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Three new species of Turcinoemacheilus from Iran and Turkey ( Teleostei : Nemacheilidae )

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Three new species of Turcinoemacheilus are described from Iran and Turkey raising the number of species in this genus to six, five of them in the Middle East. Turcinoemacheilus bahaii, new species, from the Zayandeh River in Iran is distinguished by having the anus situated in a posterior position and a dark brown blotch on each side of the anal-fin base in both sexes. Turcinoemacheilus minimus, new species, from the upper Euphrates drainage in Turkey is distinguished by having the anus situated in an anterior position, a slender and long caudal peduncle and a very small maximum size (up to 38 mm SL). Turcinoemacheilus saadii, new species, from the Karoun drainage in Iran is distinguished by having the anus situated in an anterior position and an unique colour pattern of large dark brown saddles. In the Middle East, all Turcinoemacheilus species are well distinguished by molecular characters and show between 5.3 and 8.5 % K2P sequence divergence in their COI barcode region.
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Ichthyol. Explor. Freshwaters, Vol. 24, No. 3
257
Ichthyol. Explor. Freshwaters, Vol. 24, No. 3, pp. 257-273, 16 figs., 4 tabs., March 2014
© 2014 by Verlag Dr. Friedrich Pfeil, München, Germany – ISSN 0936-9902
Three new species of Turcinoemacheilus
from Iran and Turkey
(Teleostei: Nemacheilidae)
Hamid Reza Esmaeili*, Golnaz Sayyadzadeh*, Müfit Özulug**,
Matthias Geiger*** and Jörg Freyhof***
Three new species of Turcinoemacheilus are described from Iran and Turkey raising the number of species in this
genus to six, five of them in the Middle East. Turcinoemacheilus bahaii, new species, from the Zayandeh River in
Iran is distinguished by having the anus situated in a posterior position and a dark brown blotch on each side of
the anal-fin base in both sexes. Turcinoemacheilus minimus, new species, from the upper Euphrates drainage in
Turkey is distinguished by having the anus situated in an anterior position, a slender and long caudal peduncle
and a very small maximum size (up to 38 mm SL). Turcinoemacheilus saadii, new species, from the Karoun drain-
age in Iran is distinguished by having the anus situated in an anterior position and an unique colour pattern of
large dark brown saddles. In the Middle East, all Turcinoemacheilus species are well distinguished by molecular
characters and show between 5.3 and 8.5 % K2P sequence divergence in their COI barcode region.
* Department of Biology, College of Sciences, Shiraz University, Shiraz, Iran.
E-mail: esmaeili@susc.ac.ir, golnaz_sayad@yahoo.com
** Istanbul University, Science Faculty, Department of Biology, 34134 Vezneciler, Istanbul, Turkey.
E-mail: mozulu@istanbul.edu.tr
*** Zoological Research Museum Alexander Koenig, Leibniz Institute for Animal Biodiversity, Adenauerallee
160, 53113 Bonn, Germany. E-mail: joerg.freyhof@idiv.de
Introduction
Since the description of the Zagros dwarf loach
Turcinoemacheilus kosswigi from headwaters of the
Great Zab River in Turkey (Bânârescu & Nalbant,
1964), this genus was so rarely recognized that
we learned only recently about its distribution.
Breil & Bohlen (2001) published the first records
of Turcinoemacheilus from the Euphrates drainage
(now referred to T. minimus), Golzarianpour et
al. (2009) first reported the genus from Iran (now
referred to T. hafezi), Golzarianpour et al. (2013)
recorded T. kosswigi from Sirvan, Great and Little
Zab rivers in Iraq and described T. hafezi from
Iran. This scarcity of records is surprising, since
loaches of the genus Turcinoemacheilus are now
known to be very widespread, often common,
and easy to distinguish from all other nemachei-
lids in the Middle East by the pelvic fin origin
situated in front of the dorsal fin origin and an
anterior position of the anus, a particular colour
pattern and a very slender body (Bânârescu &
Nalbant, 1964; Freyhof et al., 2011; Golzarianpour
et al., 2013). While Turcinoemacheilus was a mono-
258
Esmaeili et al.: Three new Turcinoemacheilus
typic genus for 47 years, two new species have
been described in the last years (Conway et al.,
2011; Golzarianpour et al., 2013). There were al-
ready indications that more undescribed species
might exist in the Middle East. Breil & Bohlen
(2001) observed that the examined fishes from
the Tigris drainage were relatively large, but those
from the Euphrates were much smaller. Golzari-
anpour et al. (2013) found two sympatric species
of Turcinoemacheilus in the Karoun drainage, one
with a more anterior and one with a more poste-
rior position of the anus. They did not identify
the species with the more anterior position of the
anus. Finally, fishes superficially similar to T. hafe-
zi were discovered in the Iranian endorheic Zay-
andeh drainage. In this study, the morphological
characters as well as the colour patterns of all
these populations is analysed, and we also se-
quenced their COI barcode region. These data
together supported the view that five species of
Turcinoemacheilus are present in the Middle East,
three of them still undescribed. We describe these
three new Turcinoemacheilus based on combined
findings from morphology and molecular ge-
netic characters.
Material and methods
After anaesthesia, all fishes were fixed in 5 %
formaldehyde and stored in 70 % ethanol. Mea-
surements were made with dial calliper and re-
corded to 0.1 mm. All measurements are made
point to point, never by projections. Methods for
counts and measurements follow Kottelat &
Freyhof (2007). Standard length (SL) is measured
from the tip of the snout to the end of the hypural
complex. The length of the caudal peduncle is
measured from behind the base of the last anal-fin
ray to the end of the hypural complex, at mid-
height of the caudal-fin base. The last two branched
rays articulating on a single pterygiophore in the
dorsal and anal fins are noted as 1 1
/
2. The ho-
lotype is included in the calculation of means and
SD.
Abbreviations used: SL, standard length; HL,
lateral head length; K2P, Kimura 2-parameter;
CMK, Collection of Maurice Kottelat, Cornol;
FSJF, Fischsammlung J. Freyhof, Berlin; IUSHM,
Istanbul University, Science Faculty, Hydrobiol-
ogy Museum, Istanbul; KU, University of Kansas
Natural History Museum and Biodiversity Re-
search Center, Lawrence; ZFMK, Zoological
Research Museum Alexander Koenig, Leibniz
Institute for Animal Biodiversity, Bonn; ZM-
CBSU, Zoological Museum of Shiraz University,
Collection of Biology Department, Shiraz; and
ZMH, Zoologisches Museum, Hamburg.
DNA extraction and PCR. Genomic DNA was
extracted using Machery & Nagel NucleoSpin®
Tissue kits following the manufacturers protocol
on an Eppendorf EpMotion® pipetting-roboter
with vacuum manifold. The standard vertebrate
DNA barcode region of the COI (cytochrome c
oxidase subunit 1) was amplified using a M13
tailed primer cocktail including FishF2_t1
(5' TGTAAAACGACGGCCAGTCGACTAAT-
CATAAAGATATCGGCAC), FishR2_t1 (5' CAG-
GAAACAGCTATGACACTTCAGGGTGAC-
CGAAGAATCAGAA), VF2_t1 (5' TGTAAAAC-
GACGGCCAGTCAACCAACCACAAAGACAT-
TGGCAC) and FR1d_t1 (5' CAGGAAACAGC-
TATGACACCTCAGGGTGTCCGAARAAYCAR-
AA) (Ivanova et al., 2007). Sequencing of the
ExoSAP-IT (USB) purified PCR product in both
directions was conducted at Macrogen Europe
Laboratories with forward sequencing primer
M13F (5' GTAAAACGACGGCCAGT) and re-
verse sequencing primer M13R-pUC (5' CAG-
GAAACAGCTATGAC).
Molecular data analysis. Data processing and
sequence assembly was done in Geneious (Bio-
matters, 2013) and the Muscle algorithm (Edgar,
2004) was used to create a DNA sequence align-
ment. Modeltest (Posada & Crandall, 1998), imple-
mented in the MEGA 5 software (Tamura et al.,
2011) was used to determine the most appropri-
ate sequence evolution model for the given data,
treating gaps and missing data with the partial
deletion option under 95 % site coverage cutoff.
The model with the lowest BIC scores (Bayesian
Information Criterion) is considered to best de-
scribe the substitution pattern. We generated
neighbor-joining (Saitou & Nei, 1987), maximum
parsimony (Swofford, 2002; with PAUP4b) and
maximum likelihood phylogenetic trees with 1000
bootstrap replicates to explore species phyloge-
netic affinities. Screening for diagnostic nucleotide
substitutions was performed manually from the
resulting sequence alignment.
Ichthyol. Explor. Freshwaters, Vol. 24, No. 3
259
Results
Maximum Likelihood based estimation of the
phylogenetic relationships based on the mito-
chondrial COI barcode region place the sequenced
fishes into five groups (Fig. 1), which show be-
tween 5.3 and 8.5 % K2P sequence divergence in
their COI barcode region. Two of these groups
correspond to the described species T. kosswigi
and T. hafezi. The other three groups correspond
to the new species described below. A map with
all records of Turcinoemacheilus species in the
Middle East is shown in Figure 2.
Key to species of Turcinoemacheilus
in the Middle East
1 – Anus situated behind middle between
pelvic-fin and anal-fin origins.
........................................................................2
Anus situated at or in front of middle be-
tween pelvic-fin and anal-fin origins.
........................................................................3
2 – An elongated, irregularly shaped dark
blotch on sides of anal-fin base; anal-fin
origin situated at vertical of tip of dorsal fin
when adpressed to body.
............................................................T. bahaii
– No dark blotch on side of anal-fin base;
anal-fin origin situated behind vertical of
tip of dorsal fin when adpressed to body.
............................................................T. hafezi
3 Lateral stripe or row of blotches absent along
lateral midline, 7-9 distinct dark saddles on
body.
............................................................T. saadii
Prominent row of dark brown blotches along
lateral midline, usually fused into a lateral
stripe.
........................................................................4
4 Standard length up to at least 53 mm; cau-
dal peduncle depth 1.9-2.2 times in its
length.
........................................................T. kosswigi
– Standard length up to at least 38 mm; cau-
dal peduncle depth 2.6-3.2 times in its
length.
.......................................................T. minimus
Turcinoemacheilus bahaii, new species
(Figs. 3-5)
Holotype. ZM-CBSU 7193B, 52.2 mm SL; Iran:
Esfahan prov.: Zayandeh River between Azade-
gan and Qalee Shahrokh, 32°40'54" N 50°27'47" E;
H. R. Esmaeili, A. Gholamifard, G. Sayyadzadeh,
B. Parsi, R. Zamanian Nejad, S. Mirghiasi & M.
Ghasemeian, 16 Aug 2011.
Paratypes. ZM-CBSU 7192B, 1, 38.6 mm SL; ZM-
CBSU 7194B-7208B, 15, 24.1-53.9 mm SL; same
data as holotype. – ZM-CBSU 7209B-7221B, 13,
26.2-45.1 mm SL; FSJF 3449, 4, 28.9-33.8 mm SL;
Iran: Esfahan prov.: Zayandeh River at Eskanda-
ri village, 32°49'26" N 50°25'47" E; H. R. Esmaeili,
A. Gholamifard, G. Sayyadzadeh, B. Parsi, R.
Zamanian Nejad, S. Mirghiasi & M. Ghasemeian,
16 Aug 2011.
Additional material (non types). Material used in the
molecular genetic analysis: ZM-CBSU T550F, T551F;
GenBank accession numbers: KJ179246, KJ179247; same
data as holotype.
Diagnosis. Turcinoemacheilus bahaii is the only
species of Turcinoemacheilus having a dark brown
blotch on each side of the anal-fin base (Fig. 6)
(vs. absent) and it is the only species in which the
anal-fin origin is situated at a vertical of the tip
of the dorsal fin when adpressed to the body (vs.
anal-fin origin clearly behind vertical of tip of
dorsal fin). It is further distinguished from all
other species of Turcinoemacheilus in the Middle
East except T. hafezi by the more posterior position
of the anus (distance from anus to anal-fin origin
0.1-0.4 times in distance from pelvic-fin to anal-fin
origins). Compared to all its four congeners in
the Middle East, T. bahaii is characterized by five
fixed nucleotide substitutions in the mtDNA COI
barcode region studied (Table 1).
Description. See Figures 3-5 for general appear-
ance and Table 2 for morphometric data of holo-
type and 29 paratypes. Small, very slender and
roundish bodied species with short head. Body
deepest at about pelvic-fin base, depth decreasing
towards caudal-fin base. No hump at nape. Great-
est body width at pectoral-fin base or at middle
between pectoral- and pelvic-fin bases, body al-
most equally wide until dorsal-fin origin. Section
of head roundish, flattened on ventral surface.
Caudal peduncle compressed laterally, 1.8-2.5
260
100-98-95
100-100-97
99-99-99
100-100-97
99-99-89
100-100-98
74-53-65
100-99-94
100-100-98
Paraschistura bampurensis Iran Baluchestan_KJ179269
Paraschistura bampurensis Iran Baluchestan_KJ179268
Paracobitis malapterura Iran Namak_KJ179267
Paracobitis malapterura Iran Namak_KJ179266
T. hafezi Iran Karoun_KJ179259
T. bahaii Iran Zayandeh _KJ179246
T. bahaii Iran Zayandeh _KJ179247
T. hafezi Iran Karoun_KJ179252
T. hafezi Iran Karoun_KJ179254
T. hafezi Iran Karoun_KJ179264
T. saadii Iran Karoun_KJ179257
T. saadii Iran Karoun_KJ179248
T. saadii Iran Karoun_KJ179261
T. kosswigi Iran Sirvan_KJ179258
T. kosswigi Iran Sirvan_KJ179245
T. kosswigi Iraq Little Zab_KJ179255
T. kosswigi Iraq Little Zab_KJ179262
T. kosswigi Iraq Little Zab_KJ179260
T. kosswigi Iraq Great Zab_KJ179265
T. minimus Turkey Euphrates_KJ179256
T. saadii Iran Karoun_KJ179250
T. saadii Iran Karkheh_KJ179253
T. minimus Turkey Euphrates_KJ179251
T. minimus Turkey Euphrates_KJ179263
T. minimus Turkey Euphrates_KJ179249
Fig. 1. Maximum Likelihood (ML) estimation of the phylogenetic relationships based on the mitochondrial COI
barcode region. Nucleotide positions with less than 95 % site coverage were eliminated before analysis. Numbers
of major nodes indicate bootstrap values from the Neighbour joining-, Maximum Parsimony-, and Maximum
Likelihood method from 1000 pseudo-replicates. Branch lengths are not drawn to scale.
Table 1. List of the diagnostic nucleotide substitutions for the Middle-East species of Turcinoemacheilus found in
the mtDNA COI barcode region. Nucleotide position is given with reference to the complete mitochondrial genome
of Oryzias latipes (GenBank accession number AP004421). The * denotes and amino acid change from isoleucine
to valine.
5529
5553
5593
5601
5673
5707
5715
5736
5766
5775
5793
5844
5850
5862
5874
5901
5904
5910
5919
5955
5980
6000
6012
6027
6042
6051
6066
6069
codon position 3333313333333333333913333333
T. hafezi (n = 4) GTCCCCTCCCAGAATAAGTAAAGCTACA
T. bahaii (n = 2) GTCCCCTCCCAGAATAGGCGAAACCACA
T. kosswigi (n = 6) GTCCCCTCCAAGAATAACAAACGCTACC
T. minimus (n = 4) ATCCCCTTTTAGACTAACTA
G* T GCTGTG
T. saadii (n = 5) GCTTTTCCCCGAGACGAAAAACGTTACA
Ichthyol. Explor. Freshwaters, Vol. 24, No. 3
261
(mean 2.2) times longer than deep. Pectoral fin
reaching approximately 50 % of distance from
pectoral-fin origin to pelvic-fin origin. Pelvic
axillary lobe present, fully attached to body.
Pelvic-fin origin distinctly in front of dorsal-fin
origin. Pelvic fin reaching slightly beyond or not
reaching to anus. Distance from anus to anal-fin
origin 0.1-0.4 times in distance from pelvic-fin to
anal-fin origins. Anal-fin origin at vertical of tip
of dorsal fin when adpressed to body. Anal fin
reaching beyond middle of caudal peduncle. No
adipose crest on caudal peduncle. Margin of
dorsal fin straight. Caudal fin emarginated. Larg-
est known specimen 53.9 mm SL.
Dorsal fin with 7
1
/
2 branched rays. Anal fin
with 5
1
/
2 branched rays. Caudal fin with 8 + 8
branched rays. Pectoral fin with 8-10 (usually 8)
and pelvic fin with 6 branched rays. Body without
scales. Lateral line incomplete, with 12-33 pores,
not reaching to dorsal-fin origin. Anterior nostril
opening on anterior side of a low, pointed and
flap-like tube. Nostrils separate, posterior tip of
anterior nostril not reaching posterior nostril
when folded backwards. No suborbital flap or
groove in males. Mouth small, slightly arched
(Fig. 7a). Lips moderately thick. A median inter-
ruption in lower lip. Upper lip without median
incision. Processus dentiformis small and blunt.
No median notch in lower jaw. Barbels short,
inner rostral barbel not reaching base of maxillary
barbel; outer one reaching slightly beyond base
of maxillary barbel, not reaching vertical of ante-
rior margin of eye. Maxillary barbel reaching
vertical of middle of eye. No external sexual di-
morphism observed.
Coloration. In alcohol, body pale yellow with
pale or dark brown mottled colour pattern. Ir-
regular blotches on body forming irregularly set
and shaped bars in some individuals or blotches
more prominent along lateral midline. Flank
below a line between pectoral- and pelvic-fin
bases without pigmentation. An irregularly
shaped, dark brown or black bar at posterior
extremity of caudal peduncle, usually most
prominent at the middle. In front of this bar, a
whitish or yellowish irregular quadratic or trian-
gular patch. Cheeks and ventral surface of head
E
E
u
p
h
r
a
t
e
s
T
i
g
r
i
s
Caspian
Caspian
Sea
Sea
Karkheh
Karkheh
Isfahan
Isfahan
Karun
Karun
Persian
Persian
Gulf
Gulf
E
u
p
h
r
a
t
e
s
T
i
g
r
i
s
Caspian
Sea
Karkheh
Isfahan
Karun
Persian
Gulf
Fig. 2. Records of Turcinoemacheilus species in the Middle East.
&
, T. minimus; %, T. kosswigi; (, T. saadii; , T. hafe-
zi;
)
, T. bahaii.
262
Fig. 3. Turcinoemacheilus bahaii, ZM-CBSU 7193B, holotype, 52.2 mm SL; Iran: Zayandeh River.
Fig. 4. Turcinoemacheilus bahaii, Iran: Zayandeh River; paratypes; a, ZM-CBSU 7194B, 49.1 mm SL; b, ZM-CBSU
7192B, 41.7 mm SL; and c, ZM-CBSU 7195B, 38.6 mm SL.
Fig. 5. Turcinoemacheilus bahaii, ZM-CBSU 7198b, paratype, 44.1 mm SL; Iran: Zayandeh River.
a
b
c
Ichthyol. Explor. Freshwaters, Vol. 24, No. 3
263
Fig. 6. Turcinoemacheilus bahaii, ZM-CBSU 7194B,
49.1 mm SL; diagnostic dark blotch on sides of anal-fin
origin.
cream or pale yellow, head above cheeks with
many small brown spots and blotches sometimes
fused into a marbled pattern or plain brown. An
elongated, faint, brown blotch on each side of
anal-fin base (Fig. 6). Dorsal and caudal fins hya-
line, with elongated spots on rays, forming a
mottled pattern of 1-2 dark vertical rows ap-
proximately in middle of ray length in dorsal fin
and 1-2 vertical rows in caudal fin. Last un-
branched dorsal-fin ray black at base, black at
about middle of ray and hyaline on posterior part.
Anal-, pelvic- and pectoral fins hyaline.
abc
Fig. 7. Ventral view of head of species of Turcinoemacheilus. a, T. bahaii, ZM-CBSU 7197b, paratype, 53.6 mm SL;
b, T. minimus, IUSHM 2013-1050, holotype, 35.2 mm SL; and c, T. saadii, ZM-CBSU 7170b, paratype, 46.8 mm SL.
Fig. 8. Zayandeh River between Azadegan and Qalee Shahrokh, Iran; type locality of Turcinoemacheilus bahaii.
264
In life, body creamy whitish with very faint,
pale brown colour patterns.
Distribution and habitat. Turcinoemacheilus bahaii
is known from the Zayandeh River which is an
endorheic river in Central Iran flowing from the
Zagros Mountains to the Gavkhoni wetlands. At
the type locality (Fig. 8) the river is about 20 m
wide, the substrate consists of coarse gravel and
boulders and the water is fast-running.
Etymology. The species is named for Baha
al-Dı
n
Muh
.ammad ibn H
.usayn al-A
milı
(also known
as Shaykh-i Baha
, Persian: ) a scholar,
philosopher, architect, mathematician, astrono-
mer and poet in 16th century Iran.
Remarks. The phylogenetic tree reconstruction
(Fig. 1) suggests that T. bahaii is closely related to
T. hafezi from the Karoun and Dez drainages, both
flowing to the deltaic area of the Arvand River
or Shatt al Arab, which is the lowermost part of
the joint Euphrates and Tigris drainages. Both
species are superficially similar in body shape
and colour pattern, and are separated by 5.3 %
K2P distance in the studied COI gene region,
which is proposed as a strong indicator that two
species are involved (see Herbert et al., 2003).
Beside molecular characters, T. bahaii is distin-
guished from T. hafezi by having a dark brown
blotch on the sides of the anal-fin base (Fig. 6) (vs.
absence). It is distinguished from T. hafezi, T. koss-
wigi and T. saadii by the anal fin situated in a more
anterior position, at a vertical of the tip of dorsal
fin when adpressed to the body (vs. anal-fin ori-
gin clearly behind vertical of tip of dorsal fin).
Turcinoemacheilus bahaii is further distinguished
from T. kosswigi, T. minimus and T. saadii by the
more posterior position of the anus, which is
behind the middle between the pelvic-fin origin
and the anal-fin origin (vs. at the middle or in
front of the middle). Turcinoemacheilus bahaii is
distinguished from T. himalaya from Nepal by a
completely scaleless body (vs. small cycloid scales
present on the posterior half of the body).
Table 2. Morphometric data of Turcinoemacheilus bahaii (holotype ZM-CBSU 7193B, paratypes, ZM-CBSU 7192B-
7208B, ZM-CBSU 7209B-7221B, n = 29).
holotype min max mean SD
Standard length (mm) 52.2 24.1 53.9
In percent of standard length
Head length 20.4 20.4 25.2 23.0 1.3
Body depth at dorsal-fin origin 12.3 11.5 16.2 13.5 1.2
Prepectoral length 17.8 17.8 24.7 21.6 1.7
Predorsal length 53.9 36.4 57.5 54.4 3.6
Postdorsal length 35.0 30.1 36.6 34.2 1.4
Preanal length 72.8 66.7 74.2 71.1 1.2
Prepelvic length 50.5 49.8 55.4 52.8 1.4
Distance between pectoral and pelvic-fin origins 31.8 29.0 33.9 31.4 1.2
Distance between pelvic and anal-fin origins 21.4 15.5 22.2 19.0 1.7
Distance between vent and anal-fin origin 6.7 2.4 7.9 4.2 1.3
Depth of caudal peduncle 8.8 8.0 10.6 9.0 0.5
Length of caudal peduncle 20.7 17.5 21.6 19.4 1.1
Dorsal-fin depth 14.6 13.9 18.7 16.3 1.4
Anal-fin base length 7.5 5.3 8.1 6.9 0.7
Pectoral-fin length 14.9 14.0 17.7 16.0 1.0
Pelvic-fin length 12.2 11.2 14.0 12.0 0.8
In percent of head length
Head depth at eye 46 42 53 47 2.8
Snout length 43 31 44 39 2.8
Eye diameter 13 11 17 14 1.5
Postorbital distance 53 42 55 48 3.0
Maximum head width 63 55 64 61 2.7
Interorbital width 25 21 32 28 3.1
Ichthyol. Explor. Freshwaters, Vol. 24, No. 3
265
Turcinoemacheilus minimus, new species
(Figs. 9-11)
Holotype. IUSHM 2013-1050, 35.2 mm SL; Tur-
key: Adıyaman prov.: Upper Göksu, 5 km north-
east of Gölbası, 37°50.22' N 37°41.09' E; M. Özulug
& J. Freyhof, 18 June 2008.
Paratypes. IUSHM 2013-1051, 5, 32.1-35.3 mm
SL; FSJF 2454, 4, 33-38 mm SL; same data as
holotype.
Additional material (non types). CMK 16824, 3, 25.7-
30.3 mm SL; Turkey: Erzincan prov.: Karasu near
Mercan, 39°45' N 40°14' E; M. Breil, 17 June 1999. – Ma-
terial used in the molecular genetic analysis: FSJF
DNA-968; GenBank accession numbers: KJ179249,
KJ179251, KJ179256, KJ179263; same data as holotype.
Diagnosis. Turcinoemacheilus minimus is distin-
guished from the other species of Turcinoemacheilus
in the Middle East by a combination of characters,
none of them unique. Turcinoemacheilus minimus
is superficially similar to T. kosswigi and both
share the same colour pattern. They are distin-
guished by the more slender and longer caudal
peduncle in T. minimus (caudal peduncle length
19.5-21.0 % SL vs. 17.7-19.0 in T. kosswigi). Tur-
cinoemacheilus minimus is distinguished from
T. hafezi and T. bahaii by the more anterior position
of the anus, which is in the middle (or in front)
of the distance between the pelvic-fin and the
anal-fin origins (vs. behind the middle in T. hafe-
zi and T. bahaii).
Turcinoemacheilus minimus is also distinguished
by its very small size, up to 38 mm SL observed
(vs. larger in all other species). Relative to its four
congeners in the Middle East, T. minimus is char-
acterized by ten fixed nucleotide substitutions in
the mtDNA COI barcode region studied (Table 1),
one of them non-silent and thus leading to an
amino-acid change from isoleucine to valine.
Description. See Figures 9-11 for general appear-
ance and Table 3 for morphometric data of holo-
type and 5 paratypes. Small, very slender and
roundish bodied species with short head. Body
deepest at about pelvic-fin base, depth decreasing
towards caudal-fin base. No hump at nape. Great-
Table 3. Morphometric data of Turcinoemacheilus minimus (holotype IUSHM 2013-1050; paratypes IUSHM 2013-
1051; n = 6).
holotype min max mean SD
Standard length (mm) 35.2 32.1 35.3
In percent of standard length
Head length 20.5 18.1 21.3 19.6 1.1
Body depth at dorsal-fin origin 9.8 9.8 11.6 10.9 0.5
Prepectoral length 20.4 19.0 21.4 20.2 0.7
Predorsal length 58.0 56.8 59.5 58.1 0.8
Postdorsal length 34.8 34.7 36.9 35.8 0.9
Preanal length 73.5 70.4 73.6 72.8 1.2
Prepelvic length 54.2 49.7 54.2 51.9 1.4
Distance between pectoral and pelvic-fin origins 32.0 29.2 32.9 31.1 1.4
Distance between pelvic and anal-fin origins 20.7 20.7 23.3 21.9 1.0
Distance between vent and anal-fin origin 11.5 11.5 15.2 12.8 1.2
Depth of caudal peduncle 7.0 6.3 7.0 6.8 0.3
Length of caudal peduncle 20.1 17.1 21.0 19.7 1.3
Dorsal-fin depth 14.3 12.8 17.0 14.3 1.3
Anal-fin base length 7.4 6.3 9.4 7.6 1.1
Pectoral-fin length 16.7 14.9 16.7 15.7 0.6
Pelvic-fin length 12.8 12.0 13.9 13.0 0.6
In percent of head length
Head depth at eye 35 35 42 38 2.4
Snout length 39 38 45 42 2.9
Eye diameter 16 16 21 18 1.5
Postorbital distance 47 45 50 48 1.5
Maximum head width 48 49 57 53 2.7
Interorbital width 28 25 29 27 1.4
266
Fig. 9. Turcinoemacheilus minimus, IUSHM 2013-1050, holotype, 35.2 mm SL; Turkey: Göksu River.
Fig. 10. Turcinoemacheilus minimus, Turkey: Göksu River: paratypes, IUSHM 2013-1051; a, 35.3 mm SL; b, 35.0 mm
SL; and c, 33.3 mm SL.
Fig. 11. Turcinoemacheilus minimus, collected with IUSHM 2013-1051, about 35 mm SL; Turkey: Göksu River.
a
b
c
Ichthyol. Explor. Freshwaters, Vol. 24, No. 3
267
est body width at pectoral-fin base or at middle
between pectoral- and pelvic-fin bases, body al-
most equally wide until dorsal fin-origin. Section
of head roundish, flattened on ventral surface.
Caudal peduncle compressed laterally, 2.6-3.2
(mean 2.9) times longer than deep. Pelvic axillary
lobe present, its tip not attached to body. Pelvic-
fin origin distinctly in front of dorsal-fin origin.
Pectoral fin reaching approximately 50 % of dis-
tance from pectoral-fin origin to pelvic-fin origin.
Pelvic fin reaching beyond anus. Distance from
anus to anal-fin origin 0.5-0.6 times in distance
from pelvic-fin to anal-fin origins. Anal-fin origin
behind vertical of tip of dorsal fin when adpressed
to body. Anal fin not reaching to middle of caudal
peduncle. No adipose crest on caudal peduncle.
Margin of dorsal fin straight. Caudal fin slightly
emarginated. Largest known specimen 38 mm
SL.
Dorsal fin with 6-7 1
/
2 (usually 6 1
/
2) branched
rays. Anal fin with 5 1
/
2 branched rays. Caudal fin
with 8 + 8 or 8 + 7 branched rays. Pectoral fin with
8-9 and pelvic fin with 7 branched rays. Body
without scales. Lateral line incomplete, with 15-
20 pores, reaching beyond tip of pectoral fin but
not reaching to dorsal-fin origin. Anterior nostril
opening on anterior side of a low, blunt and flap-
like tube. Nostrils closely together, posterior tip
of anterior nostril reaching into posterior nostril
when folded backwards. No suborbital flap or
groove in males. Mouth small, slightly arched
(Fig. 7b). Lips moderately thick. A median inter-
ruption in lower lip. Upper lip without median
incision. Processus dentiformis small and blunt.
No median notch in lower jaw. Barbels short,
inner rostral barbel reaching base of outer rostral
barbel; outer one reaching to base of maxillary
barbel. Maxillary barbel reaching vertical of an-
terior part of eye. No external sexual dimorphism
observed.
Coloration. In alcohol and life, body pale cream
yellow. A row of large irregular, brown, longitu-
dinally elongated blotches along lateral midline,
often fused into a prominent irregular lateral
stripe. Large, brown saddles on back, connected
to lateral blotches along whole body. On predor-
sal back, saddles sometimes dissociated into a
marbled pattern, reaching downwards to stripe.
Flank below lateral stripe without pigmentation.
Fig. 12. Upper Göksu River northeast of Gölbası, Turkey; type locality of Turcinoemacheilus minimus.
268
An irregularly shaped, dark brown or black bar
at caudal-fin base. In front of this bar, a whitish
or yellowish triangular patch on upper and
lower caudal peduncle. Cheeks and ventral sur-
face of head cream or pale yellow, head above
cheeks plain brown. Dorsal and caudal fins hya-
line, with elongated spots on rays, forming 1-2
dark vertical rows, approximately in middle of
ray length in dorsal fin and 1-2 vertical rows in
caudal fin. Last unbranched dorsal-fin ray hyaline
at base, black at anterior half and hyaline on
posterior half. Anal-, pelvic- and pectoral fins
hyaline.
Distribution and habitat. Turcinoemacheilus mini-
mus is known from two localities in the upper
Euphrates drainage in Turkey. The Göksu at the
type locality of the species (Fig. 12) is about 30 m
wide, the substrate consists of coarse gravel and
boulders and the water is fast-running.
Etymology. The species is named for its small
size. An adjective.
Remarks. The phylogenetic tree reconstruction
suggests that T. minimus is most closely related
to T. kosswigi from the Tigris drainage. It had
initially been identified as T. kosswigi by Breil &
Bohlen (2001). The most striking character of
T. minimus, the very small maximum size (up to
38 mm SL vs. 53 in T. kosswigi), was not considered
as a difference by Breil & Bohlen (2001), as only
five individuals of T. kosswigi were known then.
When MÖ & JF collected this species again in the
Euphrates, the small fishes were considered as
juveniles. When fresh materials of T. kosswigi
became available from the Iraqi Tigris drainage,
no such small juveniles were found despite the
fact that both samplings were conducted in June
and the same fishing device, with a 150 mm anode
ring that caught small juvenile cyprinids without
problems. When molecular data became available,
it turned out that both species are well distin-
guished. Based on the complete COI barcode
fragment mean smallest K2P distance of T. mini-
mus is 5.9 % to T. kosswigi, its closest relative.
While there is discussion about a priori thresholds
and their use in species delimitation and recogni-
tion (e. g. Meier et al., 2006; Srivathsan & Meier,
2012), a 5.9 % distance between two populations
is strongly indicative that two species are involved
(see Herbert et al., 2003).
Interestingly, the fishes described by Breil &
Bohlen (2001) (now CMK 16824) were kept about
one year in captivity before they were preserved,
and they did not grow during this time (J. Bohlen,
pers. comm.). This observation and the absence
of larger fishes in our samples suggest that the
Turcinoemacheilus from the upper Euphrates is a
dwarf species. Taking this information and the
molecular data together, we do not consider the
available material from the upper Euphrates as
juveniles and we treat their small size as a diag-
nostic character distinguishing it from T. koss-
wigi. Beside its small size and molecular charac-
ters, T. minimus is superficially very similar to
T. kosswigi and both share the same colour pattern.
They are distinguished by several morphometric
characters, but as no individuals of similar sizes
were available, it cannot be totally excluded that
these differences are the result of allometric
growth. The most obvious difference is the more
slender and longer caudal peduncle in T. mini-
mus.
Turcinoemacheilus minimus is distinguished
from T. hafezi and T. bahaii by the more anterior
position of the anus, which is in the middle (or
in front) of the distance between the pelvic-fin
and the anal-fin origins (distance from anus to
anal-fin origin 0.5-0.6 times in distance from
pelvic-fin to anal-fin origins) while it is behind
the middle (0.2-0.4 times) in T. hafezi and T. ba-
haii.
Turcinoemacheilus minimus is distinguished
from T. himalaya by a completely scaleless body
(vs. presence of small cycloid scales on the pos-
terior half of body).
Turcinoemacheilus saadii, new species
(Figs. 13-15)
Holotype. ZM-CBSU 7169B, 47.1 mm SL; Iran:
Fars prov.: stream Tang-e-Tizab, a tributary to
Bashar River which drains to the Karoun, 30°23'
12" N 51°46'50" E; H. R. Esmaeili, G. Sayyadzadeh,
M. Masoudi, S. H. Aminaghai, 5 Sep 2012.
Paratypes. ZM-CBSU 7166b-7175B, 9, 44.6-
63.6 mm SL; ZM-CBSU 7165B, 1, 45.4 mm SL;
same data as holotype.
Additional material (non types). FSJF 1512, 10, 41.4-
52.6 mm SL; Iran: Kohrang river at Kaj village,
32°03'16.97" N 50°34'46" E. — FSJF 3232, 2, 30-44 mm
Ichthyol. Explor. Freshwaters, Vol. 24, No. 3
269
SL; Iran: Kermanshah prov.: Sepidbarg river west of
Javanrud, 34°48'21" N 46°27'29" E. – Material used in the
molecular genetic analysis: FSJF DNA-1995; Iran: Ker-
manshah prov.: Sepidbarg river west of Javanrud,
34°48'21" N 46°27'29" E, GenBank accession numbers:
KJ179250, KJ179257. – ZM-CBSU T540, T542; same data
as holotype, GenBank accession numbers: KJ179248,
KJ179261. – ZM-CBSU T524; Iran: Gamasiab, Karkheh,
Tigris drainage, 34°23'16" N 47°41'14" E, GenBank ac-
cession number: KJ179253.
Diagnosis. Turcinoemacheilus saadii is distin-
guished from the other species of Turcinoemacheilus
in the Middle East by its unique colour pattern
organized in 7-9 distinct dark saddles never
forming a lateral stripe (vs. mottled colour pattern
or presence of a lateral stripe). It is further distin-
guished by the combination of a deeply emargin-
ated caudal fin and the distance from anus to
anal-fin origin 0.4-0.5 times in the distance from
pelvic-fin to anal-fin origins. Relative to its four
congeners in the Middle East, T. saadii is charac-
terized by 13 fixed nucleotide substitutions in the
mtDNA COI barcode region studied (Table 1).
Description. See Figures 13-15 for general ap-
pearance and Table 4 for morphometric data of
holotype and 10 paratypes. Small, very slender
and roundish bodied species with short head.
Body deepest at about pelvic-fin base, depth
decreasing towards caudal-fin base. No hump at
nape. Greatest body width at pectoral-fin base or
at middle between pectoral- and pelvic-fin bases,
body almost equally wide until dorsal fin-origin.
Section of head roundish, flattened on ventral
surface. Caudal peduncle compressed laterally,
2.4-3.2 (mean 2.7) times longer than deep. Pelvic
axillary lobe present, not attached to body. Pelvic-
fin origin distinctly in front of dorsal-fin origin.
Pectoral fin reaching approximately 40 % of dis-
tance from pectoral-fin origin to pelvic-fin origin.
Pelvic fin reaching to anus. Distance from anus
to anal-fin origin 0.4-0.5 times in distance from
pelvic-fin to anal-fin origins. Anal-fin origin be-
hind vertical of tip of dorsal fin when adpressed
to body. Anal fin not reaching to middle of caudal
peduncle. No adipose crest on caudal peduncle.
Margin of dorsal fin straight or convex. Caudal
Table 4. Morphometric data of Turcinoemacheilus saadii (holotype ZM-CBSU 7169B; paratypes ZM-CBSU 7166b-
7175B, ZM-CBSU 7165B; n = 11).
holotype min max mean SD
Standard length (mm) 47.1 44.6 56.4
In percent of standard length
Head length 19.4 17.3 20.0 19.0 0.8
Body depth at dorsal-fin origin 13.3 11.8 13.4 12.5 0.6
Prepectoral length 18.4 16.9 19.6 18.0 0.9
Predorsal length 53.5 51.6 54.7 52.7 1.0
Postdorsal length 39.6 36.5 39.6 37.9 1.0
Preanal length 73.3 70.4 74.4 72.9 1.0
Prepelvic length 48.6 48.1 51.3 49.8 1.0
Distance between pectoral and pelvic-fin origins 32.2 30.9 33.8 32.5 0.9
Distance between pelvic and anal-fin origins 23.2 20.9 24.3 22.6 1.0
Distance between vent and anal-fin origin 8.6 8.6 12.1 9.8 1.1
Depth of caudal peduncle 7.9 6.8 8.1 7.6 0.4
Length of caudal peduncle 21.1 17.8 21.9 20.6 1.2
Dorsal-fin depth 15.5 12.5 15.5 14.4 1.0
Anal-fin base length 6.7 5.9 7.2 6.5 0.5
Pectoral-fin length 15.3 13.3 16.0 14.9 0.8
Pelvic-fin length 13.8 11.7 13.8 12.4 0.7
In percent of head length
Head depth at eye 43 37 49 43 2.9
Snout length 44 38 45 42 1.9
Eye diameter 11 11 16 13 1.4
Postorbital distance 44 40 48 44 2.1
Maximum head width 57 57 66 60 3.0
Interorbital width 25 22 28 26 2.1
270
Fig. 13. Tucinoemacheilus saadii, ZM-CBSU7169B, holotype, 47.1 mm SL; Iran: stream Tang-e-Tizab.
Fig. 14. Turcinoemacheilus saadii, Iran: stream Tang-e-Tizab; paratypes; a, ZM-CBSU 7168B, 56.4 mm SL; b, ZM-
CBSU 7166B, 49.5 mm SL; c, ZM-CBSU 7171B, 47.1 mm SL.
Fig. 15. Turcinoemacheilus saadii, collected with ZM-CBSU7169B, about 50 mm SL; Iran: stream Tang-e-Tizab.
fin deeply emarginated. Largest known specimen
63.6 mm SL.
Dorsal fin with 7
1
/
2 branched rays. Anal fin
with 4 1
/
2-5 1
/
2 (usually 5 1
/
2) branched rays. Caudal
fin with 8 + 8 branched rays. Pectoral fin with 7-8
and pelvic fin with 5-6 (usually 5) branched rays.
Body without scales. Lateral line incomplete, with
18-24 pores, not reaching to dorsal-fin origin.
Anterior nostril opening on anterior side of a low,
pointed and flap-like tube. Anterior nostril open-
a
b
c
Ichthyol. Explor. Freshwaters, Vol. 24, No. 3
271
ing on anterior side of a low, blunt and flap-like
tube. Nostrils close together, posterior tip of an-
terior nostril reaching into posterior nostril when
folded backward. No suborbital flap or groove
in males. Mouth small, slightly arched (Fig. 7c).
Lips moderately thick. A median interruption in
lower lip. Upper lip without median incision.
Processus dentiformis small and blunt. No me-
dian notch in lower jaw. Barbels short, inner
rostral barbel not reaching base of maxillary
barbel; outer one reaching slightly beyond base
of maxillary barbel, not reaching vertical of ante-
rior margin of eye. Maxillary barbel reaching
vertical of middle of eye. No external sexual di-
morphism observed.
Coloration. In alcohol and in life, body pale
cream yellow. Seven to nine wide, irregularly
shaped dark brown saddles on body; three to four
in front of dorsal fin, sometimes dissociated or
partly dissociated into two narrow bars on flank.
One wide saddle starting in front of and reaching
below dorsal-fin origin and one saddle at poste-
rior part of dorsal-fin base reaching behind dorsal-
fin base. Two or three saddles on caudal pedun-
cle. Predorsal saddles reaching slightly below
lateral midline, saddles behind dorsal-fin base
reaching to ventral body. Some saddles, usually
those on predorsal flank, Y-shaped in few indi-
viduals. Saddles usually wider on back than on
flank and wider on lateral midline than below
and above lateral midline. Saddles always discon-
nected from each other, never fused to form a
lateral stripe. A very faint inner axial streak vis-
ible in preserved specimens. A dark brown or
black bar at posterior extremity of caudal pedun-
cle. In front of this bar, a whitish or yellowish
blotch on upper caudal peduncle. Cheeks and
ventral surface of head cream or pale yellow, head
above cheeks dark brown without spots or
blotches. Fins hyaline.
Distribution and habitat. Turcinoemacheilus saa-
dii is known from the Bashar River which flows
to the Karoun and from the Gamasiab River which
flows to Karkheh. At the type locality, the stream
Tang-e-Tizab (Fig. 16) is about 20 m wide, the
substrate consists of coarse gravel and boulders
and the water is fast-running.
Fig. 16. Stream Tang-e-Tizab, a tributary to Bashar River, Iran; type locality of Turcinoemacheilus saadii.
272
Etymology. The species is named for Abu
-Mu-
hammad Muslih al-Dı
n bin Abdalla
h Shı
ra
,
Saadii Shirazi (Persian:
) better known by his pen-name as Sa
(Persian: ) or, simply, Saadi, one of the major
Persian poets of the medieval period.
Remarks. The phylogenetic tree reconstruction
(Fig. 1) suggests that T. saadii is the sister species
of the pair T. kosswigi and T. minimus. It is sepa-
rated from T. kosswigi by 8.5 % and from sympat-
ric T. hafezi by 18.2 % K2P COI sequence diver-
gence, which is considered as a strong indicator
that these fishes represent different species (see
Herbert et al., 2003). Turcinoemacheilus saadii is
further distinguished from T. kosswigi and T. mini-
mus by colour pattern (7-9 distinct saddles vs.
lateral stripe made by a series of blotches con-
nected to dorsal blotches). In T. saadii, the caudal
fin is always deeply emarginated and the central
rays of the caudal fin are 60-80 % of the length
of the lower caudal-fin lobe. In T. kosswigi, the
caudal fin is only slightly emarginated in most
individuals and the central rays of the caudal fin
are 80-95 % of the length of the lower caudal-fin
lobe. However, in some individuals of T. koss-
wigi, the caudal fin is more deeply emarginate
and the ratio is down to 70-80 % and largely
overlapping with T. saadii.
Turcinoemacheilus saadii occurs in sympatry
with T. hafezi. It is distinguished from this species
by the more anterior position of the anus, which
is in front of the middle or in the middle between
the pelvic-fin origin and the anal-fin origin (dis-
tance from anus to anal-fin origin 0.4-0.5 times
in distance from pelvic-fin to anal-fin origins)
while it is behind the middle (0.2-0.4 times) in
T. hafezi. Turcinoemacheilus saadii is further distin-
guished by a free distal part of the axillary pelvic
lobe (vs. completely attached to body), a nar-
rower caudal peduncle (depth 2.4-3.2 times in its
length vs. 1.9-2.4), anterior and posterior nostrils
very closely together (vs. separated), tip of ante-
rior nostril reaching into posterior nostril when
folded down (vs. not) and differences in colour
patters (7-9 distinct saddles vs. lateral stripe
present, disconnected from dorsal blotches or
mottles colour pattern in T. hafezi).
Turcinoemacheilus saadii is distinguished from
T. himalaya by a completely scaleless body (vs.
small cycloid scales present on the posterior half
of the body).
Comparative material. Turcinoemacheilus hafezi: all
from Iran: FSJF 3222, 33, 24-46 mm SL; Yasouj prov.:
Beshar (Bashar) river at Doruhan, 30°51'07" N 51°20'
31" E. – FSJF 3002, 3, 32.4-45.8 mm SL; Sezar river, a
tributary of Dez river, 33°28'N 49°03'E. – ZFMK 48841,
holotype, 41.0 mm SL; ZFMK 48842-48848, paratypes,
7, 34.1-66.7 mm SL; FSJF 1649, 10, 34.4-54.7 mm SL;
Chaharmahal and Bakhtiari prov.: stream Shalamzar
10 km after Joneghan in the direction to Ardal, a tribu-
tary of Ab-Kohrang River, 32°05'22" N 50°39'58" E.
Material used in the molecular genetic analysis: FSJF
DNA-1985; Yasouj prov.: Bashar river at Doruhan,
30°51'07" N 51°20'31" E (GenBank accession numbers:
KJ179254, KJ179259). – ZM-CBSU T497, ZM-CBSU T539;
Fars prov.: stream Tang-e-Tizab, a tributary to Bashar
river which drains to the Karoun, 30°23'12" N 51°46'50" E
(GenBank accession numbers: KJ179252, KJ179264).
T. himalaya: paratypes, all from Nepal: KU 40557,
2, 38.0, 59.3 mm SL; Indrawati river at Melamchi town-
ship, 27°49'42" N 85°34'37" E. – KU 40568, 3, 39.0-
51.4 mm SL; Melamchi river, 3 miles upstream from
confluence with Indrawatir river, 27°57'22" N 85°32'
27" E. – KU 40280, 4, 44.9-49.7 mm SL; Gaundi river at
Gumti, 28°03'54" N 83°33'36" E. – KU 40281, 2, 36.4,
38.8 mm SL; Seti river at Khairenitar, 28°02'00" N 84°04'
00" E.
T. kosswigi: ZMH H1884, holotype, 53 mm SL; ZMH
H1885, 5 paratypes, 41-52 mm SL; Turkey: Hakkari
prov.: Kapozik Kadun. – FSJF 3345, 7, 41.3-50.8 mm SL;
Iraq: stream north-west of Saburawa, a tributary of
Tabin river, a tributary to Little Zab, 35°50'01" N 45°06'
16" E. – FSJF 3350, 7, 40.9-50.2 mm SL; Iraq: Kuna
Massi stream in Sevanja, a tributary to Little Zab,
35°47.35' N 45°24.18' E. – FSJF 3358, 1, 52.3 mm SL; Iraq:
stream Zalm at Khurmal, a tributary to Sirvan, 35°
18.38' N 45°58.26' E. – FSJF 3370, 4, 40.1-45.9 mm SL;
Iraq: stream Suraw near Suraw village, a tributary to
Little Zab, 35°45.76' N 45°59.09' E. – FSJF 3377, 3, 41.5-
46.1 mm SL; Iraq: Chami Rean river near Ziraran, a
tributary to Great Zab, 36°56.60' N 44°11.72' E. Material
used in the molecular genetic analysis: FSJF DNA-2214;
Iraq: stream Suraw near Suraw village, a tributary to
Little Zab, 35°45.76' N 45°59.09' E (GenBank accession
number: KJ179260). – FSJF DNA-2222; Iraq: stream
north-west of Saburawa, a tributary of Tabin river, a
tributary to Little Zab, 35°50'01" N 45°06'16" E (GenBank
accession number: KJ179255). – FSJF DNA-2226; Iraq:
Chami Rean river near Ziraran, a tributary to Great Zab,
36°56.60' N 44°11.72' E (GenBank accession number:
KJ179265). – FSJF DNA-2232; Iraq: Kuna Massi stream
in Sevanja, a tributary to Little Zab, 35°47.35' N 45°
24.18' E (GenBank accession number: KJ179262). – ZM-
CBSU T507, ZM-CBSU T509; Iran: Sirvan river, 35°06'
43" N 46°15'24" E (GenBank accession numbers: KJ179258,
KJ179245).
Paracobitis malapterura: material used in the mo-
lecular genetic analysis: FSJF DNA-1992; Iran: Albroz
prov.: Kordan river near Karaj, 35°57'11" N 50°50'15" E
(GenBank accession numbers: KJ179266, KJ179267).
Ichthyol. Explor. Freshwaters, Vol. 24, No. 3
273
Paraschistura bampurensis: Material used in the
molecular genetic analysis: FSJF DNA-2001; Iran: Sistan
and Baluchestan prov.: Karvander river at bridge north
of Karevander, 27°51'18" N 60°46'03" E (GenBank acces-
sion numbers: KJ179268, KJ179269).
Acknowledgments
We are pleased to thank Ralf Thiel (ZMH), Andrew
Bentley (KU) and Maurice Kottelat (CMK) for allowing
JF to examine material of Turcinoemacheilus under their
care, Rahman Patimar and Kiavash Golzarianpour (both
Gonbad) for providing JF with material of T. saadii,
P. malapterura and P. bampurensis, and Jörg Bohlen
(Libechov) for sharing unpublished observations on
T. minimus. We also thank Shiraz University for finan-
cial supports to first and second authors and Environ-
ment Department of Fars and Isfahan provinces for
their collaborations in fish collection in Iran. Support
also came from the project FREDIE (Freshwater Diver-
sity Identification for Europe, www.fredie.eu), funded
by the Joint Initiative for Research and Innovation
(PAKT) program of the Leibniz Association.
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Received 16 July 2013
Revised 11 February 2014
Accepted 12 February 2014
... Since the description of Turcinoemacheilus kosswigi from headwaters of the Greater Zab River in Türkiye by Bănărescu & Nalbant (1964), there had been little attention on this peculiar genus of nemacheilid loaches for about half a century, until Conway et al. (2011) described an additional species, T. himalaya, from Nepal, shortly followed by Golzarianpour et al. (2013) and Esmaeili et al. (2014) describing four new species (T. bahaii, T. hafezi, T. minimus, T. saadii) from the Gulf basin in Iran and Türkiye. ...
... After this study, we expected the species diversity of Turcinoemacheilus to be well understood, until Kaya et al. (2023) described another new member of this genus, T. ekmekciae, from the upper Tigris and Euphrates drainage in Türkiye. Esmaeili et al. (2014) examined the type material of T. kosswigi (ZMH H1884, ZMH H1885) and found no clear differences between these, and Turcinoemacheilus found in the Greater Zab drainage in Iraq. Fish examined from the Greater Zab in Iraq by Esmaeili et al. (2014) were collected only about 80 km south of the type locality in the area, from where Kaya et al. (2023) collected their specimens of T. kosswigi. ...
... Esmaeili et al. (2014) examined the type material of T. kosswigi (ZMH H1884, ZMH H1885) and found no clear differences between these, and Turcinoemacheilus found in the Greater Zab drainage in Iraq. Fish examined from the Greater Zab in Iraq by Esmaeili et al. (2014) were collected only about 80 km south of the type locality in the area, from where Kaya et al. (2023) collected their specimens of T. kosswigi. As populations from the Greater Zab in Iraq, and the Lesser Zab and Sirvan in Iraq and Iran, form one monophyletic clade, they were all identified as T. kosswigi by Esmaeili et al. (2014). ...
Article
Turcinoemacheilus inexpectatus, new species, is described from the Greater Zab, Lesser Zab, and Sirvan drainages, all tributaries of the middle Tigris. It belongs to the T. kosswigi species group. The new species is distinguished from its congeners by an indistinct or prominent midlateral stripe broader than the eye diameter often overlaid by a row of dark-brown blotches, greater pre-pelvic distance, deeper caudal peduncle, and wider interorbital distance.
... The specimen from the Great Zab region, specifically from the Chami Rean River in Iraq, sampled by Esmaeili et al. (2014) is of great note. It is important to note that this site is located approximately 80 km southeast of the type locality of T. kosswigi, which is Kapozik Kadun located in Hakkari (southeastern Türkiye). ...
... It is important to note that this site is located approximately 80 km southeast of the type locality of T. kosswigi, which is Kapozik Kadun located in Hakkari (southeastern Türkiye). It seems that Esmaeili et al. (2014) overlooked the possibility that a species found in a different drainage of the same river system and at such close range might not actually be T. kosswigi. However, our molecular analysis and morphological investigation provided clear insight into this matter. ...
... However, our molecular analysis and morphological investigation provided clear insight into this matter. We found that the population from the upper Great Zab region significantly diverged by the COI sequence data from the material identified as T. kosswigi by Esmaeili et al. (2014). Alongside our molecular findings, we conducted a detailed morphological examination of our T. kosswigi specimens to further support our hypothesis. ...
Article
Full-text available
Turcinoemacheilus ekmekciae, new species, from upper Euphrates and Tigris drainages is distinguished from other species of Turcinoemacheilus in Western Asia by having a dark stripe broader than the eye diameter along the lateral line, rarely possessing roundish blotches, 5–6 mandibular pores in mandibular canal, a comperatvely smaller head, a deeper body, and a greater pre‐pelvic distance. Our specimens collected from the upper Great Zab, near the type locality of Turcinoemacheilus kosswigi, showed notable genetic divergence (a minimum K2P of 3.3%) from sequences reported as T. kosswigi in previous studies. Despite morphological similarities, this molecular difference suggests that the populations analysed in previous studies may represent a potential new species of Turcinoemacheilus, which we tentatively named as Turcinoemacheius cf. kosswigi. Molecular data also suggest that T. ekmekciae is characterized by a minimum K2P distance of 3.5% from Turcinoemacheilus minimus and T. cf. kosswigi. The three methods for species delimitation (assemble species by automatic partitioning [ASAP], Poisson tree processes [PTP], and multi‐rate PTP [mPTP]) that were utilized for testing species assignments consistently identified our test group as a distinct species.
... The final alignment consisted of 701 base pairs, with 484 positions being constant, 163 being parsimony informative, and 23 being singletons (considering only the in-group). The resulting tree topology is, in general, in agreement with previously published phylogenies that included samples of Turcinoemacheilus [9,10]. Both the BI and ML methods resulted in a similar tree (Figure 2), recovering 10 well-supported clades corresponding to 10 species in the Turcinoemacheilus genus: T. ansari sp. ...
... Esmaeili et al. [9] conducted a review of Turcinoemacheilus using both morphological and molecular data. They referenced two sequences of T. saadii (KJ179257, KJ179250) from the Leyleh River, which were found to be almost identical to our samples of T. saadii from Karkheh. ...
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Nemacheilid fishes in the genus Turcinoemacheilus are physically small members of the ichthyofauna communities of high-altitude and mountainous freshwater ecosystems. They are all distributed in Western Asia apart from a single species, described in the Himalayas. They are usually very similar in appearance, which complicates their proper identification and/or description. This is why it is important to use multidisciplinary and integrative taxonomical approaches in order to study their true diversity. In this study, three new species of Turcinoemacheilus are described from Iran, raising the total number of valid species to nine. Turcinoemacheilus ansari new species, is distinguished by the anus being situated behind the midpoint of the pelvic-fin and anal-fin origins and the short anal-fin base length. Turcinoemacheilus christofferi new species, differs by the anus being situated behind the midpoint of the pelvic-fin and anal-fin origins, with a complete lateral line reaching to the anterior part of the caudal fin. Turcinoemacheilus moghbeli new species, is distinguished by the anus being situated at or in front of the midpoint of the pelvic-fin and anal-fin origins, with a great pre-pelvic distance and a caudal peduncle length 1.5-2.3 times its length. In Western Asia, all Turcinoemacheilus species are well separated by molecular characters, showing between 3.6 and 14.1% uncorrected p genetic distances in the COI barcode region. This work shows the importance of studying the hidden diversity of under-sampled and understudied groups of organisms to have a clear image of true biodiversity in order to effectively conserve and protect it.
... Turcinoemacheilus kosswigi Banarescu & Nalbant, 1964 [N]-[LC], Kosswig's loach/Çöpçü baligi/Euphrates and Tigris river basins Fricke et al. 2007;Çiçek et al. 2015). Turcinoemacheilus minimus Esmaeili, Sayyadzadeh, Özulug, Geiger & Freyhof, 2014 [E]- [NE], Ceyhan loach/-/ Göksu River, Euphrates River basin (Esmaeili et al. 2014a;Çiçek et al. 2015). ...
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The current status of the inland waters ichthyofauna of Turkey is revised, and an updated checklist of the freshwater fishes is presented. The latest checklist included all species listed in the available previous study that was published in 2015, which is now updated after a period of five years. We revised the validity of previously accepted species and added newly described and reported species in Turkey. Some previously erroneously reported species and not established alien fishes were excluded from this checklist. A total of 384 fish species belonging to 20 orders and 34 families have been reported in the inland waters of Turkey. Among these, 15 species (3.9%) are non-native and 208 species (54.2%) are considered as endemic to Turkey. A total of 119 species previously reported from Turkey have been excluded from Turkish ichthyofauna list, either in the present study or in previous studies.
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Oxynoemacheilus chaboras , new species, from the stream Beyazsu in the Euphrates drainage, belongs to the O. persa species group, being closely related to O. shehabi from the Orontes, O. sarus from the Seyhan and Ceyhan, O. euphraticus from the Euphrates and Tigris, O. karunensis from the Karkheh, and O. persa from Central Iran. The new species is distinguished from others in the O. persa group by having 8–9 pores in the supraorbital canal, two distinct black blotches at the caudal-fin base, a rudimentary and shallow pelvic axillary lobe, 6–10 irregularly shaped bars on the flank, and a deep head, body, and caudal peduncle. Oxynoemacheilus chaboras sp. nov. is most closely related to O. euphraticus , from which it is differentiated by a mean uncorrected p -distance of 3.24% (min. 3.09%) in its COI barcode gene.
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the current status of the freshwater fishes of Iraq is revised, and an updated checklist is presented. the confirmed freshwater fishes of Iraq comprise 98 species belonging to 16 orders, 28 families, and 56 genera. Among these, 21 species (21.4%) are alien, and three species (3.0%) are considered endemic to Iraq. the orders with the largest numbers of species in the ichthyofauna of Iraq are Cypriniformes (57 species), followed by Siluriformes (12 species), Mugiliformes and Cyprinodontiformes (six species in each), Acanthuriformes and Cichliformes (three species in each), Centrarchiformes and Gobiiformes (two species in each), and the others represent only one species. At the family level, the Cyprinidae have the greatest number of species (28; 28.3% of the total species), followed by Nemacheilidae (16 species), Leuciscidae (eight species), Mugilidae (six species), Sisoridae (five species), and Xenocyprididae (four species in each). A total of 20 species that have been previously reported from Iraq have been excluded from the checklist. According to IUCN Red List criteria, among 77 naturally distributed species (alien species not included), 10 species (13.0%) are categorized as threatened extinctions, with three (3.9%) as CR, one (1.3%) as EN, and six (7.8%) as vU. A total of 38 taxa were assessed (49.4%) as LC, 27 species were not assigned NE (35.1%), and two species (2.0%) were classified as DD, i.e., with insufficient knowledge.
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DNA barcodes (COI) of Troglocobitis starostini, endemic to a single site in Turkmenistan, were analysed and put into the taxonomic context of the large group of nemacheilid loaches known from Western and Central Asia. All applied phylogenetic tree-based analyses place the species into the genus Paracobitis. This finding supports previous morphological studies. While the exact position of Troglocobitis starostini within Paracobitis was not resolved unambiguously, it was constantly recovered within Paracobitis, irrespective of the tree reconstruction method applied. With a minimum interspecific K2P distance of 7.19% P. persa was the closest hit in our dataset, which comprised a total of ten species of Paracobitis, which showed an average interspecific K2P distance of 5.43% (range 2.78-9.44%).
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Five new species of Glyptothorax are described from Iran. Glyptothorax alidaeii, new species, from the Seimare in the Karkheh drainage, G. galaxias, new species, from the upper Karun drainage, G. hosseinpanahii, new species, from the Zohreh drainage, G. pallens, new species, from the Sirvan drainage, and G. shapuri, new species, from Shapur in the Helleh drainage. Glyptothorax silviae from the Jarrahi drainage is re-diagnosed. All six species are morphologically distinguishable by the structure of the thoracic adhesive apparatus, as well as morphometric characters and details in the colour pattern. They form distinct mitochondrial clades between 1.2% and 4.1% minimum K2P distance based on the mitochondrial DNA barcode region.
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Although much biological research depends upon species diagnoses, taxonomic expertise is collapsing. We are convinced that the sole prospect for a sustainable identification capability lies in the construction of systems that employ DNA sequences as taxon 'barcodes'. We establish that the mitochondrial gene cytochrome c oxidase I (COI) can serve as the core of a global bioidentification system for animals. First, we demonstrate that COI profiles, derived from the low-density sampling of higher taxonomic categories, ordinarily assign newly analysed taxa to the appropriate phylum or order. Second, we demonstrate that species-level assignments can be obtained by creating comprehensive COI profiles. A model COI profile, based upon the analysis of a single individual from each of 200 closely allied species of lepidopterans, was 100% successful in correctly identifying subsequent specimens. When fully developed, a COI identification system will provide a reliable, cost-effective and accessible solution to the current problem of species identification. Its assembly will also generate important new insights into the diversification of life and the rules of molecular evolution.
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Turcinoemacheilus himalaya, new species, is described from the Koshi and Gandaki River basins of Nepal. The new species is distinguished from its hypothesised congener, Turcinoemacheilus kosswigi, from the Euphrates, Tigris and Karoun basins of the Middle East, by the presence of small scales on the posterior half of its body (v. absence of all scales), its shorter caudal peduncle (caudal peduncle length 12–15% standard length, L S v. 16–23), its shorter snout (snout length 28–36% head length, L H v. 40–49) and by features of its colour pattern, including the presence of small irregularly shaped dark grey markings over the lateral body surface. Turcinoemacheilus himalaya is known to date only from Nepal.
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Turcinoemacheilus kosswigi (Bănărescu and Nalbant, 1964) was recorded for the first time in Iran from the River Karoun drainage, which belongs to the Euphrates-Tigris drainage. Formerly believed to be an endemic species in the basin of River Tigris, it is now recorded in the upper River Euphrates basin. This extension of its recorded range makes it likely that it has been overlooked in other parts of the Euphrates-Tigris system. The species is distinguished from all other loaches by the pelvic-fin origin in front of the dorsal-fin origin and having the anus closer to the pelvic-fin base than to the anal fin origin.
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Oxynoemacheilus is a species-rich genus of nemacheilid fishes known from Albania eastwards to Central Iran. There are 58 available species-group names and 41 species are recognized here as valid. Four species occur in Europe (O. bureschi, O. merga, O. pindus, O. theophilii) and 38 species are found in Anatolia and the Middle East. Barbatula paucilepis, Cobitis tigris, Nemacheilus tigris cyri, N. namiri, Noemacheilus tigris seyhanensis and Orthrias an-gorae ercisianus recently placed in Paracobitis as well as Nun galilaeus and Seminemacheilus tongiorgii are all transferred to Oxynoemacheilus.
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Turcinoemacheilus hafezi, new species, from Karoun and Dez River drainages, Iran, is distinguished from both other species of Turcinoemacheilus by having the anus situated in a more posterior position, a mottled colour pattern or a prominent irregular dark brown lateral stripe along the lateral midline disconnected from dark brown blotches on back, and scales completely absent.
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The loach fish Turcinoemacheilus kosswigi Bănărescu & Nalbant, 1964 (Balitoridae), formerly believed to be an endemic species in the basin of Tigris River, was recorded for the first time in the upper Euphrates River basin. This extension of its recorded range makes it likely that it has been overlooked in other parts of the Euphrates-Tigris system. T. kosswigi seems to be a habitat specialist living on coarse gravel beds in fast-running water. Its ability to attach to hard substrate even outside the water has to be understood as an adaptation to this habitat. Kurzfassung. Die Schmerlenart Turcinoemacheilus kosswigi Bănărescu & Nalbant, 1964 (Bali- toridae), bislang nur aus dem Einzugsgebiet des Tigris bekannt, wurde zum ersten Mal im Be- cken des oberen Euphrat nachgewiesen. Diese Erweiterung des belegten Verbreitungsgebietes weist wohl auf ein weites Vorkommen im Euphrat-Tigris-System hin. T. kosswigi scheint ein Ha- bitatspezialist für schnell überströmte Grobkiesbänke zu sein. Die Fähigkeit zum Anheften an den Untergrund, auch ausserhalb des Wassers, ist als Anpassung an diesen Habitat zu verstehen.
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Here we present evidence, based on 10 datasets comprising 5283 sequences for 200 genera, that the use of the Kimura-2-parameter (K2P) model in DNA-barcoding studies is poorly justified. We demonstrate that K2P is neither expected nor confirmed to be an appropriate model for closely related COI sequences. In addition, we show that the use of uncorrected distances yields higher or similar identification success rates for neighbour-joining trees and distance-based identification techniques. K2P also does not widen the barcoding gap for closely related sequences. We conclude that the spread of K2P through the barcoding literature is difficult to explain, and urge the use of evidence-based approaches to DNA barcoding. © The Willi Hennig Society 2011.
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A new method called the neighbor-joining method is proposed for reconstructing phylogenetic trees from evolutionary distance data. The principle of this method is to find pairs of operational taxonomic units (OTUs [= neighbors]) that minimize the total branch length at each stage of clustering of OTUs starting with a starlike tree. The branch lengths as well as the topology of a parsimonious tree can quickly be obtained by using this method. Using computer simulation, we studied the efficiency of this method in obtaining the correct unrooted tree in comparison with that of five other tree-making methods: the unweighted pair group method of analysis, Farris's method, Sattath and Tversky's method, Li's method, and Tateno et al.'s modified Farris method. The new, neighbor-joining method and Sattath and Tversky's method are shown to be generally better than the other methods.
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— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.