The tooth-carps, killifishes or pupfishes are small fishes found in fresh, brackish and sea water. There are 9 genera and about 105 species found in tropical to warm temperate climates almost world-wide. Iran has 7 described species, with more to be discovered.

This family is characterised by a moderately elongate and compressed body, rather cyprinid-like but with jaw teeth (hence the common and scientific names), body and head covered with scales, no spines in the fins, no barbels, the mouth is very protractile and armed with comb-like teeth, the lower jaw is strong and robust as the dentary is expanded medially, 4-6 branchiostegal rays, gill membranes free from the isthmus, lateral line absent or reduced to points, caudal fin rounded or truncate, pectoral fins set low on the body (in contrast to Poeciliidae), dorsal and anal fins short, no adipose fin, no pyloric caeca, and no gonopodium (egg layers, in contrast to the livebearer subfamily in the Poeciliidae).

Wildekamp (1993) gives a general account of cyprinodontid species. These fish can be maintained in aquaria, and are very popular with aquarists. Specific requirements where known are given under each species. Moderate to hard fresh water is used and for water with a total hardness less than 10dGH, sea salt can be added (about one teaspoon per 10 litres). Summer temperatures of 20-25°C are good and although higher temperatures make the fish more active, they also age the fish more rapidly. Winter temperatures below 20°C imitate nature and are recommended. Weekly partial water changes are suggested and live and frozen fish foods and even flake foods are eaten. Breeding occurs throughout the year in aquaria kept above 15°C, eggs are deposited on fine-leaved plants (thread algae, Java moss or yarn mops) or even on aquarium filter sponges and gravel. Egg development takes 8-20 days, fry take about 2 days to absorb their yolk sac and will then consume baby brine shrimp. These fish can be kept in outdoor ponds in milder climates (http://ark.aka.org/Eurasian.htm, downloaded 1 December 2003).

Genus Aphanius
Nardo, 1827

These tooth-carps are found around the shores of the Mediterranean, in Southwest Asia and as far as northeastern India and Somalia. There are about 23 known species.

Sethi (1960) advocated placing Aphanius in a separate family, Aphaniidae, but this did not find general acceptance.

The genus Aphanius Nardo, 1827 has been used for these tooth-carps for many years. However, Lazara (1995) designated Lebias fasciata Valenciennes, 1821 as the type species for Lebias Goldfuss, 1820, making Lebias a subjective senior synonym of Aphanius. On this basis Lebias must be used rather than Aphanius but Lazara's type species designation is invalid (Kottelat and Wheeler, 2001). Since Lazara (1995) could have chosen another species as type species, the change involves a large number of species and these species are threatened and listed in various legislation, a petition is before the International Commission on Zoological Nomenclature to suppress Lazara's designation (Kottelat, 1997; Kottelat and Wheeler, 2001; Wildekamp, 2001; Villwock et al., 2002). Wildekamp et al. (1999) present evidence that Lebias is a synonym of Cyprinodon.

Literature on these Mediterranean and Southwest Asian fishes may appear under either of these names or under Cyprinodon Lacepède, 1803, the latter now restricted to American species.

Bănărescu (1995) disagrees with Parenti's (1981) relationship of Aphanius to South American Orestias (see also Parker and Kornfield (1995), Stevenson (1997) and Costa (1997) for conflicting views).

This genus is characterised by a thick oval body, large to moderate cycloid scales, the head flattened on top, a small, superior mouth with tricuspid teeth, the upper jaw bordered by the premaxillaries only, lateral line system present only on the head, dorsal fin positioned somewhat posteriorly with1-2 unbranched rays and 7-13 branched rays, anal fin rays 1-2 unbranched and 7-14 branched, dorsal and anal fins larger in males than in females, dorsal fin inserted opposite the anal fin origin (in contrast to Gambusia), and colouration of males and females distinct. Seyfali et al. (2002) has 2n=48 for fish from the Damghan basin which Coad and Abdoli (2000b) suggest may be a distinct taxon.

Aphanius is the only genus in the family currently recognised in Iran. However Parenti (1981) distinguishes derived members of the genus Aphanius as `Aphanius' without formally describing a new genus. One of the distinguishing features of `Aphanius' is the reduction of cephalic sensory pores to neuromasts, a character found in A. mento, A. sophiae and A. vladykovi in Iran.

Wildekamp et al. (1999) review the Turkish species which show scale reductions not seen in Iranian populations. Hrbek et al. (2002) outline the historical biogeography of the species complex in central Anatolia, Turkey using mtDNA, testing the hypothesis of geographic speciation driven by early Pliocene orogenic events that may well be paralleled in Iran. Hrbek and Meyer (2003) studied the phylogeny of Eurasian tooth-carps using mtDNA and observed a western Tethys Sea clade (all species except those listed below) with a middle Oligocene divergence into Iberian Peninsula and Atlas Mountains, and Turkey and Iran sections. Late Miocene orogenic events were correlated with a large amount of genetic differentiation in Turkey (and presumably Iran). An eastern Tethys Sea clade (dispar, ginaonis, mento, sirhani) had an Oligocene divergence into a freshwater clade inhabiting the Arabian Peninsula and neighbouring areas and a euhaline clade inhabiting coastal areas from Pakistan to Somalia. Speciation is predominately vicariant-based but ecological factors played a significant role.

These fishes are known generally as gour-e khar in Farsi (= literally "striped donkey" which means zebra here although usually in Iran this term refers to the wild ass or onager), kapurdandandar or kopurdandandar (= tooth-carp), or even آفانيوس (= Aphanius). Saefali (1999) is a recent study of these fishes in Farsi.

Iranian Aphanius away from coastal drainages are thought to be relicts of the Tethys Sea (Kosswig, 1955a; 1967; Bianco, 1995), having been trapped by the rising Iranian Plateau, rather than invaders from the coast (Krinsley, 1970). Fossil tooth-carps, Aphanius kirgisicus and A. longipinnis have been described from Kyrgyzstan in Miocene deposits, far inland from the distribution of living species (Yakovlev, 1959). Priem (1908) describes a fossil cyprinodont from the Miocene of Iran.

The systematics of the Iranian populations would repay careful study using biochemical and genetic techniques coupled with aquarium studies on behaviour and cross breeding. Certain populations, isolated from others in discrete drainage basins, appear to be different on the basis of colour patterns yet show little morphological variation, e.g. the Damghan population, see Coad and Abdoli (2000b), Saefali (1999) and Seyfali et al. (2002), the most northerly and easterly population in Iran characterised by large spots in females; populations in Fars outside the Lake Maharlu and Kor River basins; populations in Khuzestan (see below); and populations in the Namak Lake basin - all yet to be investigated in detail (Coad, 2000b). Since these patterns are used in mate recognition, the populations could be distinct taxa. Huber (1996) has noted in cyprinodonts from Africa that many taxa can only be separated on the basis of colour in life, other characters overlapping. He suggests a cyprinodont species definition which includes possession of at least one stable phenotypic characteristic, and this could be colour of the male fish. He does recommend that genetic isolation be demonstrated where possible by karylogy, biochemical techniques or breeding experiments.

Male Aphanius sp. from Khuzestan

Female Aphanius sp. from Khuzestan

Saifali et al. (2004) found a suite of morphometric characters to be different between samples of A. vladykovi and fish from Qomy-Abad in Tehran Province, the Namak Lake basin.

The mosquitofish (Gambusia) is a niche competitor and may well eliminate these tooth-carps. Generally, Aphanius prefers springs, lakes, marshes, sea shores and hot springs. Flowing water is usually avoided but they can be found near the source of springs flowing into salt lakes, constrained by hypersaline waters downstream. However they are tolerant of high salinities. They occur in schools but males are aggressive to other males. Males are brightly coloured, often striped, while females are subdued with spots or faint bars.

A related Mediterranean species is reputed to be ichthyootoxic but this has not been demonstrated for Iranian species (Coad, 1979b). Symptoms of this egg poisoning are summarised under the genus Schizothorax.

These small fishes are used in the aquarium trade as they tolerate a wide range of temperatures and salinities and are particularly colourful. As early as 1912, J. P. Arnold in Germany noted the survival of imported Iranian Aphanius (possibly A. persicus) in aquaria at temperatures of 14-27°C, with a variety of plants such as Myriophyllum, Nitella, Riccia, Cabomba, Salvinia and filamentous algae, with daphnia, cyclops, enchytraeids and red mosquito larvae as food, and egg laying apparently preferred on the Salvinia with 8-12 days for hatching at 27°C.

Aphanius sp. from the Namak Lake is recorded as having the parasite Clinsotomum complanatum (Hosseinie, 1987).

Aphanius dispar
(Rüppell, 1829)

Male

Common names

gour-e khar, kopurdandandar-e balehbolband (= tooth-carp with striped fin), kapurdandan-e balmband.

[harsun. batrikh or batrikh motakayer in Arabic; nambal in Pakistan; high-finned pupfish, Arabian killifish, mother of pearl fish].

Systematics

A detailed synonymy is given by Villwock et al., (1983), Wildekamp et al. (1986) and Wildekamp (1993). Hoedeman (1951) proposed a new genus for this species, Aphaniops, based on the absence of a dermal sheath or genital pouch around the anterior anal fin rays, only 8-9 dorsal fin rays in contrast to 10-14 rays in Aphanius and 7-8 pelvic fin rays in contrast 5-7 rays. It has not found general acceptance.

The type subspecies is found in all Iranian drainages (Krupp, 1983) although Aphanius dispar stoliczkanus (Day, 1872) has been recorded in earlier literature as the subspecies of these drainages except the Tigris River basin for which Aphanius dispar richardsoni (Boulenger, 1907) is reported (Berg, 1949). Aphanius dispar richardsoni is limited to the Dead Sea valley of Israel and western Jordan by Villwock et al. (1983) and Wildekamp (1993). Cyprinodon stoliczkanus Day, 1872 is regarded as a synonym of Aphanius dispar dispar by Krupp (1983) while Berg (1949) places Bampur River, Baluchestan fish in this subspecies as a southern representative of Aphanius dispar with 6-9, rarely 10 dorsal fin rays as opposed to 9-10 rays in Aphanius d. dispar.

Hrbek and Meyer (2003) note that, based on their mtDNA study, the monophyly of A. dispar is strongly rejected and it does not constitute a species in terms of the phylogenetic species concept.

Hybrids with Aphanius dispar richardsoni are reported from Israel (Goren and Rychwalski, 1978).

The lectotype of Lebias dispar as designated by Villwock et al. (1983) is in the Senckenberg Museum Frankfurt under SMF 821 with paralectotypes SMF 1988 (10). Further paralectotypes in the Natural History Museum, London under BM(NH) 1860.11.9:152 listed as from Abyssinia (3 fish, 44.1-49.6 mm standard length) (Eschmeyer et al., 1996) but a label in the jar states that "status as types doubtful, types in SMF range from 26.1-35.8 mm SL, locality in description and types given as 'Red Sea'".

Types of Cyprinodon stoliczkanus Day, 1872 are in the Zoological Survey of India, Calcutta (ZSI 1477, ZSI 1478), the Natural History Museum, London (BM(NH) 1889.2.1.2065-74, originally 21 fish, now 14 fish, 14.4-29.4 mm standard length when examined in September 2007) and the Australian Museum, Sydney (AMS B.7730-7731) (Whitehead and Talwar, 1976; Eschmeyer et al., 1996; Ferraris et al., 2000). This species was originally described from a "stream at the village of Joorun, and also at Lodai, along the edge of the Rann" (= the salt water Rann of Cutch, Sind, India).

The lectotype of Cyprinodon richardsoni as designated by Krupp and Schneider (1989) is in the Natural History Museum, London under BM(NH) 1856.5.2:4 with paralectotypes under BM(NH) 1856.5.2:5 (8 fish) (12.9-25.0 mm standard length, the largest being the lectotype) (Eschmeyer et al., 1996; personal observations).

Krupp (1983) did not find the variation in teeth numbers between different populations and subspecies observed by Berg (1949).

Key characters

The colour pattern is distinctive.

Morphology

Scales along the flank 24-35. Scales are squarish with an almost vertical anterior margin and protruding anterior corners, parallel dorsal and ventral margins and a rounded posterior margin with tiny teeth. The anterior margin has a small central protuberance with shallow indentations above and below or almost straight margins to each anterior corner. Radii are present on the anterior field, numbering 10-20 but mostly 14-16, and are almost horizontal and parallel. The exposed part of the scale has dimples rather than circuli. Circuli are few. The focus is subcentral posterior. There is no pelvic axillary scale. Total dorsal fin rays 7-11, usually 9-10, total anal fin rays 8-12, usually 10-11 (the number of unbranched rays in the dorsal and anal fins varies from 1-3), total pectoral rays 12-18 and total pelvic rays 6-7, usually 7. Vertebrae 24-29. Al-Hassan (1982b) reports on abnormalities in the vertebrae including loss of the posterior part of the centrum and fusions. Teeth are tricuspid with the central cusp concave at its tip and only slightly longer than the lateral cusps (Goren and Rychwalski, 1978), although in some fish seen by me the tip is rounded. There are 12-20 teeth per jaw (Krupp, 1983). Gill rakers 11-20, modally 13-16, reaching the second adjacent raker when appressed. Some gill raker counts are difficult to make accurately as those at the anterior arch end are minute and those at the dorsal end are partially concealed in flesh. The gut is coiled ?see my ginaonis paper. Reichenbacher et al. (2007) give a description of otolith morphology. Chromosomes number 2n=48 (Klinkhardt et al., 1995; Esmaeili et al., 2008). Karyotype 16Sm + 32St and arm number 32 (Esmaeili et al., 2008).).

Meristic values for Iranian specimens:- total dorsal fin rays 8(1), 9(39) or 10(10); total anal fin rays 9(2), 10(42) or 11(6); total pectoral fin rays 14(2), 15(16), 16(28), 17(3) or 18(1); total pelvic fin rays 6(6) or 7(44); lateral series scales 24(2), 25(5), 26(11), 27(17), 28(7), 29(2), 30(1), 31(2), 33(2) or 35(1); total gill rakers 15(13), 16(16), 17(16) or 18(3); and total vertebrae ?.

Sexual dimorphism

Males have longer fins than females and are more brightly coloured. The dorsal fin is twice as long in the male and reaches the caudal fin when appressed. When expanded it is widely flared and distinctive as is the enlarged anal fin.

Colour

Breeding males are brown-grey, grey or black-brown with iridescent blue-white flank spots and white and brown to light orange or light blue, irregular, narrow bars. The head has blue and orange tinges and in particular there is an orange spot on the operculum postero-dorsally. Lips are blue-white. The flank over the pectoral fin has electric blue spots. The anterior belly becomes blue with pearl spots. Scales have a dark margin. The dorsal fin is spotted light blue on a bright orange background and is barred. Barring may be irregular and in overall view in breeding males appears as speckling. Pectoral, anal and pelvic fins are lemon-yellow. The anal fin has barring on the posterior 4 rays in breeding males. The caudal fin has 2-3 dark and light blue alternating bars, the last bar being yellow. The bars are crescent-shaped, with concave side posterior. Males outside the breeding season are less brightly coloured with silvery on the flanks with a grey or black-brown back and irregular flank bars. Male specimens from the Bampur River basin in Iran have a body speckled silvery-white to light blue, weak tail bars, blue and orange tinges to the head, an orange spot on the operculum and an orange-yellow edge to the anal fin.

Young have pale brown flank bars on the caudal peduncle. Some young may have a few spots on the flank rather than bars. Females are brown-grey to silvery with 8-20 narrow flank bars and a dark-brown back. The flank bars are dark brown and may have a tinge of orange, the interspaces silvery. Flank bars may number as few as 7 (Talwar and Jhingran, 1991). Females also have the orange operculum spot and the blue spots over the pectoral fin. Female fins are hyaline. The peritoneum is brown to black.

Ross (1980) notes that Arabian specimens are brighter in waters with algae and plants; those found over sand and mud are duller, an effective camouflage. He also records variant colour patterns with blue tails, or an overall gold colour, or with green or turquoise flank spots.

Size

Reaches 8.0 cm, but this may be an error as Krupp (1983) points out. He found fish only to 5.2 cm standard length although in Carpenter et al. (1997) 8 cm is given as a maximum. Wildekamp (1993) records fish to 7.0 cm for males and 6.5 cm for females in total length. Fish from Ain Al Adhari, Bahrein reach 7.3 cm total length. Generally fish in fresh water are smaller than marine specimens.

Distribution

Found from Egypt, eastern Sudan and Ethiopia, and in the Red Sea through southern Southwest Asia and east to Rajasthan. Also in the eastern Mediterranean Sea (Kornfield and Nevo, 1976). The subspecies Aphanius dispar richardsoni is found in the Dead Sea rift valley.

A specimen in the Naturhistorisches Museum Wien from the hot spring at Ginao (NMW 13799) may be mislabelled as several other collections from this hot spring are all clearly A. ginaonis.

This species is found in Iranian fresh waters from the Tigris basin to the Makran basin in water bodies draining to the Persian Gulf and Sea of Oman, islands in the Gulf such as Qeshm (OSU 8120), and in the endorheic Jaz Murian basin. It is reported from the Shapur and Dalaki rivers (Gh. Izadpanahi, pers. comm., 1995). Abdoli (2000) illustrates distributions in the Arvand, lower Zohreh (= Hendijan), lower Hilleh, Mand, Kul, Hasan Langi and Minab rivers, lower reaches of rivers in the Makran basin from the Jagin to Bahu Kalat, the lower Halil and Bampur Rivers in the Jaz Murian basin, and the endorheic Mashkid River.

Zoogeography

Villwock et al. (1983) contend that an ancestor of A. dispar was present in the middle Miocene Transgression connecting the Mediterranean Sea to the Indian Ocean. The mesohaline conditions, which developed in the Mesopotamian basin, were a good prerequisite for the development of A. dispar since, apart from some euryhaline species, there would be no competitors. A. dispar spread from Mesopotamia through the Persian Gulf to India and the Red Sea. During the Pleistocene, it colonised inland waters.

Habitat

Highly tolerant of salinity, this species is found from the sea to fresh water habitats, including landlocked basins. Slightly saline waters in aquaria seem to discourage parasites (Andrews, 1983). Salinities as high as 145‰ are tolerated (Lotan, 1971) and this fish can be moved abruptly from fresh to hypersaline water. Adaptation to high salinity waters involves a reduced osmotic permeability of the gills and an increase in transportation of sodium chloride by the intestine (Skadhauge and Lotan, (1974). Lotan (1969, 1971, 1973) and Skadhauge and Lotan (1974) describe sodium, chloride and water balance, and osmoregulation in this species. Plaut (1999) found that salinities above ~250% seawater seem to decrease swimming capabilities and routine activity rate and a decrease in resting metabolic rate indicated stress. This species has been reported to survive in waters 38.4°C in Oman and only began to die at 45.1-46.0°C (Haas, 1982). Fish survived up to 40°C in aquaria without acclimation (Lewis, 1970). Populations survive in isolated sections of streams with little food because of histological changes in the stomach (Ba-Omar et al. (1998). Aphanius dispar occurs in shallow water and among vegetation over sand, rock or soft detritus bottoms.

Frenkel (1995) describes the effects of environmental factors on growth and reproduction in this species in Israel. Frenkel and Goren (1997) showed fish to be tolerant of a wide range of environmental variables as measured against the gonado-somatic index (GSI) and ovary maturation stages. The GSI was not affected by temperature over a range of 18 to 37°C while maturation increased from 18 to 27°C but remained the same between 27 and 37°C. For salinities between 0‰ and 56‰ differences were only found in mean GSIs at the two extremes and at 0‰ ovaries contained only primordial germ cells. GSIs and maturation stages only differed in feeding experiments when fish were deprived of food or fed at equal or greater than 1% of body weight per day. A decrease in photoperiod from 14L:10D (14 hours light, 10 hours dark) to 10L:14D caused a decrease in oocyte maturation stages and ovaries were no longer suitable for spawning. In aquaria, a pH of 7.5-8.5, temperatures above 27°C and water hardness greater than 200 p.p.m. and the addition 4 heaped tablespoons of kosher salt per ten gallons of water (38 litres) favoured reproduction (Mackowiak, 1988). Alkahem (1989) found 50% mortality in 96 hours when fish were exposed to pH levels of 3.5-3.9.

Feulner (1998) reports that United Arab Emirate fish "hover" in the water column with the tail slightly curved to one side, presumably a feeding behaviour. Males and females in aquaria swam in separate schools (Al-Daham et al., 1977).

Age and growth

Sexual maturity in ideal conditions may be attained as early as 7 months but full size takes a year. The length-weight relationship in southern Iraq was W = 1.2301 x 10^-4 L^2.4544 for males and W = 2.8357 x 10^-6 L^3.5112 for females. The average condition factor for males was 1.8900 and for females 1.6952. For a given length group males are heavier than females and condition factors are higher for males (Huq et al. 1977). Muhsin (1987) studied the effect of differing food supply on the weight and chemical composition in females of this species taken from the Al-Asafiah River, Basrah, Iraq but examined in tanks. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 41 Iranian fish measuring 2.37-3.56 cm total length. The a-value was 0.0012 and the b-value 3.214 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Iranian specimens have guts packed with filamentous algae and detrital remains. Al-Daham et al. (1977) found food in southern Iraq to be predominately filamentous algae and diatoms, with some copepods, rotifers and, in one instance, a small winged insect. In contrast, Shafi and Shalli (1986) report a diet of beetles, ephemeropteran nymphs, rotifers, filamentous algae and plants in southern Iraq. Younis et al. (2001b) found Shatt al Arab, Iraq fish fed on amphipods and copepods (29.9%), organic detritus (11.6%), and algae and diatoms (7.1%). In aquaria, Al-Daham et al. (1977) found this species to prefer animal foods in contrast to Aphanius mento which preferred plant foods. Al-Akel et al. (1987) have observed the selective feeding behaviour of this species at Al-Khurj, Saudi Arabia. Certain species of filamentous algae are selected over others, e.g. the blue-green alga Oscillatoria, the green algae Ulothrix, Spirogyra and Crucigenia, the desmid Cosmarium, and the diatoms Cocconeis, Diatoma, Navicula and Rhoicosphania, and algal spores and zygotes. Haas (1982) observed this species in Oman to pick at rocks and other substrates, take items at the surface, chase small fish and eat insect larvae. Gut contents are mainly desmids and diatoms, some filamentous algae and rarely insect larvae and one snail. This species is an omnivorous surface feeder (Carpenter et al., 1997).

Reproduction

A female in spawning condition approached a male and appeared to feed on a plant or plant-encrusted rock in fish studied by Haas (1982) in Oman. Feulner (1998) reports fluttering of the tail by male fish. The male places his chin on her nape and 2-3 eggs are spawned rapidly. The female presses her vent against the plant or rock vertical surface, the male arches his back into an s-shape and presses his body and anal fin against hers and clasps her with his dorsal fin. Eggs are released singly with a quiver. The time elapsed from chin on nape to egg release is as short as 1-2 seconds. Reproduction occurs throughout the day and throughout the year in Oman with 69% of females having ripe eggs (up to 41) in April-June. Peak spawning in southern Iraq is April to June when only a small proportion of eggs in the single ovary are fully developed eggs (Al-Daham et al., 1977). Shafi and Shalli (1986) record up to 73 mature eggs in southern Iraq specimens with a diameter of 220 microns. Breeding season was May-July. In Saudi Arabia mature and developing oocytes are observed in fish during the whole year. The egg cycle may be more than one year with spawning in March and April (El-Hawawi and Al-Imam, 1984). On the Mediterranean coast of Egypt, spawning occurs from March to September with a peak in July and August. Maximum egg diameter is 2.2 mm and size at maturity for females is 30 mm total length (Lotan and Ben-Tuvia, 1996). Iranian specimens have large eggs (2.0 mm) on 16 March but young fish (8.9 mm standard length) were caught on 26 November suggesting reproduction is almost year round.

Spawning areas have some water flow and vertical surfaces of plants or algae-encrusted rocks. Males defend a territory about 30 cm wide against other males and will display with erect fins if approached. Parallel swimming, head to head confrontations, rapid circling, and flank bites were observed by Haas (1982). Two males will swim and chase each other in circles until one swims away with fins depressed. Males will change their territory after one day defending it. In Israel, males of Aphanius dispar richardsoni do not show any territorial behaviour (Goren and Rychwalski, 1978). This fish will spawn in a few months of birth in aquaria (Ross, 1987). Three pairs of fish yielded about 40 eggs per day in aquaria and these eggs hatched in two weeks at 27°C (Mackowiak, 1988).

Parasites and predators

This species is reported to be heavily infested with metacercariae of the trematode Clinostomum complanatum in eastern Saudi Arabia, up to 86.6% incidence with up to 41 metacercariae per fish, the infestation being higher in females than males (Kalantan et al., 1987).

Economic importance

Al-Akel et al. (1987) suggest the use of this species as a control agent for filamentous algae.

Steven (1913) reported that this fish was responsible for the absence of mosquito larvae from streams in Karachi which otherwise appeared to be good breeding places. Christophers and Shortt (1920-1921) thought that tooth-carps in southern Mesopotamia were a significant factor in lowering the malaria rate, presumably mostly this species. Sharma and Al-Daham (1979) compared the efficiency of this tooth-carp with the mosquitofish for mosquito control under experimental conditions. The tooth-carp is more efficient when males and females were together than mosquitofish since males of the latter expend time and energy defending territory. Mosquitofish are more active in pursuit of mosquito larvae than the tooth-carp when filamentous algae was present. Cloudy conditions and lower temperatures slow consumption and both species prefer pupae to larvae of mosquitos, with females taking more pupae than males. The tooth-carp could consume nearly twice the amount of mosquitos than the mosquitofish over a given period of time. Overall the tooth-carp compares favourably with the mosquitofish as a destroyer of the malaria-carrying mosquito. Homski et al. (1994) under laboratory conditions found that Aphanius dispar is more successful than the mosquitofish in preying on the third and fourth instars and on pupal stages of mosquitos while the mosquitofish, with a smaller relative mouth size, is more successful on the first two instars. Small fish of both species eat first instars exclusively. However, Aphanius dispar eats more second instar larvae under a vegetation cover since the larger Aphanius are more capable of penetrating shallow water than mosquitofish. Aphanius dispar occurs in shallower water and among more vegetation than the open water mosquitofish. The two species can be used to complement each other in control of mosquitos.

Ataur-Rahim (1981) found that this species will eat mosquito larvae in water storage tanks in Riyadh, Saudi Arabia. Fifteen fish consumed all larvae at a mean density of 5/100 sq cm water surface area in a tank 3 by 2 m in a day. One fish, 5.2 cm long, ate 61 mosquito larvae. Haas (1982) fed this species mosquito larvae in the presence of abundant algae as an alternative food. An average of 96 larvae are eaten each day. Etemadfar et al. (1983) studied the potential use of this species for malaria control in southern Fars, Bushehr and Hormozgan provinces. Louis et al. (1988) detail a control strategy against malaria using this species in Djibouti as do Fletcher et al. (1992) in Ethiopia and Fareed and Baban (1995) in Yemen. Fletcher et al. (1992) showed a decrease in sites harbouring mosquito larvae from 34% without the fish to 1.6% with the fish stocked. Chandra et al. (2008) briefly reviews the use of this species in biocontrol.

The studies of Frenkel and Goren (1997) in Israel show that the reproduction of this species can be controlled such that it has potential for mass production and could be used in mosquito control. Carpenter et al. (1997) state that this species is used in mosquito control. Ross (1987) gives details of aquarium care and breeding of this species.

Conservation

Al-Johany and Yousuf (1993) show that Gambusia affinis has a wider range of temperature tolerance than this species and is better suited for a desert habitat in Saudi Arabia. The exotic Gambusia is therefore a threat to A. dispar.

Further work

This widespread species has been reported from a number of springs in southern Iran and, like A. ginaonis, may be distinct at some level. Molecular work would be useful in determining the taxa involved and evolution in hot springs. The utility of this species in mosquito control in smaller water bodies of Iran where it also occurs naturally could be explored. Al-Daham et al. (1977) found that this species in aquaria ate mosquito larvae "eagerly" compared to "very eagerly" for Gambusia holbrooki and "not eaten" for Aphanius mento.

Sources

Type material:

Iranian material: NMW 13799, 1, ?; CMNFI 1979-0129, 1, 31.3 mm standard length, Fars, spring about 2 km north of Farrashband (28º54'N, 52º04'E); CMNFI 1979-0137, 106, 8.9-37.7 mm standard length, Fars, stream outside Lar (27º40'N, 54º32'30"E); CMNFI 1979-0138, 1, ? mm standard length, Fars-Hormozgan border, Rasul River drainage (ca. 27º32'N, ca. 54º58'30"E); CMNFI 1979-0139, 2, ? mm standard length, Fars-Hormozgan border, Rasul River drainage (ca. 27º25'30"N, ca. 54º59'E); CMNFI 1979-0140, 3, ? mm standard length, Hormozgan, Kul River drainage (27º14'N, 55º46'30"E); CMNFI 1979-0141, 6, ? mm standard length, Hormozgan, Kul River at road bridge (27º17'30"N, 56º03'30"E); CMNFI 1979-0142, 10, ? mm standard length, Hormozgan, Baghu River at Baghu (27º17'N, 56º28'E); CMNFI 1979-0143, 118, ? mm standard length, Hormozgan, Hasan Langi River drainage (27º21'N, 56º50'30"E); CMNFI 1979-0145, 1, ? mm standard length, Hormozgan, Geru River (26º55'N, 57º01'30"E); CMNFI 1979-0148, 21, ? mm standard length, Hormozgan, Sarzeh River (27º30'30"N, 56º15'30"E); CMNFI 1979-0151, 45, ? mm standard length, Hormozgan, Shur River drainage near Kahkom (28º02'N, 55º53'E); CMNFI 1979-0177, 37, ? mm standard length, Hormozgan, Sarzeh River drainage (27º33'N, 56º14'E); CMNFI 1979-0179, 13, ? mm standard length, Hormozgan, Sarzeh River drainage (27º36'N, 56º15'E); CMNFI 1979-0181, 9, ? mm standard length, Hormozgan, Kul River drainage (27º17'30"N, 56º03'30"E); CMNFI 1979-0182, 31, ? mm standard length, Hormozgan, stream on road to Bandar-e Lengeh (27º14'N, 55º46'30"E); CMNFI 1979-0185, 24, ? mm standard length, Hormizgan, stream in Rasul Rievr drainage (27º06'N, 55º45'E); CMNFI 1979-0187, 2, 26.2-33.2 mm standard length, Hormozgan, Sar Khun oasis (27º23'30"N, 56º26'E); CMNFI 1979-0312, 7, ? mm standard length, Baluchestan, dam on Bampur River (27º11'N, 60º36'E); CMNFI 1979-0313, 1, ? mm standard length, Baluchestan, Bampur River at Bangharabad (27º20'N, 60º46'E); CMNFI 1979-0328, 18, ? mm standard length, Baluchestan, jube 2 km south of Bampur River near Bampur (27º10'30"N, 60º21'E); CMNFI 1979-0352, 2, ? mm standard length, Khuzestan, marsh in Jarrahi River drainage (30º33'30"N, 48º48'E); CMNFI 1979-0354, 8, ? mm standard length, Khuzestan, Karun River tributary (30º31'N, 48º19'E); CMNFI 1979-0355, 1, ? mm standard length, Khuzestan, Karun River tributary (30º35'N, 48º22'E); CMNFI 1979-0362, 2, ? mm standard length, Khuzestan, jube in Karkheh River drainage (31º42'N, 48º33'E); CMNFI 1979-0379, 4, ? mm standard length, Khuzestan, Dez River (32º12'N, 48º27'E); CMNFI 1979-0381, 1, ? mm standard length, Khuzestan, stream west of Shushtar (ca. 32º10'N, ca. 48º35'E); CMNFI 1979-0384, 7, ? mm standard length, Khuzestan, Ab-e Shur drainage (32º00'N, 49º07'E); CMNFI 1979-0403, 25, check size range ?19.7-37.0 mm standard length, Bushehr, stream 23 km south of Kaki (ca. 28º11'N, ca. 51º43'E); CMNFI 1979-0406, 48, 15.6-35.8 mm standard length, Hormozgan, stream near Bandar-e Charak (26º48'N, 54º18'E); CMNFI 1979-0407, 3, ? mm standard length, Hormozgan, stream 30 km from Barvedun (26º51'N, 54º37'E); CMNFI 1979-0408, 106, 14.6-48.4 mm standard length, Hormozgan, Mehran River (27º04'N, 54º35'E); CMNFI 1979-0410, 10, ? mm standard length, Hormozgan, Mehran River (26º53'N, 55º17'E); CMNFI 1979-0414, 12, ? mm standard length, Hormozgan, pool on road to Tiab (27º05'N, 57º02'E); CMNFI 1979-0415, 1, ? mm standard length, Hormozgan, stream 18 km south of Genu (27º17'30"N, 56º20'E); CMNFI 2007-0053, 4, ? mm standard length, Hormozgan, Sarzeh River (ca. 27º36'N, ca. 56º15'E); CMNFI 2007-0057, 16, ? mm standard length, Hormozgan, Mehran River 4 km below Bastak (ca. 27º05'N, ca. 54º05'E); CMNFI 2007-0058, 3, ? mm standard length, Fars, headwaters of Gowdar River (ca. 27º24'N, ca. 54º16'E); OSU 8129, 1, 54.5 mm standard length, Hormozgan, Qeshm Island salt spring (no other locality data); BWC95-30, uncatalogued, 3, ?, Khuzestan, Kupal (31º15'N, 49º10'E); BWC98-6, uncatalogued, 24, ?, Hormozgan, Ab-e Garm-e Khamir (ca. 26º59'N, ca. 55º35'E); BWC2000-6, uncatalogued, ?, ?, Khuzestan, Dez River (32º14.663'N, 48º20.112'E); BWC2000-15, uncatalogued, ?, ?, Khuzestan, stream south of Shushtar (31º41.867'N, 48º59.581'E).

Comparative material:- BM(NH) 1920.3.3:193-202, 12, 21.0-30.5 mm standard length, Iraq, Basra (30º30'N, 47º47'E); BM(NH) 1920.3.6:9-10, 3, 17.1-26.8 mm standard length, Iraq, Basra (30º30'N, 47º47'E); BM(NH) 1949.7.21:1-11, 12, 18.4-42.4 mm standard length, Iraq, Bahr-el-Milk (32º35'N, 43º50'E); BM(NH) 1974.5.22:30, 1, 35.2 mm standard length, Iraq, Basra (30º30'N, 47º47'E); BM(NH) 1974.5.22:31-32, 2, 37.4-44.4 mm standard length, Iraq, Al Faw ().

Aphanius ginaonis
(Holly, 1929)

Male

Common names

gour-e khar, kopurdandan-e Genu or kapurdandan-e Geno (= Genow tooth-carp).

[Holly's pupfish, Genow pupfish, Gheno pupfish].

Systematics

Cyprinodon ginaonis was originally described from "Hei$e Quelle vom Djebel Ginao nordlich von Bender Abbas, südöstliches Persien". This is the Ab Garm-e Ganow at 27°26-28'N, 56°18-20'E, north of the Iranian port of Bandar Abbas at the Straits of Hormuz.

Berg (1949) places this species in the synonymy of Aphanius dispar stoliczkanus (see A. dispar above) and Villwock et al. (1983) regard it tentatively as a synonym of Aphanius dispar. Wildekamp (1993) is of the opinion that it may be a subspecies of Aphanius dispar. Hrbek and Meyer (2003) using mtDNA found this species to be deeply nested within the A. dispar clade. Villwock (2004) has used cross-breeding experiments that demonstrate this taxon and A. dispar are comparable to intraspecific crosses in other taxa. However, fin ray counts are non-overlapping and Reichenbacher et al. (2007) demonstrate differences in otolith morphology. Holly (1929b) reports a specimen of Cyprinodon dispar from the same spring. I was unable to verify the co-occurrence of two species in the hot spring through my own samples. Holly's specimen of A. dispar is in the Naturhistorisches Museum Wien (NMW 13799) but there are no field notes to confirm its locality with accuracy. It may have been collected in an adjacent stream. A. ginaonis is regarded here as a good species on morphological grounds (see below).

Four syntypes are in the Naturhistorisches Museum Wien (NMW 13800-13803). Holly (1929a) in his original description refers to 3 syntypes. One of these 4 fish was selected as a lectotype by F. Krupp (NMW 13800) and measures 21.8 mm standard length while the remainder measure 17.8-23.2 mm standard length.

Key characters

This species is distinguished from Aphanius dispar, the only other tooth-carp in this part of southern Iran by its locality and by having fewer total dorsal fin rays (5-7 as opposed to 8-11, mostly 9-10 in A. dispar from neighbouring drainages).

Morphology

Total dorsal fin rays 5(16), 6(38) or 7(6). Gut coiling is highly variable between individuals and is complex (illustrated in Coad (1980b)). Reichenbacher et al. (2007) give a description of otolith morphology. Chromosome number 2n=48, karyotype 14Sm + 34St and arm number 31 (Esmaeili et al., 2008).

Saefali (1999) has D4-7, Pelvic5-7, Pect 13-17, A6-10, ll25-32,vert 26-30, bars10-22, gr13-21 SEE DATA SHEETS

Sexual dimorphism

Females have a higher mean numbers of anal fin rays, total vertebrae and flank bars, and longer pelvic fins (Coad, 1980b). Seifali and Sheidai (2001) found rostrum to pelvic fin distance, rostrum to anal fin distance, weight, total distance (? total length), head depth, head distance (? length) and body depth to differ between the sexes.

Colour

Males have a brighter colouration than females although colouration is individually variable within sexes. The back is mottled black. The dorsal, anal, pectoral and pelvic fins are a light orange. Flank bars are alternately black tinged with orange and grey-white, light blue or light orange. The side of the head anteriorly is iridescent light blue to green and the opercle is dorsally orange, postero-ventrally blue. The iris is iridescent green and the anterior ventral head surface is light blue. The peritoneum is black.

Preserved specimens have two bars on the male caudal fin with the fin margin hyaline. These bars are usually straight. Some fish have 1-3 bars or a forked bar. Females lack these bars and their caudal fin rays are lightly speckled. The male dorsal fin has 4-5 horizontal rows of spots, fading dorsally. These are not always well-developed and then consist of irregular, black pigmentation on the rays and membranes. The dorsal fin pigmentation in females is weak. The anal and pelvic fins have little or no pigment. The pectoral fins of males carry pigment concentrated on the rays, weakly speckled in females. Flank bars in females vary from weak to obvious, the light intermediate bars not as highly coloured as in males nor as well defined.

Size

Reaches 4.5 cm total length.

Distribution

This species was described by Holly (1929a) from the Ab Garm-e Ganow at 27°26-28'N, 56°18-20'E (various reports give differing latitude-longitudes for this well-known locality, hence the range). Ab Garm literally means "hot water" in Farsi and is used to denote a hot spring. Ganow means "foul water". The spring lies at an altitude of about 135 m (Moghadam, 1974) on the slopes of the Ganow Mountain (= transliterated as Ginao in German), hence its scientific name.

Zoogeography

The occurrence of sympatric A. dispar would confirm the species distinction of this species.

Habitat

The species is common within the bounds of the hot spring stream, numbering in the thousands in the 1970s, but no population trends have been recorded (but see below). Fish are found particularly along the stream margin and in many minor subsidiary springs which emerge a few metres from the main spring. Some fish are found in water as shallow as 1 cm but temperatures are 37-40°C (when shade air temperature was 20°C at 1400 hours on 27 January). The main spring itself issues from the ground at 30 litres second^-1 and 41°C and drains as a stream about 10-15 m wide. A fault, over which the spring stream pours, isolates the fish from those downstream. Coad (1980b) summarises water chemistry. The water is clear and colourless but there is a strong sulphur odour. The stream bed is composed of stones and pebbles covered by lime-green to dark blue-green algae. Moghadam (1974) gives details of the algae species.

Age and growth

Essentially unknown although fish are larger later in the year (11.8-31.4 mm standard length in January and 15.6-34.7 mm in March). Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 33 fish measuring 1.70-3.17 cm standard length. The a-value was 0.0210 and the b-value 3.375 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Gut contents are the lime green algae typical of the hot spring and chironomids.

Reproduction

Temperature is constant and daylength may then be important in triggering reproduction, although fishes caught on 27 January and 21 March have mature eggs of similar size (ca. 1.6-1.7 mm) suggesting an extended reproductive period.

Parasites and predators

Unknown.

Economic importance

None except related species are important in the aquarium trade and this is a potential economic use.

Conservation

Coad (1980b) listed this species as rare although it should be classed as vulnerable under the revised IUCN Red List Categories (IUCN, 1996) according to Coad (1998a).

The site is easily accessible by an asphalt road and is close to the large city of Bandar Abbas. Two hammams (bath houses) are in operation, one for men and one for women, the spring water is drained off to irrigate date palms, and local people also use the site for bathing, washing kitchen utensils and food containers, and washing clothes so there is input of soap and food debris (personal observations; A. R. Zinaei and H. R. Esmaeili, Payam-e Noor University, Bandar Abbas, 6 August 1997). The area is not under direct management by environment officials although in 1999 press reports referred to planned research studies aimed at preventing extinction of this taxon (IRNA, 29 September 1999) and the spring does lie in the Genu Protected Area (Biosphere Reserve) described by Zehzad et al. (1997).

The habitat could deteriorate if too much soap and food debris enter the stream. A single catastrophic event, such as a chemical spill, water diversion or construction activity, could eliminate the species. Recent construction has severely affected the population with only 1-2 specimens being caught and the population being replaced by A. dispar (H. R. Esmaeili, pers. comm., September 2007).

Local people using the spring should be educated on its ecological significance and alternative facilities for bathing and washing established. Roadside access to the spring should be prevented. The population structure and reproductive regime of the fishes could be analyzed to determine if they are susceptible to abrupt changes.

Further work

This species differs from A. dispar in a number of characters but this may merely be a consequence of the unusual environment of the hot spring. Cross-breeding experiments would be very revealing and would demonstrate or disprove reproductive isolation and hybrid fertility. The population is very important as the only species in a unique habitat, lending itself to studies of speciation, adaptation and variation in response to high temperatures.

Sources

Type material: see above, NMW 13800-13803.

Topotypes: CMNFI 1979-0175, 77, 11.8-31.4 mm standard length. check size range; CMNFI 1979-0416, 7, ? mm standard length, Hormozgan, Ab Garm-e Ganow below falls (ca. 27º26'N, ca. 56º20'E); CMNFI 1979-0417, 44, ? mm standard length, Hormozgan, Ab Garm-e Ganow, side springs and edge of mainstream (ca. 27º26'N, ca. 56º20'E); CMNFI 2007-0054, 13, ? mm standard length, Hormozgan, Ab Garm-e Ganow (ca. 27º26'N, ca. 56º20'E);

Aphanius isfahanensis
Hrbek, Keivany and Coad, 2007

Common names

كپوردندان اصفهان (= kapurdandan-e Esfahan or Esfahan tooth-carp).

Systematics

The holotype is in the Canadian Museum of Nature, Ottawa under CMNFI 2004-0001, male, 25.0 mm SL, Esfahan Province, Zayandeh Rud (Zayandeh River) at Varzaneh Bridge, 32º25'32"N, 52º39'14"E, 1 July 2002, Y. Keivany and S. Asadollah and paratpes are under CMNFI 2004-0002, 18 males, 20.8–30.9 mm SL, 18 females, 22.0–38.4 mm SL, of 49 total (13 not used in meristic and morphometric analyses, total range in size 12.1-38.4 mm standard length), same locality as holotype; AMNH 233639, 1 male, 25.2 mm SL, 1 female, 21.6 mm SL, same locality as holotype; MRAC 2004-05-P-01-02, 1 male, 25.2 mm SL, 1 female, 21.6 mm SL, same locality as holotype; GenBank accession numbers AY593488, AY593489, AY593497, and AY593498.

Key characters

Distribution and colour pattern identify this species. Males can be distinguished from those of all other Iranian species by having distinct black edge on the dorsal, anal, and pelvic fins. The dorsal fin is covered with a high density of black blotches. Females can be distinguished from females of A. sophiae and A. vladykovi by having flank-bars rather than spots. It can also be distinguished from A. persicus by less well-defined bars terminating at a mid-flank stripe and a relatively light gray stripe at the caudal-fin base rather than a black spot or blotch. Flank-bars are also characteristic of females of Aphanius dispar, however.

Morphology

Morphometric data are given by Hrbek et al. (2007) but these characters cannot be used for a simple distinction from other Aphanius in Iran but only in multivariate space. This species is clearly distinguished at the genetic level from all other species of Aphanius. It has 82 molecular apomorphies – 19 transversions, two transversions/transitions (depending on comparison), and 61 transitions – that show fixed character state differences to homologous characters analyzed in A. sophiae, A. persicus, and A. vladykovi from Iran. Thirty-seven of these character states are also apomorphies when compared to A. anatoliae, A. danfordii, A. villwocki, A. asquamatus, and A. fasciatus from Turkey.

Dorsal fin rays 11-14, anal fin rays 10-13, pectoral fin rays 13-16, pelvic fin rays 4-6, lateral line scales 25-29, total gill rakers 10-13, precaudal vertebrae 10-13 and caudal vertebrae 15-18. Chromosome number 2n=48, karyotype 12Sm + 36St and arm number 30 (Esmaeili et al., 2008).

Sexual dimorphism

See colour below.

Colour

Male flank bars number 8-12, mean 10.2, significantly less than in A. sophiae at 10-21, mean 14.4. The bars are broad with interspaces about equal or slightly narrower. The bars extend from behind the head to the tail. Anterior bars fade on the belly, whereas, posteriorly on the caudal peduncle, they encircle the body. Dorsally, the head is dark gray and the body is lighter but still heavily pigmented with melanophores; the belly lacks pigmentation. The sides of the head are densely speckled with melanophores, more thinly on the ventral side; in most specimens the chin is darker than the rest of the ventral head surface. The eye is bounded ventrally and postero-ventrally by a thin line of black pigment. The defining male coloration is the black margins of the pelvic, anal, and dorsal fins. The dorsal and anal fins may present a halo effect, the margins being so dark in relation to the rest of these fins. The tips and outer margin of the pelvic fin are blackish. The anal fin has a broad, blackish margin with the rest of the fin light cream-colored. The dorsal fin has the blackest margin. The rest of the dorsal fin is variably blotched, the blotches being much lighter than the fin margin. Most specimens have a contrasting pigmentless area just below the fin margin. The pectoral fin has sparse pigmentation along rays and ventrally on the interradial membranes but lacks the concentrated black pigmentation seen on the pelvic fins. The caudal fin rays and membranes are sparsely pigmented, and the whole margin may be blackish but in most fish pigment is restricted to the upper and lower margins, the lower margin only, or is absent. Large females have a grayish dorsal surface to the head, a lighter back and upper flank covered with scattered melanophores, and a mid-flank stripe terminating on the caudal peduncle in a blackish, short stretch covering up to three scales (Fig. 2b). This short stretch of pigment is present in all females, faint in some, rarely forming a blotch and in some small fish tapering anteriorly. The flank stripe may be broken into a series of blotches in some smaller females, or it may be continuous as in large fish. Starting anterior to the belly there is a ventrolateral series of thin bars (up to ten) separated by cream colored interspaces 1–3 bars wide. These are absent in some smaller females, which may only have blotches at various levels present in this region. Even some large females have faint flank pigmentation so that bars and the stripe are weakly expressed. At about the origin of the anal fin, the flank bars may continue onto the caudal peduncle in regular form or become irregular, breaking up into blotches. Anteriorly the bars terminate ventrally at about the level of the lower edge of the pectoral fin. The belly and lower head have sparse pigment although the chin and sides of head are speckled with melanophores. The eye is bounded ventrally and posteroventrally by a thin line of black pigment. Fins lack any distinctive pigment pattern. Fin rays are outlined with melanophores, and interradial membranes of the caudal and anal fins have melanophores at varying degrees of density. The dorsal fin has the most interradial pigmentation, particularly near, but clear of the fin base.

Size

Attains 38.4 mm standard length.

Distribution

The type locality is near the town of Varzaneh on the lower reaches of the Zayanadeh River, about 30 km upriver from the terminal sump, the Gav Khuni marsh.

Zoogeography

An estimate of a 4.8 MYA divergence of A. isfahanensis and A. sophiae + A. persicus is given by Hrbek et al. (2007). This divergence time is in accord with the hypothesis of a near-simultaneous diversification of ~5 MYA of organisms occupying different geological units of central Iran.

Habitat

The type locality had a water temperature of 27ºC, pH was 6.7, the water was brackish, conductivity was 10.9 mS, dissolved solids were 5450 ppm, dissolved oxygen was 12.3 mg/L, river width was 50 m, and capture depth was 0.5 m. Current was slow, and there was no cover. Gambusia holbrooki was captured at the same locality.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None recorded.

Economic importance

None at present but a potential aquarium species. Breeding and aquarium conditions are similar to A. mento although it is reputedly difficult to breed.

Conservation

Known only from the type locality, which is not protected, this species needs a conservation assessment.

Further work

The biology of this species requires study.

Sources

Type material: See above, CMNFI 2004-0001, CMNFI 2004-0002, AMNH 233639, MRAC 2004-05-P-01-02.

Aphanius mento
(Heckel, 1843)

Male

Further illustrationsAphanius mento male; Aphanius mento female; after Berg (1949)?

Common names

gour-e khar, kopurdandandar-e Arvand (= Arvand River tooth-carp), kopurdand-e Irani.

[batrikh qabras in Arabic; Persian pupfish, Persian minnow, Black Persian minnow, Persian killie, Arvand pupfish, chin killifish, pearl-spotted killifish].

Systematics

Lebias Cypris Heckel, 1843 is a synonym. Krupp (1984a) gives a detailed synonymy of this species but considered cypris to have priority. Klee (1967) and Lazara (1989) point out that Garman (1895), in a work dated in July of that year, as first revisor placed cypris in mento. This decision has priority over Gaillard (1895), a work with no exact date of publication, in which mento was placed in cypris, according to the Zoological Code of Nomenclature (Ride et al., 1985). The type locality for both Lebias mento and Lebias cypris is "Mossul" according to Heckel (1843b).

Hybrids with Aphanius dispar richardsoni are reported from Israel (Goren and Rychwalski, 1978).

Parenti (1981; 1984) places this species in the genus "Aphanius", i.e. distinct from true Aphanius without defining and naming a new genus. This genus is more closely related to the Anatolian Kosswigichthys Sözer, 1942 and the South American Orestias Valenciennes in Cuvier and Valenciennes, 1846 than true Aphanius. Hrbek and Meyer (2003) note that, based on their mtDNA study, this species diverged from the ancestor of the A. dispar clade at an early date, 26.87±1.31 MYA. They speculate that the ancestor of mento invaded the northern perimeter of the Arabian plate as it was abutting Laurasia some time after the separation of their eastern and western clades of Aphanius (see above) before the closing of the Tethys Sea. It later spread through the present-day Tigris-Euphrates and into the Levant and southern Turkey.

The 5 syntypes of Lebias cypris are in the Naturhistorisches Museum Wien (NMW 59598) with 1 specimen selected as a lectotype by Krupp and Schneider (1989), the other 4 being paralectotypes. The syntypes of Lebias mento are (NMW 62105 (4 fish), 59042 (5) and 22624-22632 (9). Eschmeyer et al. (1996) list NMW 21699-704 (6) and NMW 59832 (21) as possible syntypes. The Vienna catalogue lists 5 specimens of Lebias cypris and 8 of Lebias mento.

Key characters

The adult colour pattern is distinctive and dorsal fin ray counts are usually higher than in A. dispar.

Morphology

Scales along side of body 23-28, total dorsal fin rays 9-14, usually 12-14, total anal fin rays 9-13, pectoral rays 12-16, and pelvic rays 4-6, usually 5. Frontal scalation is E-type (Hoedemann, 1958). Flank scales are squarish to a vertical oval. The former shape has a vertical anterior margin which may be wavy or slightly convex, upper and lower anterior corners rounded but square-cut, dorsal and ventral margins parallel, and the posterior margin rounded. The oval scales have all margins rounded. The focus is central to subcentral posterior, circuli are fine although coarser on the posterior field, radii are restricted to the anterior field and are almost horizontal and parallel. There is no pelvic axillary scale. Total gill rakers 11-15, rakers being spinulose and reaching beyond the adjacent raker when appressed. Vertebrae 25. Teeth are tricuspid with a long and pointed central cusp (Goren and Rychwalski, 1978). There are 14-16 teeth on the lower and 10-12 on the upper jaw (Berg, 1949). The gut is s-shaped. Reichenbacher et al. (2007) give a description of otolith morphology. Chromosome number 2n=44 or 48 (Wildekamp, 1993; Klinkhardt et al., 1995).

Sexual dimorphism

See below under Colour.

Colour

Adult, breeding males are a dark blue-black to dark brown or almost black with iridescent blue-white to silvery spots regularly-arranged on the fins as curved lines, and irregularly on the body (sometimes as irregular vertical bars and sometimes the spots are vertically elongate) (Richter, 1989; preserved material from Iraq; Goren and Rychwalski, 1978). The edges of the gill cover are orange-red (Wildekamp et al. (1999). Male colour fades in low light conditions and in winter males have silvery flanks with a dark brown back. Spots are silvery-blue on the upper flank and are not as numerous as in the spawning male. Females are grey-brown or grey-white to silvery with large golden blotches or silvery to blue spots and dark dots. Scales along the mid-flank usually have a dark border (Wildekamp et al. (1999). Fins in females are hyaline. Body colour is reportedly heightened in brackish water (Grimes, 1974). The peritoneum is silvery with dense but fine melanophores.

Size

Reaches 6.0 cm.

Distribution

Found in the Orontes (= Asi) and Tigris-Euphrates basins, the Levant in coastal and Dead Sea basins, western Jordan, and in southern Turkey in Mediterranean basins as well as in central Turkey. Tigris-Euphrates records are relatively rare (see map in Wildekamp (1993)). The species was described from Mosul in northern Iraq and there are records from the region of Basrah in southern Iraq near the border with Khuzestan. Wildekamp (1993) has one record apparently mapped in southern Iran near the border with Iraq. This is the only record from Iran although Abdoli (2000) indicates the Arvand River on a map.

Zoogeography

See Systematics above and under the genus.

Habitat

This species has been reported to survive in waters up to 38°C and as low as 0°C. pH at 7.0-8.5 has been reported as ideal for aquaria maintenance. Aquarists report it as a robust species favouring hard and alkaline water with some salt added (Thiermann, 1978; Kostich, 1979; Phillips, 1985). Grimes (1974) considers it to be quarrelsome in aquaria like other Aphanius and Allen (1988) also noted that males are very aggressive. It inhabits fresh or slightly brackish water of springs, streams and lakes, usually near shore where males establish territories in vegetation. It generally prefers habitats dense in vegetation.

Age and growth

Life span is about 3 years with maturity in aquaria at 4-6 months. The length-weight relationship in southern Iraq was W = 6.0001 x 10^-5 L^2.6403 for males and W = 2.3028 x 10^-4 L^2.235 for females. The average condition factor for males was 2.0499 and for females 2.3121. The condition factor increases with increase in length in females and decreases in males (Huq et al. 1977).

Food

Al-Daham et al. (1977) found filamentous algae and diatoms were the most important foods in southern Iraq. Shafi and Shalli (1986) report a diet of beetles, ephemeropteran nymphs and algae in southern Iraq. Aquarium specimens prefer meaty foods such as brine shrimps over flake foods (Grimes, 1974) but flake food is taken (Allen, 1988). Curiously, Al-Daham et al. (1977) for their southern Iraqi fish in aquaria found this species to prefer plant food. Males approached food items individually and prevented other fish from taking the food. Females were also solitary though less aggressive than the males. However Gambusia holbrooki out-competed male A. mento for food, on one occasion even snatching a Gambusia embryo from the mouth of an A. mento.

Reproduction

Anderson (1966), Lutman (1982) and Richter (1989) observed spawning in aquaria. There is a dominant male which is more deeply and richly coloured than other males. The dominant male defends a spawning territory, fighting other males. Males will defend separate spawning mops if these are made available in aquaria. The dominant male will pursue a female and places his larger dorsal fin over the her back. Several spawnings will follow and females will spawn with more than one male. Spawning can occur near the surface. These fish can spawn at six months of age. Peak spawning in southern Iraq is April to June when only a small proportion of eggs in the single ovary contained fully developed eggs (Al-Daham et al., 1977). However, Shafi and Shalli (1986) state that it breeds in southern Iraq in May-July. There are up to 71 highly-adhesive, eggs of 170 microns diameter. Aquarium spawned eggs hatch in 7-14 days, depending on temperature, 10 days at 25°, 3 days in the 80s°F, or 6-7 days in the low 70s°F (accounts vary). Allen (1988) found egg production to be higher in aquaria when the equivalent of seven tablespoons of salt was added per gallon at a water temperature of 25°C. A minimum of 20 eggs per day were produced.

Byniak (1979) describes the effects of day length and temperature on the reproductive cycle of females in this species in Israel.

Parasites and predators

Unknown.

Economic importance

An aquarium fish with breeding details given by Zipay (1961), Anderson (1966), Lutman (1982), Richter (1989), Semeit (1999) and www.killi-data.org/ (downloaded 21 September 2007). The species is hardy and fry are easily raised, given large enough tanks and an easily defensible area for the male to guard eggs. However, populations stop breeding in aquaria after 6-8 generations (www.killi-data.org/, downloaded 21 September 2007). Blaustein and Byard (1993) show that this species will prey on mosquito larvae in a laboratory situation and has potential in control of this vector of malaria, at least in vegetated habitats.

Conservation

The presence of this species in Iran needs confirmation by field work and until this is done and its distribution, population numbers and biology worked out, its conservation status cannot be assessed. Endangered in Turkey (Fricke et al., 2007).

Further work

See above.

Sources

Iranian material: None.

Comparative material: BM(NH) 1920.3.3:203-222, 22, 24.9-36.0 mm standard length, Iraq, Basra (30º30'N, 47º47'E); BM(NH) 1949.7.21:12, 24.4 mm standard length, Iraq, Bahr-el-Milh Lake east of Karbala (32º35'N, 43º50'E); BM(NH) 1981.10.6:19-23, 5, 26.6-31.8 mm standard length, Iraq, Qarmat Ali, Basra (30º30'N, 47º47'E); BM(NH) no catalogue number, 4, 19.7-31.7 mm standard length, Iraq, Basra Liwa (no other locality data); uncatalogued, 1, 31.2 mm standard length, Iraq, Khawr az Zubayr (no other locality data). ?Iraq fish from Hussain

Aphanius persicus
(Jenkins, 1910)

Male

Female

check male has bars on tail!

Common names

gour-e khar, kopurdandan-e Irani (= Iranian tooth-carp).

[Persian pupfish or toothcarp, but see also A. sophiae].

Systematics

Cyprinodon blanfordii Jenkins, 1910 and Cyprinodon pluristriatus Jenkins, 1910 are potentially synonyms. All are synonymised with A. sophiae by authors (see Coad, 1996a). Hanko (1924) placed Cyprinodon blanfordi and C. persicus in Cyprinodon chantrei Gaillard, 1895 but this is an error.

Jenkins (1910) described Cyprinodon blanfordii ("East of Shiraz, South Persia"), C. persicus ("Spring on the edge of Shiraz Lake, Southern Persia") and C. pluristriatus ("East of Shiraz, stream running to Fussa, Southern Persia, 5,000 feet"). The first nominal species has so generalised a locality as to be of very uncertain provenance. It could conceivably be in the Shiraz valley, the Kor River valley or outside either of these (see below under A. sophiae where it is argued that is is a synonym of this species). The second nominal species is clearly from the valley of Shiraz and its salt lake Maharlu, an endorheic basin. The last nominal species was found near Fussa (= ? Fasa) and, based on maps, is probably in the drainage of the Mand River which flows to the Persian Gulf.

The valid name for Lake Maharlu basin Aphanius is given here as persicus. The types in Calcutta have not been examined by me. Cyprinodon blanfordii has page priority but the locality is vague and the fish have a pigment pattern description and illustration which is spotted on the flank with a lozenge-shaped spot at the caudal fin base, thus appearing to be a female A. sophiae. Cyprinodon pluristriatus appears last in the descriptions and may be an available name for Aphanius outside the Lake Maharlu basin in rivers draining to the Persian Gulf should these prove distinct.

Material described and illustrated by Villwock (1960) as A. sophiae are this species.

Three syntypes of Cyprinodon blanfordi (ZSI F9416 to ZSI F9418), two syntypes of Cyprinodon persicus (ZSI F9403 and ZSI F9404) and four syntypes of Cyprinodon pluristriatus (ZSI F9408 to ZSI F9411) are in the Zoological Survey of India, Calcutta (Menon and Yazdani, 1968), although Jenkins (1910) has ZSI F9408-9412 for C. pluristriatus.

Key characters

Females of this species are barred where all other Aphanius species in this area of southern Iran have spotted females. The distribution is apparently limited to the Lake Maharlu basin near Shiraz.

Morphology

Although populations are isolated in springs and streams around Lake Maharlu, Coad (1996a) found them to be relatively homogenous.

Lateral line scales 24(3), 25(34), 26(69), 27(176), 28(100) or 29(22). Scales above the lateral line 4-6, scales between lateral line and the anal fin 4-8, scales between the lateral line and the pelvic fin 5-9, and scales around the caudal peduncle 12-17. Esmaeili et al. (2007) determined a chromosome number of 2n=48 with 11 pairs of submetacentric and 13 pairs of subtelocentric chromosomes and an arm number of NF=70. Karyotype 16Sm + 32St and arm number 32 (Esmaeili et al., 2008).

Total dorsal fin rays 8(1), 10(8), 11(97), 12(204), 13(89), or 14(7); total anal fin rays 9(3), 10(111), 11(225), 12(64) or 13(3); total pectoral fin rays 13(9), 14(82), 15(211), 16(94), 17(9) or 18(1); total pelvic fin rays 4(11), 5(181) or 6(213); total gill rakers 9(5), 10(51), 11(220), 12(108), 13(19) or 14(3); abdominal vertebrae 9(1), 10(1), 11(27), 12(355), or 13(22); and caudal vertebrae 12(1), 14(16), 15(285) or 16(104).

Sexual dimorphism

Most apparent in colour and pigmentation detailed below. Females are longer and heavier than males (Esmaeili and Shiva, 2006).

Colour

"The basic colouration is bright shining silvery white. The entire body, with the exception of the dorsal crest and the belly, is tigered with small brown spots; in some specimens these spots seem to be arranged in three irregular longitudinal rows of which the uppermost extends as a band across the operculum and to the eyes; all fins are uniformly yellow with a reddish border. In males the spots are more dense, in females they are paler and scarcer, only the 2-3 at the base of the caudal fin are dark brown" (Heckel (1846-1849b) on A. sophiae). "The colour of this small fish with its enviable crystal teeth is uniformly brown, at least it is now in ethyl alcohol, with a silvery-white throat and belly; laterally on the tail there are a few scattered, darker spots, and a deeper black spot is located on the last scales anteriorly of the caudal fin. All fins are blackish without any markings" (Heckel (1846-1949b) on A. crystallodon). "Silvery, brownish on back, brown on top of the head, with numerous small spots of brownish, arranged in three or more, more or less irregular, longitudinal series, the median of which, on the lateral line, commonly ends in a black spot on the middle base of the caudal. On some this median series is entirely composed of blackish spots. Rarely the spots are somewhat confluent into longitudinal bands. Brownish specimens show a longitudinal band of silvery along the middle of the flank. The form described as A. crystallodon has spots only towards the base of the caudal, around the black spot usually ending the median series" (Garman, 1895).

The flank in females bears numerous alternating light and dark bars, the light bars varying in width from about one half to twice that of dark bars. The bars gradually merge with background pigmentation anteriorly on the flank and while clearly defined on the rear flank are difficult to distinguish anteriorly.

The caudal fin spot in females can be oval, tear-drop shaped or elongate but is usually in the form of a lozenge. Occasionally, single, smaller, dark, subsidiary spots may be found antero-dorsally and antero-ventrally to the basal spot or scattered spots may be found irregularly before and behind the basal spot.

Males have a pigmentation very similar to that of A. sophiae and the description here is identical. Some minor observed variation is attributed to variation in size and maturity of the fish compared. Males have light flank bars half the width or much narrower than alternating dark bars. The margins of the dorsal, anal and caudal fins are clear while the rest of these fins is dark. Some fish have up to 3, but usually 2, thin, light bars on the basal half of the caudal fin; these are generally larger fish. The margin of the lower half of the pectoral fin has concentrations of pigment on the membranes such that this area is darker than the rest of the fin. The anal fin is darkest posteriorly where pigment is concentrated on membranes. The distal third of the fin is pigmented to form a dark band, becoming lighter proximally. The dorsal fin is the darkest fin (except for the clear margin) and the anterior base is the darkest part of the fin. Bands are not always evident but pigment spots are large proximally. Some fish have 2, sometimes 3, thin light bars at the base separated by thin dark bars and paralleling the body profile while others have none. The dorsal fin base may have instead a series of lighter spots, sometimes irregular and not paralleling the body profile.

Size

Reaches 59.5 mm standard length (Esmaeili and Shiva, 2006).

Distribution

This species is restricted to the Lake Maharlu basin near Shiraz.

Zoogeography

This species is found only within the endorheic Lake Maharlu basin, Coad (1996a) suggests that tooth-carps in inland waters may have risen with the post-Pliocene uplift of the Zagros Mountains rather than being the result of relatively recent inter-basin dispersal.

Habitat

This species is found in fresh streams and springs and in springs of varying saline content or saline influence from hypersaline chloride Lake Maharlu. The lake at 124‰ is an impassable barrier to dispersal but is very shallow and dries out periodically, e.g. in 1967 (Cornwallis, 1968a). The larger springs and their streams can then meet and transfer fishes on the dried-out lake bed. Some springs are 27 km apart along the lake margin and have probably had no contact or fish exchange over many generations. This is especially the case for the smaller springs which emerge from the ground with a diameter of only 1 m and restricted flow. When lake water rises, lower springs are inundated with hypersaline water and the fishes are killed. The spring is presumably recolonised from a higher spring when lake level falls.

Fish placed in pure lake water die within minutes. Springs discharging directly into the lake contain tooth-carps but the fish do not venture into the lake. Some fish, if disturbed or deliberately harassed, will swim out for a short distance into the hypersaline lake water but they blanch and rapidly retreat to fresher water (Coad, 1996a).

Esmaeili and Shiva (2006) found the bottom of these water bodies to be muddy but the water was clear and slow-running. Conditions in October at three different sites were 16.9-19.0ºC, pH 6.70-6.74, dissolved oxygen 3.96-6.11 mg/l, nitrate 0.9-1.6 mg/l, nitrite 0.029-0.062 mg/l, phosphate 0.35-0.65 mg/l and ammonium 1.55-2.60 mg/l.

Age and growth

In the studies of Esmaeili and Shiva (2006), the sex ratio was 1.67:1 for females:males, highly significant, the age groups were 0+ to 3+ years, and positive allometric growth was determined (b value significantly greater than 3).The condition factor for females was highest in February and March and lowest in September while in males it was lowest in December and increased until February. Fishes smaller than 25 mm had an equal sex ratio, suggesting selective predation on males, or better survival or longevity of females. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 62 fish measuring 1.86-4.27 cm standard length. The a-value was 0.0222 and the b-value 3.395 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Unknown but presumably similar to other tooth-carps.

Reproduction

Esmaeili and Shiva (2006) found the gonadosomatic index in females to increase from November to June, decreasing slowly from late June to November. The reproductive period lasted six months. Males had two peaks, in April and August. Egg diameters reached 1.71 mm, absolute fecundity ranged from 45 to 250 eggs (average 115.7 eggs) and relative fecundity was 21.6 to 244.1 with a mean of 90.01 per gram body weight. Even small fish had hydrated eggs and, with the extended reproductive period, shows adaptation to an unstable habitat.

Parasites and predators

Mokhayer (1989) reported metacercariae of the eye fluke, Diplostomum spathaceum from Aphanius sophiae, probably this species, in Iran. The fluke can cause complete blindness and death in commercially important species of fish. This fish was also infested with yellow grub, Clinostomum complanatum. González-Solís et al. (1997) report Contracaecum sp. larvae from this species in the Lake Maharlu drainage, Fars.

Economic importance

This species has no current economic importance although it would make an excellent aquarium fish. Villwock (1959) gives details of maintaining what seems to be this species under aquarium conditions, gradually transferring them from a mixture of salty and magnesium sulphate water to stagnant tap water. They prefer some admixture of sodium chloride and magnesium sulphate which matches their natural habitat (2% synthetic sea water plus a tablespoon of epsom salts per 25 litres water). Aquatic plants such as Myriophyllum and Cabomba are also necessary for breeding as the eggs are attached to them but artificial substitutes can be used. Yellowish transparent eggs are 1.5 mm and are deposited usually individually, rarely in small groups. Larvae feed on Artemia salina nauplii and adults on daphnia, red mosquito larvae and tubifex. Male-male interactions include mock fights with spread fins and close swimming. The loser hides in the algae and is not strongly pursued by the victor. These interactions, and those between sexes, are not strongly developed and aquaria can be stocked at 15-20 fish per 25 litres. Y. Keivany notes (pers. comm., 2004) that this species is difficult to maintain in aquaria.

Conservation

The populations are found in a variety of springs and streams around Lake Maharlu. Some of these localities are threatened by water abstraction, road construction and pollution but a number of localities are small and/or saline and not under threat.

Further work

Studies on aquaria maintenance would enable it to become established in the aquarium trade and help its conservation.

Sources

Type material: None seen.

Iranian material: CMNFI 1979-0017, ?47, ?17.9-35.5 mm standard length, Fars, stream at Pol-e Fasa (29º29'N, 52º38'30"E); CMNFI 1979-0018, 75, ? mm standard length, Fars, Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0033, 1, ? mm standard length, Fars, Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0034, ?124, 14.8-38.7 mm standard length, Fars, spring on shore of Lake Maharlu (29º27'N, 52º44'E); CMNFI 1979-0035, 17, ? mm standard length, Fars, spring south of Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0038, 16, ? mm standard length, Fars, spring near Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0039, 91, 15.9-37.9 mm standard length, Fars, spring on shore of Lake Maharlu (29º27'N, 52º44'E); CMNFI 1979-0041, 61, 20.8-41.6 mm standard length, Fars, spring on shore of Lake Maharlu (29º23'N, 52º48'E); CMNFI 1979-0042, 28, ? mm standard length, Fars, spring on shore of Lake Maharlu (29º23'N, 52º48'E); CMNFI 1979-0046, 5, 18.1-32.2 mm standard length, Fars, qanat at Barm-e Dalak (ca. 29º35'N, ca. 52º38'E); CMNFI 1979-0047, 41, 11.3-32.1 mm standard length, Fars, spring source of Ab-e Paravan marshes (ca. 29º34'N, ca. 52º42'E); CMNFI 1979-0048, ?45, ?15.3-39.9 mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º32'N, ca. 52º48'E); CMNFI 1979-0049, 12, 17.2-38.5 mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º32'N, ca. 52º48'E); CMNFI 1979-0050, 27, 18.3-28.9 mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º31'30"N, ca. 52º49'30"E); CMNFI 1979-0051, 3, ? mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º30'N, ca. 52º52'E); CMNFI 1979-0052, 16, ? mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º30'N, ca. 52º52'E); CMNFI 1979-0064, 10, ? mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º31'30'N, ca. 52º49'30"E); CMNFI 1979-0065, 8, ? mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º31'N, ca. 52º50'E); CMNFI 1979-0066, ?31, 15.6-42.9 mm standard length, Fars, spring on shore of Lake Maharlu (ca. 29º32'N, ca. 52º48'E); CMNFI 1979-0110, 2, ? mm standard length, Fars, lake in Park-e Shahr, Shiraz (29º38'N, 52º32'E); CMNFI 1979-0112, 24, 11.9-45.9 mm standard length, Fars, stream at Pol-e Fasa (29º29'N, 52º38'30"E); CMNFI 1979-0118, 12, ? mm standard length, Fars, Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0119, 75, ? mm standard length, Fars, steram at Pol-e Fasa (29º29'N, 52º38'30"E); CMNFI 2004-0006 (GenBank AY593484, AY593493), 15, ?22.2-41.8 mm standard length, Fars, Nasrabad Spring near Imamzadeh Ibrahim (29º35'09"N, 52º39'08"E);

Aphanius sophiae
(Heckel, 1849)

Male

Female

Common names

gour-e khar, kopurdandan-e safiyeh or kopurdandan-e sufieh (= sufi or sophy tooth-carp).

[batrikh sophiae in Arabic; Persian minnow, Cypris pupfish, but this seems inapt].

Systematics

The type locality of Lebias Sophiae is in "lauen Salzquellen bei Persepolis" according to Heckel (1846-1849b). Lebias punctatus Heckel, 1849 listed from "Nemek-Deria oder Salzsee, in welchen sich unter Schiraz die Quellen des Saadi ergiessen" or "the salty Maharlu Lake below Shiraz into which the Saadi springs flow" (Heckel, 1846-1849b) and Lebias crystallodon from "grossen Salzsee Nemek-Deria, unter Schiraz" (Heckel, 1846-1849b) are synonyms (Günther, 1859-70; Garman, 1895; Gaillard, 1895; Hanko, 1924; Berg, 1949; Coad, 1996a). The original catalogue handwritten by Heckel in Vienna gives the type locality as "Salzquellen bei Persepolis" which is in agreement with Heckel's publication. Lebias punctatus and Lebias crystallodon are described from localities in the Shiraz valley wherein lies the salt lake Maharlu. The catalogue is more equivocal for these two nominal species, listing only "Nemek Deria". The labels in jars containing L. crystallodon and L. punctatus were written under Franz Steindachner's curatorship and are not original (H. Ahnelt, in litt., 1987). Nemek Deria (= salt lake in Farsi) is a general term employed for the type of terminal water body seen in both the Maharlu and Kor River basins. The situation was further confused by Berg (1949) who listed the locality of punctatus as "Lake Niriz in Shiraz". Lake Niriz (= Neyriz or Bakhtegan) is the terminal sump for the Kor River.

All material of Aphanius from the Kor River basin examined fresh by me comprises fish with bars along the flank and fish with spots. The barred fish are males and the spotted fish females as confirmed by dissection. Males and females do not segregate in the field and it seems very unlikely that the material examined by Heckel from several collections would be comprised solely of males (I found sample sizes as low as 5 to contain both males and females). The males and females may have been separated and allocated different collection localities. The material described as L. punctatus is then female L. sophiae and I strongly suspect that it was misunderstood as coming from the Shiraz valley and the locality in Heckel (1846-1849b) is incorrect. All female Aphanius from the Shiraz valley are barred and the name punctatus is singularly inappropriate and would not be applied to them (see illustrations in Villwock (1960; 1977)). L. crystallodon could be a female L. sophiae with weak spotting on the flanks, again mislabelled as from the Shiraz valley. Günther (1866) considers crystallodon to be a post-spawning female of L. sophiae and Gaillard (1895) also considers it to be a female L. sophiae. Berg (1949) places L. sophiae as the male and L. punctatus and L. crystallodon the females of L. sophiae.

Cyprinodon blanfordii Jenkins, 1910 may well be a synonym of A. sophiae (see above under A. persicus for other views). The locality is vague, "East of Shiraz, South Persia" and could lie in the Lake Maharlu basin (occupied only by A. persicus, the females of which are barred on the flank), the Kor River basin (A. sophiae only), or outside either of these, in rivers draining to the Persian Gulf. The illustration shows a finely-spotted fish with a lozenge-shaped larger spot at the tail base, pigmentation found in A. sophiae. Cyprinodon pluristriatus Jenkins, 1910 is often referred to the synonmy of A. sophiae but is found "East of Shiraz, stream running to Fussa, Southern Persia, 5,000 feet". Assuming this to be Fasa and using topographic maps, the locality is in the Mand River basin draining to the Persian Gulf. The illustration and description refer to male fish and fall within the parameters of A. sophiae, a species thought to be restricted to an endorheic basin. It may be an available name for a taxon occurring outside the endorheic Kor River basin where A. sophiae is endemic.

Parenti (1981; 1984) places Aphanius sophiae in the genus "Aphanius", i.e. distinct from true Aphanius without defining and naming a new genus. This genus is more closely related to the Anatolian Kosswigichthys and the South American Orestias than true Aphanius.

The syntypes of Lebias sophiae are in the Naturhistorisches Museum Wien and include a wide range of material of which the following is part:-

Lectotype and paralectotypes (as labelled in 1997): NMW 14496 (6 fish measuring 25.8-30.4 mm standard length, one of which measuring 29.3 mm standard length is the lectotype as designated by F. Krupp, 24 October 1984 according to a jar label), NMW 22616-22623 (8, 26.1-29.0 mm standard length), NMW 60327 (8, 25.2-31.1 mm standard length), NMW 68283 (8, 18.9-27.6 mm standard length, partially dried and distorted), NMW 75067 (7, 24.7-27.4 mm standard length), and probable syntypes NMW 14760 (67) and NMW 15056 (7). The catalogue in Vienna lists 40 fish in one column and 12 fish in the adjacent column, the latter possibly meant to be the type series as it is written more boldly but which fish these were cannot now be ascertained. All these fish are males.

Eschmeyer et al., (1996) list syntypes under NMW 14496 (6), NMW 33616-23 (8) (not noted above), NMW 60327 (8), NMW 68283 (8), NMW 75067 (7) and also 9 fish in the Museum für Naturkunde, Universität Humboldt, Berlin under ZMB 31377 (not noted above). The Berlin material is listed as ex 16140, is all male and 20.3-26.7 mm standard length.

Syntypes of Lebias punctatus comprise the following material:-

Lectotype and paralectotypes (as labelled in 1997): NMW 76509 (7 fish, measuring 29.9-36.0 mm standard length, one of which measuring 36.0 mm standard length is the lectotype as designated by F. Krupp, 24 October 1984 according to a jar label), NMW 15070 (5, ca. 29.9-35.6 mm standard length, dried and distorted), NMW 15156 (14, 19.2-32.1 mm standard length), NMW 59837 (5, 26.6-35.7 mm standard length), and probable syntypes NMW 59609 (6, ca. 21.4-34.7 mm standard length, smallest dried). The Vienna catalogue lists only 12 fish opposite this name. These fish are all females.

The holotype of Lebias crystallodon measuring 40.1 mm standard length is NMW 15175 as the description and catalogue agree there is only one fish. The specimen is decoloured and in poor condition.

Key characters

The large, dark, usually lozenge-shaped spot at the caudal fin base in females with fine spotting on the flanks is distinctive for tooth-carps in this area of southern Iran.

Morphology

Lateral line scales 25(1), 27(7), 28(14), 29(17), 30(12) or 31(2). Scales above the lateral line 4-7, scales between lateral line and the anal fin 5-8, scales between the lateral line and the pelvic fin 6-9, and scales around the caudal peduncle 15-20.

Total dorsal fin rays 11(1), 12(11), 13(27), 14(12) or 15(2); total anal fin rays 10(1), 11(18), 12(27) or 13(7); total pectoral fin rays 14(1), 15(7), 16(31), 17(10), 18(3) or 19(1); total pelvic fin rays 5(9) or 6(44); total gill rakers 10(2), 11(28), 12(21) or 13(2); abdominal vertebrae 11(11), 12(37), or 13(5); caudal vertebrae 15(3), 16(33) or 17(17); and total vertebrae 27(8), 28(29) or 29(16).

Cephalic sensory pores are reduced to a series of neuromasts. Older literature on chromosome number was 2n=48 but is based on fish from outside the Kor River basin to which the species is restricted here (Klinkhardt et al., 1995). Esmaeili et al. (2007) determined a chromosome number of 2n=48 with 14 pairs of submetacentric and 10 pairs of subtelocentric chromosomes and an arm number of NF=76. Karyotype 8Sm + 40St and arm number 280 (Esmaeili et al., 2008).

Sexual dimorphism

Males have more bands on the flank and are darker and more intensely coloured than females. Male dorsal, anal and caudal fins are a deep black with narrow, silvery-white borders and the anal fin has a few silvery spots at its base. Females have brown dorsal, anal and caudal fins with 3-4 rows of black spots and the anal fin also has 2 cross rows of white silvery spots near the base. The caudal fin spot in females can be oval, tear-drop shaped or elongate but is usually in the form of a lozenge. Occasionally, single, smaller, dark, subsidiary spots may be found antero-dorsally and antero-ventrally to the basal spot or scattered spots may be found irregularly before and behind the basal spot. Male flank bars 10(4), 11(1), 12(2), 13(3), 14(5), 15(4), 16(6), 17(3), 18(1) or 21(1).

Eleven of 21 morphometric characters are significantly different between the sexes including head length, head and body depths, fin lengths, and caudal peduncle shape (Coad, 1998i).

Colour

"The body colouration is basically dark brown; the body, with the exception of the forward dorsal portion and the belly, is marked by 12-17 white, silvery shining vertical lines or narrow bands. Those individuals which have the most bands and are also darker and more intensively coloured seem to be males; their vertical fins are deep black with narrow silvery-white borders, and only the anal fin has a few silvery spots at the base. The lighter individuals which are also slightly higher and wider, we believe to be the females; they have brown vertical fins with 3-4 cross rows of black spots; their anal fins are marked with 2 cross rows of white silvery spots at the base" (Heckel, 1846-1849b).

The caudal fin spot in females can be oval, tear-drop shaped or elongate but is usually in the form of a lozenge. Occasionally single, smaller, dark, subsidiary spots may be found antero-dorsally and antero-ventrally to the basal spot or scattered spots may be found irregularly before and behind the basal spot.

The flank spots in females are much lighter than the caudal base spot and are from a half to much less in size, usually much smaller than half size. Spots are usually rounded but can be oval and vertically elongate. Spots are independent of scale arrangement on the lower half of the flank, particularly posteriorly and may form 2-3 poorly defined to distinct longitudinal rows. Upper flank scales have a crescent of pigment in mid-scale, leaving the margin and anterior base mostly free of pigment, or pigment may fill each exposed scale surface almost entriely except for the posterior margin, and thus appear as spots. Females have purplish tints on mid-flank and yellowish tints on the lower flank, upper flank, head top and chin.

Males have a pigmentation very similar to that of A. persicus and the description here is identical. Some minor observed variation is attributed to variation in size and maturity of the fish compared. Males have light flank bars half the width or much narrower than alternating dark bars. The margins of the dorsal, anal and caudal fins are clear while the rest of these fins is dark. Some fish have up to 3, but usually 2, thin, light bars on the basal half of the caudal fin; these are generally larger fish. The margin of the lower half of the pectoral fin has concentrations of pigment on the membranes such that this area is darker than the rest of the fin. The anal fin is darkest posteriorly where pigment is concentrated on membranes. The distal third of the fin is pigmented to form a dark band, becoming lighter proximally. The dorsal fin is the darkest fin (except for the clear margin) and the anterior base is the darkest part of the fin. Bands are not always evident but pigment spots are large proximally. Some fish have 2, sometimes 3, thin light bars at the base separated by thin dark bars and paralleling the body profile while others have none. The dorsal fin base may have instead a series of lighter spots, sometimes irregular and not paralleling the body profile.

Size

Reaches 5.6 cm total length.

Distribution

This species is restricted to the Kor River basin in Fars.

Zoogeography

As noted in the genus account, Aphanius species in Southwest Asia are regarded as relicts of the Tethys Sea. The mountain populations of A. sophiae (and A. vladykovi) may well have risen with the post-Pliocene uplift of the Zagros Mountains. Since Aphanius species have a low dispersal ability (Kosswig, 1967) it is considered unlikely that these small fishes dispersed from lowland populations into an existing, high mountain range represented today by the Zagros.

Bobek (1963) suggests that there may have been an outflow from the Kor River basin to the Persian Gulf at the south-east corner which was cut off at the end of the Pleistocene by alluvial fans. Krinsley (1970) maintains that any outlet was closed by the late Pliocene. Evidence for the isolation time of Kor River basin tooth-carps is equivocal.

Habitat

Tooth-carps identified as this species has been reported to survive in waters up to 39°C and salinities as high as 130‰. In aquaria lives at 5-37°C. It is found in springs, streams and small pools and lakes in fresh and saline waters.

Age and growth

Unknown. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 64 fish measuring 2.18-4.64 cm standard length. The a-value was 0.0343 and the b-value 2.787 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Unknown in detail.

Reproduction

Unknown. Incubation is said to be 10 days at 25°C but this is a general citation for Aphanius spp. in aquaria.

Parasites and predators

Unknown.

Economic importance

This species has importance as an aquarium species, being colourful and hardy.

Conservation

Construction of irrigation canals and drying of some springs because of man-induced changes in the water table may threaten populations of this species.

Further work

The distribution of this species and its population numbers should be documented by field work. Molecular and chromosomal techniques would prove useful information for relating this species with others from neighbouring basins and throughout central Iran. Its biology is poorly known under natural conditions.

Sources

Type material: See above, Lebias sophiae (NMW 14496, 22616-22623, 60327, 68283, 75067, 14760 and 15056, Lebias crystallodon (NMW 15175), and Lebias punctatus (NMW 76509, 15070, 15156, 59837 and 59609).

Iranian material:- CMNFI 1979-0025, 1, 31.6 mm standard length, Fars, Kor River at Marv Dasht (29º51'N, 52º46'30"E); CMNFI 1979-0059, 30, 18.1-39.3 mm standard length, Fars, Pulvar River 8 km south of Sivand (30º01'30"N, 52º57'E); CMNFI 1979-0061, 3, 23.2-32.1 mm standard length, Fars, stream tributary to Pulvar River (30º04'N, 53º01'E); CMNFI 1979-0062, 6, 24.0-42.0 mm standard length, Fars, spring 17 km south of Sa'adatabad (30º05'N, 53º00'E); CMNFI 1979-0067, 1, 25.7 mm standard length, Fars, qanat at Zarqan (ca. 29º46'N, ca. 52º43'E); CMNFI 1979-0071, 6, ? mm standard length, Fars, qanat stream on road to Ramjerdi (ca. 30º00'N, ca. 52º38'E); CMNFI 1979-0091, 3, 23.2-32.1 mm standard length, Fars, stream tributary to Pulvar River (30º04'N, 53º01'E); CMNFI 1979-0117, , mm standard length, ; fish? CMNFI 1979-0155, 2, ? mm standard length, Fars, spring at Gavonoo (28º47'N, 54º22'E);ID? CMNFI 1979-0292, 6, 26.5-34.4 mm standard length, Fars, Lapu'i spring near Zarqan (29º48'N, 52º39'E); CMNFI 1979-0342, 1, 39.4 mm standard length, Fars, Kor River at Band-e Amir (29º49'N, 52º51'E); CMNFI 1979-0498, 6, 17.4-25.2 mm standard length, Fars, spring in Kor River basin (30º05'N, 52º27'E); CMNFI 2004-0003 (GenBank AY593483, AY593492), 47, ?20.7-28.8 mm standard length, Fars, Abdolmahdi Spring (30º06'12"N, 52º58'38"E); CMNFI 2004-0004 (GenBank AY593482, AY593492), 38, ?19.5-34.0 mm standard length, Fars, Malasskuh Spring (29º52'04"N, 52º29'20"E); uncatalogued (GenBank AY593481, AY593490), 6, 25.7-37.0 mm standard length, Fars, Dolatabad Spring (29º43'05"N, 52º50'11"E);

Aphanius vladykovi
Coad, 1988

Male

Female

Common names

gour-e khar, mahi-e gour-e khari (= "striped donkey" or zebra fish), kopurdandan-e Zagros (= Zagros tooth-carp), mahi parchami (= flag fish).

[Zagros pupfish].

Systematics

The type locality is in the Shahrestan-e Bakhtiari va Chahar Mahall in a large pool, 3 km west of Boldaji at 31°57'N, 51°01'E. The male holotype, 36.6 mm standard length, is in the Canadian Museum of Nature, Ottawa under CMNFI 1979-0247 with 35 male and 16 female paratypes from the same locality under CMNFI 1979-0247A and 1 male paratype from a stream 3 km east of Boldaji at 31°55'N, 51°05'E under CMNFI 1979-0248. Villwock (2004) has used cross-breeding experiments that demonstrate this species, A. sophiae and A. dispar are genetically isolated and distinct species.

Key characters

This species is distinguished from all other Aphanius in Iran and Southwest Asia by the higher number of normal-sized lateral line scales (36-47). Other members of Aphanius, and the related Kosswigichthys, have 35, usually 30, or less in the lateral series. The high number of scales is matched only by the genus Anatolichthys Kosswig and Sozer, 1945 of southwest Anatolia. Anatolichthys is polymorphic for scale counts which reach as high as 55 and shows scale loss, perhaps associated with the bitter, salty lakes in which it lives (Coad, 1988i). Male dorsal fin colour is much darker and the anal fin is light compared to A. sophiae, which most closely resembles A. vladykovi. Females lack the typical large spot at the base of the caudal fin found in A. sophiae.

Morphology

Scales are relatively small and numerous over the whole body as exemplified by scale counts. Scales are regularly arranged, embedded and imbricate. Anterior flank scales are a vertical oval with circuli and radii restricted to the posterior field. Numbers of circuli and radii are body and scale dependent and so are fewer in this species than in A. sophiae. There are up to 21 teeth in the upper jaw and teeth are in a single row in each jaw. Gill rakers are short, just reaching the adjacent raker when appressed. All fins are rounded distally and the anal fin is enclosed by a fleshy sheath, most apparent in large females but is also present in males. The gut has a single large loop. Cephalic sensory pores are reduced to a series of neuromasts. Keivany (2003) gives details of cephalic osteology. Pazooki et al. (2008) examined interlocality morphological variation, finding one of four populations to be more distinctive.

Lateral line scales 36(2), 37(8), 38(15), 39(6), 40(10), 41(6), 42(3), 43(1), 46(1) or 47 (1). Scales above the lateral line 6-9, scales between lateral line and the anal fin 8-12, scales between the lateral line and the pelvic fin 11-15, and scales around the caudal peduncle 20-25.

Total dorsal fin rays 11(4), 12(21), 13(27) or 14(1), total anal fin rays 11(13), 12(25) or 13(15), total pectoral fin rays 14(4), 15(18), 16(25) or 17(6), total pelvic fin rays 5(9), 6(42), 7(1) or 8(1), total gill rakers 10(8), 11(41), or 12(4), and total vertebrae 27(2), 28(24), 29(26) or 30(1). Saefali (1999) records 8-11 gill rakers, 9-14 anal rays, and 9-15 dorsal rays.

Sexual dimorphism

Colour is sexually dimorphic and is outlined below. Ten of 21 morphometric characters are significantly different between the sexes including head length, head and body depths, fin lengths, and caudal peduncle shape (Coad, 1998i). Females attain larger sizes than males (76 mm versus 58 mm total length (Keivany and Soofiani, 2004a; b)).

Colour

Live adult males bear creamy bars on yellowish flanks. The pelvic fins and the edge of the caudal fin are yellow, the anal fin has an orange-yellow edge and the pectoral fins are orange-yellow. The dorsal fin is white with a wide blue-black band in the centre and a narrow blue-black band at the base. In some males the basal band is absent. Live adult females have a slight bluish tinge to their flanks and flank spots are brown.

In preserved specimens, the dorsal fin bands are dark and remain darker in preservative than the pigmentation seen in Aphanius sophiae which is more diffuse, tending to several horizontal rows of speckles, and which is not formed into a strongly distinct band or bands. Other fins are lightly pigmented, being finely speckled with melanophores with the dorsal and pectoral fins the darkest. The anal fin is fleshy and not hyaline but pigmentation is light and not dark as in A. sophiae. There is some tendency to formation of one or two narrow bars on the anal fin, paralleling the base, but these are weakly developed or absent in most fish. A mid-dorsal stripe is not apparent, perhaps because the dorsal flanks and back are darker than in females. Flank bars are light alternating with darker broad bars. The lighter bars become broader posteriorly. There is a light, wide bar at the base of the caudal fin which is not included in the count of male flank bars. Male flank bars 9(3), 10(8), 11(8), 12(10), 13(3), or 14(5). The sides of the head and upper and lower jaws are pigmented with scattered melanophores while the underside of the head is not. The belly is also free of extensive melanophores.

The most characteristic feature of females is the large number of scattered spots on the flank, extending from behind the head to the base of the tail. These spots do not extend to the dorsal flank or the back. The distinctive, large and often lozenge-shaped spot on the central caudal fin base of A. sophiae is absent. The upper back and flank are more heavily pigmented than the flanks (except for the flank spots). The lower surface of the head and the belly are melanophore free as in males and the sides of the head and the upper and lower jaws are heavily pigmented. The dorsal fin is finely pigmented on the rays and membranes and is generally darker than the caudal fin which is also finely pigmented. The other fins have very little pigment but all are fleshy and not hyaline. There is some suggestion of a thin horizontal stripe near the base of the dorsal fin in the largest specimens but this is never well developed.

Teeth are hyaline or discoloured with brown in patches. The peritoneum is dark brown.

Size

Reaches 7.6 cm (Keivany and Soofiani, 2004b).

Distribution

This species is found near Boldaji and in the Chaghakor Wetland in the upper reaches of the Karun River basin. Abdoli (2000) indicates its presence on a map in the upper Marun River and the upper Khersan River (the latter in the Karun basin).

Zoogeography

The origins of this mountain species have been discussed above under the account for A. sophiae.

It is found only in the uppermost reaches of the Karun River basin which drains to the head of the Persian Gulf. In this locality it is isolated from populations of Aphanius in the lowlands by a series of tangs which block gene flow. Oberlander (1965) in studying the origin of Zagros drainage patterns found that streams did not follow the line of least resistance but often incise the higher parts of mountain ridges which are anticlines composed of Miocene Asmari limestone. This situation arises because the Zagros ridges were eroded by a superposed drainage pattern already antecedent to the exhumation of overlying sediments of later Miocene marls and evaporites. The trend of the Zagros ridges is in a NW-SE direction and streams often cross this direction at right angles. This has resulted in the development of the tangs, narrow clefts in the Zagros ridges with very swift waters, which may be up to 2400 m deep with vertical sides rising from the water surface for 300 m, effective barriers to small fishes.

Habitat

This species is found high in the Zagros Mountains (the type locality lies at an altitude of about 2380 m). It is found in small pools, streams and marshes, in fresh water. The type locality is an artificially dammed pool about 300 m wide with cloudy water and a mud and pebble bottom. Capture depth was 40 cm. The shore was grassy and the pool contained large amounts of Myriophyllum and marginal rushes. At 1515 hours on a warm, windy and sunny day, water temperature was 29°C, pH was 6.5 and conductivity was 0.2 mS. A second locality in Coad (1988i) was a mud-bottomed stream 3-4 m wide with pools up to 1 m deep. At 1545 hours water temperature was 22°C, pH was 6.2 and conductivity was 0.45 mS. Current was slow and the stream had moderate amounts of Myriophyllum. Keivany and Soofiani (2004b) reporting on the habitat at the Madar-Dokhtar spring near Gandoman, found water conductivity of 240-280 μS, pH 6.9-8.5 and total dissolved solids125-138 mg/l. The bottom was muddy, interspersed with pebbles, and water is clear (but cloudy in the river). Myriophyllum and Potamogeton were used for spawning. Fish survive 30‰ and 30°C in aquaria.

Pazooki et al. (2008) found females to be more numerous than males, and the sexes lived in mixed schools. Occasionally larger males would chase smaller males but then both would rejoin the school. Males are territorial and defended against fish and beetles up to twice their size. The species preferred areas free from vegetation. These authors found the species easily tolerated to 33°C in aquaria for long periods, and the optimum temperature for the species is probably 21-24°C.

A mass mortality of this species was observed by A. Abdoli in late March or early April 1994 at Chagh-khor wetland near Boldaji.

Bagheri (1999) describes the macrofauna and environment of the Chaghakhour Lagoon where this species occurs.

Age and growth

Keivany and Soofiani (2004b) show maximum age is 2+ years. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 319 fish measuring 1.46-4.64 cm standard length. The a-value was 0.0309 and the b-value 3.062 (a b-value < 3 indicating a fish that becomh increaseand a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Abdoli (2000) reports Daphnia and Gammaridae as food. The main food in the Madar-Dokhtar spring is Gammarus, but a specimen from a river also had some algae, Cyclops sp., snail eggs and fish larvae. Mouth structure would lead to the conclusion that this species is a surface feeder but it concentrated on the dominant crustacean in the Madar-Dokhtar spring (Keivany and Soofiani, 2004b).  Pazooki et al. (2008) found chironomids, dipterans, Diaptomus, Daphnia, diatoms and filamentous algae in guts and considered this species to prefer an animal diet.

Reproduction

Mature males chase the female in aquaria, pushing her into a corner, and showing pectoral fin flipping and shivering. Eggs were laid in a batch of 3-10. Absolute fecundity in the Madar-Dokhtar spring examined by Keivany and Soofiani (2004b) was 220-650 eggs, relative fecundity was 45-155 per gram body mass and working fecundity (free and ripe eggs) was 110 eggs absolute and 73 relative. Size of ripe eggs was 0.8-1.2 mm. Eggs are adhesive and attch to plants in small patches, from 3 to 30. Early April was the spawning peak. Eggs hatched at 9-13 days at 21-22ºC. Young reached 11.0-27.2 mm after three months in an aquarium.

Parasites and predators

Barzegar et al. (2004) examined this species for parasites in fish from the Beheshtabad river in Chahar Mahall va Bakhtiari Province and found Lernaea cyprinacea. Keivany and Soofiani (2004b) found it to be susceptible to trichodiniasis and ichthyophthiriasis in aquaria.

Economic importance

This pretty little fish has great potential as an aquarium species (Coad and Keivany, 1998; Keivany and Soofiani, 2004b). It has been maintained in an aquarium for several months, tolerating temperatures of 30°C but probably favouring 20-22°C. pH values were 8-9. Water changes with their associated variations in conditions had no affect on the fish. The pupfish survived 3-4 hours longer than a Capoeta species when the latter was treated with formalin in a shared tank for a Saprolegnia infection and the tank was inadvertently left for 6-8 hours without a water change. Male pupfish are very aggressive and frequently attack other fishes of the same size or even larger. They ate all the coexisting guppy larvae in the aquarium, even those aged 2-3 weeks. The pupfish was fed crushed, dried freshwater shrimp and sometimes live shrimp although they had difficulty handling live shrimp. They were also observed biting and destroying vegetation in the aquarium, although it is unclear whether they actually ate any. It is now available in aquarium stores in Germany (Oliver Lucanus, pers. comm., 23 January 2004) and is kept during summer months in garden ponds, reproducing there successfully (Thomas Schulz, pers. comm., 14 January 2005).

Conservation

The distribution of the species is apparently quite limited, in waters neighbouring the towns of Boldaji and Shahr-e Kord. This should be confirmed by extensive field work in this poorly explored area of the Zagros Mountains. A limited distribution renders this species liable to significant loss or even extinction if habitats are disturbed or destroyed. The type locality, it should be noted, is an artificially dammed pond.

Significantly, a mass mortality of this species was observed by A. Abdoli in late March or early April 1994 at Chagha Khur wetland although the cause is unknown.

An additional threat is escapes of cultured rainbow trout (Oncorhynchus mykiss). Trout culture is being rapidly developed in this region and the trout are known predators on native fishes.

Further work

The distribution of this species in the Zagros needs to be clarified.

Sources

Type material: See above, CMNFI 1979-0247, CMNFI 1979-0247A and CMNFI 1979-0248.

Iranian material: CMNFI 2004-0005, ?, ?, mm standard length, ; uncatalogued (GenBank AY593486, AY593495), 4, 25.6-37.0 mm standard length, Chahar Mahall va Bakhtiari, Ebrahimabad (31º52'30"N, 51º10'10"E); uncatalogued (GenBank AY593487, AY593496), 2, 21.5-28.7 mm standard length, Chahar Mahall va Bakhtiari, Taqanak Bridge (32º12'35"N, 50º49'29"E); uncatalogued (GenBank AY593485, AY593494), 5, 31.0-49.8 mm standard length, Chahar Mahall va Bakhtiari, Madar-Dokhtar spring (31º52'12"N, 51º08'29"E).

Top