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A Review of Vertebral and Fin-Ray Counts in the Genus Alburnoides (Teleostei: Cyprinidae) with a Description of Six New Species

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A Review of Vertebral and Fin-Ray Counts in the Genus Alburnoides (Teleostei: Cyprinidae) with a Description of Six New Species ZOOSYSTEMATICA ROSSICA, 18(1): 126-173 3 JULY 2009 A review of vertebral and fin-ray counts in the genus Alburnoides (Teleostei: Cyprinidae) with a description of six new species N.G. BOGUTSKAYA & B.W. COAD N.G. Bogutskaya, Zoological Institute, Russian Academy of Sciences, Universitetskaya nab. 1, St. Petersburg 199034, Russia. E-mail: [email protected] B.W. Coad, Canadian Museum of Nature, Ottawa, Ontario K1P 6P4, Canada. E-mail: [email protected] Variations in the different vertebral patterns and unpaired fin counts in the genus Alburnoides are discussed based on an examination of over 1100 specimens from a wide area of distribution of the genus. Though it was commonly considered that the morphological differences between subspecies and local forms of A. bipunctatus auctorum appear slight, our study shows that there are pronounced differences between many of them in vertebral and dorsal and anal fin counts. To estimate reliability of differences, statistical tests were used and a cluster analysis (UPGMA – Average Linkage clustering method) was performed based on calculated values, and tree diagrams are presented and analyzed in their taxonomic context. The differences in fin ray and vertebral counts in combination with some other morphological characters, dis- tinguished some of the former subspecies or local forms as distinct species. Along with five nominal taxa re-established to a species level (A. rossicus, A. kubanicus, A. maculatus, A. eich- waldii, A. fasciatus), we describe six new species: A. gmelini (type locality Sunzha River, Terek River drainage, Eastern Ciscaucasia, Russia), A. varentsovi (type locality Ashkhabadka River, northern slope of the Kopetdag, Turkmenistan), A. petrubanarescui (type locality Qasemlou Chay, Orumiyeh Lake basin, Iran), A. namaki (type locality qanat at Taveh, Namak Lake basin, Iran), A. nicolausi (type locality Simareh River in Karkheh River system, Tigris River drain- age, Iran), and A. idignensis (type locality Bid Sorkh River, Gav Masiab River system, Tigris River drainage, Iran). Key words: freshwater fishes, Cyprinidae, Alburnini, Alburnoides, new species, morphology, axial skeleton, unpaired fins INTRODUCTION Anaecypris Collares-Pereira, 1983 (Gilles et al., 2001; Cunha et al., 2002; Durand et Over 30 species, subspecies and local al., 2002; Freyhof et al., 2005). The latter forms of the leuciscine cyprinid fishes were genus was also shown to be morphologi- considered by Bogutskaya (1990a, 1990b, cally close to the Alburnini (Bogutskaya 1997b). They represent a monophyletic & Collares-Pereira, 1997). taxon, the tribe Alburnini, characterized The uncertainty of the generic defini- by a set of derived characters, some of them tions of Alburnus, Chalcalburnus and Al- unique, in external morphology, sensory burnoides was discussed earlier (Bianco, canals and skeleton. Alburnus Rafinesque, 1980; Krupp, 1985) and an opinion was 1820, Chalcalburnus Berg, 1932, Alburn- proposed that they may be synonyms oides Jeitteles, 1861, Tropidophoxinellus (Coad, 1991). Later, Alburnus and Chal- Stephanidis, 1974 and Leucaspius Heckel calburnus were synonymized by Boguts- & Kner, 1858 were included in the tribe. kaya (1997a) since the distinguishing Later, a number of molecular studies re- characters of Chalcalburnus according vealed that the group does form a clade to Berg (1932b, 1949) – a partly scaled and includes, if representatives geneti- ventral keel, numerous long gill rakers cally examined are considered, at least and slightly serrated or unserrated pha- Alburnus, Chalcalburnus, Leucaspius and ryngeal teeth – have no diagnostic value © 2009 Zoological Institute, Russian Academy of Scienсes N.G. BOGUTSKAYA & B.W. COAD. A REVIEW OF ALBURNOIDES 127 being variably present in most Alburnus, considered a complex species with a number Alburnoides and Chalcalburnus species. On of subspecies found from France through the other hand, according to Berg (1949), Europe north of the Alps eastwards to the the only difference between Alburnus and Black, Caspian and Aral Sea basins (e.g. Alburnoides is unserrated pharyngeal Berg, 1949; Bogutskaya & Naseka, 2004; teeth. Synonymization of Alburnus and Coad, 2009). Alburnoides ohridanus (Kara- Chalcalburnus was supported by molecu- man, 1928) and A. prespensis (Karaman, lar studies (e.g. Durand et al., 2002) and 1924) were recently given a rank of species is widely accepted (e.g. Bogutskaya & (Kottelat & Freyhof 2007) as well as A. Naseka, 2004; Kottelat & Freyhof, 2007). eichwaldii (De Filippi, 1863) (Fricke et al., In contrast, Alburnoides stands apart from 2007), and a new species from the Pulvar R. Alburnus in molecular trees (Hänfling & system in southern Iran will be described in Brandl, 2000; Gilles et al., 2001; Cunha et a paper by Coad & Bogutskaya (2009). al., 2002; Freyhof et al., 2005). However, The following nominal taxa, which have morphological definitions of the two gen- appeared in the literature as subspecies or era appear to be quite obscure. They both distinct forms, have been historically synon- include species which are characterized ymized with A. bipunctatus: Alburnoides bi- by a more or less pronounced ventral keel punctatus armeniensis Dadikyan, 1972 (type (from scaled to completely scaleless), a locality: rivers Arpa, Vorotan, Vedi, Marma- slightly to considerably elongated anal fin, rik, Kasakh with tributaries of the Aras R. pharyngeal teeth commonly 2.5-5.2 or 2.5- system in the Kura R. drainage, Caspian 4.2, an absolutely and relatively elongated Sea basin, Armenia), Alburnus eichwaldii De predorsal vertebral subregion (to 15-17 Filippi, 1863 (Kura R. at Tiflis, Caspian Sea vertebrae which is commonly over 38% of basin, now Georgia), Aspius fasciatus Nor- total and 70% of abdominal vs. 33-37% and dmann, 1840 (rivers of the western coast of 60-66%, respectively, in most other groups the Black Sea eastward to Mingrelia, now of the Leuciscinae), a tendency to equal- Russia and Georgia), Alburnoides bipunc- ity of the numbers of vertebrae in abdomi- tatus rossicus natio kubanicus Berg, 1932 nal and caudal regions (modal difference (unavailable name; Kuban’ R., Sea of Azov between abdominal and caudal numbers basin, Russia), Alburnus maculatus Kessler, decreases from 4 or 3 to 1, 0, and [-1] vs. 1859 (Salgir R., Sea of Azov basin, Crimea modes 4 to 6 in the most Leuciscinae), and Peninsula, now Ukraine), Alburnus bipunc- a large orbit with respective configuration tatus ohridanus Karaman, 1928 (Ohrid L., of all the cranial elements it is formed from Adriatic Sea basin, now the Former Yugo- (Bogutskaya, 1990a, 1990b; Bogutskaya slav Republic of (FYRO) Macedonia), Al- et al., 2000). Actually, the only character burnus bipunctatus var. prespensis Karaman, easy to use for distinguishing Alburnus and 1924 (Prespa L. and its tributaries, no di- Alburnoides is the pattern of pigmenta- rect link to any basin, now FYRO Macedo- tion. In Alburnoides, small black spots are nia), Alburnoides bipunctatus rossicus Berg, located on each side of the lateral line ca- 1924 (Dnieper, Black Sea basin, and Volga, nal pore outlining the canal at least along Caspian Sea basin, now Ukraine and Rus- its anterior portion, and a dark stripe goes sia), Alburnoides bipunctatus var. smyrnae from behind the eye to the caudal fin base Pellegrin, 1927 (Mélèl stream near Smyrna, though both characters may be variably Aegean Sea basin, Turkey), Alburnoides developed in live fish or are absent. bipunctatus strymonicus Chichkoff, 1940 Besides Alburnoides oblongus Bulgakov, (Struma R. drainage, Bulgaria), Alburnoi- 1923 and Alburnoides taeniatus (Kessler, des bipunctatus tzanevi Chichkoff, 1933 1874) from the Aral Sea basin, Alburnoides (Rezova R., Black Sea basin, Bulgaria and bipunctatus (Bloch, 1782) has long been Turkey), and Alburnoides bipunctatus subsp. © 2009 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 18(1): 126-173 128 N.G. BOGUTSKAYA & B.W. COAD. A REVIEW OF ALBURNOIDES (Berg, 1932a, 1932b) (Kuma, Terek, Sulak with regard to the dispersion, we calculated rivers, Caspian Sea basin, Russia). Kul’bak’s Divergence that has a meaning The most comprehensive review of A. bi- of average information measure of the dif- punctatus s.l. is still that by Berg (1949) who ference between two empiric distributions successfully used the number of pharyngeal (Kulbak, 1967) and a cluster analysis (UP- teeth and the number of branched anal-fin GMA – Average Linkage clustering meth- rays for distinguishing six subspecies and od), were performed based on calculated infrasubspecies forms (one unnamed) with- values. Similarity Indices (r) (Zhivotovs- in the species. Thought it was later con- kiy, 1991) were calculated for each charac- sidered that the morphological differences ter separately, for five vertebral characters between subspecies and local forms of A. and for all seven characters under consid- bipunctatus auctorum appear slight (Kotte- eration, and tree diagrams (UPGMA – Av- lat & Freyhof, 2007), our study shows that erage Linkage clustering method) are pre- there are pronounced differences between sented and analyzed. Abbreviations used: n, many of them in vertebral and dorsal and number of specimens; min, minimum value; anal fin counts, and these differences be- max, maximum value; avg, average (mean); tween the different vertebral patterns are m, standard error of the mean; std, standard discussed below. In combination with some deviation; Me – median; Mo – mode. other morphological characters this gave Below we provide two lateral
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