Corresponding author: Dušan Jelić (
Academic editor: M. Bichuette
Distribution data on many freshwater fish species in Croatia are scarce and species identifications are difficult, requiring further detailed studies. This paper presents a report of the Italian gudgeon
Jelić D, Jelić M, Žutinić P, Šimunović I, Zupančič P, Naseka AM (2018) Distribution of endangered Italian gudgeon
The richness of Croatian freshwater ichthyofauna manifests in at least 147 native fish and lamprey species, many of which are endemic (
Another example is the Italian gudgeon
Based on some diagnostic morphological characters, the Italian gudgeon is considered more similar to
For nearly two centuries after its description,
In recent years,
Although four species of the subfamily
Map of distribution of
Measurements were made according to
Morphometric characters in
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69.5–72.4 | 37.6–93.6 | 70.7 | 90.8–98.3 | 94.7 | 2.91 | 62.3–83.5 | 76.1 | 49.0–94.7 | 67.7 | ||||
% |
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Body depth at dorsal-fin origin | 21.6–23.5 | 17.9–23.8 | 21.0 | 2.01 | 21.2–24.7 | 23.3 | 1.04 | 24.4–27.7 | 26.0 | 1.53 | 21.7–28.3 | 24.7 | 1.65 |
Caudal peduncle depth | 8.7–9.6 | 8.2–10.9 | 9.7 | 0.75 | 9.7–11.0 | 10.2 | 0.48 | 10.4–11.2 | 10.8 | 0.33 | 9.0–11.5 | 10.5 | 0.61 |
Body width at dorsal-fin origin | 13.9–16.5 | 11.4–16.5 | 13.9 | 1.45 | 14.7–17.7 | 16.1 | 1.09 | 12.8–17.9 | 15.3 | 2.38 | 11.9–16.7 | 14.1 | 1.71 |
Width of caudal peduncle | 3.0–4.0 | 3.0–5.9 | 4.0 | 0.75 | 4.2–5.6 | 4.7 | 0.52 | 3.5–5.0 | 4.3 | 0.64 | 3.3–6.5 | 4.5 | 0.79 |
Predorsal length | 50.0–50.2 | 46.3–50.2 | 48.8 | 1.19 | 47.1–50.3 | 48.6 | 1.12 | 46.8–51.4 | 48.4 | 2.03 | 47.3–53.0 | 49.4 | 1.46 |
Postdorsal length | 40.3–42.4 | 40.1–42.7 | 41.3 | 0.87 | 41.5–43.4 | 42.7 | 0.82 | 41.0–41.5 | 41.3 | 0.24 | 38.5–43.7 | 41.3 | 1.37 |
Prepelvic length | 50.1–50.4 | 47.3–52.4 | 50.2 | 1.37 | 48.2–51.3 | 49.6 | 0.99 | 49.9–51.4 | 50.7 | 0.72 | 48.3–53.1 | 50.5 | 1.22 |
Preanal length | 70.8–71.7 | 68.4–74.1 | 71.7 | 1.46 | 70.1–73.5 | 71.6 | 1.17 | 67.1–70.6 | 69.3 | 1.62 | 68.9–74.0 | 70.7 | 1.45 |
Distance between pectoral fin and pelvic-fin origin | 24.3–24.8 | 21.8–27.2 | 24.9 | 1.61 | 24.7–26.4 | 25.2 | 0.57 | 24.8–30.4 | 27.5 | 2.37 | 21.3–28.3 | 25.0 | 1.40 |
Distance between pelvic fin and anal-fin origin | 20.7–22.3 | 20.7–23.3 | 21.8 | 0.87 | 21.3–23.9 | 22.6 | 0.87 | 18.7–22.2 | 20.4 | 1.80 | 18.7–23.1 | 21.1 | 1.22 |
Distance between anus and anal-fin origin | 6.3–7.1 | 5.1–10.1 | 6.8 | 1.21 | 5.8–8.0 | 6.6 | 0.72 | 2.5–4.5 | 3.4 | 0.88 | 3.1–6.2 | 4.4 | 0.90 |
Caudal peduncle length | 20.0–22.4 | 17.8–22.6 | 20.6 | 1.36 | 18.2–22.4 | 20.9 | 1.37 | 19.7–23.8 | 21.3 | 1.75 | 18.3–22.5 | 20.5 | 1.13 |
Dorsal-fin length | 13.1–14.3 | 11.9–14.3 | 13.3 | 0.81 | 12.1–13.6 | 13.0 | 0.53 | 13.8–16.3 | 15.0 | 1.26 | 12.1–15.8 | 14.3 | 0.93 |
Dorsal-fin depth | 24.2–24.9 | 21.0–26.2 | 23.4 | 1.72 | 20.9–22.0 | 21.5 | 0.48 | 21.7–25.6 | 23.7 | 1.78 | 22.2–26.3 | 23.9 | 1.09 |
Anal-fin length | 8.2–8.7 | 7.7–9.5 | 8.6 | 0.51 | 6.9–8.7 | 8.0 | 0.66 | 10.0–13.1 | 10.8 | 1.51 | 8.6–11.6 | 10.0 | 0.8 |
Anal-fin depth | 19.0–19.4 | 15.8–20.8 | 18.4 | 1.50 | 15.8–17.2 | 16.5 | 0.45 | 17.9–20.5 | 18.9 | 1.10 | 17.3–21.6 | 19.1 | 1.2 |
Pectoral-fin length | 20.1–22.1 | 18.7–23.2 | 21.3 | 1.32 | 18.6–21.5 | 19.7 | 1.07 | 21.0–22.4 | 21.8 | 0.65 | 19.7–24.5 | 21.8 | 1.41 |
Pelvic-fin length | 17.1–18.3 | 15.8–18.3 | 17.2 | 0.92 | 15.9–16.4 | 16.1 | 0.20 | 16.0–18.5 | 17.5 | 1.07 | 15.8–20.3 | 17.8 | 1.04 |
Head length | 27.7–27.8 | 24.4–29.9 | 27.4 | 1.82 | 25.6–28.1 | 26.7 | 0.91 | 23.9–26.1 | 25.2 | 0.99 | 25.0–29.1 | 26.9 | 1.11 |
Eye diameter | 6.4–6.8 | 4.9–6.8 | 6.1 | 0.62 | 5.3–5.9 | 5.5 | 0.23 | 5.6–6.6 | 6.0 | 0.42 | 5.8–8.0 | 6.6 | 0.54 |
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Head depth at nape | 55.9–56.7 | 53.3–62.5 | 57.5 | 2.61 | 58.5–64.1 | 61.6 | 2.12 | 60.0–62.1 | 61.0 | 0.88 | 55.8–67.1 | 61.7 | 2.52 |
Snout length | 40.4–41.1 | 36.7–41.6 | 38.8 | 1.67 | 40.0–43.8 | 42.1 | 1.16 | 43.2–44.1 | 43.5 | 0.39 | 37.0–45.0 | 40.4 | 2.34 |
Eye diameter | 23.1–24.4 | 19.1–26.2 | 22.8 | 1.84 | 19.2–21.6 | 20.7 | 0.90 | 22.8–26.2 | 23.9 | 1.51 | 21.0–27.9 | 24.6 | 1.6 |
Postorbital distance | 42.1–42.6 | 42.1–46.8 | 44.7 | 1.64 | 44.3–48.7 | 46.3 | 1.58 | 42.3–45.4 | 43.8 | 1.67 | 40.7–47.5 | 44.4 | 1.78 |
Maximum head width | 50.4–54.8 | 48.8–55.8 | 51.7 | 2.31 | 56.7–63.2 | 59.8 | 2.60 | 56.7–58.6 | 57.8 | 0.78 | 49.8–64.2 | 55.1 | 3.87 |
Interorbital width | 27.9–30.7 | 26.1–34.5 | 29.5 | 2.10 | 32.1–36.3 | 34.1 | 1.25 | 31.6–35.9 | 33.5 | 1.88 | 27.5–34.9 | 30.9 | 1.73 |
Length of upper jaw | 24.7–25.9 | 20.3–26.5 | 24.5 | 1.75 | 25.2–28.4 | 26.8 | 1.17 | 25.4–28.1 | 26.7 | 1.36 | 21.9–31.5 | 25.2 | 2.41 |
Length of lower jaw | 36.0–36.1 | 32.2–36.1 | 34.9 | 1.22 | 33.1–36.1 | 34.4 | 1.10 | 33.1–39.9 | 35.3 | 3.13 | 28.8–38.5 | 34.5 | 2.51 |
Barbel length | 26.3–27.6 | 24.3–34.3 | 27.5 | 2.63 | 21.8–28.4 | 25.4 | 2.21 | 30.3–39.7 | 35.1 | 4.48 | 24.6–42.7 | 33.4 | 4.59 |
Caudal peduncle depth | 31.4–34.7 | 31.4–41.2 | 36.0 | 2.58 | 34.4–40.7 | 38.1 | 2.24 | 41.5–44.5 | 42.8 | 1.48 | 33.1–42.8 | 39.0 | 2.47 |
Meristic characters in
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Unbranched dorsal-fin rays | 3 | 3 | 3 | 3 | 3 | 4 | 4 | 3–4 | 3.7 | 0.48 | |||
Branched dorsal-fin rays | 7½ | 7½ | 7½ | 7½ | 7½ | 7½ | 7½ | 7½ | 7½ | ||||
Branched anal-fin rays | 6½ | 6½ | 6½ | 6½ | 6½ | 6½ | 6½ | 6½ | 6½ | ||||
Branched pectoral-fin rays | 15–16 | 15–16 | 15.6 | 0.51 | 15–18 | 15.6 | 0.84 | 13–15 | 13.8 | 0.96 | 12–15 | 13.3 | 0.67 |
Branched pelvic-fin rays | 7 | 7 | 7.0 | 7 | 7.0 | 7 | 7.0 | 7 | 7.0 | ||||
Scales in lateral row | 42 | 41–43 | 41.4 | 0.79 | 40–42 | 41.3 | 0.73 | 38–39 | 38.5 | 0.58 | 37–40 | 38.8 | 0.73 |
Total lateral-line scales | 42 | 41–42 | 41.3 | 0.49 | 39–42 | 41.0 | 1.0 | 37–39 | 38.3 | 0.96 | 37–40 | 38.5 | 0.87 |
Lateral-line scales to posterior margin of hypurals | 39–42 | 38–40 | 39 | 0.58 | 38–40 | 38.9 | 0.81 | 35–36 | 35.8 | 0.50 | 34–37 | 36.4 | 0.80 |
Scales above lateral line | 6 | 6 | 6.0 | 6 | 6.0 | 6 | 6.0 | 6 | 6.0 | ||||
Scales below lateral line | 4 | 4–5 | 4.1 | 0.38 | 4 | 4.0 | 3–4 | 3.8 | 0.50 | 3–4 | 4.1 | 0.28 | |
Scales between anus and anal-fin origin | 6–9 | 4–7 | 5.4 | 1.27 | 4–7 | 6.1 | 0.71 | 2–3 | 2.3 | 0.50 | 1–5 | 3.7 | 0.85 |
Circumpeduncular scales | 16 | 15–16 | 15.7 | 0.49 | 13–16 | 15.3 | 0.84 | 13–16 | 14.3 | 1.26 | 12–15 | 13.6 | 0.79 |
Predorsal scales | 18–19 | 15–21 | 17.9 | 2.27 | 14–20 | 17.3 | 1.64 | 13–17 | 15.5 | 1.91 | 14–18 | 15.8 | 1.15 |
Total vertebrae | 39–40 | 38–40 | 39.3 | 0.5 | 38–41 | 39.1 | 0.79 | 36–38 | 37.0 | 0.82 | 36–38 | 37.2 | 0.4 |
Abdominal vertebrae | 20 | 20–21 | 20.6 | 0.53 | 20–22 | 20.5 | 0.54 | 19–20 | 19.5 | 0.59 | 18–20 | 19.1 | 0.46 |
Caudal vertebrae | 19–20 | 18–19 | 18.7 | 0.49 | 17–20 | 18.5 | 0.68 | 16–18 | 17.5 | 1.0 | 17–18 | 17.9 | 0.32 |
Predorsal abdominal vertebrae | 11 | 11 | 11 | 0.0 | 10–11 | 10.8 | 0.43 | 10–11 | 10.5 | 0.58 | 9–11 | 10.4 | 0.60 |
Preanal caudal vertebrae | 2–3 | 1–2 | 1.3 | 0.49 | 1–3 | 1.6 | 0.58 | 0–2 | 1.3 | 0.96 | 1–2 | 1.4 | 0.50 |
(Abbreviations:
Besides species identification based on morphological features, species were characterised using sequences of cytochrome b gene (
The obtained sequences in this study (1141 base pairs long, bp) were examined using Nucleotide Basic Local Alignment Search Tool (Nucleotide BLAST;
The Median-Joining (
Phylogenetic tree reconstructions were conducted using the
Three user trees (“Tree 1:
Comparative morphological analysis of
Lateral view of
The body is relatively deep, the depth at the dorsal-fin origin is 24–28, averaging 26%
The anus is located close to the anal-fin origin, the distance between the anus and the anal-fin origin is 3–5, averaging 3%
The dorsal fin has 4 unbranched rays in all specimens from Mirna and Soča rivers and in 14 (of 19) specimens from the Po drainage (Table
Radiographs of unbranched dorsal-fin rays. Specimens as
In
Number of specimens of
Species and locality | In front of pectoral-fin base | Anterior end of pectoral-fin base | Middle of pectoral-fin base | Posterior end of pectoral-fin base | Behind pectoral-fin base |
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1 | 1 | ||||
1 | 3 | 3 | |||
1 | 13 | 3 | |||
4 | |||||
1 | 4 | 10 | 4 |
The lateral line is complete, with 37–39 total scales averaging 38.6 (36–40 averaging 38.8 in the Po samples). These counts are lower than in
The barbel is reaching the vertical through the middle of the pupil to the posterior margin of the eye (Table
Character states of the position of the posteriormost extremity of the barbel in
Species and locality | Barbel reaching to vertical of: | |||||
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anterior margin of pupil | middle of pupil | posterior margin of pupil | between pupil and posterior margin of eye | posterior margin of eye | behind posterior margin of eye | |
2 | ||||||
4 | 1 | 2 | ||||
3 | 6 | 2 | 6 | |||
1 | 2 | 1 | ||||
4 | 4 | 8 | 1 | 1 |
Total vertebrae are 36–38, 19–20 abdominal, and (16)18 caudal including 0–2 preanal. 10–11 predorsal vertebrae. The vertebral counts in the Mirna samples of
Laterally (examined on both sides) with 7–8, usually 7 roundish dark blotches. The size of a blotch varies but it is relatively large: the size of the blotch below the dorsal-fin origin is about (close to) or larger than the horizontal eye diameter. The same pattern of the blotches, 5–8, usually 6 or 7, is found in the examined specimens from the Po and Soča drainages. In
List of species used for phylogenetic tree inference on
Species | Catalogue no. | Locality | Accession No. | Sequence length (bp) | Reference |
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Croatia: Mirna River, Kamenita Vrata | xxx | 1141 | This study | |
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Croatia: Mirna River, Kamenita Vrata | xxx | 1141 | This study | |
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Croatia: Mirna River, Kamenita Vrata | xxx | 1141 | This study | |
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Croatia: Mirna River, Kamenita Vrata | xxx | 1141 | This study | |
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Bosnia and Herzegovina: Boračko Lake | xxx | 1141 | This study | |
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MEL | Italy: Meletta River |
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342 |
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TAG | Italy: Tagliamento River |
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342 |
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ASS | Italy: Assino River |
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342 |
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OMB | Italy: Ombrone River |
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342 |
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BAD | Italy: Badolato River |
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342 |
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France: Rhone River |
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1141 |
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Spain: Tajo River |
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1141 |
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Greece: Gallikos River |
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1141 |
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Greece: Aliakmon River |
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1141 |
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Gobio_ciscaucasicus | Russia: Kuma River |
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1141 |
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G.ura.34 | Romania: Valsan River/Valsanesti |
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1141 |
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G.go.13FR.ADOUR | France: Adour River |
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1141 |
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G.go.33Czech.R | Czech Republic |
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1141 |
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Czech Republic: Plana |
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1141 |
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Gobio_gobio (2) | Germany: Rhine River |
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1141 |
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Gobio_gobio (1) | Romania |
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1141 | Luca et al. (direct submission) |
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MNCN_AT4759 | Slovenia: Sevnica River |
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1141 |
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WL*0653a | Ukraine: middle Dniestr River close to type locality |
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1141 | Witte (direct submission) |
WL*0626a | Romania: middle Nera River (Danube drainage) |
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1141 | Witte (direct submission) | |
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WL*0626b | Romania: middle Nera River (Danube drainage) |
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1141 | Witte (direct submission) |
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WL*0624a | Romania: middle Nera River (Danube drainage) |
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1141 | Witte (direct submission) |
WL*0628a | Turkey: Aras River |
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1141 | Witte (direct submission) | |
Gobio_macrocephalus | China: Yanji, Tumenjiang River |
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1141 |
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Gobio_cynocephalus | China: Fuyuan, Amur River |
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1141 |
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Romanogobio_tenuicorpus | China: Yellow River |
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1141 |
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Gobio_huanghensis | China |
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1141 | Qi et al. (direct submission) | |
IHCAS:0210055 | China: Kaiyuan, Liahe River |
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1141 |
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CTOL00130 | n/a |
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1141 |
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CTOL00128 | n/a |
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1141 |
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CTOL00129 | n/a |
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1141 |
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CTOL00112 | n/a |
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1141 |
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CTOL00375 | n/a |
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1141 |
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Bosnia and Herzegovina: Trebišnjica River, Ravno |
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1141 |
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Czech Republic: Libechovka River, Elbe River |
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1141 |
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Turkey: Kizilirmak River, Kirsehir |
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1141 |
|
A Mann-Whitney U test revealed seven morphometric and nine meristic characters different on a statistically significant (0.01%) level between the samples of typical
These 16 distinguishing characters were used for a DFA in order to classify the Reka and Mirna samples into one of the two species. DFA statistics values are as follows: Wilks’ Lambda 0.00721, approx. F (45, 78) =7.3894, p<0.0000. The Mirna specimens are the closest to Italian
Two unique
All topology tests (Table
Comparison of user trees (“Tree 1:
Tree | log L | difference | S.E. | p- |
p- |
c- |
|
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1 | -8489.12 | 0.00 | <---- best | 1.0000 + | 1.0000 + | 0.7078 + | best |
2 | -8491.32 | 2.20 | 2.8840 | 0.2040 + | 0.5230 + | 0.2036 + | + |
3 | -8499.39 | 10.27 | 8.1588 | 0.1010 + | 0.1250 + | 0.0887 + | + |
Morphological data in this study confirm observations of the previous authors (
Some of the morphological characters of
The
Result of DFA performed on 15 (8 meristic and 7 morphometric) distinguishing characters to classify Reka and Mirna samples.
Lateral view of the head of
As shown above in the results of the morphological analysis, the examined sample from the Reka River is
The morphological data discussed above confirmed that gudgeons from the Mirna River can be assigned to
Phylogenetic reconstruction in this study (Fig.
Phylogenetic tree inferred by
The Mirna population represents a single area of occurrence of
Only four specimens of
Therefore, having in mind that small-sized populations are more prone to extirpation due to genetic drift, extinction vortex, etc., it is necessary to implement systematical monitoring on the present
This research has been undertaken through research programmes of Croatian Biology Research Society (