Genus Acanthobrama
Heckel, 1843

Howes (1981) placed Acanthobrama Heckel, 1843 in the genus Rutilus Rafinesque, 1820 on osteological grounds but most other authors retain Acanthobrama as a distinct genus (Coad, 1984a; Krupp, 1985c; Eschmeyer, 1990; Bănărescu, 1992b) based on the scale, keel and anal fin characters listed below. Durand et al. (2002) include this genus in the Abramis clade based on cytochrome b data. The genus Trachibrama Heckel, 1843 is a lapsus (Krupp and Schneider, 1989).

This genus is characterised by a compressed, deep body of small to moderate size, no barbels, relatively small scales with reduced numbers of radii, a fleshy keel between the base of the pelvic fins and the vent, the last unbranched dorsal fin ray is thickened, spine-like and smooth, and the anal fin is long (9-22 branched rays). Pharyngeal teeth are usually in a single row on each arch. Gut short.

There are 8 species endemic to Southwest Asia with 1 found in southwestern Iran (Goren et al., 1973; Coad et al., 1983; Krupp, 1985c).

Acanthobrama marmid
Heckel, 1843

Common names

كلاش پا (= kalashpa), شبه ساردين (shebeh sardin = pseudo-sardine or resembling sardine), شبه نازي (= shebeh nazy), mahi sim nama (= bream-like fish).

[semnan arrez; samnan areed; arath (Rahemo et al., 2006); marmid, marmid handscherli (= marmid armed with a dagger), marmid abbiad (= white marmid), marmid asphar (= yellow marmid) or marmid mablue (= swallowing or devouring marmid) at Aleppo, arrhada (= dove, lion!) at Mosul (all these latter Arabic names after Heckel (1843b; 1846-1849a), the conflicting names for arrhada included, and are probably antiquated; Tigris bream].

Systematics

Acanthobrama Arrhada Heckel, 1843, Acanthobrama cupida Heckel, 1843, Acanthobrama marmid morpha elata Berg, 1949 and Acanthobrama marmid orontis Berg, 1949 are synonyms.

The type locality for Acanthobrama Marmid is "Gewässern bei Aleppo", for Acanthobrama arrhada "in Mossul", and for Acanthobrama cupida "in Aleppo" according to Heckel (1843b) and "Flusse Kueik bei Aleppo" in Heckel (1846-1849a). The type locality of Acanthobrama marmid morpha elata is Lake Balikli, 12 km from Erzurum, 8 km from the Karasu River, upper Euphrates, in Turkey. The type locality of Acanthobrama marmid orontis is the upper Euphrates region according to Eschmeyer et al. (1996) (but this is an error, see below).

Details on the syntypes of this species and its synonyms arrhada and cupida in the Naturhistorisches Museum Wien are given by Krupp (1985c). Eight syntypes of marmid measuring 41-144 mm standard length are listed from Mosul (in contrast to Heckel's papers where the type locality is Aleppo), the number of fish agreeing with the catalogue in Vienna. These 8 fish are under NMW 55334. Eschmeyer et al. (1996) do not list these fish as types and the card index in Vienna in 1997 concurs. A further 15 fish are listed by Krupp from the Quwayq near Aleppo: 1 fish, 102 mm standard length (NMW 55342 - not in the 1997 card index; but the following NMW fish are listed), 2, 113-139 mm standard length (NMW 55345), 2, 86-121 mm standard length (NMW 55346), 2, 98-126 mm standard length (NMW 55347), 2, 113-132 mm standard length (NMW 55348), 2, 114-138 mm standard length (NMW 79068), and 4 fish in the Senckenberg Museum Frankfurt, 82-112 mm standard length (SMF 543, formerly NMW). Eschmeyer et al. (1996) list NMW 55345-48 (8), NMW 79068 (2), SMF 543 (4) and in the Rijksmuseum van Natuurlijke Historie, Leiden RMNH 2537 (4) and RMNH 2539 (2) (both formerly NMW) as the type series.

Two syntypes of A. arrhada from Mosul, 85-92 mm standard length, are in the Senckenberg Museum Frankfurt (SMF 411, formerly NMW) (F. Krupp, pers. comm., 1985; 85.7-89.0 mm standard length) while 2 others are in the Naturhistorisches Museum Wien, ca. 150 mm standard length (NMW 55335) and 141 mm standard length (NMW 55336) (Krupp, 1985c). However, the Vienna catalogue lists 6 specimens of A. arrhada and in addition to the above material there is also NMW 55334 (8 fish) tagged as syntypes so there is some confusion in what constitutes the type series. Two possible syntypes are in the Rijksmuseum van Natuurlijke Historie, Leiden (RMNH 2538) (Eschmeyer et al., 1996).

Krupp (1985c) records syntypes of A. cupida, 151 mm standard length, (NMW 55340) and 152 mm standard length (NMW 55341). The Vienna catalogue lists 4 A. cupida which agrees with Heckel's description although I observed only NMW 55340 (1 fish), NMW 55341 (1) and also NMW 55342 (1). Eschmeyer et al. (1996) list NMW 55340-43 (1, 1, 1) as syntypes but the numbers indicate 4 fish. The card index in Vienna in 1997 also lists 55505 (5 fish), one of which is designated as the lectotype.

The 2 syntypes of Acanthobrama marmid orontis are in the Zoological Institute, St. Petersburg under ZISP 6720 from "L. Antioch, 1884, Lortet" according to Berg (1949). This subspecies is distinguished only by larger scales from the typical form but the 2 syntypes examined by me had lost their scales and were difficult to count; one seemed to have a count around 64, not as low as 54-55 recorded by Berg (1949). Krupp (1985c) examined type material and new specimens from the Orontes and found them not to differ from A. marmid from the Quwayq and Tigris-Euphrates basins. He accordingly synonymises Acanthobrama marmid orontis with the type subspecies.

Karaman (1972) considered Acanthobrama arrhada to be a subspecies of A. marmid rather than a synonym based on an unusually strongly ossified spiny dorsal fin ray in the former. Since A. marmid was described from Aleppo (= Halab, Syria) and A. arrhada from Mosul, the synonomy of these two taxa may warrant re-examination.

The fish reported from the Tigris River basin of Iran by Nümann (1966) as Xenocypris macrolepidotus was this species (Zoologisches Institut und Zoologisches Museum, Hamburg catalogue number ZMH H2700 examined by me). Saadati (1977) thought it a new species of Acanthobrama but I disagree.

A hybrid with Chalcalburnus (= Alburnus) mossulensis was reported from the Hawr al Hammar in southern Iraq by Krupp et al. (1992).

Key characters

The characters of the genus distinguish this species from all other Iranian cyprinids.

Morphology

Mouth nearly horizontal to oblique, equal or lower jaw slightly behind the upper. The belly has a fleshy keel where the ventral scales do not meet along the mid-line between the pelvic base and the anus. The last unbranched dorsal fin ray is a thickened, stiff and smooth spine, the rigid part varying from 15 to 26% of standard length. The spine may be strong for much of its length and then abruptly become thin and flexible or it may taper gradually to a flexible tip. Some small fish lack an enlarged dorsal fin spine.

Lateral line scales 53-72, scales above the lateral 10-14, scales between the pelvic fin and lateral line 4-7. There is a pelvic axillary scale. Radii are restricted to the posterior field on scales and are few in number. The focus is subcentral anterior to almost central. Dorsal fin with 3 unbranched rays and 7-9 branched rays. Anal fin unbranched rays 3, branched rays 13-22. Pectoral fin branched rays 12-18, pelvic fin branched rays 7-9. Total vertebrae 38-43 (38(3), 39(3), 40(7), 41(5), 42(7) or 43(1) combining Iranian and Iraqi material).. Gill rakers short with a basal swelling, 2-4 on the upper arch, 0-1 at the flexure and 9-12 on the lower arch. Total rakers 12-17. The rakers reach the one below or to its further base end when appressed. Pharyngeal teeth usually 5-5, with the anterior tooth compressed and bluntly pointed, the remainder bevelled with a cutting edge and a hooked tip. The two anterior teeth are more rounded than the others although the second one may have a slight hook and is bevelled. Tigris River basin fish may have 1-2 teeth in a second row. The gut is an elongate s-shape with a large anterior loop in larger fish. The diploid chromosome number is 2n=50, with the karyotype consisting of 8 metacentric, 13 submetacentric and 4 pairs of subtelocentric to acrocentric chromosomes. The karyotype is nearly identical to other Eurasian leuciscine cyprinids (Gaffaroğlu et al., 2006)..

Different body forms occur in slow-flowing and fast-flowing waters. In the former habitat fish have a deep body, often humped behind the head, while in the latter the body is more streamlined (Karaman, 1972). It seems that A. marmid is founded on the humped form and A. arrhada and A. cupida on the streamlined one.

Meristic values for Iranian specimens are:- dorsal fin branched rays 7(1) or 8(8); anal fin branched rays 13(2), 14(2), 15(4) or 17(1); pectoral fin branched rays 13(2), 14(3), 15(3) or 18(1); pelvic fin branched rays 7(1), 8(7) or 9(1); lateral line scales 54(1), 55(2), 56(2), 58(1), 59(1) or 63(1); total gill rakers 12(1), 14(6) or 17(1); pharyngeal teeth 5-4(1) or 5-5(7); and total vertebrae 38(3), 39(1), 41(2), 42(2) or 43(1).

Sexual dimorphism

Fine tubercles are found over the top, sides and bottom of the head in males. Tubercles line the first, unbranched pectoral fin ray irregularly with up to 2 branching rows. Very fine tubercles are found on the adjacent membrane and on the lower pectoral fin surface. Tubercles line the pelvic fin rays in branching rows. The lower caudal fin rays are lined with tubercles. Anterior upper flank scales, all belly scales and lower caudal peduncle scales have their margin lined with tubercles, the peduncle with some tubercles on the mid-scale and the belly with a concentration on the scale base.

Colour

The overall colour is silvery to whitish with the head and back reddish-brown. The flanks can be greyish to blackish from numerous melanophores. There may be a well-developed mid-flank stripe or it may be poorly developed or evident only posteriorly. The pelvic fins are bright red, the pectoral and anal fins less red and the dorsal and caudal fins reddish proximally and black distally. Fin colours may be more orange or yellow than red. All fin rays and membranes have melanophores and these can be quite concentrated such that some fish have dark fins. Young fish in preservative have numerous, distinctive, small to minute, rounded, square or oblong patches of pigment in 1-3, irregular, mid-flank rows. Peritoneum black, silvery with a dorsal concentration of melanophores or with widely scattered melanophores so it appears silvery.

Size

Reaches 20.8 cm (Berg, 1949)

Distribution

This species is found in the Tigris-Euphrates basin of Turkey, Syria, Iraq and Iran, the Quwayq (= Kueik) and Orontes rivers, and possibly the Amik Lake and the Bardan suyu (= stream) near Tarsus (Ladiges, 1960; - Krupp (1985c) suggests these latter should be checked). In Iran it is found in the Tigris River basin including the upper reaches of the Karkheh, the Qara Su, and in marshes such as the Hawr Al Azim.

Zoogeography

The majority of species are found in the Levant which once had connections to the Tigris-Euphrates basin (Krupp, 1985c).

Habitat

Hussain et al. (1997) report this species to be dominant in the small fish assemblages in the Shatt Al-Arab near Basrah, Iraq at 70.8% of 14,084 fish caught. It favours side branches off the Shatt al Arab, presumably to avoid predators which are found in deeper water. Younis et al. (2001b) noted that this species dominated in the polluted and disturbed environment of a dockyard on the Shatt al Arab. This was one of the most abundant species in the recovering marshes of southern Iraq in 2005-2006 (Hussain et al., 2006) and is also known from large rivers and dams. Also recorded from the Hawr al Azim marsh in Iran.

Age and growth

Al-Nasiri and Salman (1977) studied this species in the Little Zab River, Iraq. Their largest specimen was 13.7 cm. They described length-weight relationships and condition factors but some important length groups were missing from their samples. Condition factor showed a gradual decrease with increasing length and the means for actual and calculated weights were 1.141 and 1.118 respectively. Relative condition factor was 1.0009. Younis et al. (2001) examined three populations of this species in the Shatt al Arab, Iraq and found the 0+ age group to be represented by fish 2.1-11.0 cm long and 1+ age group by fish 8.3-14.1cm. The length-weight relationship was W = -3.821 L^2.32. Four age groups with a length range of 4-19 cm were found in the Qarmat Ali River of southern Iraq, with maturity in the first year (Saud, 1997).

Ünlü et al., (1994) examined a population of this species in the Tigris River, Turkey and gave figures for growth in length and in weight. Females grew faster and are larger in size than males at the same age, particularly for age groups III and IV. Condition factor for males was 1.554 and for females 1.550. They found 5 age groups with age group III dominant for both sexes. Overall sex ratio was 1.83 females:1 male. Sexual maturity was attained by 75% of females and 85% of males in the second year of life and all fish in age group III were mature.

Food

Heckel (1843b) suggests that they are ravenous feeders based on the name "swallowing marmid". Gut contents are crustaceans, insects, and plant and gastropod shell fragments in Iranian specimens. Younis et al. (2001a; 2001b) found Shatt al Arab, Iraq fish to be detritivores, having organic detritus as the dominant gut content, followed by phytoplankton (blue-green algae and diatoms), small crustaceans (ostracods, cyclopoids, cladocerans), and aquatic plants, with dominance varying by month. Gut contents were crustaceans, insects, and plant and gastropod shell fragments in fish from Iran examined by me. In a study of the recovering Hammar Marsh, Iraq, diet was 70.77% insects and 9.81% algae with diatoms, plants, crustaceans and snails at less than 10% each, in the Hawr al Hawizah 66.4% insects and 14.1% algae, with amounts of diatoms and various crustaceans being less than 10% each, and in the Al Kaba'ish (= Chabaish) Marsh 62.7% insects and 17.7% algae with diatoms, plants and various crustaceans at less than 10% each (Hussain et al., 2006).

Reproduction

Younis et al. (2001) found most females to be ripe in March and July samples, and some were spent. Well-developed testes are noted in fish caught on 16 May in Turkey and 7 July near Ravansar, Kermanshahan indicating either a prolonged breeding season or local variations.

Ünlü et al., (1994) report spawning in May to late June for their Tigris River, Turkey population. They cite data for a Keban Dam population (on the Euphrates River in Turkey) where the spawning season is extended and runs from April to August. Egg diameter exceeds 1.2 mm and egg numbers reach 8125, and elsewhere may reach 11,000 eggs. In the Qarmat Ali River in southern Iraq, fecundity reached 1759-9293 eggs.

Parasites and predators

None reported from Iran.

Economic importance

None in Iran. In the early 1990s in Iraq, this species was used for human consumption and for fish meal (Younis et al., 2001).

Conservation

This species is rarely reported from Iranian waters and its status needs to be assessed through further field work. Endangered in Turkey (Fricke et al., 2007).

Further work

Additional field work is required to secure more materials and assess conservation status and biology.

Sources

Type material: See discussion above. Syntypes of Acanthobrama marmid (NMW 55345, NMW 55346, NMW 55347, NMW 55348, NMW 79068, SMF 543); syntypes of Acanthobrama marmid orontis (ZISP 6720), syntypes of A. arrhada (SMF 411, NMW 55335, NMW 55336, NMW 55334); syntypes of A. cupida (NMW 55340, NMW 55341, NMW 55342, NMW 55505).

Iranian material: CMNFI 1979-0287, 2, 89.9-92.1 mm standard length, Kermanshahan, spring near Ravansar (ca. 34º42'N, ca. 46º40'E); CMNFI 1979-0360, 1, 40.6 mm standard length, Khuzestan, Karkeheh River canal (31º40'N, 48º35'E); CMNFI 1979-0377, 2, 28.5-34.6 mm standard length, Khuzestan, Karkheh River (ca. 32º57'N, ca. 47º50'E); CMNFI 1979-0384, 1, 23.1 mm standard length, Khuzestan, Ab-e Shur drainage (32º00'N, 49º07'E); CMNFI 1991-0154, 1, 113.6 mm standard length, Khuzestan, Hawr-al-Azim (ca. 31º45'N, ca. 47º55'E); CMNFI 1993-0128, 1, 113.6 mm standard length, Kermanshahan, Sarab-e Sabz `Ali Khan (34º25'N, 46º32'E); CMNFI 2007-0114, Kermanshahan, Qareh Su basin (ca. 34º28'N, ca. 46º54'E); ZMH H2700, 1, 145.0 mm standard length, Kermanshahan, Gharasu-Gamasiab-Seymarreh (Qareh Su, Gav Masiab and Simareh rivers, no other locality data); uncatalogued, 1, 101.7 mm standard length, Kermanshahan, sarabs near Kermanshah (no other locality data).

Comparative material: BM(NH) 1931.12.21:22-25, 4, 65.7-84.6 mm standard length, Iraq, Mosul (ca. 36º20'N, ca. 43º08'E); BM(NH) 1974.2.22:1084-1091, 7, 105.1-118.3 mm standard length, Iraq, Najab Bazar (no other locality data); BM(NH) 1974.2.22:1094, 109.3 mm standard length, Iraq, Great Zab River at Aski Kalak (36º16'N, 43º39'E); BM(NH) 1971.4.2:7, 96.5 mm standard length, Iraq, River Tigris near Mosul (ca. 36º20'N, ca. 43º08'E); BM(NH) 1974.2.22:1078-1083, 6, 105.2-122.8 mm standard length, Iraq, Najab Bazar (no other locality data); BM(NH) 1974.2.22:1092, 109.5 mm standard length, Iraq, Najab Bazar (no other locality data); CMNFI 1987-0017, 3, 83.8-108.3 mm standard length, Iraq, Hawr al Hammar (no other locality data); BM(NH) 1920.3.3:147-156, 15, 29.5-102.0 mm standard length, Syria, Ouadi Khneizer (no other locality data); BM(NH) 1968.12.13:108-112, 1 (of 5), 112.6 mm standard length, Syria, Ouadi Khneizer, Khabour (no other locality data); BM(NH) 1968.12.13:113-118, 6, 56.5-117.4 mm standard length, Syria, River Euphrates at Houreira (no other locality data); ZSM 26136, 5, 55.3-80.3 mm standard length, Syria, Assad Reservoir, Euphrates basin (no other locality data); CMNFI 1980-0810, 2, 114.8-118.3 mm standard length, Turkey, Göksu in Tigris River basin (no other locality data); CMNFI 1980-1036, 1, 101.5 mm standard length, Turkey, Keban Dam on Murat Nehri near Elâzığ (no other locality data).

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