materials that have been used for comparisons from the collection under
their care. We appreciate valuable comments on an earlier version of the manuscript

Two new freshwater sh species of the genus Telestes (Actinopterygii, Cyprinidae)... 79
and linguistic revision by two anonymous reviews, the editor Carole Baldwin, and
Brian Coad (Canadian Museum of Nature). Contribution by NB and AN to the study
was supported by a grant from the Russian Foundation for Basic Research No. 10-04-
01178, All Cypriniformes Species Inventory Project (ACSI-2), and a grant to ZISP
from Ministry of Education and Science of Russian Federation.
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