Review of the genus Alburnoides Jeitteles, 1861 (Actinopterygii, Cyprinidae) from Iran with description of three new species from the Caspian Sea and Kavir basins

[Research]

Review of the genus Alburnoides Jeitteles, 1861 (Actinopterygii, Cyprinidae) from Iran with description of three new species from the Caspian Sea and Kavir basins

H. Mousavi-Sabet^{1, 2}*, S. Vatandoust³, I. Doadrio⁴

1. Department of Fisheries, Faculty of Natural Resources, University of Guilan, Sowmeh Sara, Guilan, Iran

2. The Caspian Sea Basin Research Center, University of Guilan, Rasht, Iran

3. Department of Fisheries, Babol Branch, Islamic Azad University, Babol, Iran

4. Biodiversity and Evolutionary Group. National Museum of Natural Sciences. CSIC. C/ José Gutiérrez Abascal, 2. 28006. Madrid. Spain

* Corresponding author’s E-mail: mousavi-sabet@guilan.ac.ir

ABSTRACT

The genus Alburnoides Jeitteles, 1861 in Iran is reviewed, and diagnoses are presented for all eight recognized species. Alburnoides eichwaldii, A. holciki, A. idignensis, A. namaki, A. nicolausi, A. petrubanarescui, A. qanati and A. tabarestanensis are considered valid and three new species are described: Alburnoides coadi sp. n., from Namrud River in Kavir basin, A. parhami sp. n., from Atrak River in the south-eastern Caspian Sea basin, and A. samiii sp. n., from Sefidroud River in the southern Caspian Sea basin. All species, except unstudied A. petrubanarescui are illustrated and characterized morphologically.

INTRODUCTION

The genus Alburnoides Jeitteles, 1861, is characterized by presenting small black spots near the pores located on each side of the lateral line outlining the canal at least along its anterior portion (Bogutskaya & Coad, 2009). Until recently, Alburnoides bipunctatus (Bloch, 1782) was the name applied to most populations of riffle minnows across Europe and the Middle East from France north of the Alps eastwards to the Black, Caspian and Aral seas basins (e.g. Berg, 1949; Coad & Bogutskaya, 2009), but ongoing research is revealing a greater diversity (Bogutskaya & Coad, 2009; Coad & Bogutskaya, 2009; Coad & Bogutskaya, 2012; Bogutskaya et al., 2010; Mousavi-Sabet et al., 2015; Stierandová et al., 2016). Recent studies have shown that at least eight species of the genus Alburnoides are found in Iran (Coad & Bogutskaya, 2009; Coad & Bogutskaya, 2012; Jouladeh-Roudbar et al., 2015), namely Alburnoides eichwaldii De Filippii (1863), found in river drainages of the south-western Caspian Sea basin, Alburnoides holciki Coad & Bogutskaya, 2012 known from north-western Afghanistan and north-eastern Iran, Alburnoides idignensis Bogutskaya & Coad, 2009 recordedfrom upper reaches of tributaries of Karkheh [Qareh Su] River in the Zagros Mountains, Alburnoides namaki Bogutskaya & Coad, 2009 recorded from Hamedan Province, in Qareh-Chay River drainage north of Razan, Alburnoides nicolausi Bogutskaya & Coad, 2009 found in Simareh River drainage at Nurabad which flows into the Karkheh River, Alburnoides petrubanarescui Bogutskaya & Coad, 2009 from Qasemlou-Chay, Orumiyeh [Urmia] Lake basin, Alburnoides qanati Coad & Bogutskaya, 2009 from Pulvar River drainage, Kor River basin, and A. tabarestanensis Mousavi-Sabet, Anvarifar & Azizi, 2015 from Tajan River in the south-eastern Caspian Sea basin. Alburnoides populations are widely distributed along the southern Caspian Sea basin from the Aras-Kura to the Atrak rivers (Jouladeh-Roudbar et al., 2015; Mousavi-Sabet et al., 2015). Recently, some researchers tried to clarify the taxonomic status of the genus Alburnoides in the region (Bogutskaya & Coad, 2009; Seifali et al., 2012; Mousavi-Sabet et al., 2015). As mentioned, Alburnoides bipunctatus was the name applied to populations of riffle minnows along the southern Caspian Sea basin. However, Seifali et al. (2012) sequenced the mitochondrial cytochrome b of Alburnoides populations from this region and concluded that A. bipunctatus does not exist in the area. Bogutskaya & Coad (2009) confirmed the presence of A. eichwaldii in the region and stated that the species is restricted to the south-western Caspian Sea basin at the Aras-Kura drainage. They also suggested that the riffle minnow from Atrak and Safid Rud (or correctly Sefidroud) rivers may represent distinct taxa from A. eichwaldii. Seifali et al. (2012) stated the Alburnoides populations occurring in the southern Caspian Sea basin, are distinct from the remaining species of spirlin in Iran, and that the western and eastern Alburnoides populations in the basin are distinct taxa. Seifali et al. (2012) data also suggested that the Alburnoides populations from Sefidroud and Gorganroud rivers belong to unnamed species, which was suggested previously by Coad & Bogutskaya (2012), based on meristic and vertebral counts. Similar data are showed in a recent phylogenetic study of the genus Alburnoides (Stierandová, et al., 2016). Our morphological and vertebral comparisons of Alburnoides specimens from the Atrak and Sefidroud rivers in the southern Caspian Sea basin and Namroud River in Kavir basin, with their nominal congeners in Iranian river basins, demonstrates that they clearly represents three unnamed species, which are described here.

MATERIALS AND METHODS

Sample collection and measurements: A total of 466 spirlins specimens were collected by electrofishing throughout Iran during 2010-2014 (Table 1). Caught specimens fixed in 10% formaldehyde and stored in 70 % ethanol or directly fixed in 99 % ethanol. Counts and measurements follow Hubbs & Lagler (1958) and Mousavi-Sabet et al., (2015). Measurements were made point to point to the nearest 0.1 mm. The standard length (SL) is measured from the tip of the snout to the end of the hypural complex. Head length and interorbital width were measured to their bony margins. 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 the mid-height of the caudal-fin base. The last two branched rays articulated on a single pterygiophore in dorsal and anal-fins are noted as “1½”. Mean and standard deviation were calculated without the “½”. Lateral-line scales count includes pored scales, from the first one just behind the supracleithrum to the posteriormost one at the base of the caudal-fin rays (i.e. posterior margin of hypurals) excluding 1 or 2 scales located on the bases of the caudal-fin rays. Osteological characters were examined from radiographs. Diagnosis for all the eight existent Iranian spirlin species are based on original descriptions, however detailed comparisons with congeners found in Iranian waters are given in the comparative remarks of each new described species. No material of A. petrubanarescui was available for this  study  and

Table 1. Examined Alburnoides specimens from Iranian water basins (n = number of specimens).

we based our comparisons with this species on the original description by Bogutskaya & Coad (2009). Statistical analyses were performed by Microsoft Excel 2010 package.

Abbreviations used: GUIC, the collection of the ichthyology museum of the university of Guilan, Iran; VMFC, Vatandoust and Mousavi-Sabet fish collection, Tehran, Iran. The type series is deposited in VMFC and GUIC.

RESULTS AND DISCUSSION

Obtained results showed that the genus Alburnoides in Iran contains at least eleven species, including eight described; Alburnoides eichwaldii De Filippii (1863), A. holciki Coad & Bogutskaya, 2012, A. idignensis Bogutskaya & Coad, 2009, A. namaki Bogutskaya & Coad, 2009, A. nicolausi Bogutskaya & Coad, 2009, A. petrubanarescui Bogutskaya & Coad, 2009, A. qanati Coad & Bogutskaya, 2009 and A. tabarestanensis Mousavi-Sabet, Anvarifar & Azizi, 2015, and three unnamed species. A map with all records of Alburnoides species considered in this study is shown in Fig. 1.

Alburnoides eichwaldii (De Filippi, 1863)

(Fig. 2)

Alburnus eichwaldii De Filippi, 1863: 392 [18 of separatum] (Kura R. at Tiflis [Tbilisi]).

Alburnoides bipunctatus armeniensis Dadikyan, 1972: 566 (Marmarik R., Aras R. system).

Material examined. GUIC AL201EI; 55 specimens, 48.3-67.9 mm SL, Iran: Ardebil Province, Balekhlu-Chay Stream, Aras-Kura drainage, the Caspian Sea basin, 38°02ʹ N, 48°03ʹ E; coll. H. Mousavi-Sabet, S. Vatandoust & A. Jouladeh-Roudbar.

Diagnosis.Original diagnosis of Alburnoides eichwaldii (De Filippi, 1863): the body is deep, its length exceeds the depth in four times; eye large; dorsal-fin rays branched rays 8 [=8½]; branched anal-fin rays 12 [=12½]; scales in the lateral series 50, 11 scales above and 7 scales below lateral line.

According to Bogutskaya & Coad (2009) and our examined materials, A. eichwaldii is distinguished from the other species of Alburnoides in Iran by a combination of characters, none of them unique. Alburnoides eichwaldii is distinguished from the congeners in Iran by moderately rounded caudal fin lobes, not deeply forked caudal fin; commonly scaleless ventral keel; commonly deep head and slightly to markedly rounded snout; the tip of the mouth cleft is slightly below the level of the middle of the eye or at about the lower margin of pupil; 8½ dorsal-fin rays (rarely 7½ or 9½); (10)11-14½ branched anal-fin rays (commonly 12-13½); commonly 2.5-4.2 pharyngeal teeth; 44-56 total lateral line scales; 6-10 gill rakers; (38, 39)40-43 total vertebrae; and the most common vertebral formulae are 21 + 21, 21 + 20 and 20 + 21 (Tables 2-4).

See Fig. 2 for general appearance, table 5 for morphometric data, and tables 2-4 for meristic and vertebral counts. See below for more details to distinguish A. eichwaldii from the other Iranian  Alburnoides  species  examined  for  this

study.

Distribution. Alburnoides eichwaldii is distributed in river drainages of the southwestern Caspian coast from Samur (according to Berg, 1932, 1949) down to rivers of the Lenkoran. In Iran this species is restricted to the Aras River drainage, in the southwestern Caspian Sea basin. See Fig. 1 for the distribution of the species in Iran.

Fig. 1. Records of Alburnoides species in Iran; (1) A. eichwaldii, (2) A. holciki, (3) A. idignensis, (4) A. namaki, (5) A. nicolausi, (6) A. petrubanarescui, (7) A. qanati, (8) A. tabarestanensis, (9) A. coadi sp. n., (10) A. parhami sp. n., (11) A. samiii sp. n.

Fig. 2. Alburnoides eichwaldii; Iran, Aras River, GUIC AL201EI, (a) 66 mm SL, (b) 68 mm SL.

Alburnoides holciki Coad & Bogutskaya, 2012

(Fig. 3)

Material examined. VMFC AL201HO, 36 specimens, 44.6-66.2 mm SL, Iran: Khorasan-e-Razavi Province, Hariroud River, Hariroud River basin, 35°57ʹ N, 61°07ʹ E; coll. H. Mousavi-Sabet, S. Vatandoust, B. Ganjbakhsh & A. Jouladeh-Roudbar.

Diagnosis.According to original diagnosis, Alburnoides holciki is distinguished from the other species of Alburnoides in Iran by a well-defined, sharp, scaleless or only slightly scaled ventral keel; a short, slightly pointed snout; a terminal mouth with the tip of the mouth cleft, on a level with the upper half of the pupil; a large eye (orbit width about equal to interorbital width); (46)47–51(55) lateral-line scales to posterior margin of hypurals; 2.5–4.2 pharyngeal teeth; usually 8½ branched dorsal-fin rays; usually 13–16½ branched anal-fin rays; 40–42, usually 41, total vertebrae; caudal vertebral region longer than abdominal region (most frequent vertebral formulae 20 + 21, 20 + 22 and 19 + 21); and usually 13 or 14 predorsal vertebrae. See Fig. 3 for general appearance, table 5 for morphometric data, and tables 2-4 for meristic and vertebral counts. See below for more details to distinguish A. holciki from the other Iranian Alburnoides species examined for this study.

Table 4. Frequency of occurrence of variants of the number of abdominal and caudal vertebrae in the Alburnoides species in Iran (n = number of specimens; n^*: present study, n^**: Bogutskaya & Coad (2009), n^***: Coad & Bogutskaya (2012)).

Distribution. Alburnoides holciki is distributed in the Hariroud River basin at the border of three countries including northwestern Afghanistan, northeastern Iran and southern Turkmenistan. See Fig. 1 for the distribution of the species in Iran.

Alburnoides idignensis Bogutskaya & Coad, 2009

(Fig. 4)

Material examined. VMFC AL201ID; 52 specimens, 49.0-82.4 mm SL, Iran: Kermanshah Province, Bid-e-Sorkh Stream, Gamasiab River, the Tigris River basin, 34°26ʹ N, 47°46ʹ E; coll. H. Mousavi-Sabet, S. Vatandoust, H. Bagherpour & A. Jouladeh-Roudbar.

Diagnosis.According to original diagnosis (Bogutskaya & Coad, 2009), Alburnoides idignensis is distinguished from the other species of Alburnoides in Iran by a combination of characters, none of them unique. An unbranched pectoral fin ray strongly lined with melanophores on its inner margin; an eye of an average size, the orbit diameter larger than the snout length and markedly smaller than the interorbital width; caudal fin lobes rounded and fin shallowly forked; a variably scaled ventral keel though most commonly scaled along about 1/3-2/3 of its length; a deep head with a markedly rounded, stout snout; a small mouth which is between terminal and subterminal; a tip of the mouth cleft on a level from the lower margin of the pupil; commonly 8½ branched dorsal-fin rays; 10-12(13-14)½ branched anal-fin rays; 41-49(50-51) total lateral line scales (39-49 scales to posterior margin of hypurals); commonly 2.5-4.2 or 2.4-4.2 pharyngeal teeth; (37)38-40, with a mode of 39, total vertebrae; 11-13(14) predorsal vertebrae, (18)19-20 abdominal vertebrae; (18)19-20 caudal vertebrae; a caudal vertebral region most commonly one vertebra shorter or one vertebra longer than the abdominal region; the most common vertebral formulae are 20 + 19 and 19 + 20, and the difference between the abdominal and caudal counts averaging 0.

See Fig. 4 for general appearance, table 6 for morphometric data, and tables 2-4 for meristic and vertebral counts. See below for more details to distinguish Alburnoides idignensis from the other Iranian Alburnoides species examined for this study.

Fig. 3. Alburnoides holciki; Iran, Hariroud River, VMFC AL201HO, (a) 64 mm SL, (b) 58 mm SL.

Distribution. Alburnoides idignensis is known from some upper reaches of tributaries of Karkheh [Qareh Su] River in the Zagros Mountains. The Karkheh is falling into the Tigris just below its confluence with the Euphrates. Alburnoides idignensis is normally found in western Iran, in Kermanshah and Ilam provinces. This species is found in springs and qanats in the region. Other species found sympatrically together with Alburnoides idignensis are Barbus lacerta, Alburnus mossulensis, Cobitis avicennae, Capoeta aculeata, Cyprinion macrostomum, Garra rufa, Oxynoemacheilus kiabii, Oxynoemacheilus kermanshahensis. This species is restricted to the Tigris River basin. See Fig. 1 for the distribution of the species in Iran.

Alburnoides namaki Bogutskaya & Coad, 2009

(Fig. 5)

Material examined. VMFC AL201NA; 21 specimens, 44.8-94.4 mm SL, Iran: Markazi Province, Qareh-Chay River, Namak Lake basin, 34°53′ N, 050°02′ E; coll. H. Mousavi-Sabet, S. Vatandoust, H. Bagherpour & A. Jouladeh-Roudbar. VMFC AL201NA2; 20 specimens, 46.5-77.2 mm SL, Iran: Markazi Province, Qareh-Chay River, Namak Lake basin, 34°53′ N, 050°02′ E; coll. H. Mousavi-Sabet & S. Vatandoust. VMFC AL201NA3; 10 specimens, 48.0-81.5 mm SL, Iran: Markazi Province, Qareh-Chay River, Namak Lake basin, 34°53′ N, 050°02′ E; coll. H. Mousavi-Sabet.

Table 5. Morphometric data of Alburnoides eichwaldii (GUIC AL201EI, n = 55), and A. holciki (VMFC AL201HO, n = 36).

Diagnosis.According to original diagnosis (Bogutskaya & Coad, 2009), Alburnoides namaki is distinguished from the other species of Alburnoides in Iran by a combination of characters, none of them unique. Alburnoides namaki is distinguished from the congeners in Iran by the lack of strong spots or dark outline to the lateral line canal (however, some specimens have strong pigmentation above and below the lateral line pores, forming an evident pale line margined with dark. As mentioned by Bogutskaya & Coad (2009), a broad mid-flank stripe can be well developed or weakly expressed and, on the caudal peduncle, obscures the lateral line pigment pattern. The lateral line pattern can be weak and this can be seen over the anal fin where the flank stripe does not extend down to the decurved lateral line); a small eye, the orbit width about equal to the snout length but markedly smaller than the interorbital width; caudal fin lobes rounded and fin shallowly forked; a sharp scaleless ventral keel behind the pelvic fins along the abdomen to the anus; a deep head with a stout snout which is markedly rounded; a small, almost subterminal mouth with the tip of the mouth cleft on a level from the lower margin of the eye or below; commonly 8½ branched dorsal-fin rays; 10-13½, commonly 11-12½, branched anal-fin rays; (43)44-50(52) total lateral line scales (42-51 scales to posterior margin of hypurals); 2.5-4.2 pharyngeal teeth (or other variants with four teeth on the right ceratobranchial); commonly 39-41 total vertebrae; 11-13(14); a low number of predorsal vertebrae, commonly 12-13; 19-20(21) abdominal vertebrae; 19-21 caudal vertebrae; a caudal vertebral region most commonly equal to the abdominal region; and the most common vertebral formulae are 20 + 20, 20 + 19 and 19 + 20. See Fig. 5 for general appearance, table 6 for morphometric data, and tables 2-4 for meristic and vertebral counts. See below for more details to distinguish Alburnoides namaki from the other Iranian Alburnoides species from the related water basins in northern and eastern parts, examined for this study.

Fig. 4. Alburnoides idignensis; Iran, Gamasiab River, VMFC AL201ID, (a) 51 mm SL, (b) 55 mm SL.

Fig. 5. Alburnoides namaki; Iran, Qareh-Chay River, VMFC AL201NA, (a) 71 mm SL, (b) 67 mm SL.

Distribution. Alburnoides namaki is known from the qanat in the Qareh-Chay drainage of the Namak Lake basin.

This species is restricted to the Namak Lake basin. About 25 years ago, Tave qanat has been dried and now there is no river or qanat around the type locality. This species is restricted to the Namak Lake basin. See Fig. 1 for the distribution of the species in Iran.

Alburnoides nicolausi Bogutskaya & Coad, 2009

(Fig. 6)

Material examined. VMFC AL201NI; 38 specimens, 35.4-67.8 mm SL, Iran: Lorestan Province, Nurabad Stream, Gamasiab River, the Tigris River basin, 34°05ʹ N, 48°00ʹ E; coll. S. Vatandoust, A. Jouladeh-Roudbar & H. R. Bagherpour.

Diagnosis.According to original diagnosis, Alburnoides nicolausi is distinguished from the other species of Alburnoides in Iran by a combination of characters, none of them unique. Alburnoides nicolausi is distinguished by an eye of an average size, the orbit diameter larger than the snout length and smaller than the interorbital width; caudal fin lobes rounded and fin shallowly forked; a variably scaled ventral keel though most commonly scaled only along about 1/3 of its length or scaleless; a deep head with a moderately stout snout which is slightly pointed; a tip of the mouth cleft on the level about the lower margin of the pupil, commonly

Table 6. Morphometric data of Alburnoides idignensis (VMFC AL201ID, n = 52), and A. namaki (VMFC AL201NA, n = 51).

Fig. 6. Alburnoides nicolausi; Iran, Nurabad Stream, VMFC AL201NI, (a) 67 mm SL, (b) 46 mm SL (Photograph by Arash Jouladeh-Roudbar).

7½ branched dorsal-fin rays; 8-11½ branched anal-fin rays; (43)43-47(48-50) total lateral line scales (42-48 scales to posterior margin of hypurals); commonly 2.5-4.2 or 2.4-4.2 pharyngeal teeth; commonly 39-40 total vertebrae. See Fig. 6 for general appearance, table 7 for morphometric data, and tables 2-4 for meristic and vertebral counts. See below for more details to distinguish A. nicolausi from the other Iranian Alburnoides species examined for this study.

Distribution. Alburnoides nicolausi is only known from its type locality, a stream in the Simareh River drainage at Nurabad. This species is restricted to the Tigris River basin. See Fig. 1 for the distribution of the species in Iran.

Alburnoides petrubanarescui Bogutskaya & Coad, 2009

Material examined. None.

Diagnosis.According to original diagnosis, Alburnoides petrubanarescui is distinguished from the other species of Alburnoides in Iran by a combination of characters, none of them unique. Alburnoides petrubanarescui is distinguished from the congeners in Iran by a small eye; the orbit width about equal to the snout length but markedly smaller than the interorbital width; caudal fin lobes rounded and the fin shallowly forked; a scaled ventral keel behind the pelvic fins along the abdomen to the anus, a deep head with a stout snout which is markedly rounded; a tip of the mouth cleft on the level below the lower margin of the eye; commonly 7½ (less frequently 8½) branched dorsal-fin rays; 8-10½, commonly 9½, branched anal-fin rays; 44-51 total lateral line scales (42-49 scales to posterior margin of hypurals); 2.5- 4.2 pharyngeal teeth

Table 7. Morphometric data of Alburnoides nicolausi (VMFC AL201NI, n = 38), and A. qanati (VMFC AL201QA, n = 50).

(or other variants with four teeth on the right ceratobranchial); commonly 40-41 total vertebrae. See tables 2-4 for meristic and vertebral counts. See below for more details to distinguish A. petrubanarescui from the other Iranian Alburnoides species examined for this study.

Distribution. Alburnoides petrubanarescui is only known from Qasemlou-Chay, Orumiyeh [Urmia] Lake basin.

This species is an endemic species to the Orumiyeh Lake basin and restricted to the basin. See Fig. 1 for the distribution of the species in Iran.

Alburnoides qanati Coad & Bogutskaya, 2009

(Fig. 7)

Material examined. VMFC AL201QA1; 21 specimens, 45.7-70.3 mm SL, Iran: Fars province, Kor River, Kor River basin, 30°17ʹ N, 52°18ʹ E; coll. H. Mousavi-Sabet, S. Vatansoust. VMFC AL201QA2; 29, 44.1-71.0 mm SL, Iran: Fars province, Kor River, Kor River basin, 30°28ʹ N, 52°05ʹ E; coll. H. Mousavi-Sabet, S. Eagderi.

Diagnosis.According to original diagnosis, Alburnoides qanati is distinguished from the other species of Alburnoides in Iran by a combination of characters, none of them unique.

Fig. 7. Alburnoides qanati; Iran, Kor River, VMFC AL201QA, (a) 65 mm SL, (b) 59 mm SL.

Alburnoides qanati is distinguished from the congeners in Iran by a large eye, the orbit width exceeding both the snout length and the interorbital width; a scaled ventral keel behind the pelvic fins along the abdomen to the anus; commonly 43-47 lateral line scales to posterior margin of hypurals; 2.5-4.2 pharyngeal teeth; commonly 8½ branched dorsal-fin rays; 10½-12½ branched anal-fin rays; 40-41 total vertebrae. See Fig. 7 for general appearance, table 7 for morphometric data, and tables 2-4 for meristic and vertebral counts. See below for more details to distinguish A. qanati from the other Iranian Alburnoides species examined for this study.

Distribution. Alburnoides qanati is only known from Pulvar River drainage, Kor River endorheic basin. The type series was collected from a small stream which issues from a qanat near the archaeological site of Naqsh-e-Rostam to the northeast of Persepolis. This species is an endemic species and restricted to the Kor River basin. See Fig. 1 for the distribution of the species in Iran.

Alburnoides tabarestanensis Mousavi-Sabet, Anvarifar & Azizi, 2015

(Fig. 8)

Material examined. VMFC AL201MH, 68.3 mm SL; Holotype. VMFC AL2049P, 45 specimens, 33.7-83.0 mm SL; GUIC ALT-P, 4 specimens, 36.4-65.2 mm SL Iran, Paratypes. Iran: Mazandaran Province, Tajan River, the southern Caspian Sea basin, 36°11ʹ N, 53°19ʹ E; coll. H. Mousavi-Sabet, S. Vatandoust & H. Anvarifar.

Diagnosis.According to original diagnosis, A. tabarestanensis is distinguished from the other species of Alburnoides in Iran by a combination of characters, none of them unique. Alburnoides tabarestanensis is distinguished from the congeners in Iran by a completely scaleless ventral keel (most common); the lack of well-marked spots or dark pigmentation in the lateral line canal; terminal mouth with the tip of the mouth cleft between level of middle of pupil and lower margin of pupil; 47–52 lateral line scales to posterior margin of hypurals; typically 9 or 10 scale rows between lateral line and dorsal fin origin; 2.5-4.2 pharyngeal teeth; usually 4 or 5 scale rows between lateral line and anal fin origin; commonly 16-18 scales around caudal peduncle; typically 39-41 total vertebrae; usually 8½ branched dorsal-fin rays; commonly 12½ or 13½ branched anal fin rays; usually 7 branched pelvic fin rays; commonly 11-13 branched pectoral fin rays; usually 8 or 9 gill rakers in the outer row on first left arch. See Fig. 8 for general appearance, table 8 for morphometric data, and tables 2-4 for meristic and vertebral counts. See below for more details to distinguish A. tabarestanensis from the other Iranian Alburnoides species examined for this study.

Distribution. Alburnoides tabarestanensis is only known from Tajan River in the south-eastern Caspian Sea basin. This species is an endemic species and restricted to the Caspian Sea basin. See Fig. 1 for the distribution of the species in Iran.

Alburnoides coadi, new species

(Figs. 9-11)

Holotype. VMFC-ALK2-H, 107.9 mm SL; Iran, Tehran Province, Namroud River, Hableroud River drainage, Kavir basin, 35°43ʹ N, 52°39ʹ E; coll. H. Mousavi-Sabet & S. Vatandoust, 15 September 2012.

Paratypes. VMFC-ALK2-P1 to VMFC-ALK2-P45, 45 specimens, 55.2-81.4 mm SL; collected with holotype. GUIC-ALK2-P1 to GUIC-ALK2-P4, 4 specimens, 58.2-66.0 mm SL; collected with holotype.

Fig. 8. Alburnoides tabarestanensis; Iran, Tajan River, (a) VMFC AL201MH, 68 mm SL, (b) VMFC AL2049P, 66 mm SL.

Diagnosis: Alburnoides coadi, sp. n., is distinguished from the other species of Alburnoides in Iran diagnosed above by a combination of the following characters: an eye of a relatively small size, the orbit diameter larger than the snout length and markedly smaller than the interorbital width; caudal fin lobes rounded and fin moderately forked; a variably scaled ventral keel though most commonly scaled along about 2/3 of its length, or a completely scaled ventral keel; a deep head with a markedly rounded and stout snout; a small mouth which is between terminal and subterminal; a tip of the mouth cleft on a level from the lower margin of the pupil or somewhat below it; the lack of well-marked spots or dark pigmentation in the lateral line canal; 47–52 lateral line scales to posterior margin of hypurals; typically 9 or 10 scale rows between lateral line and dorsal fin origin; usually 5 or 6 scale rows between lateral line and anal fin origin; 2.5-4.2 pharyngeal teeth; commonly 17-19 scales around caudal peduncle; typically 39-41, with a mode of 40, total vertebrae; 13-14 predorsal vertebrae, 19-20 abdominal vertebrae, 19-21 caudal vertebrae; caudal vertebral region

equal or slightly longer than an abdominal region (vertebral formulae 19 + 20, 20 + 20 and 20 + 21); usually 8½ branched dorsal-fin rays; commonly 12½-13½ branched anal fin rays; usually 8 or 9 gill rakers in the outer row on first left arch. Detailed comparisons with congeners found in Iranian waters are given in the comparative remarks.

Table 8. Morphometric data of Alburnoides tabarestanensis (paratypes: VMFC AL2049P, n = 45; GUIC ALT-P, n = 4), and A. coadi sp. n. (holotype VMFC-ALK2-H, paratypes: VMFC-ALK2-P, n = 45; GUIC-ALK2-P, n = 4). H, holotype.

Fig. 9. Alburnoides coadi sp. n.; Iran, Tehran Province, Namroud River, holotype, VMFC-ALK2-H, 107.9 mm SL.

Description: General appearance shown in Figs. 9-11. Morphometric data presented in table 8. Meristic and vertebral counts presented in tables 2-4. Body is moderately compressed, upper body profile convex to slightly straight; lower profile convex. The ventral keel between the pelvics and anal fin is variably scaled: scaled along about 2/3 of its length or completely scaled. Anal-fin origin is in front of a vertical from posterior end of the dorsal fin base. Snout markedly stout and rounded. Mouth terminal to subterminal; tip of the mouth cleft on a level from the lower margin of the pupil or somewhat below it. Mouth cleft oblique, upper jaw slightly to moderately longer than lower jaw, junction of lower jaw and quadrate at vertical through the lower part of anterior eye margin. Dorsal-fin origin behind a vertical through pelvic-fin origin. Pectoral fin short, not reaching pelvic fin origin (pectoral fin 3-5 scale rows shorter than pectoral-pelvic distance), its outer margin convex. Pelvic fin rounded, its tip not reaching to anus. Caudal fin moderately forked, its lobes rounded.

Body depth enters standard length 3.0-3.5 times, head length enters 3.7-4.2, predorsal length 1.7-1.9, caudal peduncle depth 8.0-8.5, and caudal peduncle length 4.8-5.8. Orbit width enters head length 3.5-4.7 times, snout length enters 3.6-4.6, and interorbital width 2.6-2.9. Pectoral fin length enters pectoral fin origin to pelvic fin origin distance 0.7-0.9 times, and pelvic fin length enters pelvic fin origin to anal fin origin distance 0.7-0.8 times.

Dorsal fin unbranched rays 3, dorsal fin branched rays 7½ (4) or 8½ (46) (mean 7.92, standard deviation 0.27), anal fin unbranched rays 3, anal fin branched rays 11½ (2), 12½ (26), 13½ (19) or 14½ (3) (mean 12.46 ± 0.68), pectoral fin branched rays 11(8), 12(17), 13(23) or 14(2) (mean 12.38 ± 0.81), pelvic fin branched rays 6(3) or 7(47) (mean 6.94 ± 0.24). Pelvic axillary scale present, its length is about 3-4 scale rows. Dorsal and anal fins outer margins truncate to slightly concave.

Lateral line complete with 0, 1 or 2 unpored scales at the posterior end of the lateral series; lateral line scales to posterior margin of hypurals 47(2), 48(18), 49(19), 50(6) or 51(5) (mean 48.88 ± 1.02); scales above lateral line to dorsal fin origin 9(10), 10(38) or 11(4) (mean 9.92 ± 0.49); scales below lateral line to anal fin origin 4(19), 5(24) or 6(7) (mean 4.76 ± 0.69); scales below lateral line to pelvic fin origin 4(29),

Fig. 10. Alburnoides coadi sp. n.; Iran, Tehran Province, Namroud River, paratype; a, VMFC-ALK2-P1, 81 mm SL; b, VMFC-ALK2-P3, 76 mm SL; c, VMFC-ALK2-P4, 71 mm SL; d, VMFC-ALK2-P5, 64 mm SL.

5(20) or 6(1) (mean 4.44 ± 0.54); total scale radii 16(7), 17(10), 18(28) or 19(5) (mean 17.62 ± 0.85). Total gill rakers at outer row on first left arch number 8(25), 9(23) or 10(2) (mean 8.54 ± 0.58). Total vertebrae including four Weberian vertebrae and last hypural complex centrum 39(15), 40(26) or 41(9) (mean 39.88 ± 0.69). Thirteen to 14 predorsal vertebrae; 19-20 abdominal vertebrae, 19-21 caudal vertebrae; caudal vertebral region most commonly equal to the abdominal region; most common vertebrae formulae 19 + 20, 20 + 20, or 20 + 21. Pharyngeal teeth 2.5-4.2 in 10 examined specimens.

Colouration: In living specimens (Fig. 11), overall colouration is silvery, with the bases of the pectoral (markedly), pelvic and anal fins (slightly) orange. Dorsum and top of head are light to dark grey, with a pale olive hue. Facial bones and opercle silvery, top of head light grey. The lower portion of head and body are pearly-white. The flanks above lateral line may have a golden hue. Faint yellow spots occur in rows along the flanks in some specimens. Pigmentation of preserved specimens fixed in 10% formalin and stored in 70% alcohol (Figs. 9-10) overall tan, darker dorsally; horizontal rows of dark blotches formed by dark pigmentation concentrated on middle of scales, moderately conspicuous, above the lateral line; lateral line with some scales with pores outlined with dark pigmentation, especially the anterior-most scales; narrow dark mid-lateral stripe along lateral septum, more discernible from vertical through dorsal fin onwards; a faint, broad grey mid-lateral stripe; fins mostly hyaline, with some black pigmentation lining caudal fin rays.

Distribution and notes on biology: Alburnoides coadi is known only from Namroud River, Hableroud River drainage, in Kavir basin, northern part of Iran (Fig. 1). The upstream portion of Namroud River at the type locality at time of the collection of the type specimens (Fig. 12) had clear water, with medium to fast water flow. The stream width was about 3 m and maximum depth was up to 1 m, with grassy shores, submergent plants, and the stream bed was gravel and mud. Other species collected syntopically were Barbus sp.,Capoeta aculeata (Valenciennes, 1844), Capoeta buhsei Kessler, 1877, Carassius auratus (Linnaeus, 1758), Carassius gibelio (Bloch, 1782), Squalius cf. orientalis (Nordmann, 1840), Paracobitis malapterura (Valenciennes, 1846), and Oncorhynchus mykiss (Walbaum, 1792).

Fig. 11. Alburnoides coadi sp. n., holotype, VMFC-ALK2-H, 107.9 mm SL.

Fig. 12. Namroud River, Kavir basin; type locality of A. coadi sp. n.

Etymology: The species name coadi is in honor of Brian W. Coad (the Canadian Museum of Nature, Ottawa), the most famous ichthyologist who studied Iranian freshwater fishes, especially the genus Alburnoides.

Comparative remarks: Alburnoides coadi differs from A. eichwaldii, A. holciki, A. namaki, A. nicolausi and A. tabarestanensis by its completely or almost completely scaled ventral keel (vs. most commonly 2/3 or completely scaleless ventral keel).Alburnoides coadi is further distinguished fromA. holciki, A. parhami and A. qanati by having a rounded snout (vs. pointed), and the tip of the mouth cleft on a level with the lower margin of the pupil to below the lower margin of the eye (vs. a tip of the mouth on a level with the upper half of the pupil).

Alburnoides coadi is further distinguished from A. eichwaldii by having less scales above lateral line to dorsal fin origin (commonly 9-10 vs. 11), less scales below lateral line to anal fin origin (typically 4-5 vs. 7), precaudal and caudal vertebrae equal in number or precaudal one vertebrae longer than it (vs. the caudal region is commonly one vertebrae shorter than the abdominal region), mid fins located at more posterior levels (predorsal 52.2-57.8, mean 54.0 in SL, vs. 47.5-53.7, mean 50.6 in SL, preanal 64.8-69.0, mean 66.8 in SL, vs. 60.2-65.8, mean 62.8 in SL, prepectoral 23.0-29.8, mean 26.8 in SL, vs. 17.2-23.5, mean 20.8 in SL, and preventral 47.3-52.0, mean 49.5 in SL, vs. 41.6-47.3, mean 44.2 in SL), and tip of the mouth cleft commonly at level of the lower margin of the eye or below (vs. at level of the middle of the eye).

Alburnoides coadi further differs from A. holciki by having fewer anal fin branched rays (commonly 12½-13½ vs. 14½–16½), less numerous total vertebrae (commonly 39-40 vs. 41-42), shorter anal fin base (14.3-19.8, mean 17.5 in SL, vs. 17.4-21.8, mean 19.4 in SL), mid fins located at more posterior levels, and the tip of the mouth cleft commonly at level of the lower margin of the eye or below (vs. at level with the upper half of the pupil).

Alburnoides coadi is further distinguished from A. idignensis by having more anal fin branched rays (commonly 12½-13½ vs. 11½–12½), 39-41 total vertebrae (vs. 38-40), 47-51 lateral line scales (vs. 41-46 lateral line scales), total scale radii 16-19, median 18 (vs. 12-17, median 15), mid fins located at more posterior levels, and normally 12½-13½ anal fin branched rays (vs. 11½-12½).

Alburnoides coadi is further distinguished from A. namaki by having less branched anal fin rays (commonly 11½-12½, vs. 12½-13½), smaller scales (commonly 47-51, median 49, vs. 46-49, median 47), more numerous gill rakers (commonly 8-9 vs. 6-8), scales above lateral line to dorsal fin origin (commonly 9-10 vs. 10-11), a relatively smaller eye (21.4-30.5, mean 27.4 in HL, vs. 24.6-35.5, mean 31.4 in HL), deeper caudal peduncle (caudal peduncle depth 11.7-12.6, mean 12.0 in SL, vs. 9.2-12.1, mean 10.5 in SL), and longer post orbital distance (47.6-56.4, mean 52.4 in HL, vs. 43.1-48.5, mean 45.0 in HL).

Alburnoides coadi is further differs from A. nicolausi by having more branched dorsal-fin rays (commonly 8½ vs. 7½), more branched anal-fin rays (12½-13½ vs. 9½-11½), more gill rakers (8-9 vs. 7-8), smaller scales (commonly 47-51, median 49, vs. 42-50, median 44), total scale radii 16-19, median 18 (vs. 13-17, median 15), and completely or almost completely scaled ventral keel (vs. most commonly 2/3 or completely scaleless ventral keel).

Alburnoides coadi is further distinguished from A. petrubanarescui by having more branched dorsal-fin rays (commonly 8½ vs. 7½), more branched anal-fin rays (commonly 12½-13½ vs. 9½-10½), and more gill rakers counts (8-9 vs. 7-8).

Alburnoides coadi further differs from A. qanati by having more branched anal-fin rays (commonly 12½-13½ vs. 11½-12½), smaller scales (commonly 48-49 vs. 43-47), more gill rakers (commonly 8-9 vs. 6-8), and total scale radii commonly 16-19 (vs. 11-16).

Alburnoides coadi is further distinguished from A. tabarestanensis by having 13-14, median 13 predorsal vertebrae (vs. 12-13, median 12), shorter snout (21.6-29.4, mean 25.7 in HL, vs. 24.8-31.9, mean 28.9 in HL), and completely or almost completely scaled ventral keel (vs. most commonly 2/3 or completely scaleless ventral keel). See below for more details to distinguish A. coadi from the other Iranian Alburnoides species described in this study.

Alburnoides parhami, new species

(Figs. 13-15)

Holotype. VMFC-ALP3-H, 66.5 mm SL; Iran, Khorasan-e-Shomali Province, Baba-Aman Stream, Atrak River drainage, the south-eastern Caspian Sea basin, 37°29' N, 57°26' E; coll. H. Mousavi-Sabet, B. Ganjbakhsh & A. Jouladeh-Roudbar, 10 Noveber 2013.

Paratypes. VMFC-ALP3-P1 to VMFC-ALP3-P45, 45 specimens, 39.9-62.0 mm SL; collected with holotype. GUIC-ALP3-P1 to GUIC -ALP3-P4, 4 specimens, 43.1-53.5 mm SL; collected with holotype.

Diagnosis: Alburnoides parhami is distinguished from the other species of Alburnoides in Iran diagnosed above by a combination of the following characters: most commonly a sharp, scaleless ventral keel; a long, slightly pointed snout; a terminal mouth with the tip of the mouth cleft on a level with the upper half of the pupil; the lack of well-marked spots or dark pigmentation in the lateral line canal; a large eye

Fig. 13. Alburnoides parhami sp. n.; Iran, Khorasan-e-Shomali Province, Baba-Aman Stream, Atrak River drainage, holotype, VMFC-ALP3-H, 66.5 mm SL.

(orbit width about equal to interorbital width); 46–51 lateral line scales to posterior margin of hypurals; commonly 2.5-5.2 pharyngeal teeth; commonly 8½ branched dorsal-fin rays; commonly 12½ or 13½ branched anal fin rays; typically 10 scale rows between lateral line and dorsal fin origin; 39–41, usually 40-41, total vertebrae; caudal vertebral region equal or slightly longer than abdominal region (most frequent vertebral formulae 20 + 20 and 20 + 21); and usually 12 or 13 predorsal vertebrae. Detailed comparisons with congeners found in Iranian waters are given in the comparative remarks.

Description: General appearance shown in Figs. 13-15. Morphometric data presented in table 9. The body is markedly compressed. The upper body profile is moderately rounded while the lower profile is more convex or both profiles are about equally convex. The ventral keel between the pelvic and anal fins is well-developed, sharp, protruding, and most commonly scaleless (rarely scaled along 1/4 of its length). The eye is large, orbit width about equal to interorbital width. The snout is relatively long, and slightly pointed. The mouth is terminal, with the tip of the mouth cleft on a level with the upper half of the pupil. The mouth cleft is always turned upward, and the junction of the lower jaw and the quadrate is on about a vertical through the anterior eye margin. Anal-fin origin at vertical through dorsal-fin base end, its outer margin truncate to slightly concave. Dorsal fin located behind vertical through pelvic fin insertion, its outer margin truncate to slightly round.

Pectoral fin long, commonly reaching pelvic fin insertion, its outer margin convex.

Pelvic fin rounded, its tip normally reaching or slightly behind anus, in some specimens reaching anal fin base. Pelvic axillary scale present. Caudal-fin lobes are pointed, the fin is clearly forked.

Body depth enters standard length 3.4-4.0 times, head length enters 3.6-4.0, predorsal length 1.9-2.1, caudal peduncle depth 8.7-9.8, and caudal peduncle length 4.6-4.9. Orbit width enters head length 3.1-3.5 times, snout length enters 3.1-3.7, and interorbital width 3.0-3.5. Pectoral fin length enters pectoral fin origin to pelvic fin origin distance 0.9-1.2 times, and pelvic fin length enters pelvic fin origin to anal fin origin distance 0.9-1.1 times.

Dorsal fin unbranched rays 3, dorsal fin branched rays 7½ (2), 8½ (47) or 9½ (1) (mean 7.98, standard deviation 0.25), anal fin unbranched rays 3, anal fin branched rays 11½ (4), 12½ (25), 13½ (19) or 14½ (2) (mean 12.38 ± 0.70), branched pectoral fin rays 11(18), 12(24), 13(7) or 14(1) (mean 11.82 ± 0.75), pelvic fin branched rays 6(6) or 7(44) (mean 6.88 ± 0.33).

Lateral line complete with 0, 1 or 2 unpored scales at the posterior end of the lateral series; lateral line scales to posterior margin of hypurals 46(1), 47(4), 48(23), 49(12), 50(5), 51(4) or 52(1) (mean 48.64 ± 1.21); scales above lateral line to dorsal fin origin 8(1), 9(8), 10(39) or 11(2) (mean 9.84±0.51); scales below lateral line to anal fin origin 3(1), 4(47) or 5(2) (mean 4.02 ± 0.25); scales below lateral line to pelvic fin origin 3(3), 4(45) or 5(2) (mean 3.98 ± 0.32). Total scale radii 16(7), 17(26), 18(14) or 19(3) (mean 17.26 ± 0.78). Total gill rakers at outer row on first left arch number 6(2), 7(18), 8(27) or 9(3) (mean 7.62 ± 0.67). Total vertebrae including four Weberian vertebrae and last hypural complex centrum 39(3), 40(32) or 41(15) (mean 40.24 ± 0.56). Twelve to 13 predorsal vertebrae; 19(1), 20(57) or 21(2) abdominal vertebrae, 19(2), 20(42) or 21(16) caudal vertebrae; caudal vertebral region most commonly equal to the abdominal region; most common vertebrae formulae 20 + 20 or 20 + 21. Pharyngeal teeth 2.5-5.2 (8) and 2.5-4.2 (2) in 10 examined specimens.

Colouration: In living specimens, overall colouration is silvery; darker at upper flank; facial bones and opercle silvery; the bases of the pectoral, pelvic and anal fins orange. Dorsum and top of head are light to dark grey, with an olive hue. The lower portion of head and body are pearly-white. The flanks above lateral line may have a golden hue. Faint yellow spots occur in rows along the flanks. Rows of dark blotches along scales rows, formed by dark pigmentation concentrated on center of scales, more conspicuous above lateral line and present only as scattered dark pigmentation below it; a diffuse, relatively narrow dark stripe at caudal peduncle; lateral-line outlined by dark pigmentation, forming a weakly developed, often interrupted but conspicuous longitudinal dark stripe. Faint dark pigmentation on flank scales above the lateral line forming diffuse stripes. Dorsal and caudal fins either with some grey pigmentation or dark grey. Pigmentation of preserved specimens fixed in 10% formalin and stored in 70% alcohol (Figs. 13-14) overall tan, darker dorsally; horizontal rows of dark blotches formed by dark pigmentation concentrated on middle of scales, moderately conspicuous, above the lateral line; lateral line with some scales with pores outlined with dark pigmentation, especially the most anterior scales; narrow dark middle lateral stripe along lateral septum, more discernible from vertical through dorsal fin onwards. Fins mostly hyaline, with some black pigmentation lining dorsal and caudal fins rays, dorsal most pectoral-fin rays and the most anterior anal fin rays.

Distribution and notes on biology: Alburnoides parhami is known from Baba-Aman Stream, Atrak River drainage, in the south-eastern Caspian Sea basin (Fig. 1). The stream Baba-Aman at the type locality at time of the collection of the type specimens (Fig. 16) had none-clear water, water flow was medium to fast, the stream width was about 3-5 m and maximum depth was up to 1.5 m, with grassy shores, submergent plants, and the stream bed was gravel and mud. Other species collected syntopically were Capoeta gracilis (Keyserling, 1861), Luciobarbus mursa (Guldenstadt, 1773), and Paracobitis atrakensis, Esmaeili et al., 2014.

Etymology: The species name parhami is generally in honor of all the Iranian conservative officers who sacrificed their lives in order to keep the wild environment, and especially for Mr. Saeid Parham (1980 – 2009), the conservative officer in Khorasan-e-Shomali Province, who was killed in a battle with illegal hunters near the type locality of Alburnoides parhami (near the border of Turkmenistan).

Comparative remarks: Alburnoides parhami differs from A. coadi, A. idignensis,A. petrubanare-

Fig. 14. Alburnoides parhami sp. n.; Iran, Khorasan-e-Shomali Province, Baba-Aman Stream, Atrak River drainage, paratype; a, VMFC-ALP3-P1, 62 mm SL; b, VMFC-ALP3-P2, 61 mm SL; c, VMFC-ALP3-P3, 56 mm SL; d, VMFC-ALP3-P5, 41 mm SL.

Fig. 15. Alburnoides parhami sp. n., above: holotype, VMFC-ALP3-H, 66.5 mm SL; below: paratype, VMFC-ALP3-P7, 60 mm SL.

-scui and A. qanati by its completely or almost completely scaleless ventral keel (vs. most commonly scaled ventral keel).Alburnoides parhami is distinguished from A. coadi, A. idignensis, A. namaki, A. nicolausi and A. petrubanarescui by having a short, slightly pointed snout (vs. stout and rounded), a terminal mouth with the tip of the mouth cleft on a level with the upper half of the pupil (vs. almost or completely subterminal with a tip of the mouth on a level with the lower margin of the pupil to below the lower margin of the eye), a clearly forked caudal-fin with pointed lobes (vs. shallowly forked with rounded lobes), and orbit width about equal to interorbital width (vs. smaller). Alburnoides parhami is further distinguished from A. coadiby an upturned mouth with a tip of the mouth cleft on a level with the upper half of the pupil (vs. a more horizontal, slightly curved or straight but never upturned mouth, with a tip of the mouth cleft on a level with the middle of the eye or below); a pointed snout (vs. slightly to markedly rounded); 12–13 with a mode of 12 predorsal vertebrae (vs. 13–14, median 13); shallower head (head depth 61.3-74.9, mean 67.9 in HL, vs. 67.1-80.7, mean 75.5 in HL); and orbit width about equal to interorbital width (vs. smaller).

Alburnoides parhami can be further distinguished from A. eichwaldii by having an upturned mouth with a tip of the mouth cleft on a level with the upper half of the pupil (vs. a more horizontal, slightly curved or straight but never upturned mouth, with a tip of the mouth cleft on a level with the middle of the eye or below), a pointed snout (vs. slightly to markedly rounded), less scales above lateral line to dorsal fin origin (commonly 10 vs. 11), less scales below lateral line to anal fin origin (commonly 4 vs. 7), abdominal and caudal vertebrae equal in number or abdominal one vertebrae shorter than it vs. the caudal region is commonly one vertebrae shorter than the abdominal region, 12-13 predorsal vertebrae (vs. 13-15), longer head (19.6-25.0, mean 21.2 in SL, vs. 25.1-29.9, mean 27.2 in SL), and longer paired fins (pectoral and pelvic fins reaching pelvic and anal fins respectively, vs. not reaching).

Alburnoides parhami is further distinguished from A. holciki by having commonly 5 pharyngeal teeth in the long row on the 5th ceratobranchial (vs. 4); 12½–13½ branched anal-fin rays (vs. 14½–15½); 39–41, usually 40, total vertebrae (vs. 40–42, usually 41); 12–13 with a mode of 12 predorsal vertebrae (vs. 13–14, with a mode of 13); a shorter caudal vertebral region, containing 19–21, with a mode of 20 vertebrae (vs. 20–22, with a mode of 21); and 20 + 20 most usual vertebral formula (vs. 20 + 21).

Alburnoides parhami is further distinguished from A. idignensis by having 39–41, usually 40, total vertebrae (vs. 37–40, usually 39); commonly 12½-13½ anal fin branched rays (vs. 11½–12½); 46-51 lateral line scales (vs. 41-46); and 16-19 total scale radii, median 17 (vs. 12-17, median 15).

Alburnoides parhami is further distinguished from from A. namaki by having commonly 40–41 total vertebrae (vs. 39–40); commonly 7-8 gill rakers, median 8 (vs. 6-8, median 7); smaller scales (commonly 48-49, median 48 lateral line scales, vs. 46-49, median 47); commonly 9-10 scales above lateral line to dorsal fin origin (vs. 10-11); and commonly 4 scale rows between lateral line and anal fin origin (vs. 5).

Alburnoides parhami is further distinguished from A. nicolausi by having commonly 8½ branched anal-fin rays (vs. 7½); commonly 12½-13½ branched anal-fin rays (vs. 9½-11½); commonly 40–41 total vertebrae, median 40 (vs. 38–40, median 39); and relatively longer snout (21.9-32.4, mean 28.6 in HL, vs. 18.7-27.2, mean 22.2 in HL).

Alburnoides parhami is further easily distinguishable from A. petrubanarescui by having a usually scaleless ventral keel (vs. completely scaled); commonly 12½-13½ branched anal-fin rays (vs. 8½-10½); 12–13 with a mode of 12 predorsal vertebrae (vs. 13–14, with a mode of 13); and an abdominal region equal with or shorter than a caudal vertebral region, most frequent vertebral formulae 20 + 20 or 20 + 21 (vs. longer, usually 21 + 19 and 21 + 20).

Alburnoides parhami is further distinguished from A. qanatiby having commonly 12½-13½, median 12½ branched anal-fin rays (vs. 11½-12½, median 11½); smaller scales (commonly 48-49 lateral line scales, vs. 43-47); 12–13 with a mode of 12 predorsal vertebrae (vs. 13–14, median 13); 16-19 total scale radii, median 17 (vs. 11-16, median 12); and scaleless ventral keel (vs. completely scaled).

Alburnoides parhami is further distinguished from A. tabarestanensis by the shape of the mouth which is upturned with a pointed snout and a tip of the mouth cleft on a level with the upper half of the pupil (vs. a more horizontal, slightly curved mouth with a rounded snout and a tip of the mouth on a level with the lower margin of the pupil); and 5 pharyngeal teeth in the long row on the 5th ceratobranchial (vs. usually 4).

See below for more details to distinguish A. parhami from the other Iranian Alburnoides species described in this study.

Fig. 16. Baba-Aman Stream, Atrak River, the southeastern Caspian Sea basin; type locality of A. parhami sp. n.

Alburnoides samiii, new species

(Figs. 17-19)

Holotype. VMFC-ALS4-H, 75.9 mm SL; Iran, Guilan Province, upper Sefidroud River drainage, Tutkabon Stream, 36º50.756′N, 49º35.021′E; coll. H. Mousavi-Sabet, 15 October 2013.

Paratypes. VMFC-ALS4-P1 to VMFC-ALS4-P45, 45 specimens, 52.9-81.5 mm SL; collected with holotype. GUIC-ALS4-P1 to GUIC -ALS4-P4, 4 specimens, 56.2-63.5 mm SL; collected with holotype.

Diagnosis: Alburnoides samiii is distinguished from the other species of Alburnoides in Iran diagnosed above by a combination of the following characters: a terminal mouth, with a tip of the mouth cleft on a level with the middle of the eye or below; a mostly or completely scaled ventral keel; the lack of well-marked spots or dark pigmentation in the lateral line canal; the lack of strong spots or dark outline to the lateral line canal; dorsal fin outer margin truncate to slightly concave; anal fin outer margin markedly concave; a large eye (eye diameter about equal to interorbital width); 2.5-4.2 pharyngeal teeth; a long and shallow caudal peduncle (its length 22.8-26.5, mean 24.5 in SL); a shallow head (head depth 61.2-74.6, mean 68.9 in HL); caudal-fin lobes are pointed, the fin is clearly forked; commonly 8½ branched dorsal fin rays; commonly 12½-14½, branched anal-fin

Fig. 17. Alburnoides samiii sp. n.; Iran, Guilan Province, Sefidroud River, holotype, VMFC-ALS4-H, 75.9 mm SL.

rays; commonly 40-41 total vertebrae; commonly 12-13 predorsal vertebrae; 19-20 abdominal vertebrae and 20-21 caudal vertebrae.

Description: General appearance shown in Figs. 17-19. Morphometric data presented in table 9. Body compressed, upper body profile slightly convex to straight, lower profile convex. The body is relatively shallow (depth at the dorsal fin origin is 26.5-29.7, mean 28.1 in SL). Snout short, slightly to markedly rounded. Mouth terminal, slightly curved or straight, tip of the mouth cleft on a level with the middle of the eye or below. Junction of lower jaw and quadrate at vertical through the lower part of anterior pupil margin. Ventral keel between the pelvic and the anal fins is mostly or completely scaled. Pelvic axillary scale developed, its length about three scale rows. Anal-fin origin at vertical through dorsal-fin base end. Dorsal fin located behind vertical through pelvic fin origin. Dorsal fin outer margin truncate to slightly concave; anal fin outer margin markedly concave. Pectoral fin long, commonly reaching pelvic fin insertion, its outer margin slightly convex. Pelvic fin rounded, its tip normally not reaching to anal fin origin, slightly behind anus. Caudal fin clearly forked, lobes slightly to markedly pointed.

Body depth enters standard length 3.4-3.8 times, head length enters 3.8-4.0, predorsal length 1.9-2.1, caudal peduncle depth 8.5-9.7, and caudal peduncle length 4.3-5.2. Orbit width enters head length 2.8-3.5 times, snout length enters 3.6-4.4, and interorbital width 2.8-3.4. Pectoral fin length enters pectoral fin origin to pelvic fin origin distance 1.0-1.2 times, and pelvic fin length enters pelvic fin origin to anal fin origin distance 1.1-1.3 times.

Dorsal fin unbranched rays 3, dorsal fin branched rays 7½ (2), 8½ (46) or 9½ (2) (mean 8.00, standard deviation 0.29), anal fin unbranched rays 3, anal fin branched rays 11(2), 12(18), 13(19), 14(10) or 15(1) (mean 12.80 ± 0.88), branched pectoral fin rays 11(8), 12(19), 13(22) or 14(1) (mean 12.32 ± 0.77), pelvic fin branched rays 6(5) or 7(45) (mean 6.90 ± 0.30). Lateral line complete with 0, 1 or 2 unpored scales at the posterior end of the lateral series; lateral line scales to posterior margin of hypurals 47(3), 48(19), 49(14), 50(7), 51(5) or 52(2) (mean 48.96±1.24); scales above lateral line to dorsal fin origin 8(3), 9(35) or 10(12) (mean 9.18±0.52); scales below lateral line to anal fin origin 3(8), 4(38) or 5(4) (mean 3.92 ± 0.49); and scales below lateral line to pelvic fin origin 3(3), 4(43) or 5(4) (mean 4.02 ± 0.38). Total scale radii 15(4), 16(14), 17(14), 18(16) or 19(2) (mean 16.96 ±

Fig. 18. Alburnoides samiii sp. n.; Iran, Guilan Province, Sefidroud River, paratype; a, VMFC-ALS4-P2, 75 mm SL; b, VMFC-ALS4-P3, 76 mm SL; c, VMFC-ALS4-P4, 68 mm SL; d, VMFC-ALS4-P5, 53 mm SL.

Table 9. Morphometric data of Alburnoides parhami sp. n. (holotype VMFC-ALP3-H, paratypes: VMFC-ALP3-P, n = 45; GUIC-ALP3-P, n = 4), and A. samiii sp. n. (holotype VMFC-ALS4-H, paratypes: VMFC-ALS4-P, n = 45; GUIC-ALS4-P, n = 4). H, holotype.

1.05). Total gill rakers at outer row on first left arch number 7(2), 8(19), 9(25) or 10(4) (mean 8.62±0.70). Total vertebrae including four Weberian vertebrae and last hypural complex centrum 39(1), 40(32) or 41(17) (mean 40.32 ± 0.51).  Twelve  to 13  predorsal  vertebrae;  19-20

abdominal vertebrae, 20-21 caudal vertebrae; caudal vertebral region most commonly equal to the abdominal region or one vertebrae longer than it; most common vertebrae formulae 19 + 20, 19 + 21, 20 + 20, or 20 + 21. Pharyngeal teeth 2.5-4.2 in 10 examined specimens.

Colouration: In living specimens, overall colouration is silvery, with the bases of the pectoral, pelvic and anal fins orange. Dorsum and top of head are light to dark grey, with an olive hue. The lower portion of head and body are pearly-white. Facial bones and opercle silvery.

The flanks above lateral line may have golden and/or cobalt-blue hues. Faint yellow spots occur in rows along the flanks. Faint dark pigmentation on flank scales above the lateral line forming diffuse stripes, and present only as scattered dark pigmentation below it. Dorsal, anal and caudal fins either with some grey pigmentation or dark grey. Pigmentation of preserved specimens fixed in 10% formalin and stored in 70% alcohol (Figs. 17 & 18) overall tan to cream, darker dorsally; horizontal rows of dark blotches formed by dark pigmentation concentrated on middle of scales, moderately conspicuous, above the lateral line; lateral line with some scales with pores outlined with dark pigmentation, especially the anterior-most scales; a relatively thick dark stripe at caudal peduncle; narrow dark mid-lateral stripe along lateral septum, less discernible from vertical through dorsal fin onwards; fins mostly hyaline, with some black pigmentation lining dorsal and

Fig. 19. Alburnoides samiii sp. n., none-types, above: 67 mm SL; below: 59 mm SL.

caudal fins rays, dorsal-most pectoral-fin rays and anterior-most anal fin rays.

Distribution and notes on biology: Alburnoides samiii is known only from Sefidroud River drainage in the south-eastern Caspian Sea basin (Fig. 1). The upstream portion of Sefidroud River at the type locality (Fig. 20) at time of the collection of the type specimens had clear water, water flow was medium, the stream width was about 5-15 m and maximum depth was up to 1.5 m, and the stream bed was rocky and gravel. Other species collected syntopically were Capoeta gracilis (Keyserling, 1861), Luciobarbus mursa (Guldenstadt, 1773), Barbus lacerta Heckel (1843),Luciobarbus capito (Guldenstadt, 1773), Squalius cf. orientalis (Nordmann, 1840), Cobitis keyvani Mousavi-Sabet et al., 2012, Oxynoemacheilus bergianus (Derzhavin, 1934) and Ponticola iranicus Vasil′eva, Mousavi-Sabet, Vasil′ev, 2015.

Etymology: The species name samiii is in honor of “Professor Doctor Majid Samii” the world famous Iranian neurosurgeon and medical scientist. He is currently the President of the International Neuroscience Institute (INI), also in 2014 he was named the world top neurosurgeon garnered Golden Neuron Award by World Academy of Neurological Surgery. Professor Majid Samii born (19 June 1937) in Rasht, the capital city of Guilan Province, the region where the type locality (Sefidroud River) of the new species is located.

Comparative remarks: Alburnoides samiii differs from A. eichwaldii, A. holciki, A. namaki, A. nicolausi, A. parhami and A. tabarestanensis by its completely or almost completely scaled ventral keel (vs. most commonly 2/3 or completely scaleless ventral keel). Alburnoides samiii is further distinguished fromA. holciki, A. parhami and A. qanati by having a rounded snout (vs. pointed), and a subterminal mouth with the tip of the mouth cleft on a level with the lower margin of the pupil to below the lower margin of the eye (vs. almost or completely terminal with a tip of the mouth on a level with the upper half of the pupil).

Alburnoides samiii is further distinguished from A. coadiby having 12–13 predorsal vertebrae (vs. 13–14); shallower head (head depth 61.2-74.6, mean 68.9 in HL, vs. 67.1-80.7, mean 75.5 in HL); shallower body (body depth 26.5-29.7, mean 28.1 in SL, vs. 28.9-33.8, mean 30.8 in SL); eye located in the mid of head vs. in anterior half (postorbital distance 40.8-53.0, mean 45.9 in HL, vs. 47.6-56.4, mean 52.4 in HL); and orbit width about equal to interorbital width (vs. smaller).

Alburnoides samiii is further distinguished from A. eichwaldii by having a larger eye (eye diameter about equal to interorbital width vs. significantly smaller than interorbital width); commonly 40-41, median 40 total vertebrae (vs. 41-42, median 41); commonly 12-13, median 13 predorsal vertebrae (vs. 13-14, median 14); commonly 19-20, median 20 abdominal vertebrae (vs. 20-21, median 21); less scales above lateral line to dorsal fin origin (commonly 9-10 vs. 11); less scales below lateral line to anal fin origin (typically 4 vs. 7); precaudal and caudal vertebrae equal in number or precaudal 1-2 vertebrae longer than it vs. the caudal region is commonly one vertebrae shorter than the abdominal region; longer head (HL 25.1-26.5, mean 25.8 in SL, vs. 19.6-25.0, mean 21.2 in SL); and shallower head (head depth 61.2-74.6, mean 68.9 in HL, vs. 79.1-92.7, mean 87.6 in HL).

Alburnoides samiii is further differs from A. holciki by having fewer anal fin branched rays (commonly 12½-14½, median 13½ vs. 14½-15½, median 15½); less numerous total vertebrae (commonly 40-41, median 40 vs. 41-42, median 41); shorter anal fin base (15.5-18.3, mean 17.0 in SL, vs. 17.4-21.8, mean 19.4 in SL); and the tip of the mouth cleft commonly at level of the lower margin of the eye or below (vs. at level with the upper half of the pupil).

Alburnoides samiii is further distinguished from A. idignensis by having more anal fin branched rays (commonly 12½-14½, median 13½ vs. 11½–12½, median 11½); 40-41, median 40 total vertebrae (vs. 38-40, median 39); 47-52 lateral line scales (vs. 41-46 lateral line scales); and total scale radii 16-19, median 18 (vs. 12-17, median 15).

Alburnoides samiii is further distinguished from A. namaki by having less branched anal fin rays (commonly 12½-14½, median 13½ vs. 14½-15½, median 15½); smaller scales (commonly 47-52, median 48 lateral line scales, vs. 46-49, median 47); more numerous gill rakers (commonly 8-9 vs. 6-8), scales above lateral line to dorsal fin origin (commonly 9-10 vs. 10-11); longer medial fins; longer caudal peduncle (caudal peduncle length 19.3-24.3, mean 21.9 in SL, vs. 14.9-22.3, mean 18.1 in SL); and shallower head (head depth 61.2-74.6, mean 68.9 in HL, vs. 68.0-82.6, mean 78.1 in HL). Alburnoides samiii is further differs from A. nicolausi by having more branched dorsal-fin rays (commonly 8½ vs. 7½); more branched anal-fin rays (12½-14½ vs. 9½-11½); more gill rakers (8-9 vs. 7-8); smaller scales (commonly 47-52, median 48 lateral line scales, vs. 42-50, median 44); total scale radii 15-19, median 18 (vs. 13-17, median 15); and completely or almost completely scaled ventral keel (vs. most commonly 2/3 or completely scaleless ventral keel).

Alburnoides samiii is further distinguished from A. petrubanarescui by having more branched dorsal-fin rays (commonly 8½ vs. 7½); more branched anal-fin rays (commonly 12½-14½ vs. 9½-10½); commonly 12-13 predorsal vertebrae (vs. 13-14); commonly 19-20, median 20 abdominal vertebrae (vs. 20-22, median 21); commonly 19-20 caudal vertebrae (vs. 20-21); and more gill rakers (8-9 vs. 7-8).

Fig. 20. Sefidroud River, the southern Caspian Sea basin; type locality of A. samiii sp. n.