Diagnosis.

Medium-sized cyprinids with an ossified, smooth last unbranched dorsal-fin ray; 9 or 10 branched dorsal-fin rays and 6 branched anal-fin rays; large, shield-shaped scales with numerous parallel radii; the lateral line with 25 to 39 scales; the pharyngeal teeth hooked at their tips, their count being 2.3.5-5.3.2 or 2.3.4-4.3.2; 1 or 2 pairs of barbels present.

Carasobarbus species are evolutionarily hexaploid (Machordom and Doadrio 2001, Gorshkova et al. 2002, Leggatt and Iwama 2003, Tsigenopoulos et al. 2010).

Remarks and discussion.

‘Barbus’ grypus, Mesopotamichthys sharpeyi and ‘Barbus’ reinii from the Middle East have five branched rays in the anal fin. The hexaploid species from Africa (Labeobarbus and Varicorhinus), which are the sister group to Carasobarbus and the other species from the Middle East (Tsigenopoulos et al. 2010, KB unpublished), have five branched rays in the anal fin. The Asian species (Tor and Neolissochilus) are sister group to the species from Africa and the Middle East (Tsigenopoulos et al. 2010, KB unpublished) and have five branched rays in the anal fin. By application of the parsimony principle the possession of six branched anal-fin rays is a synapomorphy of the genus Carasobarbus. The possession of nine to 10 branched rays in the dorsal fin is synapomorphic for Carasobarbus, because the closely related Middle-Eastern species ‘Barbus’ grypus, Mesopotamichthys sharpeyi and ‘Barbus’ reinii as well as many African hexaploids have the plesiomorphic state of eight branched rays in the dorsal fin. However, in some African species the number of branched dorsal-fin rays is increased convergently. These two synapomorphies establish Carasobarbus as a monophyletic group. Analyses of the mitochondrial cytochrome b gene confirm the monophyly of this genus (Durand et al. 2002, Tsigenopoulos et al. 2010, KB unpublished data). Colli et al. (2009) found Carasobarbus to be monophyletic in their maximum likelihood analysis but not in their neighbour joining analysis. ‘Barbus’ grypus Heckel, 1843 is the sister taxon of the genus Carasobarbus (Tsigenopoulos et al. 2010).

Out of the generic names Barbellion, Barbus, Barynotus, Capoeta, Carasobarbus, Cyclocheilichthys, Kosswigobarbus, Labeobarbus, Luciobarbus, Pseudotor, Puntius, Systomus, Tor, and Varicorhinus that were used for this taxon − or its parts − by previous authors, only Carasobarbus, Kosswigobarbus and Pseudotor are available for the genus in question. All other generic names have not been considered, because their type species are not closely related to the species under discussion here (Durand et al. 2002, Tsigenopoulos et al. 2010, KB unpublished data) or do not share the characters mentioned above. Carasobarbus, Kosswigobarbus and Pseudotor are subjective synonyms. They all were established in the same publication (Karaman 1971) and thus none of them has priority. We, acting as First Reviser, select Carasobarbus to have priority in accordance with article 24.2 of the International Code for Zoological Nomenclature (ICZN 2012). Thus Carasobarbus is the valid name for this genus.

Within the genus, several species share characters that are potentially synapomorph and elucidate sister group relations. Carasobarbus fritschii and Carasobarbus harterti both have pharyngeal bones with four teeth in the medial row. This character is probably synapomorph, because all other congeners have five teeth in the medial row. This group corresponds to Pseudotor. Carasobarbus kosswigi and Carasobarbus sublimus share the possession of a spatulate lower jaw and a median lobe on the lower lip. The spatulate lower jaw is synapomorph, because no congener and no other closely related species shares this character. The close phylogenetic relationship between Carasobarbus kosswigi and Carasobarbus sublimus is confirmed by genetic analysis (Borkenhagen et al. 2011). These two species correspond to Kosswigobarbus.

Material.

Type material. Holotype of Barbus apoensis: BMNH 1976.4.7:166, Saudi Arabia, permanent stream near Khamīs Mushayt (18°17'N, 42°34'E), F. Tippler, 12 Dec 1968.

Paratypes of Barbus apoensis: BMNH 1976.4.7:167-171, 5, same data as holotype. - BMNH 1976.4.7:172-175, 4, Saudi Arabia, upper Wādī Turabah near Aţ Ţā’if (22°56'N, 40°54'E), G. Popov. - BMNH 1971.2.11:1-2, 2, Saudi Arabia, intermittent watercourse in Wādī Adamah (19°53'N, 41°57'E), J. P. Mandaville, 27 Oct 1969.

Non-type material. Endorheic darinages. BMNH 1980.7.1:15, 1, Saudi Arabia, Wādī Habayaba between Aţ Ţā’if and Ash Shafā [N21°11’, E 40°24'], A. Farag, 1980. - SMF 30167, 3; SMF 30170, 10 Saudi Arabia, Wādī Būwah (20°45'N, 41°8'E), F. Krupp and W. Schneider, 21 Mar 1990. - SMF 30169, 6; SMF 33147, 4, Saudi Arabia, Wādī Būwah (20°44'N, 41°7'E), F. Krupp and W. Schneider, 21 Mar 1990. - SMF 30168, 6; SMF 30171, 9, Saudi Arabia, Wādī Turabah (20°32'N, 41°17'E), F. Krupp and W. Schneider, 20 Mar 1990.

Streams draining towards the Red Sea. CMNFI 87-0135, 1; CMNFI 87-0137, 4, Saudi Arabia, Wādī Hadīyah (25°34'N, 38°41'E). - SMF 33149, 1, Saudi Arabia, Wādī Ḩaqqaq (22°49'N, 39°22'E), W. Büttiker, 5/6 May 1983. - SMF 33148, 2, Saudi Arabia, Wādī ‘Ilyab (20°5'N, 40°54'E), H. Felemban and J. Gasparetti, 28 Oct 1983. - SMF 33539, 3, Saudi Arabia, Wādī ‘Ilyab (20°7'N, 40°57E), W. Büttiker, 10−11 Nov 1983.

Unknown drainage system. SMF 33146, 4, Saudi Arabia, Al Ḩijāz, W. Büttiker.

Diagnosis.

One pair of barbels, usually 10 branched rays in the dorsal fin, 27 to 32 scales in the lateral line, usually 12 scales around the least circumference of the caudal peduncle, last unbranched ray of dorsal fin shorter than head.

Description.

The body depth is comparatively low and a nuchal hump is present in adults but not developed in juveniles. The height of the caudal peduncle is relatively low (Table 1). The dorsal and ventral fins are usually positioned behind the middle of the body. The head is elongate with a straight or slightly concave dorsal profile. The ventral profile of the head is slightly convex. (Figs 1, 2). The head length is about equal to the body depth. The mouth is broad and terminal or slightly sub-terminal with one pair of barbels (Fig. 3, Table 2). Only one out of 65 specimens had two pairs of barbels and in one specimen a single anterior barbel was present. The eyes are in the anterior half of the head and slightly protuberant. The morphometric characters are summarised in Table 1.

The dorsal fin and its base are rather short. It usually has four unbranched and 10 branched rays (Table 3). The last unbranched ray is considerably shorter than the head (Fig. 4), weakly ossified, and its distal part is flexible. The anal fin has three unbranched and six branched rays (Table 4). Pectoral and ventral fins are relatively short (Table 1).

Carasobarbus apoensis has 27 to 32 scales in the lateral line (Table 5), usually 4.5 scales above the lateral line (Table 6), 3.5 or 4.5 scales below the lateral line (Table 7) and 12 scales around the least circumference of the caudal peduncle (Table 8). The scales are shown in Fig. 5.

The pharyngeal teeth count is 2.3.5- in 12 specimens, -5.3.2 in two specimens and 1.3.5- in one specimen. The pharyngeal teeth are hooked at their tips (Fig. 6).

Live colouration is golden with olive fins. The upper side is darker than the belly (Fig. 2). In ethanol-preserved specimens the upper side is dark, the belly yellow and the fins are grey or yellow (Fig. 1). Juveniles have a dark lateral spot on the caudal peduncle.

The maximum length observed in the material examined is 288 mm SL.

Carasobarbus apoensis differs from all congeners, except Carasobarbus luteus, by having one rather than two pairs of barbels. For a comparison with Carasobarbus luteus populations see below.

Distribution. Carasobarbus apoensis occurs in the Al Ḩijāz mountain range in wadis draining either inland or towards the Red Sea (Fig. 7). It is endemic to Saudi Arabia.

Habitats and biology.

This species inhabits the upper courses of wadis, which are characterised by strong seasonal fluctuations in water levels, temperature and other physiochemical parameters.

Conservation status.

Carasobarbus apoensis is rated Least Concern and still occurs in large numbers, but abstraction of large specimens by recreational fishing, water abstraction and habitat loss might become problematic for this species (BCEAW 2002).

Remarks and discussion.

Carasobarbus apoensis was originally described from Khamīs Mushayt, Wādī Turabah and Wādī Adamah as a member of the genus Barbus (Banister and Clarke 1977). It was later transferred to the genus Carasobarbus (Ekmek-çi and Banarescu 1998). Alkahem and Behnke (1983) reported an unknown Barbus and tentatively considered these specimens to be atypical Carasobarbus apoensis. We did not find any evidence of an undescribed Carasobarbus species that occurs sympatrically with Carasobarbus apoensis, thus we agree with their conclusion.

Carasobarbus apoensis is very closely related to Carasobarbus luteus (KB unpublished data).

Material.

Type material. Lectotype of Barbus canis: MNHN 0000-1413, 1, Jordan River [31°46'N, 35°33'E], Bové, 1833 (designated by Krupp and Schneider 1989).

Paralectotype of Barbus canis: MNHN 0000-3944, 1, same data as lectotype.

Holotype of Barbus beddomii: BMNH 1863.11.3:5, 1, Lake Tiberias [32°48'N, 35°35'E], T. W. Beddome.

Non-type material. Jordan River Drainage. SMF 14075, 2, Lake Tiberias (32°48'N, 35°35'E), M. Goren, 15 Mar 1968. - SMF 33134, 16, Syria, Nahr al Yarmūk near Jallayn (32°44'21"N, 35°58'56"E), N. Alwan et al., 16 Oct 2008. - SMF 24464, 1, Jordan, Nahr al Yarmūk near Maqārin (32°43'N, 35°53'E), F. Krupp and W. Schneider, 23 Sep 1985. - SMF 30175, 11, Syria, Lake Muzayrīb [32°42'40"N, 36°1'39"E], F. Krupp and W. Schneider, 12 Apr 1989. - SMF 33135, 17, Jordan, Wadi al-‘Arab near the dam (32°37'6"N, 35°37'46"E), N. Alwan et al., 25 Oct 2008. - SMF 17123, 16, Wādī al Yābis (32°24'N, 35°36'E), F. Krupp and W. Schneider, 23 Jul 1980. - ZMH H 2343, 3, Jordan, Wādī Kufrinjah (32°16'25"N, 35°33'42"E). - SMF 24344, 3; SMF 24345, 17, Jordan, Nahr az Zarqā’ (32°12'N, 35°50'E), F. Krupp and W. Schneider, 22 Jul 1980. - SMF 24339, 3, Jordan, Nahr az Zarqā’ (32°10'N, 35°37'E), F. Krupp and W. Schneider, 21 Jul 1980. - SMF 24340, 1, Jordan, Nahr az Zarqā’ near Sadd al Malik Talal (32°10'N, 35°49'E), F. Krupp and W. Schneider, 22 Jul 1980. - SMF 24331, 7; SMF 24346, 3, Jordan, Nahr al Yarmūk channel (32°08'N, 35°36'E), F. Krupp and W. Schneider, 21 Jul 1980. - NMW 53961, 1, Jordan River [31°46'N, 35°33'E], Cenoni, Dec 1898.

Azraq Oasis. BMNH 1956.2.24:15-16, 2; BMNH 1965.11.24:2, 1, Jordan, wetland near Azraq ash Shīshān [31°50'N, 36°49'E].

Coastal rivers of the Mediterranean Sea. BMNH 1949.9.16:124, 1, Israel, Naẖal Na‘aman [32°54'42"N, 35°4'50"E]. - NMW 22367, 1, Israel, Naẖal Na‘aman [32°54'42"N, 35°4'50"E], H. Steinitz, 21 Oct 1955. - SMF 9229, 1, Israel, Naẖal Yarqon [32°6'7"N, 34°46'32"E].

Diagnosis.

Two pairs of barbels, 29 to 35 scales in the lateral line and usually 12 scales around the least circumference of the caudal peduncle, last unbranched ray of dorsal fin shorter than head.

Description.

The body is low. A nuchal hump is present in adults but absent in juveniles. The largest body depth is at the origin of the dorsal fin. The head is long, rather low and fairly narrow with straight dorsal and convex ventral profile (Figs 8, 9). The head length approximately equals the body depth. The mouth is terminal or slightly subterminal. Two pairs of barbels are present (Table 2). The lips are smooth and thin (Fig. 3). The eyes are at the end of the anterior half of the head. The morphometric characters are summarised in Table 1.

Pectoral, ventral, dorsal and anal fins are comparatively short (Table 1). The dorsal fin usually has four unbranched and 10 branched rays (Table 3). The last unbranched ray is ossified and its distal part is flexible. It is usually markedly shorter than the head (Fig. 4). The anal fin usually has three unbranched and six branched fin rays (Table 4).

There are 29 to 35 scales in the lateral line (Table 5), usually 4.5 or 5.5 scales above the lateral line (Table 6), usually 4.5 scales below the lateral line (Table 7) and usually 12 scales around the least circumference of the caudal peduncle (Table 8). The scales are shown in Fig. 5.

The pharyngeal teeth count is 2.3.5-5.3.2 in 23 specimens, 2.3.3-5.3.2 in one specimen, 2.3.5- in one specimen and -5.3.2 in one specimen. The pharyngeal teeth are hooked at their tips (Fig. 6).

Live specimens are silvery to bronze coloured. The posterior third of the body and the fins are distinctly yellow in many specimens (Fig. 9). Ethanol-preserved specimens are brownish yellow and the back is only slightly darker than the rest of the body (Fig. 8). The fins are brownish yellow. Juveniles have a dark lateral spot on the caudal peduncle.

Carasobarbus canis differs from Carasobarbus apoensis and Carasobarbus luteus in having two pairs of barbels vs. one, from Carasobarbus kosswigi and Carasobarbus sublimus in having a crescent-shaped lower lip without median lobe vs. a spatulate lower lip with median lobe, from Carasobarbus exulatus in modally having 10 branched dorsal-fin rays vs. nine and from Carasobarbus chantrei, Carasobarbus fritschii and Carasobarbus harterti in modally having 10 scales around the least circumference of the caudal peduncle vs. 14 or 16.

Distribution.

Carasobarbus canis occurs in the Jordan River system (Fig. 7). There are only few records from coastal rivers of the Mediterranean Sea (Naẖal Na‘aman and Naẖal Yarqon). A recent treatment of the inland water fish communities of Israel does not report Carasobarbus canis from coastal rivers (Goren and Ortal 1999). The population in the Azraq Oasis was introduced by humans (Krupp and Schneider 1989). Since the year 2000 this species was not found in Azraq (Hamidan 2004) and the population may have disappeared due to drought. Records from the Tigris-Euphrates system (Banister 1980) are based on misidentifications.

Habitats and biology.

Carasobarbus canis inhabits a wide range of rivers, lakes and ponds (Goren 1974) with clean as well as polluted water (Mir 1990). Adults reach a length of about 40 cm (max. 66 cm) and are of economic importance locally (annual catch in Israel 1970-85 about 50 t, Fishelson et al. 1996). It feeds on fish, aquatic invertebrates, algae and detritus (Ben-Tuvia 1978, Spataru and Gophen 1985, Krupp and Schneider 1989). The relative proportion of fish in the diet increases with body length and small cyprinids of the genus Mirogrex are their most important prey (Spataru and Gophen 1985). The spawning grounds are (among others) at the shore of Lake Tiberias where the spawning occurs in shallow water over hard bottom in December and January, one month after the start of the rainy season (Fishelson et al. 1996). The sticky eggs attach to the substrate. Winter spawning is seen as evidence for an origin in cooler areas (Fishelson et al. 1996).

Conservation status.

Catches in Lake Tiberias are declining (Fishelson et al. 1996). The species is rated Least Concern by the IUCN (Crivelli 2006a). The population in Lake Tiberias does not face serious threats; the riverine populations are declining and threatened by pollution, water extraction, drought and fragmentation due to damming (Crivelli 2006a).

Remarks and discussion.

Carasobarbus canis was described from the Jordan River as a member of the genus Barbus (Cuvier and Valenciennes 1842). Later it was assigned to Luciobarbus (Heckel 1843), and Labeobarbus (Günther 1864). Subsequently it was transfered back to Barbus (Günther 1868) and then placed in Tor (Karaman 1971, Banarescu 1977), Barbus (Banister and Clarke 1977, Krupp 1983a), Carasobarbus (Ekmekçi and Banarescu 1998) and Barbus subgenus Carasobarbus (Tsigenopoulos et al. 2010). MNHN 0000-1413 was designated as lectotype (Krupp and Schneider 1989). Barbus beddomii is considered to be a junior synonym of Carasobarbus canis (Berg 1949, Karaman 1971, Krupp and Schneider 1989).

Material.

Type material. Lectotype of Labeobarbus chantrei: MNHN A-3866, Turkey, Amik Gölü [36°12'24"N, 36°9'26"E], H. Chantre, 1881 (designated by Krupp 1985a).

Paralectotypes of Labeobarbus chantrei: MNHN A-3937, 1, same data as lectotype. - MNHN A-3938, 2; MNHN A-3939, 3; MNHN A-3940, 1, Syria, Ḩamāh [35°9'0"N, 36°43'59"E], H. Chantre, 1881.

Non-type material. Orontes River drainage. MNHN B-2977, 1, Syria, Orontes, A. Gruvel, 1829. - BMNH 1934.1.25:4, 1, Syria, Orontes. - FSJF 2311, 11, Turkey, Karasu Çayı below dam of Tahtaköprü Barajı (36°51'7"N, 36°41'10"E), M. Özulug and J. Freyhof, 7 Nov 2007. - SMF 17115, 8, Turkey, Orontes, 8 km E of Hatay (36°17'N, 36°11'E), J. Winkler and B. Koster, 20 Sep 1982. - CMNFI 88-0019, 1, Turkey, 8 km southwest of Hatay (36°11'N, 36°3'E). - SMF 17110, 4, Turkey, tributary to Orontes (36°11'N, 36°3'E), F. Krupp, 23 Aug 1978. - SMF 17122, 2, Turkey, 2 km southeast of Samandağı (36°6'N, 35°58'E), F. Krupp, 23 Aug 1978. - FSJF uncatalogued, 16, Turkey, at Sinanlı (36°5'51"N, 36°4'43"E), M. Özulug and J. Freyhof, 8 Nov 2007. - SMF 33130, 40, Syria, near Mashra’a el Būz (35°57'3"N, 36°23'45"E), N. Alwan et al., 8 Oct 2008. - SMF 33131, 58, Syria, ‘Ayn az Zarqa (35°56'40"N, 36°24'9"E), N. Alwan et al., 8 Oct 2008. - SMF 17107, 1, Syria, Jisr ash Shughūr (35°48'N, 36°19'E), F. Krupp, 20 Aug 1980. - SMF 17109, 2, Syria, main bridge at Jisr ash Shughūr (35°48'N, 36°19'E), F. Krupp, 19 Aug 1978. - CMNFI 88-0018, 4, Syria, ‘Ayn Zaqa (35°27'N, 36°23'E). - SMF 17114, 1; SMF 17121, 7, Syria, ‘Ayn Zaqa (35°27'N, 36°23'E), F. Krupp, 25–27 Mar 1979. - BMNH 1968.12.13:188-190, 3, Syria, spring lake at Qal‘at al Maḑīq [35°25'N, 36°23'E]. - SMF 17120, 7, Syria, aquaculture pond near Qal‘at al Maḑīq (35°25'N, 36°23'E), F. Krupp, 8 Aug 1978. - SMF 33132, 5, Syria, stream at Qal‘at al Jarras (35°19'49"N, 36°18'38"E), N. Alwan et al., 12 Oct 2008. - SMF 17111, 6, Syria, ‘Ašārna (35°17'N, 36°19'E), F. Krupp, 11 Aug 1978. - SMF 17117, 5, Syria, near Shayzar (35°16'N, 36°34'E), F. Krupp, 27 Mar 1979. - SMF 24349, 3, Syria, Shayzar (35°16'N, 36°34'E), F. Krupp and W. Schneider, 17 Aug 1980. - SMF 17118, 1, Syria, 200 m below western outlet of Lake Homs (34°40'N, 36°37'E), F. Krupp and W. Schneider, 3 Aug 1978. - SMF 17119, 5, Syria, western outlet of Lake Homs (34°40'N, 36°37'E), F. Krupp and W. Schneider, 3 Aug 1978. - SMF 33133, 24, Syria, Lake Homs at Qaţţīnah (34°39'43"N, 36°37'6"E), N. Alwan et al., 13 Oct 2008.

Mediterranean coastal rivers. SMF 31669, 1; SMF 31670, 1, Syria, Nahr Marqīyah (35°1'50"N, 35°54'18"E), N. Alwan et al., 10 Oct 2008.

Tigris-Euphrates system. SMF 12966, 1, Turkey, Balıklıgöl at Şanlıurfa [37°8'52"N, 38°47'4"E], L. Lortet, 1884.

Diagnosis.

Two pairs of barbels, 31 to 38 scales in the lateral line and usually 14 to 16 scales around the least circumference of the caudal peduncle, last unbranched dorsal-fin ray equal to or shorter than head.

Description.

The body is comparatively high-backed and laterally compressed in mid-sized specimens but low-backed and almost cylindrical in large specimens. In large specimens a pronounced nuchal hump is present, in smaller specimens it is only weakly developed or absent. The maximum body depth is at the origin of the dorsal fin. The head is short and blunt with a convex ventral profile and a slightly convex to straight dorsal profile (Figs 10, 11). The mouth is terminal or slightly sub-terminal with two pairs of short barbels (Table 2).The body depth is usually greater than the head length (Fig. 12).The eyes are slightly protuberant and lie at the end of the anterior half of the head. The morphometric characters are summarised in Table 1.

The dorsal fin usually has four unbranched and nine to 11 branched rays (Table 3). The last unbranched ray is ossified but not very thick and flexible in its distal part. It is usually shorter than the head (Fig. 4). The anal fin usually has three unbranched and five or six branched rays (Table 4).

There are 31 to 38 scales in the lateral line (Table 5), 4.5 to 6.5 scales above the lateral line (Table 6), four to six scales below the lateral line (Table 7) and 12 to 16 scales around the least circumference of the caudal peduncle (Table 8). The scales are shown in Fig. 5.

The pharyngeal teeth count is 2.3.5-5.3.2 in two specimens, 2.3.5- in 11 specimens, -5.3.2 in two specimens and 1.3.5- in one specimen. The pharyngeal teeth are hooked at their tips (Fig. 6).

Small live specimens are silvery; larger specimens are silvery or bronze coloured and sometimes have yellow pectoral and ventral fins (Fig. 11). Small ethanol-preserved specimens are silvery with a somewhat darker back and a salmon pink hue. Juveniles have a dark lateral spot on the caudal peduncle. Ethanol-preserved adults are yellow-brown and the back is only slightly darker than the rest of the body (Fig. 10).

The maximum length observed in the material examined is 385 mm SL.

Carasobarbus chantrei differs from Carasobarbus apoensis, Carasobarbus canis, Carasobarbus exulatus, Carasobarbus luteus and Carasobarbus sublimus in having 31 to 38 scales in the lateral line vs. 27 to 32, 29 to 35, 26 to 32, 25 to 33 and 27 to 29 respectively and modally 14 scales around the least circumference of the caudal peduncle vs. 12. It differs from Carasobarbus kosswigi and Carasobarbus sublimus in having a crescent-shaped lower lip without median lobe vs. a spatulate lower lip with median lobe and from Carasobarbus exulatus, Carasobarbus fritschii and Carasobarbus harterti in modally having 10 branched dorsal-fin rays vs. nine.

Distribution.

Carasobarbus chantrei occurs in the Orontes river drainage system (Fig. 7). Two juvenile specimens where collected in Nahr Marqīyah, a coastal river in Syria. This species had never before been reported from this location (Krupp 1985a) and it has most likely been introduced by humans. Two potential records from Nahr Quwayq (MNHN A-3861, MGHN 3554) are discussed in Krupp (1985a, c). Locality data for MHNL 3554 are ambiguous (Krupp 1985a). The locality for MNHN A-3861 is given as “Syria, Aleppo” in Krupp (1985a) and considered to be from Nahr Quwayq. The collection database of the MNHN gives “Origine: Syrie, localité: Alep, Milieu: Continent, Bassin hydrologique: Asi, Cours d’eau: Asi” as locality. As these data are contradictory, it is likely that the specimens do not come from the Nahr Quwayq, but from the Orontes (=Asi) and Carasobarbus chantrei does probably not occur in the Nahr Quwayq. A record from the Ceyhan Nehri (Krupp 1985c) is not backed by specimens. Records from the Tigris-Euphrates basin are misidentified Carasobarbus luteus (Krupp 1985a, Krupp and Schneider 1991) and the specimen from Balıklıgöl at Şanlıurfa in Turkey (SMF 12966) is probably mislabelled or was introduced there (Krupp 1985a). It is not included in the map.

Habitats and biology.

Carasobarbus chantrei occurs in a wide range of habitats stretching from stagnant waters of lakes to rapidly flowing river courses.

Conservation status.

Carasobarbus chantrei is utilised as food fish locally but is increasingly replaced by carp (Krupp 1985a). During a field survey in Syria in 2008, the species was still abundant in parts of the Orontes. However, large stretches of this river, especially in the Al Ghāb area, suffer heavily from water abstraction and pollution by sewage and domestic waste and are devoid of fish. The species is rated “Endangered B1ab(ii, iii)” by the IUCN (Crivelli 2006b). The main threat is habitat degradation due to water extraction, pollution and drought (Crivelli 2006b).

Remarks and discussion.

Carasobarbus chantrei was described from the Orontes and placed in Labeobarbus by Sauvage (1882). He transferred it to Barbus two years later (Sauvage 1884). In 1942 Barynotus verhoeffi was described from Amik Gölü, Turkey (Battalgil 1942). Ladiges (1960) erroneously synonymised Barynotus verhoeffi with Carasobarbus canis. Karaman (1971) synonymised Carasobarbus chantrei with Carasobarbus canis and thus transferred it to the genus Tor (sensu Karaman 1971). Fowler (1976) transferred Barynotus verhoeffi to the genus Barbellion. In 1985 Krupp redescribed Carasobarbus chantrei as a valid species and provisionally placed it into the genus Barbus sensu lato. He found the type series to be inhomogeneous (MNHN B-2889 are ‘Barbus’ grypus) and designated MNHN A-3866 as lectotype of Carasobarbus chantrei (Krupp 1985a). The ‘Catalog of Fishes’ does not list MNHN B-2889 as types for Carasobarbus chantrei (Eschmeyer 2011). Ekmekçi and Banarescu (1998) transferred the species to the genus Carasobarbus. Tsigenopoulos et al. (2010) used Barbus subgenus Carasobarbus.

Material.

Type material. Holotype of Barbus exulatus: BMNH 1976.4.7:299, Yemen, Wādī Ḩaḑramawt at Qasam (16°10'N, 49°4'E), W. A. King-Webster.

Paratypes of Barbus exulatus: BMNH 1976.4.7:308, 1; BMNH 1976.4.7:300-307, 8, same data as holotype. - BMNH 1976.4.7:328-329, 2; BMNH 1976.4.7:330-331, 2, Yemen, Wādī ‘Idim/Wādī Ḩaḑramawt at Ghuraf (16°0'N, 49°0'E), W. A. King-Webster. - BMNH 1976.4.7:309, 1; BMNH 1976.4.7:310-318, 9; BMNH 1976.4.7:319-327, 9, Yemen, Wādī Ḩaḑramawt at Ghayl ‘Umar (15°44'N, 48°51'E), W. A. King-Webster. - BMNH 1976.4.7:332-333 probably Wādī Marrān in Wādī Aḩwar system [13°53'51"N, 46°05'14"E], G. Popov, 2 Aug 1962.

Non-type material. Wādī Ḩaḑramawt/al Masīlah drainage. BMNH 1976.5.17:9-10, 2, Yemen, Wādī al Khūn (16°10'N, 49°10'E). - SMF 33108, 10, Yemen, Wādī al Khūn (16°9'51"N, 49°6'2"E), F. Krupp et al., 3 Jun 2005. - SMF 33109, 17, Yemen, Wādī al Khūn (16°9'45"N, 49°4'46"E), F. Krupp et al., 3 Jun 2005. - SMF 33110, 14, Yemen, Wādī al Masīlah near Fughmah (16°8'36"N, 49°27'7"E), F. Krupp et al., 4 Jun 2005. - SMF 33111, 1, Yemen, Wādī al Masīlah at al Hind (15°44'53"N, 50°24'32"E), F. Krupp et al., 5 Jun 2005. - SMF 33106, 8, Yemen, Wādī ‘Idim at Ghayl ‘Umar near Arḑ ar Raydah (15°40'51"N, 48°51'59"E), F. Krupp et al., 2 Jun 2005. - SMF 33107, 11, Yemen, Wādī ‘Idim near Ghayl ‘Umar (15°40'10"N, 48°51'4"E), F. Krupp et al., 2 Jun 2005. -SMF 33105, 13, Yemen, Wādī Mara in Wādī Daw‘an system (15°8'36"N, 48°26'58"E), F. Krupp et al., 31 May 2005.

Diagnosis.

Dorsal fin with 9 branched rays in most specimens; last unbranched ray of dorsal fin as long as or longer than head; 2 pairs of barbels; 26 to 32 scales in the lateral line and usually 12 scales around the least circumference of the caudal peduncle.

Description.

The body is not particularly high backed and the maximum body depth is at the origin of the dorsal fin or slightly in front of it (Fig. 13). A nuchal hump is present in adult specimens (Fig. 14) but absent in juveniles (Fig. 15). The caudal peduncle is slender. The head profile is convex ventrally and straight dorsally. The body depth is about the same as the head length (Fig. 12). In specimens below 100 mm SL, the head is rather narrow, in larger specimens it becomes wider. The mouth is subterminal and comparatively narrow. Two pairs of barbels are present (Table 2), the posterior one is rather long. The eyes are at the end of the anterior half of the head and slightly protuberant. The morphometric characters are summarised in Table 1.

The dorsal fin is long and usually has four unbranched and eight to 10 branched rays (Table 3). The last unbranched ray is strongly ossified and only the tip is flexible. Its length is about the same as the head length (Fig. 4). The anal fin is long, usually has three unbranched and five or six branched rays (Table 4).

There are 26 to 32 scales in the lateral line (Table 5), 4 to 5.5 scales above the lateral line (Table 6), 3.5 to five scales below the lateral line (Table 7) and 10 to 12 scales around the least circumference of the caudal peduncle (Table 8). The scales are shown in Fig. 5.

The pharyngeal teeth count is 2.3.5-5.3.2 in one specimen, 2.3.5- in 16 specimens, -5.3.2 in one specimen and 2.3.4- in one specimen. The pharyngeal teeth are hooked at their tips (Fig. 6).

In live specimens and freshly preserved specimens the back and the sides are grey to golden, the belly is yellowish white and the fins are sometimes golden to orange (Fig. 15). Preserved specimens have a dark back and a lighter belly, the fins are whitish or greyish. Juveniles have a dark spot on the sides of the caudal peduncle.

The maximum length observed in the material available is 288 mm SL.

Carasobarbus exulatus differs from all congeners, except Carasobarbus fritschii and Carasobarbus harterti in modally having nine instead of 10 branched dorsal-fin rays. It differs from Carasobarbus fritschii and Carasobarbus harterti in modally having 12 scales around the least circumference of the caudal peduncle vs. 16 and in having 26 to 32 scales the lateral line vs. 30 to 39 and 31 to 38 respectively.

Distribution.

This species is endemic to Yemen and occurs in Wādī Ḩaḑramawt / Wādī al Masīlah and its pleistocene tributaries (Banister and Clarke 1977, Krupp 1983a, Fig. 7). It is also known from Sadd Ma’rib (Al-Safadi 1995), a dam lake at 15°23'46"N, 45°14'37"E and Wādī Ḩajr (14°02'42"N, 48°40'27"E), where they are “found throughout the whole year and are distributed all over the stream” (Attaala and Rubaia 2005).

Locality data for BMNH 1976.4.7:332-333 is given as “Wadi Maran, E. Yemen” (Banister and Clarke 1977), which is most likely Wādī Marrān [13°53'51"N, 46°05'14"E], representing the westernmost record of this species that is backed by specimens.

Habitats and biology.

The biology of this species is mostly unknown.

Conservation status.

During a field expedition in 2005 one of the authors saw large, continuous water bodies in the Wādī Ḩaḑramawt / Wādī al Masīlah area. The species is rated as “Endangered B1a, b; B2a, b” and water extraction is identified as the main threat (BCEAW 2002).

Discussion.

Carasobarbus exulatus was described from Wādī Ḩaḑramawt and Wādī Maran in Yemen and placed in Barbus (Banister and Clarke 1977). Later it was transferred to Carasobarbus (Ekmekçi and Banarescu 1998).

Material.

Type material. Syntypes of Barbus fritschii: BMNH 1874.1.30:27-31, 5, Morocco, Oued Ksob in Oued Igrounzar drainage [31°28'59"N, 9°46'3"W], K. v. Fritsch and J. Rein, 1872.

Syntypes of Barbus paytonii: BMNH 1903.10.29:17-20, 7, Morocco, Oued Oum er Rbia [33°19'40"N, 8°20'2"W], F. W. Riggenbach.

Syntypes Barbus riggenbachi: BMNH 1902.7.28:20-21, 2, Morocco, Oued Oum er Rbia [33°19'40"N, 8°20'2"W], F. W. Riggenbach. - BMNH 1902.7.28:19, 1, Morocco, Oued Talmest [31°52'15"N, 9°18'31"W], F. W. Riggenbach.

Syntypes Barbus rothschildi: BMNH 1901.7.26:6-7, 2, Morocco, Oued Oum er Rbia [33°19'40"N, 8°20'2"W], E. Hartert.

Syntypes of Capoeta atlantica: BMNH 1902.1.4:18-19, 2, Morocco, Oued Nfis at Trigadir-el-hor (Tagadirt n’Bour?) [31°9'21"N, 8°6'2"W], E. G. B. Meade-Waldo.

Syntypes of Capoeta waldoi: BMNH 1902.1.4:16-17, 2, Morocco, Oued Nfis at Trigadir-el-hor (Tagadirt n’Bour?) [31°9'21"N, 8°6'2"W], E. G. B. Meade-Waldo.

Non-type material. Oued al Maleh drainage. SMF 33412, 5; SMF 33510, 1; SMF 33511, 1; SMF 33512, 1, Morocco, Oued al Maleh above the dam (33°33'53"N, 7°22'3"W), K. Borkenhagen and J. Freyhof, 19 Apr 2011. - MNHN 1919-0365, 1; MNHN 1919-0366, 1, Morocco, Oued Bou Asseïla near Chaouia [33°19'34"N, 7°16'46"W], H. Millet, 1919.

Oued Bou Regreg drainage. MNHN 1939-0124, 1, Morocco, Oued Akrech [33°56'7"N, 6°47'41"W], J. M. Pérès, 1939. - SMF 33411, 10; SMF 33503, 1; SMF 33504, 1; SMF 33505, 1, Morocco, Oued Korifla above the dam lake (33°44'0"N, 6°43'43"W), K. Borkenhagen and J. Freyhof, 18 Apr 2011.

Oued Igrounzar drainage. BMNH 1889.7.19:9, 1, Morocco, near Essaouira [31°30'45"N, 9°46'12"W], C. Payton. - SMF 636, 4; SMF 952, 6, Morocco, Oued Ksob [31°28'59"N, 9°46'3"W], K. v. Fritsch and J. Rein, 1872. - SMF 33405, 19; SMF 33446, 1; SMF 33450, 1; SMF 33451, 1, Morocco, Oued Ksob near Essaouira (31°28'0"N, 9°45'32"W), A. Azeroual et al., 11 Apr 2011. - SMF 33388, 1; SMF 33389, 1; SMF 33390, 1; SMF 33404, 20, Oued Igrounzar between Ounara and El Ghazouane (31°27'21"N, 9°41'4"W), A. Azeroual et al., 10 Apr 2011. - SMF 33406, 2, Oued Igrounzar near El Khemis des Meskala (31°21'31"N, 9°24'20"W), A. Azeroual et al., 11 Apr 2011.

Oued Iqem drainage. SMF 33509, 1, Morocco, Oued Iqem near Skhirat (33°53'22"N, 6°59'56"W), K. Borkenhagen and J. Freyhof, 19 Apr 2011.

Oued Kiss drainge. MNHN 1924-0174, 1, Algeria, Oued Kiss at Marsa Ben Mehid (35°4'59"N, 2°10'1"W), C. A. Alluaud, 1924.

Oued Moulouya drainage. SMF 33407, 8; SMF 33408, 4; SMF 33479, 1; SMF 33481, 1; SMF 33484, 1, Morocco, Oued Za near Guefaït (34°13'36"N, 2°23'34"W), K. Borkenhagen and J. Freyhof, 15 Apr 2011. - MNHN 1926-0070, 1, Morocco, Oued Melloulou near Guercif [34°13'32"N, 3°21'13"W], P. M. Pallary, 1926. - NMW 19533, 1, Morocco, Ras el Aïn near Aïn Beni Mathar (=Berguent) [34°0'41"N, 2°1'47"W], F. Werner. - NMW 19532, 1, Morocco, Oued Za [32°57'0"N, 5°12'0"W], F. Werner.

Oued Oum er Rbia drainage. BMNH 1902.7.28:22-26, 5, Morocco, Oued Oum er Rbia [33°19'40"N, 8°20'2"W], F. Riggenbach. - BMNH 1903.7.1:8, 2, Morocco, El Jadida [33°15'18"N, 8°30'22"W], F. Riggenbach. - MNHN 1927-0099, 1; MNHN 1927-0100, 1; MNHN 1989-0535, 1, Morocco, Oued Oum er Rbia near Khenifra [32°56'21"N, 5°40'7"W], A. Gruvel and R. Dollfus, 1927. - MNHN 1928-0054, 1; MNHN 1928-0055, 1, Morocco, Oued Oum er Rbia near Khenifra [32°56'21"N, 5°40'7"W], P. Pallary, 1928. - SMF 33513, 1; SMF 33514, 1; SMF 33515, 1, Morocco, Oued Oum er Rbia near Boulaouane (32°51'33"N, 8°2'41"W), K. Borkenhagen and J. Freyhof, 20 Apr 2011. - SMF 33344, 1; SMF 33345, 1; SMF 33346, 1; SMF 33394, 12, Morocco, Oued Srou at bridge between Tighassaline and Khenifra (32°49'51"N, 5°36'36"W), A. Azeroual et al., 7 Apr 2011. - SMF 33360, 1; SMF 33361, 1; SMF 33362, 1; SMF 33395, 17, Morocco, Oued Derra near Oulad Yaïch (32°26'23"N, 6°19'24"W), A. Azeroual et al., 9 Apr 2011. - SMF 33363, 1; SMF 33364, 1; SMF 33365, 1; SMF 33397, 22, Morocco, Oued Oum er Rbia (32°18'53"N, 6°54'33"W), A. Azeroual et al., 9 Apr 2011.

Oued Sebou drainage. SMF 33410, 9; SMF 33494, 1; SMF 33495, 1; SMF 33496, 1, Morocco, Oued Ouergha between Sidi Qacem and Ouazzane (34°27'52"N, 5°30'39"W), K. Borkenhagen and J. Freyhof, 17 Apr 2011. - MNHN 1939-0125, 1; MNHN 1939-0126, 1; MNHN 1939-0127, 1, Morocco, El Gharb [34°25'N, 6°20'W], J. M. Pérès, 1939. - MNHN 1939-0122, 2; MNHN 1939-0123, 2; MNHN 1939-0145, 1, Morocco, Oued Sebou [34°15'53"N, 6°41'5"W], J. M. Pérès, 1939. - SMF 33409, 15; SMF 33489, 1; SMF 33491, 1; SMF 33493, 1, Morocco, Oued Lahdar near Taza (34°14'35"N, 4°3'55"W), K. Borkenhagen and J. Freyhof, 16 Apr 2011. - MNHN 1924-0191, 3, Morocco, Oued Beth near Dar Bel Hamri [34°11'14"N, 5°57'54"W], C. A. Alluaud, 1924. - MNHN 1920-0061, 1; MNHN 1920-0062, 1, Morocco, Oued Bou Hellou [34°9'19"N, 4°25'33"W], P. M. Pallary, 1920. - MNHN 1922-0065, 1, Morocco, Moulay Yacoub [34°5'17"N, 5°10'54"W], C. A. Alluaud, 1922. - MNHN 1920-0202, 1, Morocco, Faraoun near Volubilis [34°4'25"N, 5°33'25"W], C. A. Alluaud, 1920. - MNHN 1939-0128, 1; MNHN 1939-0129, 1, El Mabbabat [?], J. M. Pérès, 1939.

Oued Tennsift drainage. BMNH 1904.11.28:60, 1; BMNH 1905.11.28:60-63 and BMNH 1904.11.28:57-58, 6, Morocco, Oued Chichaoua [31°43'48"N, 8°49'48"W], F. Riggenbach. - MNHN 1919-0379, 1; MNHN 1919-0380, 1; MNHN 1919-0381, 1; MNHN 1919-0382, 1, Morocco, Oued Nfis near Dar Goundafi [31°43'41"N, 8°21'1"W], P. M. Pallary, 1919. - MNHN 1988-1146, 4, Morocco, Oued Nfis [31°43'41"N, 8°21'1"W], Goubier, VI.1988. - MNHN 1922-0066, 1; MNHN 1922-0067, 1; MNHN 1922-0068, 1, Morocco, Oued Chichaoua near Chichaoua [31°32'37"N, 8°45'46"W], C. A. Alluaud, 1922. - SMF 33371, 1; SMF 33372, 1; SMF 33373, 1; SMF 33374, 1; SMF 33398, 3, Morocco, Oued Nfis near Tameslouht (31°27'2"N, 8°8'22"W), A. Azeroual et al., 10 Apr 2011. - SMF 33378, 1; SMF 33379, 1; SMF 33380, 1; SMF 33399, 14; SMF 33403, 22, Morocco, Oued Nfis near Ouirgane (31°13'24"N, 8°6'50"W), A. Azeroual et al., 10 Apr 2011. - MNHN 1925-0371, 1, Morocco, Oued Nfis near Ouirgane [31°10'40"N, 8°4'24"W], J. Pellegrin, 1925.

Diagnosis.

Two pairs of barbels, 30 to 39 scales in the lateral line and 14 to 20 scales around the least circumference of the caudal peduncle; dorsal fin usually shorter than anal fin and more than 15 % of its last unbranched ray flexible, dorsal profile of the head convex.

Description.

The body is of moderate height and sometimes has a small nuchal hump in larger specimens. The head is round with a convex dorsal profile and convex or straight ventral profile (Figs 16, 17). The head length is shorter than the body depth (Fig. 12), the mouth is inferior with two pairs of barbels (Table 2). The lower lip is crescent shaped and sometimes weakly keratinised. The eyes are in the anterior half of the head. The morphometric characters are summarised in Table 1.

The dorsal fin is short and weakly ossified and more than 15 % of the length of its last unbranched ray is flexible. Its last unbranched ray is about as long as the head (Fig. 4). It usually has four unbranched and seven to 10 branched rays (Table 3). The anal fin usually has three unbranched and five or six branched rays (Table 4). Its length is rather variable in adult specimens. It reaches the base of the caudal fin in some specimens.

Carasobarbus fritschii has 30 to 39 scales in the lateral line (Table 5), usually 5.5 scales above the lateral line (Table 6), usually 4.5 or 5.5 scales below the lateral line (Table 7), and 14 to 20 scales around the least circumference of the caudal peduncle (Table 8). The scales are shown in Fig. 5.

The pharyngeal teeth count is 2.3.4-4.3.2 in two specimens, 2.3.4- in one specimen and -4.3.2 in eight specimens. Pharyngeal teeth are hooked at their tips (Fig. 6).

Live specimens are silvery and usually have a dark longitudinal band above the lateral line. Fins are hyaline to slightly orange (Fig. 17). Ethanol-preserved specimens are yellow-brown, the back is usually distinctly darker than the belly and flanks.

The maximum length observed in the material available is 180 mm SL.

Carasobarbus fritschii differs from all congeners except Carasobarbus exulatus and Carasobarbus harterti in having nine instead of 10 branched dorsal-fin rays. It differs from Carasobarbus exulatus in having 30 to 39 scales in the lateral line vs. 26 to 32 and modally 16 scales around the least circumference of the caudal peduncle vs. 12. It differs from Carasobarbus harterti in having a convex dorsal head profile and a last unbranched dorsal-fin ray that is weakly ossified and flexible for more than 15 % of its length vs. a straight dorsal head profile and a strongly ossified last unbranched dorsal-fin ray that is flexible in less than 15 % of its length.

Distribution.

Carasobarbus fritschii is widespread and abundant in Northern and Central Morocco (Fig. 18). It occurs in the Oued al Maleh, Oued Bou Regreg, Oued Igrounzar, Oued Moulouya, Oued Oum er Rbia, Oued Sebou and Oued Tennsift drainage systems, and in numerous small coastal rivers. Most records are from Morocco, but one specimen is from the Oued Kiss in Algeria.

Habitats and biology.

Carasobarbus fritschii occurs in a wide range of running water courses and dam lakes.

Conservation status.

Carasobarbus fritschii is a hardy species and occurs in near-natural as well as heavily modified habitats. It is tolerant against pollution, damming and the presence of several exotic species (KB pers. obs.). The IUCN rates Carasobarbus fritschii as “Least Concern” and Barbus paytonii (which is treated as a junior synonym in this study) as “Vulnerable B2ab(iii)” (Crivelli 2006c, Crivelli 2006e). According to the latter assessment the population in the lower Oued Oum er Rbia is adversely affected by agricultural pollution (Crivelli 2006e).

Discussion. tle>

Carasobarbus fritschii was described from the Oued Ksob as a member of the genus Barbus (Günther 1874). The same author described Barbus rothschildi from the Oued Oum er Rbia (Günther 1901). It is a junior synonym of Carasobarbus fritschii. One year later Günther (1902) described Barbus riggenbachii from Oued Oum er Rbia and Oued Talmest. It is a junior synonym of Carasobarbus fritschii. In the same year Capoeta atlantica and Capoeta waldoi were described from Oued Nfis (Boulenger 1902). These two species were placed into Capoeta, based on the keratinised lower lip that occurs in some specimens of Carasobarbus fritschii. Both are junior synonyms of Carasobarbus fritschii. Barbus paytonii was described from Oued Oum er Rbia (Boulenger 1911). It is a junior synonym of Carasobarbus fritschii. In the same publication Boulenger transferred Capoeta waldoi to the genus Barbus. The junior synonyms listed above were described, based on slight differences in mouth and lower lip shape or the degree of ossification of dorsal-fin rays. Sample sizes were usually very small. The examination of a large number of specimens revealed high variability and a continuous distribution of these characters. Boulenger (1919) transferred all species to the genus Barbus subgenus Labeobarbus, based on the possession of scales with parallel radii and an unserrated last unbranched dorsal-fin ray. Pellegrin (1919) listed the species in the genus Barbus but later (Pellegrin 1921, 1939) accepted the subgenus Labeobarbus. Pellegrin (1939) synonymised Barbus riggenbachi with Barbus rothschildi and did not list Capoeta atlantica. Karaman (1971) created the genus Pseudotor and synonymised Capoeta atlantica and Capoeta waldoi with Pseudotor fritschii fritschii. Fowler (1976) accepted all previously described species and transferred Capoeta atlantica and Capoeta waldoi to the genus Varicorhinus. Berrebi (1981) used the genus Barbus subgenus Labeobarbus and found no relevant differences between Barbus fritschii and Barbus paytonii in his morphometric and biochemical analysis. El Gharbi et al. (1993) highlighted the African distribution of the subgenus Labeobarbus. Doadrio (1994) and Tsigenopoulos et al. (2010) used Labeobarbus. Subsequent authors used the genus Barbus (Azeroual et al. 2000, Machordom and Doadrio 2001, Leggatt and Iwama 2003, Colli et al. 2009) or the provisional genus ‘Barbus’ (Borkenhagen et al. 2011). We transfer this species to the genus Carasobarbus, based on the possession of a smooth last unbranched dorsal-fin ray, modally nine branched dorsal-fin rays, six branched rays in the anal fin and shield-shaped scales with numerous parallel radii. Analysis of molecular genetic characters (Durand et al. 2002, Tsigenopoulos et al. 2010, KB unpublished data) support this decision.

The name of this species is frequently misspelled “Barbus fritschi”.

The ‘Catalog of Fishes’ lists SMF 636 and SMF 952 as types for Carasobarbus fritschii (Eschmeyer 2011). Both lots where collected by K. v. Fritsch and J. Rein in Oued Ksob in 1872, together with the types of ‘Barbus’ reinii Günther, 1874, Luciobarbus nasus (Günther, 1874) and the syntypes of Carasobarbus fritschii. SMF 636 contains seven specimens: one Luciobarbus nasus, one Luciobarbus ksibi (Boulenger, 1905), one ‘Barbus’ reinii and four Carasobarbus fritschii. SMF 952 contains eight specimens: two ‘Barbus’ reinii and six Carasobarbus fritschii. In the original description Günther (1874) did not state the number of type specimens on which he based the description of Carasobarbus fritschii, but in the same paper he described Luciobarbus nasus (as Barbus nasus), based on two specimens and ‘Barbus’ reinii, based on three specimens. It is likely that Günther never saw the lots SMF 636 and SMF 952, because all syntypes of Luciobarbus nasus and ‘Barbus’ reinii are in the BMNH. The collectors, K. v. Fritsch and J. Rein probably deposited these samples immediately in the SMF and we conclude that SMF 636 and SMF 952 are not part of the type series of Carasobarbus fritschii.

Material.

Type material. Syntypes: BMNH 1901.7.26:4-5, 2, Morocco, Oued Oum er Rbia [33°19'40"N, 8°20'2"W], E. Hartert.

Non-type material. Oued Oum er Rbia drainage. BMNH 1902.7.28:27-33, 7; BMNH 1903.10.29:11-15, 8, Morocco, Oued Oum er Rbia [33°19'40"N, 8°20'2"W], F. Riggenbach. - BMNH 1903.7.1:5-7, 3, Morocco, Oued Oum er Rbia near El Jadida [33°15'18"N, 8°30'22"W], F. Riggenbach. - MNHN 1912-0089, 1; MNHN 1912-0090, 1; MNHN 1912-0091, 1; MNHN 1912-0092, 1; MNHN 1912-0093, 1, Morocco, Oued Oum er Rbia near Azemmour [33°17'22"N, 8°20'33"W], C. du Gast, 1912. - SMF 33366, 1; SMF 33368, 1; SMF 33370, 1, Morocco, Oued Oum er Rbia (32°18'53"N, 6°54'33"W), A. Azeroual et al., 9 Apr 2011.

Oued Tennsift drainage. BMNH 1902.7.28:34, 1, Morocco, Oued Talmest [31°52'15"N, 9°18'31"W], F. Riggenbach.

Diagnosis.

Two pairs of long barbels; 31 to 38 scales in the lateral line and 13 to 17 scales around the least circumference of the caudal peduncle; dorsal fin longer than anal fin and less than 15 % of the length of its last unbranched ray is flexible, dorsal profile of the head straight.

Description.

The body is of moderate height and without a nuchal hump. The head is triangular with almost straight dorsal and ventral profile (Figs 19, 20). The head length is shorter than the body depth (Fig. 12). The mouth is subterminal with two pairs of long barbels (Table 2). The eyes are in the anterior half of the head and relatively big. The morphometric characters are summarised in Table 1.

The dorsal fin is long and strongly ossified and less than 15 % of the length of its last unbranched ray is flexible. Its last unbranched ray is as long as or longer than the head (Fig. 4). It usually has four unbranched and nine branched rays (Table 3). The anal fin usually has three unbranched and six or seven branched rays (Table 4). It does not reach the caudal fin origin.

Carasobarbus harterti has 31 to 38 scales in the lateral line (Table 5), usually 5.5 or 6.5 scales above the lateral line (Table 6), 4.5 to 6.5 scales below the lateral line (Table 7) and 13 to 17 scales around the least circumference of the caudal peduncle (Table 8). The scales are shown in Fig. 5.

The pharyngeal teeth count is -4.3.2 in four specimens examined. The pharyngeal teeth are hooked at their tips (Fig. 6).

Live specimens are silvery with an olive tinge and orange fins (Fig. 20). Ethanol-preserved specimens are yellow-brown, the back is darker than the belly and flanks.

The maximum length observed in the material examined is 250 mm SL.

Carasobarbus harterti differs from all congeners except Carasobarbus exulatus and Carasobarbus fritschii in having nine rather than 10 branched dorsal-fin rays. It differs from Carasobarbus exulatus in having 31 to 38 scales in the lateral line vs 26 to 32 and modally 16 scales around the least circumference of the caudal peduncle vs. 12. It differs from Carasobarbus fritschii in having a straight dorsal head profile and a last unbranched dorsal-fin ray that is strongly ossified and flexible for less than 15 % of its length vs. a convex dorsal head profile and a last unbranched dorsal-fin ray that is weakly ossified and flexible for more than 15 % of its length.

Distribution.

Carasobarbus harterti occurs in the rivers of the Oued Oum er Rbia and Tennsift drainage systems in Morocco (Fig. 18).

Habitats and biology.

Carasobarbus harterti is less common than Carasobarbus fritschii and inhabits only the lower and middle course of big rivers.

Conservation status.

The IUCN rates this species as “Vulnerable A2ace“ (Crivelli 2006d). The population has declined more than 30 % in the time from 1996 to 2006 due to urban, agricultural and industrial pollution (Crivelli 2006d).

Discussion.

Carasobarbus harterti was described from Oued Oum er Rbia as Barbus harterti (Günther 1901). Some authors placed this species in the genus Barbus subgenus Labeobarbus (Boulenger 1919, Pellegrin 1921) while others continued using the genus Barbus (Pellegrin 1919, 1939). Karaman (1971) synonymised it with Carasobarbus fritschii, but regarded it as a distinct subspecies. He incorrectly synonymised Barbus rothschildi, Barbus riggenbachi and Barbus paytonii with this subspecies and placed it in his newly erected genus Pseudotor. Subsequent authors did not accept Karaman’s proposal and continued using Barbus (Fowler 1976, El Gharbi et al. 1993, Azeroual et al. 2000, Leggatt and Iwama 2003, Colli et al. 2009, Borkenhagen et al. 2011) or proposed using Labeobarbus (Doadrio 1994, Tsigenopoulos et al. 2010). We transfer this species to the genus Carasobarbus, based on the possession of a smooth last unbranched dorsal-fin ray, nine branched dorsal-fin rays, six branched rays in the anal fin and shield-shaped scales with numerous parallel radii. Analysis of molecular genetic characters (Durand et al. 2002, Tsigenopoulos et al. 2010, KB unpublished data) support this decision.

Material.

Type material. Holotype of Cyclocheilichthys kosswigi: ZMH H 1148, Turkey, Batman Çayı [37°47'16"N, 41°0'51"E], C. Kosswig, IV.1939.

Non-type material. Tigris-Euphrates system. NMW 90369, 1, Turkey, Batman Çayı near Baschkaja [37°53'15"N, 41°7'56"E], V. Pietschmann, 15 Jul 1914. - NMW 90805, 1, Turkey, Gökçesu Çayı (37°45'N, 41°45'E), 26 Sep 1985. - ZMH 9548, 2, Turkey, Ceylanpınar [36°50'50"N, 40°3'0"E]. - SMF 33119, 1, Syria, Nahr al Khābūr at Al Ḩasakah [36°30'9"N, 40°44'52"E], F. Krupp. - SMF 30172, 1, Syria, Nahr al Khābūr near Tall Budayrī (36°24'N, 40°52'E), F. Krupp, 2–4 Nov 1986. - SMF 30173, 1, Syria, Nahr al Khābūr near Nahāb (36°23'N, 40°50'E), F. Krupp, 23–27 May 1989. - SMF 30174, 1, Syria, Nahr al Khābūr near Nahāb (36°23'N, 40°50'E), F. Krupp, 28 Sep–8 Oct 1988. - CMNFI 79-0290, 2, Iran, Qaşr-e Shīrīn (34°31'N, 45°35'E). - CMNFI 79-0289, 1, Iran, 25–30 km from Qaşr-e Shīrīn (34°28'N, 45°52'E). - BMNH 1974.2.22:1292-1296, 4; BMNH 1974.2.22:1281, 1, Iraq, Euphrates at Ḩadīthah [34°8'23"N, 42°22'41"E], 19 Oct 1953. - CMNFI 79-0275, 1, Iran, Rūdkhāneh-ye Kashgān, 2 km from Ma‘mūlān (33°25'N, 47°58'E). - SMF 33129, 3, Iran, Rūdkhāneh-ye Karkheh at Pol-e Dokhtar (33°9'36"N, 47°43'12"E), N. Alwan et al., 3 Mar 2008. - ZM-CBSU 4153, 1; ZM-CBSU 4154, 1, Iran, Rūdkhāneh-ye Dez at Dezfūl [32°22'57"N, 48°24'7"E], F. Bossaghzadeh, 8 Jun 2005.

Diagnosis.

Two pairs of barbels; 32 to 38 scales in the lateral line, usually 14 to 16 scales around the least circumference of the caudal peduncle; last unbranched dorsal-fin ray markedly longer than head; mouth narrow, lower lip spatulate and median lobe present.

Description.

Body moderately high, laterally compressed and without a nuchal hump. The greatest body depth is at the point of the origin of the dorsal fin. The ventral profile of the head is straight, its dorsal profile has a slight to pronounced hump near the nostrils (Figs 21, 22). The head is short and narrow. The mouth is inferior. The maximum body depth is bigger than the head length (Fig. 12). The lips are comparatively thick and the lower jaw is narrow with a sharp horny sheath and a median lobe. The two pairs of barbels (Table 2) are stout and the anterior pair is quite long. The eyes are rather high in the middle of the head and rather small. The morphometric characters are summarised in Table 1.

The dorsal fin is long and usually has four unbranched and nine or 10 branched rays (Table 3). The last unbranched ray is long and well ossified; only the tip is flexible. It is considerably longer than the head (Fig. 4). The anal fin usually has three unbranched rays and six branched rays (Table 4). Its base is long. The bases of the dorsal and anal fin have a sheath of scales.

There are 32 to 38 scales in the lateral line (Table 5), 5.5 to seven scales above the lateral line (Table 6), 4.5 to 6.5 scales below the lateral line (Table 7) and (12) 14 to 16 scales around the least circumference of the caudal peduncle (Table 8). The scales are shown in Fig. 5.

The pharyngeal teeth count is 2.3.5-5.3.2 in seven specimens, 2.3.5- in one specimen and -4.3.2 in one specimen. The pharyngeal teeth are hooked at their tips (Fig. 6).

Live specimens are silvery. The back is darker than the belly, which is almost white (Fig. 22). Fixed specimens are yellow-brown and some have a darker back.

Carasobarbus kosswigi differs from all congeners, except Carasobarbus sublimus, by having a spatulate lower jaw with a median lobe on the lower lip vs. a crescent-shaped lower jaw and a lower lip without median lobe. It differs from Carasobarbus sublimus by having 32 to 38 scales in the lateral line vs. 27 to 29 and modally 14 scales around the least circumference of the caudal peduncle vs. 12 and by having a longer and more ossified last unbranched ray in the dorsal fin.

Distribution.

Carasobarbus kosswigi occurs in the Euphrates-Tigris system (Fig. 7).

Habitats and biology.

Carasobarbus kosswigi is rare, inhabits fast-flowing reaches of rivers and feeds on small animals (Krupp and Schneider 2008). The maximum length is about 150 mm SL and this species has no economic importance (Krupp and Schneider 2008).

Conservation status.

Little information is available, but because this species is dependent on fast-flowing water, it is probably impacted by the construction of dams.

Discussion.

Carasobarbus kosswigi was described from the Batman Çayı and placed in the genus Cyclocheilichthys (Ladiges 1960). Karaman erected the new genus Kosswigobarbus for this species (Karaman 1971). Coad gave a detailed re-description of this species and transferred it to the genus Barbus (Coad 1982). Kosswigobarbus was revalidated (Ekmekçi and Banarescu 1998) and sometimes used as a subgenus of Barbus (Tsigenopoulos et al. 2010). Later the species was placed in Carasobarbus (Borkenhagen et al. 2011).

Carasobarbus kosswigi is paraphyletic with respect to Carasobarbus sublimus (Borkenhagen et al. 2011).

Material.

Type material. Nahr Quwayq basin. Paralectotypes of Systomus luteus: NMW 54248, 1; NMW 54250:1-2, 2; NMW 54254:1-3, 3; SMF 6784, 1, Syria, Nahr Quwayq near Aleppo [36°12'10"N, 37°9'31"E], T. Kotschy, 17 May 1842.

Syntypes of Systomus albus: NMW 53674-53677, 4; NMW 53680, 1; SMF 812, 1, Syria, Nahr Quwayq near Aleppo [36°12'10"N, 37°9'31"E], T. Kotschy, 18 May 1842.

Rūd-e Mand basin. Syntypes of Systomus albus alpina: NMW 53678, 5; NMW 53679:1-2, 2; NMW 53681:1-2, 2, Iran, Rūdkhāneh-ye Qarah Āghāj near Shīrāz [29°31'3"N, 52°15'0"E], 2 Jan 1844.

Rūdkhāneh-ye Ḩelleh basin. Syntypes of Systomus albus alpina: NMW 53682:1-2, 2, Iran, Daryācheh-ye Parīshān [29°31'7"N, 51°47'47"E].

Tigris-Euphrates system. Lectotype of Systomus luteus (by present designation): NMW 54253:2, Iraq, Tigris near Mosul [36°20'6"N, 43°7'8"E], T Kotschy, 10 Apr 1843.

Paralectotypes of Systomus luteus: NMW 54247:1-2, 2; NMW 54249, 1; NMW 54253:1, 1; NMW 54255:1-2, 2; NMW 80043, 2 same data as lectotype.

Syntype of Systomus albus: NMW 91400, 1, Iraq, Tigris near Mosul [36°20'6"N, 43°7'8"E], 11 Apr 1843.

Unknown drainage system. Paralectotype of Systomus luteus: NMW 10827, 1, Syria, “Damascus”, T. Kotschy, 1837.

Non-type material. Daryācheh-ye Mahārlū basin. CMNFI 79-0047, 1, Iran, source of Ab-e Paravan marshes 19.9 km from Shīrāz University [29°36'N, 52°32'E]. - FSJF 2232, 2, Iran, Pirbano spring about 10 km south of Shīrāz (29°31'8"N, 52°27'56"E), A. Abdoli and J. Freyhof, 21 Apr 2007. - ZM-CBSU 3439, 1; ZM-CBSU 3449, 1; ZM-CBSU uncatalogued, 1, Iran, Pol-e Berenji, southwest of Shīrāz [29°27'30"N, 52°32'0"E], H. R. Esmaeili et al. - CMNFI 79-0347, 1, Iran, Solţānābād marshes near Pol-e Berenji (29°27'30"N, 52°32'0"E).

Orontes basin. MNHN 1977-0255, 1, Syria, Orontes, Gruvel, 1929, only one of two specimen examined. - MNHN 1977-0257, 1, Syria, Orontes, Gruvel, 1930. - SMF 24341, 1, Syria, Orontes at Jisr ash Shughūr (35°48'N, 36°19'E), F. Krupp, 21 Mar 1979 (aberrant specimen).

Rūd-e Mand basin. CMNFI 79-0206, 1, Iran, Qanat 41 km from Estahbān on road to Kharāmeh (29°12'N, 53°40'E). - CMNFI 79-0160, 1, Iran, cement pool near spring along road to Neyrīz (29°9'N, 53°37'E). - ZM-CBSU 4934-4942, 9, Iran, Dareh Daarveshan between Rudbal and Simakan (28°39'10"N, 52°2'27"E), H. R. Esmaeili et al. - ZM-CBSU 101-103, 3; ZM-CBSU 110, 1; ZM-CBSU uncatalogued, 1, Iran, Rūdkhāneh-ye Sīmakān near Jahrom [28°30'0"N, 53°33'38"E], H. R. Esmaeili et al.

Rūdkhāneh-ye Ḩelleh basin. CMNFI 79-0026, 1, Iran, Rūdkhāneh-ye Shāhpūr near Shahr-e Tārīkhī-ye Neyshābūr (29°47'N, 51°35'E). - ZM-CBSU 5180-5190, 10; ZM-CBSU 5192, 1, Iran, Kāzerūn, Sarab Dokhtar [29°37'10"N, 51°39'15"E], H. R. Esmaeili et al. - ZM-CBSU 6508-6517, 10; ZM-CBSU 6574, 1; ZM-CBSU 6602-6607, 6; ZM-CBSU 6610, 1; ZM-CBSU 6614+6615+6617-6619, 5; ZM-CBSU uncatalogued, 12, Iran, Daryācheh-ye Parīshān [29°31'7"N, 51°47'47"E], H. R. Esmaeili et al. - CMNFI 79-0240, 2; CMNFI 79-0304, 3, Iran, Daryācheh-ye Parīshān (29°31'N, 51°50'E). - CMNFI 79-0125, 1, Iran, Rūdkhāneh-ye Dālakī near Dālakī (29°28'N, 51°21'E). - ZM-CBSU 2650-2651, 2; ZM-CBSU 2654-2655, 2, Iran, spring at Palangī Dādīn, near Kāzerūn, Rūdkhāneh-ye Dālakī [29°25'20"N, 51°43'54"E], H. R. Esmaeili et al.

Rūdkhāneh-ye Kol basin. ZM-CBSU 3219-3229, 11; ZM-CBSU 3252-3260, 9, Iran, Golabi spring north of Dārāb [28°47'15"N, 54°22'19"E], H. R. Esmaeili et al. - FSJF 2253, 6, Iran, Golabi spring 35 km north of Dārāb (28°47'15"N, 54°22'19"E), A. Abdoli and J. Freyhof, 21 Apr 2007. - CMNFI 79-0155, 1, Iran, spring at Gavanoo, east of Ḩasanābād [28°47'N, 54°22'E]. - CMNFI 79-0154, 2, Iran, Korsia village on Dārāb-Fasā road (28°45'30"N, 54°24'0"E). - ZM-CBSU 5622-5626, 5, Iran, Tang-e Khūr near Lār [27°36'N, 54°17'E], H. R. Esmaeili et al.

Rūdkhāneh-ye Naband basin. CMNFI 79-0187, 10, Iran, stream and pools at Sarkhūn, Rūdkhāneh-ye Sarzeh (27°23'30"N, 56°26'0"E).

Tigris-Euphrates system. SMF 30208, 1, Turkey, Tigris at Diyarbakır (37°53'N, E40°14’), R. Kinzelbach, 1982. - SMF 30176, 11, Syria, Nahr al Khābūr at Ra’s al ‘Ayn (36°51'N, 40°4'E), F. Krupp, 24–26 May 1989. - SMF 30186, 12, Syria, ‘Ayn Sālūba and ‘Ayn Hamza near Ra’s al ‘Ayn (36°51'N, 40°4'E), F. Krupp, 3 Oct 1988. - SMF 30200, 2, Syria, ‘Ayn Sālūba at Ra’s al ‘Ayn (36°51'N, 40°4'E), F. Krupp, 3 Oct 1988. - SMF 30190, 7, Syria, Nahr al Khābūr 2 km East of Tall Junaydīyah (36°44'N, 40°6'E), F. Krupp, 26 May 1989. - SMF 30197, 2, Syria, Nahr al Khābūr 2 km East of Tall Junaydīyah (36°44'N, 40°6'E), F. Krupp, 5 Oct 1988. - SMF 30179, 3, Syria, Nahr al Khābūr at Tall ʿAtaš (36°42'N, 40°11'E), F. Krupp, 26 May 1989. - SMF 30188, 3, Syria, Nahr al Khābūr at Tall ʿAtaš (36°42'N, 40°11'E), F. Krupp, 6 Oct 1988. - SMF 31317, 1; SMF 33139, 7, Syria, Nahr al Khābūr at Tall Tamr (36°39'7"N, 40°21'51"E), N. Alwan et al., 29 Oct 2008. - SMF 30199, 1, Syria, Nahr al Khābūr at Tall Naşrī (36°37'N, 40°23'E), F. Krupp, 6–7 Oct 1988. - SMF 30178, 1; SMF 30202, 10, Syria, Nahr al Khābūr near Tall Bāz (36°35'N, 40°27'E), F. Krupp, 7 Oct 1988. - SMF 30184, 1; SMF 30193, 3, Syria, Nahr al Khābūr at Tall Bāz (36°35'N, 40°27'E), F. Krupp, 26 May 1989. - SMF 30181, 1; SMF 30192, 3, Syria, Nahr al Khābūr at Tall Umm al Māʿaz (36°34'N, 40°35'E), F. Krupp, 27 May 1989. - SMF 30183, 3, Syria, Nahr al Khābūr at Umm al-Māʿaz (36°34'N, 40°35'E), F. Krupp, 7 Oct 1988. - SMF 30182, 2, Syria, Nahr al Khābūr at Al Ḩasakah (36°30'N, 40°44'E), F. Krupp, 27 May 1989. - SMF 30195, 1, Syria, Nahr al Khābūr at Al Ḩasakah (36°30'N, 40°44'E), F. Krupp, 7 Oct 1988. - SMF 30185, 1; SMF 30213, 6, Syria, Nahr al Khābūr and Wādī Furātī at Tall Tayyiǧ (36°26'N, 40°52'E), F. Krupp, 8 Oct 1988. - SMF 30189, 4, Syria, Nahr al Khābūr at Baḩrat Khātūnīyah (36°24'N, 41°13'E), F. Krupp, 23–24 May 1989. - SMF 30214, 5, Syria, Nahr al Khābūr at Tall Budayrī (36°24'N, 40°49'E), F. Krupp, 26 Sep–8 Oct 1988. - SMF 30206, 7, Syria, Nahr al Khābūr at Tall Budayrī (36°24'N, 40°52'E), F. Krupp, 2–4 Nov 1986. - SMF 30177, 3, Syria, Nahr al Khābūr at Nahāb (36°23'N, 40°50'E), F. Krupp, 28 Sep–8 Oct 1988. - SMF 30201, 23, Syria, Nahr al Khābūr at ‘Ayn Ţābān (36°22'N, 40°50'E), F. Krupp, 28 Sep 1988. - SMF 30191, 2, Syria, Nahr al Khābūr at mouth of Wādī ar Raml (36°15'N, 40°48'E), F. Krupp, 8 Oct 1988. - SMF 30196, 1, Syria, Nahr al Khābūr at Umm Rukaybah (36°8'N, 40°42'E), F. Krupp, 8 Oct 1988. - SMF 30194, 3, Syria, Nahr al Khābūr at Ash Shaddādah (36°4'N, 40°44'E), F. Krupp, 9 Oct 1988. - SMF 31316, 1; SMF 33138, 2, Syria, Nahr al Khābūr at Ash Shaddādah (36°3'46"N, 40°44'30"E), N. Alwan et al., 28 Oct 2008. - SMF 33152, 6, Syria, Jisr Shānīn (36°3'4"N, 39°5'10"E), F. Krupp and W. Schneider, 19 Aug 1980. - SMF 31308, 1, Syria, Mamlaḩat al Jabbūl (36°3'36"N, 37°33'1"E), N. Hamidan, 23 Jun 2008. - SMF 28707, 18, Syria, Euphrates down stream Buḩayratt al Asad (35°51'48"N, 39°0'34"E), R. Beck, Jun 1998. - SMF 30198, 2, Syria, Nahr al Khābūr at Tall ash Shaykh Ḩamad (35°37'N, 40°45'E), F. Krupp, 21 Sep–14 Oct 1988. - SMF 30204, 1; SMF 30205, 4, Syria, Nahr al Khābūr at Tall ash Shaykh Ḩamad (35°37'N, 40°45'E), F. Krupp, 20 Oct–9 Nov 1986. - SMF 33140, 1; SMF 33141, 37, Syria, Euphrates at Harmūshīyah (35°35'52"N, 39°51'25"E), N. Alwan et al., 31 Oct 2008. - SMF 30203, 2, Syria, Nahr al Khābūr 8 km South of Tall ash Shaykh Ḩamad (35°33'N, 40°43'E), F. Krupp, 24 Oct 1986. - SMF 28737, 5, Syria, Euphrates between Ḩalabīyah-Zalābīyah and Dayr az Zawr, R. Beck, Jun 1998. - SMF 28630, 3, Syria, Euphrates upstream Dayr az Zawr (35°31'N, 39°54'E), R. Beck, 23 May 1998. - SMF 28674, 41, Syria, Euphrates upstream Dayr az Zawr [35°31'N, 39°54'E], R. Beck, 30 May 1998. - SMF 33153, 1, Syria, Nahr al Khābūr at Aş Şuwar (35°30'N, 40°38'E), F. Krupp, 15 Mar 1979. - SMF 31315, 1; SMF 33137, 1, Syria, Nahr al Khābūr at Ghawat (35°28'51"N, 40°39'54"E), N. Alwan et al., 28 Oct 2008. - SMF 30187, 2, Syria, Nahr al Khābūr near Ḩarījīyah (35°27'N, 40°38'E), F. Krupp, 10 Oct 1988. - SMF 30180, 5, Syria, Nahr al Khābūr at Mashikh (35°14'N, 40°31'E), F. Krupp, 10 Oct 1988. - SMF 28663, 6, Syria, Euphrates at Qal‘at aş Şāliḩīyah (Dura Europos) [34°45'0"N, 40°43'30"E], R. Beck, 28 May 1998. - SMF 28758, 2, Syria, Euphrates at Abū Kamāl at mouth of Wādī Ratqah [34°26'45"N, 40°56'0"E], R. Beck, 9 Jul 1998. - NMW 93019:1-2, 2, Iraq, Tigris at Baghdād [33°20'26"N, 44°24'3"E], V. Pietschmann, Aug 1910. - SMF 33127, 4, Iran, Rūdkhāneh-ye Bālārūd (32°35'19"N, 48°17'11"E), N. Alwan et al., 3 Mar 2008. - BMNH 1980.8.28:6, 1, Iran, Rūdkhāneh-ye Dez at Dezfūl [32°25'N, 48°13'E]. - SMF 33125, 1, Iran, Rūdkhāneh-ye Dez at Dezfūl (32°22'40"N, 48°22'58"E), N. Alwan et al., 2 Mar 2008. - SMF 33121, 5, Iran, Rūdkhāneh-ye Dez at Dezfūl (32°21'49"N, 48°21'28"E), K. Borkenhagen et al., 3 Nov 2006. - SMF 17303, 1, Iraq, Hawr al Ḩammār (30°50'N, 47°10'E), L. A. J. Al-Hassan, 1986. - SMF 30211, 1, Iraq, ‘Ayn Zālah 50 km west of Mosul, Z. Rahemo, 1990.

Unknown drainage system. SMF 33120, 2, Syria, fish market in Damascus (reported to be from Buḩayratt Ar Rastan [34°56'N, 36°44'E] in Orontes drainage), F. Krupp. - CMNFI 79-0687, 4, Iran, Shīrāz bazar (probably from Rūd-e Mand basin or Daryācheh-ye Mahārlū basin).

The lectotype (NMW 54253:2) is a specimen of 211 mm SL, collected in the Tigris near Mosul on 10 Apr 1843 by T. Kotschy (Fig. 23). It has four unbranched and 10 branched rays in the dorsal fin, three unbranched and six branched rays in the anal fin, 27 scales in the lateral line and one pair of barbels. A bigger specimen (216 mm SL) from the same lot (NMW 54253:1) was not selected as lectotype, because it is atypical in having 11 branched rays in the dorsal fin and two pairs of barbels. The designation of a lectotype became necessary to fix the type locality of Systomus luteus (see Discussion).

Diagnosis.

One pair of barbels; 25 to 33 scales in the lateral line, and typically 12 scales around the least circumference of the caudal peduncle; last unbranched ray of the dorsal fin about as long as the head or slightly shorter.

Description.

Specimens from Rūdkhāneh-ye Naband basin were excluded from this species description (see below).

The dorsal profile is convex up to the origin of the dorsal fin and a nuchal hump is present in specimens longer than about 100 mm SL. This species has a high back and caudal peduncle (Figs 23, 24). The ventral profile of the head is convex, its dorsal profile is almost straight to convex and has a hump near the nostrils in juvenile specimens. The mouth is sub-terminal. The barbels are short and stout. The maximum body depth is usually greater than the head length (Fig. 12). Usually one pair of barbels is present, but about 10 % of the specimens have two pairs of barbels (Table 2). The eyes are at the back of the anterior half of the head. They are big and slightly protuberant. The morphometric characters are summarised in Table 1.

The dorsal fin usually has four unbranched and eight to 11 branched rays (Table 3). In specimens from the Tigris-Euphrates drainage system the last unbranched ray of the dorsal fin is strong with only the tip being flexible and it is about as long as the head. It is shorter and less ossified in Iranian populations (Fig. 4). The anal fin usually has three unbranched rays and five to seven branched rays (Table 4).

There are 25 to 33 scales in the lateral line (Table 5), 3.5 to 6 scales above the lateral line (Table 6), 3 to 5.5 scales below the lateral line (Table 7) and 10 to 13 scales around the least circumference of the caudal peduncle (Table 8). The scales are shown in Fig. 5.

The pharyngeal teeth count is 2.3.5-5.3.2 in 26 specimens, 2.3.4-5.3.2 in two specimens, 2.3.5-4.3.2 in one specimen, 2.3.5-5.3.3 in one specimen, 1.3.5-5.3.2 in one specimen, 2.3.5- in one specimen and 2.3.4- in one specimen. The pharyngeal teeth are hooked at their tips (Fig. 6).

Live specimens are silvery to olive and sometimes have yellowish fins (Fig. 24). Ethanol-preserved specimens are light yellowish brown to grey. In most cases the back is darker than the rest of the body. Some of the lighter coloured specimens have a salmon hue, others are silvery. The fins are yellowish brown to grey. Juveniles have a dark spot on the sides of the caudal peduncle.

Carasobarbus luteus from Ḩelleh, Kol, Mahārlū and Mand populations: The last unbranched ray of the dorsal fin is shorter and less well ossified. It is pronouncedly shorter than the head (Fig. 4). The mouth is wider and the body is not as high-backed as in specimens from the Tigris-Euphrates system (Fig. 12).

Carasobarbus luteus from Rūdkhāneh-ye Naband basin: In this population all specimens examined had two pairs of barbels (Table 2). The anterior pair is longer than in specimens from Tigris-Euphrates system with two pairs. The last unbranched ray in the dorsal fin is considerably shorter than the head (Fig. 4) and comparatively weak. Compared with specimens from Tigris-Euphrates system, the dorsal and ventral fins tend to be slightly further away from the head. The head is longer and the body not as high backed as in specimens from Tigris-Euphrates system (Fig. 12). The general body shape (Fig. 25) resembles that of Carasobarbus apoensis and Carasobarbus canis. Some of the gill rakers are y-shaped in the largest specimen examined.

Carasobarbus luteus, except the population from Rūdkhāneh-ye Naband, differs from all congeners, except Carasobarbus apoensis, in having one instead of two pairs of barbels. It differs from Carasobarbus apoensis, Carasobarbus canis, Carasobarbus chantrei, Carasobarbus fritschii, Carasobarbus harterti and Carasobarbus kosswigi in modally having 28 scales in the lateral line vs. 30, 32, 34, 34, 34 and 33 respectively. It differs from Carasobarbus kosswigi and Carasobarbus sublimus in having a crescent-shaped lower lip without median lobe vs. a spatulate lower lip with median lobe and from Carasobarbus exulatus, Carasobarbus fritschii and Carasobarbus harterti in modally having 10 rather than nine branched dorsal-fin rays. All populations, except the one from Rūdkhāneh-ye Naband differ from Carasobarbus apoensis in having a shorter head and a higher back. The population from Rūdkhāneh-ye Naband is very similar to Carasobarbus apoensis in body shape, but differs in having two as compared to one pair of barbels.

Distribution.

Carasobarbus luteus has a much greater range than any of its congeners and its distribution area is fragmented, resulting in several isolated populations. It is widespread all over the Tigris-Euphrates drainage system, and occurs in the rivers of south-western Iran (Fig. 7). The Nahr al Quwayq population, from one of the sites of the type locality, is probably extirpated due to drought and pollution (Krupp 1980, Krupp 1983b). There are only few, mostly older, records from the Orontes (Krupp 1985c, Krupp 1987). During recent fieldwork Carasobarbus luteus was not found there. Because Carasobarbus chantrei is still widespread and abundant in many parts of the Orontes, it is unlikely that Carasobarbus luteus disappeared due to habitat degradation. It might have been driven out by competition with Carasobarbus chantre i or records were based on misidentifications or mislabelled specimens. One specimen (NMW 10827) is reported from Damascus. Because Carasobarbus luteus does not occur in the Damascus basin and it is highly unlikely that it ever occurred there, the origin of this specimen is unclear.

Habitats and biology.

Carasobarbus luteus is mainly herbivorous. It feeds on algae, aquatic plants, detritus and small invertebrates, the main feeding period is at noon, but food is also taken at night (Naama and Muhsen 1986). The intestine is long (Ali 1986). The maximum size is 38 cm total length and 750 g, but normally they are smaller than 35 cm and weight less than 500 g (Ahmed 1982). They reach maturity at the age of one or two years and at a size of about 14 cm; the spawning period is June and July in the Tigris-Euphrates system, the eggs are spawned among reeds, roots or other aquatic vegetation and fecundity is high (Al Hazzaa and Hussein 2003a).

This species can tolerate saline waters to some degree (Al-Hassan and Muhsin 1986, Mohamed et al. 1993) and is of commercial importance due to its size and abundance (Ahmed 1982, Barak and Mohamed 1983, Krupp and Schneider 2008).

There are attempts on aquaculture of this species. The stickiness of the eggs can be lowered by several chemical treatments for this purpose (Al Hazzaa and Hussein 2003b). During spawning males get reddish brown in the anterior part of the body and greenish at the caudal peduncle while females are less colourful (Al Hazzaa and Hussein 2003a). Males can produce series of sharp clicking noises which do not seem to be associated with aggressive behaviour (Al Hazzaa and Hussein 2003a).

Larvae hatch at 64 degree-days in well oxygenated water and the eyes are still without pigments at this stage. The development is similar to that of other cyprinids (Al Hazzaa and Hussein 2003a). Ahmed et al. (1984) studied the reproductive biology of Carasobarbus luteus.

Conservation status. Carasobarbus luteus is widespread and abundant in the Tigris-Euphrates system. Peripheral populations, like those in smaller Iranian rivers and the Nahr al Quwayq in Syria are more threatened or have already been extirpated (see above).

Discussion.

Carasobarbus luteus was described as Systomus luteus by Heckel (1843). Heckel (1843) listed Orontes, Tigris, Aleppo and Mosul as type localities. As all but one of the type specimens are either from the Tigris-Euphrates system or from the Nahr al Quwayq and Aleppo is located on the Nahr al Quwayq and not on the Orontes, Heckel may have confused these two rivers. One of the type specimens (NMW 10827) is from “Damascus” and can not be attributed to any of the relevant drainage systems. By designating NMW 54253:2 as lectotype we fix the Tigris near Mosul as type locality for Systomus luteus. The same confusion exists for the type localities of Systomus albus, which was also described from Tigris and Orontes in the same publication. A few years later Systomus albus var. alpina was described from the Daryācheh-ye Parīshān (Heckel 1847). These three taxa where later synomymised and placed in the genus Barbus (Günther 1868). Sauvage (1882, 1884) accepted Carasobarbus luteus and Carasobarbus albus as valid species and transferred them to the genus Barynotus. Later, both species where synonymised again and transferred to the genus Barbus, subgenus Puntius (Misra 1947) or the genus Puntius (Menon 1956). Ladiges (1960) synonymised both species under the name Barynotus albus. Because Günther (1868) had previously selected luteus as the valid species name, he is to be considered the first revising author and Ladiges’ action is not valid. Kähsbauer (1963) lists the species under two different generic names: Barbus (as Barbus luteus) and Systomus (as Systomus albus var. alpina). Karaman (1971) erected the new genus Carasobarbus for this species. This met mixed acceptance. While some authors accepted the new taxonomic position (e.g. Wossughi 1978, Bianco and Banarescu 1982, Ahmed et al. 1984, Naama and Muhsen 1986), others did not embrace it (e.g. Banister and Clarke 1977, Krupp 1985a, c, Coad 1995, Coad 1996) until the revision by Ekmekçi and Banarescu (1998). Fowler (1976) placed Carasobarbus luteus in the genus Barbellion. Tsigenopoulos et al. (2010) used Barbus subgenus Carasobarbus. Barbus parieschanica was described from Daryācheh-ye Parīshān (Wossughi et al. 1983). In the same publication the species name is also spelled B. parschanica, but Barbus parieschanica is probably the intended spelling (Coad 1995). Coad (1995) as the first revising author fixed Barbus parieschanica as the correct original spelling. Barbus parieschanica is a synonym of Carasobarbus luteus. The ‘Catalog of Fishes’ lists RMNH 2463 as possible syntype of Systomus luteus and RMNH 2464 of Systomus albus var. alpina (Eschmeyer 2011). We did not examine these specimens.

We do not think that the population at Rūdkhāneh-ye Naband should be elevated to specific rank, because the number of specimens available is too low. We provisionally consider it an atypical population of Carasobarbus luteus that might have been affected by bottleneck effects and accelerated morphological change, due to the restricted size and extreme conditions (high salinity and temperature) of its habitat. It would be very interesting to collect more samples for morphological studies and molecular sequence analysis.

In spite of some morphometric differences, Carasobarbus luteus populations of Tigris-Euphrates system and Iran belong to the same species (Borkenhagen et al. 2011); specimens from Rūdkhāneh-ye Naband were not included in that study.

Carasobarbus luteus and Carasobarbus apoensis are closely related to each other (KB, unpublished data) and Carasobarbus apoensis might be the ecologically specialised sister species of Carasobarbus luteus, that is adapted to the environmental conditions of the wadi ecosystems of the Al Ḩijāz mountains.

Material.

Type material. Holotype of Barbus sublimus: CMNFI 1995-0009, Iran, Rūdkhāneh-ye A‘lā near Pol-e Tīghen (31°23'30"N, 49°53'0"E), B. W. Coad et al., 20 Sep 1995, not examined.

Paratypes of Barbus sublimus: CMNFI 95-0009a, 1, same data as holotype. - CMNFI 95-0010, 1, same data as holotype, not examined. - CMNFI 95-0011, 3, Iran, Rūdkhāneh-ye A‘lā near Pol-e Tīghen (31°23'30"N, 49°53'0"E), G. Eskanderi, Dec 1994, only one specimen examined.

Non-type material. Rūdkhāneh-ye Kashgān. CMNFI 79-0277, 1, Iran, Rūdkhāneh-ye Kashgān at Harpul Kashkow, 50 km from Khorramābād (33°30'0"N, 47°59'30"E), K. Evans and H. Assadi, 5 Jul 1977.

Rūdkhāneh-ye Zohreh drainage. ZM-CBSU 5781-5786, 6, Iran, Rūdkhāneh-ye Fahlīān at Nūrābād [30°6'51"N, 51°31'18"E], H. R. Esmaeili et al. - SMF 33117, 3, Iran, Rūdkhāneh-ye Fahlīān (30°11'10"N, 51°31'14"E), K. Borkenhagen et al., 29 Nov 2007. - SMF 33118, 6, Iran, Rūdkhāneh-ye Fahlīān (30°11'9"N, 51°31'15"E), N. Alwan et al., 29 Feb 2008.

Diagnosis.

Two pairs of barbels; 27 to 29 scales in the lateral line, 12 scales around the least circumference of the caudal peduncle; last unbranched dorsal-fin ray about as long as the head; mouth narrow, lower jaw spatulate and median lobe present on lower lip.

Description.

A nuchal hump is not developed. The maximum body depth is at the anterior end of the dorsal fin base. The ventral profile of the head is almost straight; the dorsal profile is convex and evenly curved (Figs 26, 27). The maximum body depth is greater than the head length (Fig. 12). The mouth is inferior, narrow, the lips are thick and the lower jaw is spatulate with a horny sheath and a median lobe on the lower lip. The two pairs of barbels (Table 2) are well developed. The eyes are at the posterior end of the anterior half of the head. Some morphometric characters are summarised in Table 1.

The dorsal fin usually has four unbranched and nine or 10 branched rays (Table 3). The last unbranched ray of the dorsal fin is weakly ossified and about as long as the head (Fig. 4). The anal fin usually has three unbranched and six branched rays (Table 4) and its base is surrounded by a sheath of scales. Pectoral, ventral and anal fins are longer than in all other Carasobarbus species (Table 1).

There are 27 to 29 scales in the lateral line (Table 5), 4.5 or 5.5 scales above the lateral line (Table 6), 3.5 to 5.5 scales below the lateral line (Table 7) and 12 scales around the least circumference of the caudal peduncle (Table 8). The scales are shown in Fig. 5.

The pharyngeal teeth count is 2.3.4-5.3.2, 2.3.4-5.3.1 or 3.3.4-4.3.3 (Coad and Najafpour 1997). The pharyngeal bones available were too small for photography but are very similar to those of Carasobarbus kosswigi (Fig. 6).

Live specimens from Rūdkhāneh-ye Fahlīān are silvery with hyaline fins (Fig. 27). Live specimens from Rūdkhāneh-ye A‘lā are silvery with a slightly darker back, the scales have dark pigments on their hind margin; pectoral, ventral and anal fins have a yellow to orange hue, which is most obvious with fins folded back; dorsal and caudal fins are grey or hyaline (Coad and Najafpour 1997). Ethanol-preserved specimens are yellowish brown with a somewhat darker back and juveniles have a dark spot on the sides of the caudal peduncle.

Carasobarbus sublimus differs from all congeners, except Carasobarbus kosswigi, by having a spatulate lower jaw with a median lobe on the lower lip vs. a crescent shaped lower jaw and a lower lip without median lobe. It differs from Carasobarbus kosswigi by having 27 to 29 scales in the lateral line vs. 32 to 38 and modally 12 scales around the least circumference of the caudal peduncle vs. 14 and by having a shorter and less ossified unbranched last dorsal-fin ray.