Research Article |
Corresponding author: Cüneyt Kaya ( cnytkaya@yahoo.com ) Academic editor: Nicolas Hubert
© 2022 Davut Turan, Cüneyt Kaya, İsmail Aksu, Yusuf Bektaş.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Turan D, Kaya C, Aksu İ, Bektaş Y (2022) Paracapoeta, a new genus of the Cyprinidae from Mesopotamia, Cilicia and Levant (Teleostei, Cypriniformes). Zoosystematics and Evolution 98(2): 201-212. https://doi.org/10.3897/zse.98.81463
|
The molecular and morphological studies carried out within the scope of this study revealed that the scrapers, known as the Mesopotamian group, belong to a different genus. The Paracapoeta gen. nov., from the Mesopotomia and Levant, is distinguished from Capoeta and Luciobarbus species by the presence of a strong ligament between the base of the last simple and the first branched rays of the dorsal-fin (vs. no or a very weak ligament). The Paracapoeta further differs from Capoeta by the last simple dorsal-fin ray strongly ossified in adult specimens (more than 75%, vs. less than 75%). The Paracapoeta further differs from Luciobarbus by the lower lip with horny layer (vs. fleshy lips). The molecular phylogeny based on the combined dataset (COI + Cytb, 1312 bp.) showed that the genus Paracapoeta was recovered from the other groups in the subfamily Barbinae with high bootstrap and posterior probability values (BP: 94%, PP: 0.96). Also, Paracapoeta and Capoeta are well differentiated by an average genetic distance of 8.02±0.78%. The morphological and molecular findings have largely overlapped each other. Besides, Capoeta turani is treated as a synonym of Capoeta erhani.
Capoeta, Mesopotamia, molecular phylogeny, scrapers, taxanomy
Cyprinid genus Capoeta Valenciennes, 1842 has a wide distribution in the Mediterranean, Middle East, Caucasus and South-West Asia. Even though the members of the genus occur in lakes and spring waters, they generally prefer fast-flowing streams (
Molecular phylogenies based on nuclear and mitochondrial molecular markers unambiguously demonstrated that Capoeta is clustered together with the Western Palaearctic barbels of the genera Barbus Daudin, 1805 and Luciobarbus Heckel, 1843 (
The mtDNA sequences of the protein-coding, cytochrome oxidase subunit I (COI) and cytochrome b (Cytb) genes, are powerful markers for deducing evolutionary relationships at the species, genera, family, and higher levels (
In recent years, various ideas have been put forward among researchers about the validity of Mesopotamian group species.
Here (i) we discussed the validity of the Capoeta turani and (ii) proposed a new generic name, Paracapoeta, for the scrapers, formerly known as the Mesopotamian Capoeta group based on morphological and molecular analysis.
See the list of materials examined in
The care of experimental animals was consistent with the Republic of Turkey’s animal welfare laws, guidelines and policies approved by Recep Tayyip Erdogan University Local Ethics Committee for Animal Experiments (permit reference number 2014/77).
Samples were collected by electro-shocker. After anaesthesia, fish were fixed in 4% formaldehyde. Methods for counting followed
For osteological preparation (last simple dorsal-fin ray), one specimen of each selected species of Paracaopeta and Capoeta (Paracaopeta trutta [200 mm SL], P. erhani [190 mm SL] P. barroisi [190 mm SL], Capoeta damascina (Valenciennes, 1842) [205 mm SL] C. tinca (Heckel, 1843) [190 mm SL] and C. pestai (Pietschmann, 1933) [195 mm SL]) were cleared and stained with alizarin red S, according to the protocol of
The COI (577 bp.) and Cytb (735 bp.) fragments of the seventy-one samples for Capoeta, (Cytb: Hashemzadeh Segherloo et al. unpublished;
For genera Capoeta, Luciobarbus, Barbus, Cyprinion, Scaphiodonichthys Vinciguerra, 1890, Aulopyge Heckel, 1841 and Paracapoeta gen. nov., the average intra- and intergeneric distances were computed by the General Reversible Time (GTR) model with gamma distributed invariant sites (G+I) in MEGA X (
FFR Recep Tayyip Erdogan University Zoology Museum of the Faculty of Fisheries, Rize;
SL standard length; BI, Bayesian inference;
ML maximum likelihood;
mtDNA mitochondrial deoxyribonucleic acid;
Cytb cytochrome b;
COI cytochrome c oxidase subunit 1;
AIC Akaike Information Criteria;
BIC Bayesian Information Criteria;
bp base pai;
BP Bootstrap Percentage;
PP Bayesian Posterior Probability.
In the original description, C. erhani (Ceyhan River) is distinguished from C. turani (Seyhan River) by having numerous spots (vs. few) and a brown back, lateral head and body (vs. silvery). Besides, the spots in C. erhani are large and often fused into blotches giving the fish a mottled appearance in individuals smaller than 120 mm SL (vs. spots are always small and never fused into blotches in C. turani. Also, caudal peduncle and operculum of C. erhani are always densely spotted (vs. few isolated spots or no spots).
Indeed, the silvery body color and the shape, form and number of the spots used in the original description were useful to distinguish both species. However, the fact is that Çakıt (type locality) is a constantly turbid stream (confirmed by Jörg Freyhof, pers. comm., 2019) which has possibly caused these differences in body color and pattern. These changes have also been observed in other species (Capoeta damascina, Oxynoemacheilus sarus, Garra turcica, Squalius adanaensis, Chondrostoma ceyhanensis, Salariopsis sp.) co-occurring with Capoeta turani.
Besides, in the original description, C. turani is distinguished from C. erhani by having more lateral line scales (64–70, vs. 69–78). However, our examination of lateral line scales of both species completely overlaps (63–70 in stream Aksu, Ceyhan; 63–76 in stream Çakıt, Seyhan; 71–79 in stream Üçürge, Seyhan).
On the other hand, recent molecular studies have stated that there is no difference (0.35%) between these two species at species level (
1 | Lower lip with horny layer; no fleshy lips | 2 |
– | Lower lip without horny layer on; fleshy lips | Luciobarbus |
2 | The presence of a strong ligament between the base of the last simple and the first branched rays of the dorsal-fin | Paracapoeta gen. nov. |
– | No, or very weak ligament between the base of the last simple and the first branched rays of the dorsal-fin | Capoeta |
The body fusiform and slightly compressed laterally. In adult individuals, the general body color is brownish, and without dark brown or blackish spots (except C. pestai). The head plain brownish, and no black spots on head and cheek. The mouth inferior, mouth transversely slit or horseshoe-shaped. Lips not developed and lower lip with keratinize edge. One or two pairs of barbel around the mouth. The last simple dorsal-fin slightly or moderately ossified (less than %75) and its posterior edge serrated (except C. antalyensis). No or very weak ligament between the base of the last simple and the first branched rays of the dorsal-fin. There are melanophore rows on the posterior edge of the flank scales. There is no keel in predorsal area, in front of dorsal-fin.
Cyprinus capoeta Güldenstädt, 1773 [actual status of the type species is Capoeta capoeta (Güldenstädt, 1773)].
Capoeta aculeata, C. antalyensis, C. aydinensis, C. banarescui, C. bergamae, C. buhsei, C. caelestis, C. capoeta, C. coadi, C. damascina, C. ekmekciae, C. ferdowsii, C. fusca, C. gracilis, C. heratensis, C. kaput, C. macrolepis, C. oguzelii, C. pestai, C. pyragyi, C. razii, C. saadii, C. sevangi, C. shajariani, C. sieboldii, C. tinca, C. umbla.
Afghanistan, Armenia, Azerbaijan, Georgia, Iran, Iraq, Jordan, Syria, Pakistan, Kazakhstan, Palestine, Tajikistan, Turkey, Turkmenistan and Uzbekistan: The genus Capoeta has a wide distribution in the Mediterranean, Middle East, Caucasus and South-West Asia.
Scaphiodon trutta Heckel 1843 [actual status of the type species is Paracapoeta trutta (Heckel, 1843)].
The new genus Paracapoeta is distinguished from other genus of Capoeta and Luciobarbus by having a strong ligament between the base of the last simple and the first branched rays of the dorsal-fin (Fig.
Nucleotide positions for some genera within the subfamily Barbinae. Diagnostic and distinctive nucleotide positions are represented in bold font and gray background, respectively.
Genera | Diagnostic nucleotide positions | |||||||||||||||||||||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | ||||||||||||||||||||||||||
1 | 1 | 2 | 2 | 3 | 3 | 3 | 4 | 4 | 4 | 6 | 6 | 7 | 7 | 8 | 8 | 8 | 8 | 8 | 9 | 9 | 9 | 9 | 0 | 0 | 0 | 1 | 2 | 2 | 2 | 2 | 2 | |||
6 | 8 | 0 | 8 | 0 | 9 | 4 | 7 | 8 | 1 | 7 | 9 | 1 | 4 | 1 | 8 | 4 | 5 | 5 | 8 | 9 | 4 | 9 | 7 | 9 | 3 | 4 | 4 | 5 | 0 | 1 | 4 | 8 | 9 | |
0 | 0 | 6 | 4 | 5 | 5 | 6 | 0 | 5 | 8 | 8 | 3 | 4 | 9 | 5 | 4 | 1 | 3 | 9 | 3 | 2 | 1 | 4 | 0 | 7 | 6 | 3 | 6 | 3 | 4 | 4 | 0 | 5 | 7 | |
Paracapoeta | C | A | T | C | C | T | G | A | T | T | T | T | T | T | T | C | C | G | G | C | C | T | T | G | C | T | T | T | C | G | T | C | T | T |
Capoeta | T | G | C | T | A/G | C | A | G | C | C | C | C | C | C | C | T | T | T | T | T | T | C | G/A | A | T | C | C | C | T | A | C | A/G | C | C |
Luciobarbus | T | G | A/C/T | T | A/G | C | A | G | T/C | T/C | C | C | T | C | C | C | C | A | T | C | C | C | G/A | G/A | T | C/T | C | C/T | C | A | C/A | A/G | C/T | C/T |
Barbus | T | G | C/T | T | A/G | C | G | G | T/C | T/C | T | T | T | C | C | C | C | A/G | T | C | C | C | G/A | A | T | T | C | C | C | A | C | A/G | C | C/T |
Presence (a, b) and absence (c, d) of strong ligament between the base of the last simple and the first branched rays of the dorsal-fin; a. Paracapoeta trutta, 227 mm SL, Euphrates River; b. P. erhani, 265 mm SL, Ceyhan River; Capoeta damascina, 250 mm SL, Euphrates River; Luciobarbus pectoralis, 227 mm SL, Orontes River.
Melanophores on the free part of the flank scales: upper row from left, P. trutta 146 mm SL; P. erhani, 201 mm SL; P. barroisi, 155 mm SL; middle row from left, C. capoeta; 198 mm SL; C. banarescui; 181 mm SL; C. damascina, 181 mm SL; lower row from left, Luciobarbus lydianus, 192 mm SL; L. barbulus, 155 mm SL; L. capito, 220 mm SL.
Lateral view of Paracapoeta erhani; a. Not preserved, about 155 mm SL, stream Aksu at Kuyumcular, Ceyhan drainage; b. FFR 1952, 201 mm SL, stream Aksu at Pazarcık, Ceyhan drainage; c. Not preserved, about 200 mm SL, stream Üçürge at Karaisali, Seyhan drainage; d. FFR 1955, 130 mm SL, stream Çakıt at Salbaş (type locality of C. turani), Seyhan drainage.
Paracapoeta anamisensis, P. barroisi, P. erhani, P. mandica, P. trutta.
Turkey, Iran, Iraq and Syria: Seyhan, Ceyhan and Orontes rivers, Levant drainages; Tigris, Euphrates, Mond and Minab River, Persian Gulf drainages.
The name of the new genus is formed by combining the words “Para” and “Capoeta”. “Para” means “beside” or “near”, and “Capoeta” is the available name of the closest genus of Paracapoeta, deriving from the local vernacular name “kapwaeti” used in Georgia and Azerbaijan.
Phylogenetic analyses using BI and ML methods provided similar topologies for the western Palearctic Barbinae genera, with high posterior probability (PP = 0.80–1.00) and high bootstrap (BP = 94–98%) values (Fig.
Phylogenetic tree generated based on the mitochondrial combined dataset. ML and BI methods recovered similar topologies, and therefore only the ML tree is presented here. The bootstrap percentage values (BP) ≥ 50% from ML analysis and Bayesian posterior probabilities (PP) ≥ 0.90 are shown on the nodes (BP/PP)
The K2P-based distance analyses found a maximum intrageneric distance was 7.1%, in Luciobarbus (Lineage III) while the minimum intrageneric distance was 1.1% in Paracapoeta gen. nov. (Lineage II) (Table
In an effort to re-evaluate the generic structure of the scrapers, Paracapoeta gen. nov., formerly known as the Mesopotamian Capoeta group, was assessed based on morphological and molecular data.
In agreement with previous mitochondrial and nuclear markers-based phylogenies (
For a combined dataset of Cytb (735 bp) and COI (577 bp), the intergeneric genetic distances for Paracapoeta and Capoeta lineages (mean 8.02±0.78%) represent the lowest limit of the predicted intergeneric genetic distances estimated for especially the western Palearctic Barbinae genera, while it corresponds to the lower limits of intergeneric distance (8–10%;
Consistent with previous studies (
The authors would like to thank Ülgen Aytan, Başak Esensoy and Sedanur Uğur (Rize) for taking the scale photos. We are also thankful to Dr. Münevver Oral (Rize) for discussing and editing this manuscript.
Table S1
Data type: excel file
Explanation note: List of Genbank accession numbers of sequences used in molecular analyzes in this study.