Corresponding author: Hamid Reza Esmaeili (
Academic editor: R. Mayden
As presently recognized, the genus
Alwan NH, Zareian H, Esmaeili HR (2016)
The Middle East is a transition zone between three major biogeographic units, the Palaearctic, the Afrotropical, and the Oriental realms. It served as an important crossroad of biotic exchange resulting in an outstanding biological diversity of freshwater fishes (
Members of the genus
As presently recognized, the genus
Members of the cytochrome c oxidase subunit I
Traditionally,
After anaesthesia, fishes were either fixed in 5% formaldehyde, and stored in 70% ethanol, or directly fixed in 99% ethanol (for molecular studies). Measurements were made with a digital caliper and recorded to 0.01 mm. All measurements were made point to point, and never by projections. Methods for counts and measurements follow Standard length
Abbreviations used: , standard length , lateral head length
Abbreviations used for museum collections: Zoological Museum of Shiraz University, Collection of Biology Department, Shiraz, Iran Senckenberg Research Institute and Natural History Museum Jörg Freyhof
For DNA sequencing, specimens were directly fixed in 99% molecular grade ethanol. Mitochondrial DNA was extracted using Salt method (
Purification and sequencing of the PCR products were conducted at Macrogen Korea Laboratories using the aforementioned primer sets.
Data processing and sequence assembly was done in BioEdit 7.2.5 ( (Bayesian Information Criterion) Bayesian analysis Markov Chain Monte Carlo method
General body shape and appearance are shown in Figs
Live specimen of
Morphometric data of
Holotype | Paratypes (n=33) | |||||||||
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Range | Mean | SD | Range | Mean | SD | Range | Mean | SD | ||
Standard length (mm) | 157.64 | 67.23–157.64 | 110.67 | 74.30–149.30 | 112.56 | 51.31–231 | 109.30 | |||
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Head length | 23.76 | 22.87–26.33 | 24.5 | 0.80 | 21.47–25.98 | 23.56 | 0.98 | 24.28–29.62 | 26.84 | 1.32 |
Body depth at dorsal-fin origin | 21.82 | 21.33–25.04 | 23.15 | 0.98 | 19.78–24.55 | 21.82 | 1.19 | 19.58–27.78 | 23.32 | 2.11 |
Predorsal length | 49.07 | 47.75–53.43 | 50.23 | 1.31 | 48.85–55.05 | 51.59 | 1.34 | 44.33–55.12 | 51.93 | 2.51 |
Postdorsal length | 54.13 | 54.13–63.19 | 57.53 | 1.88 | 48.20–60.13 | 55.24 | 2.78 | 50.79–59.64 | 55.06 | 2.60 |
Preanal length | 72.45 | 70.22–76.14 | 72.8 | 1.36 | 71.34–76.34 | 74.01 | 1.28 | 69.37–78.38 | 75.61 | 2.07 |
Prepelvic length | 53.74 | 50.22–55.90 | 52.84 | 1.31 | 50.17–56.64 | 53.58 | 1.49 | 51.23–61.21 | 56.22 | 2.36 |
Distance between pectoral and pelvic-fin origins | 32.42 | 27.81–32.42 | 30.19 | 1.13 | 29.07–33.64 | 31.30 | 1.04 | 25.55–32.66 | 30.87 | 2.15 |
Distance between pelvic and anal-fin origins | 21.48 | 19.31–23.17 | 21.12 | 0.88 | 19.90–23.65 | 21.60 | 0.91 | 18.32–23.41 | 20.83 | 1.56 |
Depth of caudal peduncle | 10.37 | 10.03–11.61 | 10.65 | 0.37 | 8.58–10.84 | 10.05 | 0.50 | 8.98–11.15 | 10.43 | 0.60 |
Length of caudal peduncle | 20.73 | 17.16–22. 35 | 19.85 | 1.14 | 18.64–22.01 | 19.81 | 0.91 | 15.19–20.11 | 17.67 | 1.30 |
Dorsal-fin base length | 12.71 | 12.27–16.17 | 14.41 | 0.89 | 11.75–15.28 | 13.51 | 0.80 | 10.46–14.39 | 12.98 | 1.11 |
Anal-fin base length | 6.78 | 6.38–8.85 | 7.39 | 0.55 | 6.96–8.80 | 7.88 | 0.58 | 6.24–8.27 | 7.17 | 0.60 |
Pectoral-fin length | 17.32 | 16.68–20.46 | 18.43 | 0.89 | 16.39–20.96 | 18.38 | 0.99 | 16.15–19.16 | 17.86 | 1.01 |
Pelvic-fin length | 15.05 | 14.24–16.96 | 15.61 | 0.68 | 13.85–18.08 | 15.61 | 1.08 | 13.58–16.23 | 15.08 | 0.82 |
Length of the longest dorsal fin ray | 18.90 | 14.92–21.58 | 19.57 | 1.27 | 16.42–21.22 | 18.78 | 1.06 | 16.35–21.53 | 19.03 | 1.47 |
Mouth width | 8.65 | 7.48–9.77 | 8.63 | 0.51 | 6.49–8.89 | 7.87 | 0.57 | 6.51–9.38 | 8.1 | 0.73 |
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Head depth at eye | 56.88 | 49.05–61.87 | 54.21 | 2.73 | 48.01–56.63 | 67.01 | 2.26 | 49.17–57.97 | 65.47 | 3.96 |
Snout length | 38.32 | 31.60–47.70 | 38.08 | 2.60 | 32.69–38.89 | 35.55 | 1.67 | 32.65–40.61 | 36.18 | 2.41 |
Postorbital distance | 48.83 | 33.82–51.84 | 48.01 | 3.01 | 47.66–56.59 | 51.57 | 1.89 | 46.58–54.84 | 51.05 | 2.28 |
Interorbital width | 40.04 | 34.62–42.81 | 38.19 | 1.98 | 33.88–41.49 | 37.15 | 1.93 | 30.90–40.16 | 35.57 | 2.70 |
Eye diameter | 15.97 | 15.07–23.57 | 18.52 | 2.36 | 13.91–24.44 | 17.36 | 2.11 | 11.95–26.18 | 18.23 | 3.43 |
Maximum head width | 60.53 | 51.75–66.89 | 59.60 | 3.99 | 57.83–69.68 | 62.76 | 3.04 | 47.38–59.00 | 54.62 | 3.39 |
Barbel length | 15.14 | 13.30–20.20 | 16.25 | 1.66 | 15.66–24.60 | 19.71 | 2.27 | 13.34–24.64 | 18.11 | 2.73 |
Dorsal-fin origin anterior to pelvic-fin origin, its outer margin usually straight to concave with 3–5 unbranched and 8–9 branched rays (3 and 8 in holotype, respectively); last unbranched dorsal-fin ray weakly to moderately ossified, flexible and soft at the tip, serrated in 1/2–2/3 of its length (Fig.
Dorsal fins of
Number of pectoral and pelvic fin rays in examined
Pectoral fin rays | Pelvic fin rays | |||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | 22 | 7 | 8 | 9 | 10 | 11 | ||
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2 | 10 | 4 | 6 | 2 | 14 | 10 | |||||||||
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6 | 10 | 8 | 11 | 7 | 1 | 3 | 1 | 14 | 16 | 12 | 8 | ||||
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1 | 7 | 2 | 1 | 9 | 2 | ||||||||||
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3 | 12 | 4 | 1 | 9 | 10 | 1 | |||||||||
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2 | 8 | 17 | 8 | 5 | 22 | 17 | 1 |
Number of branched caudal fin rays in examined
Branched caudal fin rays | 15 | 16 | 17 | 18 | 19 | 20 |
---|---|---|---|---|---|---|
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3 | 21 | 3 | |||
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1 | 29 | 2 | 1 | ||
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2 | 8 | 1 | |||
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1 | 9 | 16 | 11 | 3 |
Scales small, total lateral-line scales 70–84; 12–17 scales between dorsal-fin origin and lateral line (Table
Number of scales above (between dorsal-fin origin and lateral line) and below (between dorsal-fin origin and lateral line) lateral line in examined
Above lateral line | Below lateral line | ||||||||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | |
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3 | 6 | 4 | 12 | 3 | 13 | 7 | 3 | 1 | ||||||||||||
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1 | 9 | 9 | 15 | 15 | 1 | 11 | 20 | 18 | ||||||||||||
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1 | 10 | 4 | 5 | 2 | ||||||||||||||||
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1 | 2 | 1 | 8 | 7 | 1 | 2 | 4 | 3 | 9 | 2 | ||||||||||
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2 | 1 | 1 | 7 | 16 | 7 | 3 | 3 | 7 | 19 | 8 | 6 |
Number of circum-pendicular scales in examined
19 | 21 | 22 | 23 | 24 | 25 | 26 | 27 | 28 | 29 | 30 | 31 | 32 | 33 | |
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1 | 3 | 2 | 5 | 5 | 3 | 4 | 3 | ||||||
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3 | 11 | 7 | 14 | 4 | 7 | 5 | |||||||
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1 | 6 | 2 | 4 | 1 | 1 | ||||||||
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5 | 1 | 3 | 1 | 1 |
Dorsum, head and sides grey or brownish-grey dorsally and beige or yellow ventrally; dorsal and caudal fins dusky grey; pectoral, pelvic and anal fins white or beige with or without grey tinge; blotches and spots well discernible (Figs
Breeding tubercles present in both sexes, being bigger and more pronounced in males. Tubercles present on the sides of the snout but may also cover the entire body surface, on and above the lateral line with one or two tubercles per scale but not on each scale, below the lateral line especially in the area above the anal fin and on the branched anal-fin rays; tip of anal fin reaching to or beyond the vertical of the caudal-fin base in females and to about 2/3 of the caudal peduncle in males.
Beshar River at Taleh Gah village, Karun River drainage, type locality of
The new species is named after Brian W. Coad, a well-known ichthyologist for his valuable contribution to the knowledge of freshwater fishes of Iran.
The presence of one pair of barbels in
Number of caudal-peduncle scales in examined
10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | 22 | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
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6 | 3 | 7 | 9 | 1 | 1 | |||||||
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1 | 1 | 2 | 8 | 15 | 2 | 3 | 1 | |||||
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4 | 2 | 4 | 1 | |||||||||
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2 | 1 | 10 | 6 | 1 | ||||||||
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1 | 1 | 2 | 3 | 5 | 14 | 8 | 1 | 4 | 1 |
Number of lateral-line scales in examined
58 | 59 | 61 | 62 | 63 | 64 | 65 | 66 | 67 | 68 | 69 | 70 | 71 | 72 | 73 | 74 | 75 | 76 | 77 | 78 | 79 | 80 | 81 | 82 | 83 | 84 | 85 | 87 | 89 | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
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1 | 3 | 1 | 2 | 1 | 2 | 4 | 3 | 1 | 2 | 4 | 1 | 1 | ||||||||||||||||
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2 | 1 | 2 | 6 | 1 | 4 | 4 | 1 | 5 | 3 | 5 | 6 | 5 | 1 | 1 | ||||||||||||||
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1 | 1 | 2 | 1 | 2 | 1 | 2 | 1 | |||||||||||||||||||||
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1 | 2 | 1 | 2 | 2 | 1 | 2 | 3 | 1 | 1 | 2 | 2 | |||||||||||||||||
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1 | 2 | 1 | 3 | 1 | 5 | 4 | 2 | 2 | 2 | 5 | 3 | 2 | 4 | 2 | 1 |
Gill rakers on the lower limb of the first gill arch in studied
GR | 8 | 9 | 10 | 11 | 12 | 13 | 17 | 18 | 19 | 20 | 22 | 24 |
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2 | 19 | 6 | |||||||||
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1 | 9 | 19 | 20 | ||||||||
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1 | 2 | 3 | 1 | 3 | 1 |
Number of total gill rakers on the first gill arch in examined
12 | 13 | 14 | 15 | 16 | 17 | 18 | 21 | 22 | 23 | 24 | 25 | 26 | 27 | 28 | 31 | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
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10 | 13 | 6 | |||||||||||||
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1 | 7 | 14 | 6 | 5 | |||||||||||
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2 | 2 | 2 | 1 | 4 | |||||||||||
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1 | 9 | 6 | 1 | 2 | 1 | ||||||||||
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1 | 1 | 9 | 11 | 7 | 5 | 4 | 1 | 1 |
It is distinguished from
Compared to other Iranian species of
Bayesian tree inferred from
Bayesian tree inferred from
We generated
List of species used for molecular analysis for
Species | Accession Number | Locality |
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Stream Sangan, Karun River basin, Tigris basin, Iran |
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Sevah River, Kor basin, Iran |
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Beshar River, Karun basin, Tigris basin, Iran |
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Sevah River, Kor basin, Iran |
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Pozanti River, Mediterranean Sea basin, Turkey |
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Boga Cayi River, Mediterranean Sea basin, Turkey |
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Kizilirmak River, Black Sea basin, Turkey |
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Biggest tributary of Kurtbog˘azi dam lake, Sakarya River basin, Turkey |
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Stream Cakirca, Lake Iznik basin, Turkey |
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Unknown |
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Unknown |
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Kizilirmak River, Black Sea basin, Turkey |
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Unknown |
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Unknown |
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Bakacak stream, Marmara Sea basin, Turkey |
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Bakircay River, Turkey |
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Stream Guzelhisar, Aegean Sea basin, Turkey |
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Taghra Rud stream, Namak Lake basin, Iran |
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Kordan River, Namak Lake basin, Karaj, Iran | |
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Kordan River, Namak Lake basin, Karaj, Iran | |
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Ilica stream, Gulf of Antalya, Mediterranean Sea basin,Turkey |
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Goksu River, Mediterranean Sea basin, Turkey |
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Kargi Cayi River, Mediterranean Sea basin, Turkey |
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Beshar River, Tigris River basin, Iran | |
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Beshar River, Tigris River basin, Iran | |
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Beshar River, Tigris River basin, Iran | |
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Beshar River, Tigris River basin, Iran | |
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Karadut River, Euphrates basin, Turkey |
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Stream Arsuz, Iskenderun Gulf basin, Mediterranean Sea, Turkey |
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Yocalti River, Turkey |
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Spring Incesu, Orontes basin, Mediterranean Sea, Turkey |
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Yocalti River, Mediterranean Sea basin, Turkey |
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Unknown |
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Keltechinar River, Turkmenistan |
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Yanbash River, Turkmenistan |
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Yanbash River, Turkmenistan |
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Murgab River, Turkmenistan |
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Deli Cayi River, Van Lake basin, Turkey |
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Deli Cayi River, Van Lake basin, Turkey |
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Ghare Aghaj River, Mond River basin, Khaneh Zanian , Iran | |
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Ghare Aghaj River, Mond River basin, Khaneh Zanian , Iran | |
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Ghare Aghaj River, Mond River basin, Khaneh Zanian , Iran | |
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Eflatum spring, Beysehir Lake basin, Turkey |
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Sarioz stream, Beysehir Lake basin, Turkey |
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Ghare Aghaj River, Mond River basin, Firuzabad, Iran | |
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Ghare Aghaj River, Mond River basin, Firuzabad , Iran | |
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Saadii Tomb Spring, Maharlu basin, Iran | |
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Helleh River, Helleh basin, KohmarSorkhi, Iran | |
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Kor River, Kor basin, Kamfirouz, Iran | |
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Kor River, Kor basin, Kamfirouz, Iran | |
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Rodan River, Makran basin, Iran |
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Spring Golabii, 35 km north from Darab, Hormuz basin, Iran |
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Kizilirmak River, Black Sea basin, Turkey |
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Kelkit Cayi River, Black Sea basin, Turkey |
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Unknown |
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Unknown |
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Dez River, Karun River basin, Iran |
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Sultansuyu River, Euphrates basin, Turkey |
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Gelal River, Ab e Seymareh, Tigris River basin, Iran |
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Çatkit River, Mediterranean Sea basin, Turkey |
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Unknown |
List of species used for molecular analysis for
Species | Accession num. | Locality |
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Gilas spring, Tedzen basin, Iran | |
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Gilas spring, Tedzen basin, Iran | |
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Gilas spring, Tedzen basin, Iran | |
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Bezangan Lake, Tedzen basin, Iran | |
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Seyhan, Turkey |
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Seyhan, Turkey |
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Aksu, Turkey |
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Aksu, Turkey |
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Orontes, Turkey |
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Orontes, Turkey |
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Orontes, Turkey |
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Orontes, Turkey |
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Bakir, Turkey |
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Bakir, Turkey |
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Biga, Turkey |
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Biga, Turkey |
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Kordan River, Namak Lake basin, Karaj, Iran | |
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Kordan River, Namak Lake basin, Karaj, Iran | |
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Roudbar River, Kavir basin,Iran | |
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Roudbar River, Kavir basin,Iran | |
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Göksu, Turkey |
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Ilica, Turkey |
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Göksu, Turkey |
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Göksu, Turkey |
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Arsuz, Turkey |
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Orontes, Turkey |
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Orontes, Turkey |
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Orontes, Turkey |
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Orontes, Turkey |
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Ceyhan, Turkey |
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Orontes, Syria |
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Orontes, Syria |
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Ceyhan, Turkey |
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Ceyhan, Turkey |
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Ceyhan, Turkey |
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Jordan River Drainage, Syria |
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Orontes, Turkey |
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Orontes, Turkey |
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Ceyhan, Turkey |
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Ceyhan, Turkey |
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Ceyhan,Turkey |
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Ceyhan,Turkey |
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Ghare Aghaj River, Mond River basin, Khaneh Zanian, Iran | |
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Ghare Aghaj River, Mond River basin, Khaneh Zanian, Iran | |
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Ghare Aghaj River, Mond River basin, Khaneh Zanian, Iran | |
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Egirdir, Turkey |
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Egirdir, Turkey |
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Egirdir, Turkey |
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Egirdir, Turkey |
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Egirdir, Turkey |
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Simav, Turkey |
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Simav, Turkey |
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Karkheh River, Tigris River basin, Seymareh, Iran | |
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Gavi River, Tigris River basin, Illam, Iran | |
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Ceyhan Nehri, Turkey |
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Saadii Tomb Spring, Maharlou basin, Iran | |
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Spring Pirbanoo, Maharlou basin, Iran | |
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Helleh River, Helleh basin, KohmarSorkhi, Iran | |
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Helleh River, Helleh basin, KohmarSorkhi, Iran | |
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Kor River, Kor basin, Kamfiruz, Iran | |
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Kor River, Kor basin, Kamfiruz, Iran | |
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Kor River, Kor basin, Kamfiruz, Iran | |
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Beshar River, Tigris River basin, Iran | |
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Beshar River, Tigris River basin, Iran | |
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Beshar River, Tigris River basin, Iran | |
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Beshar River, Tigris River basin, Iran | |
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Beshar River, Tigris River basin, Iran | |
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Dalaman, Turkey | |
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Büyük Menderes, Turkey | |
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Dalaman, Turkey | |
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Unknown |
Diagnostic nucleotide substitutions found in mtDNA
Species | 6545 | 6620 | 6626 | 6665 | 6683 | 6713 | 6749 | 6758 | 6761 | 6770 | 6818 | 6845 | 6875 | 6887 | 6905 | 6986 | 6995 | 7076 | 7088 |
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C | A | A | T | G | G | T | C | G | G | G | A | A | C | C | G | A | G | T |
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T | G | C | T | G | A | G | T | G | A | A | C | G | T | T | G | G | G | C |
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C | A | A | T | A | G | T | C | G | G | G | A | A | C | C | A | A | G | T |
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T | A | G | T | G | A | G | T | A | A | G | C | A | T | T | G | G | A | C |
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T | A | A | C | G | G | T | C | A | A | G | A | A | C | C | T | A | G | T |
Diagnostic nucleotide substitutions found in
Species | 15430 | 15451 | 15457 | 15463 | 15472 | 15526 | 15550 | 15610 | 15670 | 15682 | 15760 | 15814 | 15925 | 16011 | 16027 | 16039 | 16045 | 16063 |
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C | G | T | A | A | G | G | T | G | T | G | G | G | C | C | A | G | C |
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T | A | C | A | A | G | G | A | A | C | A | A | G | T | T | G | A | T |
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T | G | C | A | G | G | A | C | G | T | G | G | G | C | C | A | G | C |
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T | A | C | G | A | A | A | T | A | T | G | A | A | C | T | G | A | T |
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T | G | C | G | A | A | A | T | A | T | G | A | A | C | T | G | A | T |
For inter-specific differences, the greatest pairwise genetic divergence between
Mean genetic distance for
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3.8 | |||||||||||||
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5.1 | 4.6 | ||||||||||||
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5.3 | 5.4 | 4.8 | |||||||||||
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4.4 | 4.7 | 4.5 | 5.7 | ||||||||||
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4.3 | 4.1 | 4.1 | 4.9 | 4.6 | |||||||||
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5.3 | 5.6 | 5.3 | 6.6 | 1.0 | 5.6 | ||||||||
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5.7 | 4.9 | 4.9 | 6.0 | 4.9 | 4.4 | 4.9 | |||||||
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8.1 | 8.4 | 9.3 | 9.5 | 8.5 | 8.6 | 8.8 | 11.0 | ||||||
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8.7 | 8.7 | 9.1 | 9.9 | 9.1 | 9.2 | 9.4 | 10.9 | 1.2 | |||||
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4.3 | 2.6 | 4.1 | 5.6 | 4.3 | 4.0 | 5.1 | 4.7 | 9.3 | 9.5 | ||||
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4.2 | 2.6 | 4.5 | 6.0 | 5.0 | 4.3 | 5.8 | 5.4 | 9.4 | 9.6 | 1.5 | |||
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8.5 | 8.8 | 9.0 | 9.9 | 8.7 | 9.5 | 9.2 | 11.4 | 1.5 | 1.1 | 9.6 | 9.7 | ||
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4.8 | 3.3 | 4.8 | 4.6 | 4.7 | 4.8 | 5.4 | 5.8 | 8.5 | 8.9 | 2.8 | 2.7 | 9.1 | |
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6.5 | 6.6 | 7.2 | 7.7 | 7.4 | 6.5 | 8.0 | 7.5 | 9.2 | 9.2 | 6.8 | 7.0 | 9.7 | 6.8 |
Mean genetic distance for
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7.15 | |||||||||||||||
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6.02 | 5.10 | ||||||||||||||
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6.01 | 5.23 | 0.44 | |||||||||||||
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6.22 | 5.16 | 1.12 | 1.42 | ||||||||||||
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5.82 | 5.07 | 3.83 | 3.60 | 3.82 | |||||||||||
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6.03 | 4.54 | 2.61 | 3.10 | 2.88 | 4.01 | ||||||||||
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5.65 | 4.31 | 2.56 | 3.05 | 2.49 | 3.65 | 1.24 | |||||||||
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0.57 | 6.72 | 5.99 | 6.52 | 6.17 | 6.29 | 5.93 | 5.52 | ||||||||
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6.56 | 4.87 | 3.95 | 3.81 | 3.98 | 4.33 | 3.64 | 3.45 | 6.75 | |||||||
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4.39 | 5.32 | 4.50 | 4.77 | 4.49 | 5.02 | 4.44 | 4.01 | 4.21 | 4.47 | ||||||
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0.94 | 6.74 | 6.03 | 6.55 | 6.20 | 6.18 | 5.83 | 5.27 | 0.99 | 6.35 | 4.18 | |||||
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6.37 | 4.68 | 3.28 | 3.41 | 3.02 | 3.97 | 1.91 | 0.74 | 6.27 | 3.78 | 4.46 | 5.97 | ||||
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5.25 | 4.15 | 2.58 | 2.71 | 2.92 | 2.91 | 2.42 | 2.76 | 5.34 | 2.73 | 3.64 | 5.25 | 3.08 | |||
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5.82 | 5.07 | 3.83 | 3.60 | 3.82 | 0.00 | 4.01 | 3.65 | 6.29 | 4.33 | 5.02 | 6.18 | 3.97 | 2.91 | ||
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0.42 | 7.31 | 6.22 | 6.34 | 6.39 | 5.99 | 6.20 | 5.83 | 0.73 | 6.54 | 4.58 | 1.18 | 6.54 | 5.43 | 5.99 | |
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15.32 | 14.89 | 15.13 | 14.97 | 14.23 | 15.85 | 16.32 | 15.39 | 15.53 | 15.57 | 15.45 | 16.06 | 15.34 | 15.61 | 15.85 | 15.87 |
The two different phylogenetic analyses produced similar topologies. Both analyses produced a tree with 3 major clades (Figs
The Iranian members of the
Based on morphological and molecular results,
The populations of
We are pleased to thank A. Gholamhosseini, A. Teimori, M. Masoudi, H. Mehraban, G. Sayyadzadeh, R. Sadeghi, A. Khajepanah & M. Razbani for their help during field work. The research work was funded by the Shiraz University and was approved by the Ethics Committee of Biology Department (ECBD-SU-909789).