Morphological development and growth of Bunni, Mesopotamichthys sharpeyi (Gunther, 1874), larvae reared in the laboratory.
The early development of the endemic cyprinid, Bunni Mesopotamichthys sharpeyi (Gunther, 1874) larvae has been determined according to morphological changes and total length, standard length, head length, thickness of larvae, eye diameter and snout length measurements. The results showed that the initial period of Bunni larval life can be divided into two phases: early stages dependent upon endogenous nutrient sources, and a second phase of stages dependent upon exogenous food sources. In the first three days of larvae development there was a gradual yolk sac reduction after which there was a switch to exogenous feeding. From the fourth to eleventh day, the final development of heart, gill, air bladder, Fins and intestine were observed. The newly hatched larvae and the fifteen day old larvae were 6.26 [+ or -] 0.14 and 8.35 [+ or -] 0.17 mm in mean total length (TL), respectively. The mouth opened 2-4 days after hatching (DAH). The larvae started to swim actively within 2-3 days and the yolk sac had been totally absorbed at 4-5 DAH. Notochord flexion began at 11 DAH. Compare to other cyprinids, the larval development of Mesopotamichthys sharpeyi is similar to other Mesopota-michthys species.
Die fruhe Entwicklung der Larven der endemischen Barbe Mesopotamichthys sharpeyi (Gunther, 1874) (Cyprinidae) wurde nach folgenden Kriterien dokumentiert: morpholo-gische Veranderungen und Gesamdange, Standardlange, Kopflange, Dicke der Larven, Augendurchmesser und Schnauzenlange. Die Ergebnisse zeigen, dass sich das Larvenleben dieser Barbe am Anfang in zwei Phasen aufteilen lasst: ein erstes Stadium, das aus dem Korperinneren ernahrt wird, und eine zweite Phase, die von au[beta]eren Nahrungs-quellen abhangt. An den ersten drei Tagen ist eine allmahliche Verminderung des Dottersacks festzustellen, danach wird auf au[beta]ere Nahrung umgestellt. Vom vierten bis elften Tag lasst sich die endguldge Entwicklung von Herz, Kiemen, Schwimmblase, Flosscn und Darm beobachten. Die frisch geschlupften und die 15 Tage alten Larven hatten eine mittlere Gesamtlange cm von 6,26 [+ or -] 0,14 bzw. 8,35 [+ or -] 0,17 mm. Das Maul offnete sich 2 bis 4 Tage nach dem Schlupf (DAH). Nach 2 bis 3 Tagen begannen die Lar-yen aktiv zu schwimmen, und nach 4-5 DAH war der Dot-tersack vollig absorbiert. Die Notochord-Krummung begann mit 11 DAH. Im Vergleich mit anderen Zypriniden verlauft die Larvenentwickl ung von Mesopotamichthys sharpeyi ahnlich der anderer Mesopotamichthys-Arten.
Le premier developpement des larves du Cyprinide endemique, Mesopotamichthys sharpeyi (Gunther, 1874) a ete determine en fonction de modifications mor-phologiques et de la longueur totale, de la longueur standard, de la longueur de la tete, de la grosseur des larves, du diametre de red et de la longueur du rostre. Les resultats ont montre que la periode initiate de la vie de ces larves peat etre divisee en deux phases: les premiers stades en rapport avec les sources de nutriments endogenes et une se-conde phase en rapport avec les sources exogenes d'ali-mentation. Les trois premiers jours du developpement des larves ont montre une reduction progressive du sac vitellin suivie d'un passage a la nourriture exogene. Du quatrierne au onzieme jour, on a observe le developpement final du coeur, des branchies, de la vessie natatoire, des nageoires et de l'intestin. Les larves nouvellement ecloses et les larves de quinze jours mesuraient en longueur totale (LT) respec-tivement 6,26 +/- 0,14 et 8,35 +/- 0,17 mm. La bouche s'ouvrait 2-4 jours apres l'eclosion (JAE). Les larva ont commence a nager activement en 2-3 jours et le sac vitellin a ete resorbe a 4-5 JAE. La flexion notocorde a commence au 1 le JAE. Compare aux autres Cyprinides, le diveloppement larvaire de Mesopotamichthys sharpeyi est similaire a celui des autres especes de Mesopotamichthys.
Lo sviluppo larvale del ciprinidc endemico Mesopotamichthys sharpeyi (Gunther, 1874), noto comunemente come Bunni, e stato monitorato misurando la lunghezza to-tale, la lunghezza standard, la lunghezza della testa, lo spes-sore delle larve, ii diametro dell'occhio, la lunghezza del muso e altri parametri morfologici. I risultati hanno mo-strato che ii periodo di vita larvale pub essere diviso in due fasi: una prima fase dipendente da fonti nutrizionali endo-gene e una seconda fasc dipendentc da fonti di cibo eso-gene. Nei primi tre giorni di sviluppo larvale c'e una grad-uale ridu.zione del sacco vitellino, poi si passa all'alimen-tazione esogena. Dal quarto all'undicesimo giorno sono stati osservati il completamento dello sviluppo del cuore, delle branchie, della vescica natatoria, delle pinne e dell'in-testino. Le larve neonate e le larve al quindicesimo giorno avevano una lunghezza totale di 6.26 [+ or -] 0.14 e 8.35 [+ or -] 0.17 mm, rispettivamente. La bocca si apriva 2-4 giorni dopo la schiusa (DAH). Le larve iniziavano a nuotare activamente entro 2-3 giorni e ii sacco vitellino era totalmente riassor-bito a 4-5 DAH. La flessione della notocorda iniziava all' 11 DAH. Confrontato con altri ciprinidi, lo sviluppo larvale di Mesopotamichthys sharpeyi e simile a quello di altre specie di Mesopotamichthys.
The study of embryonic development of fishes is necessary for the identification of eggs and larvae collected in the natural environment. Additionally, with clarification of embryonic stages we can classify fishes according to ecologic, reproductive and ontogenetic (fish development from embryo to adult) factors (Reynalte-Tataje 2001). Understanding the growth processes under laboratory conditions is essential for successful aquaculture (Pyka et al. 2001). The Bunni, Mesopotamichthys shopeyi (Gunther, 1874) is a member of Cyprinidae family found in the Tigris-Euphrates Rivers basin, including its Iranian portion, in such marshes as the Hurol'azim, and in the northern Persian Gulf basin in the Zohreh River (Marammazi 1995; Abdoli 2000). In Iran this fish is reported in the Karun and Karkheh rivers (Nikpei 1996), and Hurol'azim and Shadegan lagoons (Najafpur 1996).
Although Bunni is one of the most economically important endemic bony fishes in Khtizestan province (southwestern Iran) and there are some plans to culture the fish, there is no information about their physiologic characteristics, particularly of the primary stages of development (Najafpur 1996).
The most sensitive growth stages in fishes are the embryonic and larval development periods which are affected by biological and non-biological conditions, in addition to genetic and physiologic factors (Penaz 2001).
Previous larval development studies of Iranian endemic fishes have been mostly on sturgeons and marine fishes. For example in the southern Caspian Sea, Behzadi (1991), Shafizadeh (1993) and Parandavar (2004) studied the embryonic and larval development of Rutilus frisii kutum, Acipenser persicus and Acipenser stellatus, respectively. In the Persian Gulf area, Sarvi-Ghiasabadi (2008) studied the embryonic and larval development of Acanthopagrus latus. In addition, Kimmel et al. (1995), Reynalte-Tataje et al. (2001), Firat et al. (2003; 2005) and Liew et al. (2006) studied the embryonic and larval development of Danio rerio, Leporinus macrocephalus, Den tex dentex and Sparus aurata, and Amphiprion ocellaris, respectively.
Larval development in some cyprinid files is well-documented (Sado & Kimura 2002, 2005, 2006; Al-Hazza & Hussein 2007), but is poorly understood in M shopeyi, a challenge to our effective evaluation of production capacity in aquaculture and stock management (Al-Nasih 1992; Mulchaysin & Jawad 2012). It is clear that information about the larval development of this species would be of great value to the artificial culture and management of Bunni. The present study is done for the first time on Bunni in Iran, while some related reports about the species are available in other areas (Al-Nasih 1992; Mulchaysin & Jawad 2012). Herein we provide a morphological and quantitative description of the larval period of Bunni under controlled conditions, recording morphological changes and growth factors like total length, standard length, eye diameter, head length and other measurements of newly hatched to 15 days old larvae.
MATERIALS AND METHODS
The investigation was done at the Khuzestan Endemic Fish Development Center from April to May 2008. The average of total weight and length of females and males were 1000 [+ or -] 127 and 690 [+ or -] 154 g, and 41 [+ or -] 4.5 and 43.5 [+ or -] 3.7 cm, respectively. Broodstocks were kept in concrete ponds with dimensions of 3x0.5x1.5m with a constant flow of water at the rate of 5 kg/[m.sup.3]. Pond surfaces were covered. Dissolved oxygen, temperature and EC were measured two times per day (Calta 1996). Dissolved oxygen, pH, temperature and EC were recorded 6.5-7.3 mg/l, 7-7.3, 22-28[degrees]C and 0.5-0.6 pm/[m.sup.3], respectively.
The water exchange rate was 30% per hour before injection of pituitary hormone and was reduced to 15% per hour after the injection (Buke et al. 2003). To stimulate the broodstocks to spawn, pituitary hormone of common carp was used at a rate of 4mg per kg body weight for females in two injections, and 2mg per kg body weight for males in one injection (Al-Hazza & Hussein 2007).
After hatching, larvae were transferred to Zoug incubators with a constant flow of water. Feeding began from 8 to 12 hours after hatching (Conides & Glamuzina 2001). The larvae food included boiled egg yolk (Calta 1996) and milk powder. Larvae were randomly sampled from newly hatched to 15 days old stage, twice a day by siphon method. The samples were fixed in formaldehyde 2% (Buke et al. 2003).
During the incubation period, morphological and anatomical characteristics of living larvae were studied by light microscope and information for each period was recorded. After transferring samples to the laboratory, these samples were photographed by stereomicroscope, then total length, standard length, head length, snout length, larvae thickness, eye diameter and tail length were measured by Axio Vision software with 15 times scale and their time of appearance was recorded (Firat et al. 2005, Tachihara & Kawaguchi 2003).
Volume of the yolk sac was estimated according to the formula below which assumes an oval-shaped yolk sac (Blaxter & Hempel 1966; Cetta & Capuzzo 1982);
V = 4/3[pi]L/2[(H/2).sup.2]
L is the longest and H is the shortest axis of oval.
Relative growth for total length, head length, eye diameter and tail length were determined according to the equation below. In this equation total length is assumed as a fixed datum (Huxley 1932):
Y = [X.sup.b]
Where Y is the dependent variable, X is the independent variable and b is the growth coefficient. Isometric growth occurs when growth coefficient is about 1. Positive growth is when growth coefficient is more than 1 and negative growth is when growth coefficient is less than 1. Minitab statistical software was used to measurement and drawing diagrams of relative growth. To obtain average and standard deviation (SD), 30 larvae were used per each sampling.
Based on morphological characteristics and development, embryonic development of Bunni is divided into two main phases. In the first phase the nutritional requirements of larvae are provided by an internal source--the yolk sac. In this short stage because of the food availability, larval growth and development are fast. In the second phase, the yolk sac has been completely absorbed and the larvae need external sources for feeding. Because of the feeding source change, growth and development are slower in the second phase compared to the first phase.
General morphology, fins development and pigmentation: Morphological and anatomical characters were classified into 5 steps based on developmental state as follows:
1) From 0 (newly hatched) to 2 DAH: Yolk sac is big and spherical, digestive system is not completely developed, heart appeared and is beating. The larvae are bent and crescent, without dorsal, caudal, pectoral and anal fins. Fin wrinkle is like a narrow fine continuous bar extending almost from mid-dorsum and completely surrounding the tail. Larvae swim in an inharmonic wave motion. Notochord is straight, mouth is closed, body has blood circulation and blood flows in two main vessels through heartbeat. One of the vessels is in the dorsal part of the body, which catches blood from the heart and transfers it to dorsal part of the body. Another vessel is pelvic and transfers blood from this part to the heart (Figs 1A & B).
2) From 2 to 4 DAB: Volume of yolk sac is reduced, the buds of pectoral fins grow, gills can be seen clearly, dorsal fin grows, swim bladder forms, blood is flowing between gill filaments and respiratory system. Pigment concentrations are in eye lenses, head and notochord. Pigments of head are brown, big and patchy. Mouth is open and represents the beginning of exogenous feeding. Exogenous feeding begins from the third day. There is blood circulation in all parts of the body (Fig. 1C).
3) From 4 to 7 DAH: Yolk sac is completely absorbed and feeding source is 100% exogenous. Melanophore concentrations in eyes can be clearly observed, air bladder is completely formed, anus is forming, fin rays are forming and melanophores are apparent in dorsal part of head and body (Figs 1D & E).
4) From 7 to 11 DAH: Opercula are formed, caudal fin is well-developed, the mouth opening and closing indicates formed jaws, trunk muscles can be observed in cross bands formed in sides of the body, single pore air bladder is oval and air filled, the mouth movements are slow (Figs 1F & G).
5) From 11 to 15 DAH: Larvae swim well and have maneuverability, dorsal end of notochord is a little bent upward, there are bony rays in homocercal caudal fin, a mass of melanophores increases on caudal fin, dorsal fin is developed from embryonic fin wrinkle. In this stage, total length is 8.35[+ or -]0.17 mm (Figs 1H, I &J).
Proportions: The average of yolk sac length showed a reduction from 0 to 4 DAH (Table I). As the larvae don't have any ability to use exogenous foods, the yolk sac was absorbed quickly with a severe slope in the first few days (Fig. 2). Concurrent with the larval development, yolk sac diameter reduced and its length increased. In newly hatched larva the head is short (12% standard length), but this proportion increasing with growth, reached 17% standard length in 15 days old larvae. Eye diameter is 52% head length initially, decreasing to 45% at 5 DAH, and then increased to 48% at 6 DAH, a decrease was observed to 42% at 13 DAH, subsequently increasing and reaching to 46% head length at 15 DAH. Snout is small in newly hatched larvae (4% standard length), but rapidly increases with growth to 7% standard length at 15 DAH. Table II, shows total length, standard length, head length, snout length, larvae thickness, eye diameter and tail length of newly hatched larvae to 15 days after hatching.
Table I. Average ([+ or -] SD) volume of yolk sac in Bunni fish larvae in 0 to 4 days after hatching. Age (day) Average [+ or -] SD ([mm.sup.3]) 0 0.264 [+ or -] 0.034 1 0.136 [+ or -] 0.021 2 0.071 [+ or -] 0.031 3 0.024 [+ or -] 0.015 4 0.017 [+ or -] 0.005 Table II. Morphological parameters (average [+ or -]SD) of 0 (newly hatched larvae) to 15 day-old larvae of Bunni Mesopota-michthys sharpeyi Age Total Standard Head Snout Larvae Eye (day) length legth length length thickness diameter (mm) (mm) (mm) (mm) (mm) (mm) 0 6.26 [+ 5.95 [+ or 0.69 [+ 0.27 [+ 1.01 [+ or 0.36 [+ or -] or -] -] 0.16 or -] or -] -] 0.07 0.04 0.14 0.09 0.04 1 7.09 [+ 6.52 [+ or 0.93 [+ 0.32 [+ 1.08 [+ or 0.45 [+ or -] or -] -] 0.29 or -] or -] -] 0.09 0.03 0.29 0.11 0.05 2 7.53 [+ 6.94 [+ or 1.00 [+ 0.38 [+ 1.12 [+ or 0.45 [+ or -] or -] -] 0.36 or -] or -] -] 0.08 0.06 0.30 0.13 0.05 3 7.71 [+ 7.08 [+ or 1.03 [+ 0.40 [+ 1.15 [+ or 0.48 [+ or -] or -] -] 0.33 or -] or -] -] 0.10 0.04 0.29 0.12 0.04 4 7.89 [+ 7.22 [+ or 1.04 [+ 0.45 [+ 1.20 [+ or 0.50 [+ or -] or -] -] 0.42 or -] or -] -] 0.07 0.01 0.35 0.13 0.05 5 8.03 [+ 7.30 [+ or 1.13 [+ 0.47 [+ 1.25 [+ or 0.51 [+ or -] or -] -] 0.23 or -] or -] -] 0.07 0.03 0.16 0.14 0.06 6 8.10 [+ 7.35 [+ or 1.19 [+ 0.50 [+ 1.26 [+ or 0.51 [+ or -] or -] -] 0.53 or -] or -] -] 0.06 0.04 0.43 0.12 0.03 7 8.19 [+ 7.36 [+ or 1.21 [+ 0.50 [+ 1.29 [+ or 0.52 [+ or -] or -] -] 0.30 or -] or -] -] 0.10 0.03 0.33 0.07 0.02 8 8.25 [+ 7.39 [+ or 1.25 [+ 0.51 [+ 1.31 [+ or 0.53 [+ or -] or -] -] 0.21 or -] or -] -] 0.20 0.02 0.35 0.13 0.05 9 8.27 [+ 7.41 [+ or 1.25 [+ 0.51 [+ 1.06 [+ or 0.53 [+ or -] or -] -] 0.22 or -] or -] -] 0.07 0.02 0.23 0.06 0.04 10 8.27 [+ 7.45 [+ or 1.25 [+ 0.51 [+ 1.07 [+ or 0.53 [+ or -] or -] -] 0.33 or -] or -] -] 0.10 0.03 0.27 0.01 0.04 11 8.29 [+ 7.50 [+ or 1.25 [+ 0.51 [+ 1.11 [+ or 0.53 [+ or -] or -] -] 0.36 or -] or -] -] 0.07 0.02 0.27 0.01 0.03 12 8.30 [+ 7.51 [+ or 1.25 [+ 0.51 [+ 1.12 [+ or 0.53 [+ or -] or -] -] 0.23 or -] or -] -] 0.09 0.02 0.17 0.10 0.03 13 8.32 [+ 7.52 [+ or 1.25 [+ 0.52 [+ 1.10 [+ or 0.53 [+ or -] or -] -] 0.22 or -] or -] -] 0.11 0.03 0.25 0.13 0.05 14 8.34 [+ 7.53 [+ or 1.26 [+ 0.52 [+ 1.12 [+ or 0.55 [+ or -] or -] -] 0.28 or -] or -] -] 0.08 0.04 0.25 0.11 0.04 15 8.35 [+ 7.55 [+ or 1.26 [+ 0.52 [+ 1.22 [+ or 0.58 [+ or -] or -] -] 0.24 or -] or -] -] 0.20 0.05 0.17 0.16 0.05 Age Tail (day) length (mm) 0 1.79 [+ or -] 0.23 1 1.94 [+ or -] 0.17 2 1.99 [+ or -] 0.41 3 2.05 [+ or -] 0.50 4 2.30 [+ or -] 0.31 5 2.42 [+ or -] 0.13 6 2.50 [+ or -] 0.23 7 2.55 [+ or -] 0.13 8 2.56 [+ or -] 0.29 9 2.56 [+ or -] 0.16 10 2.56 [+ or -] 0.22 11 2.56 [+ or -] 0.21 12 2.57 [+ or -] 0.19 13 2.58 [+ or -] 0.19 14 2.58 [+ or -] 0.22 15 2.60 [+ or -] 0.26
Growth patterns: Tail length and head length showed ascending growth rates (2.63 and 2.25, respectively). Growth of eye diameter was almost isometric (b = 1.08) (Fig. 3).
Larval development in Bunni is similar to most members of the Cyprinidae family, however this species has some special characteristics which haven't been observed in other studied cyprinids. Development can be accelerated or delayed by environmental factors (such as temperature, dissolved oxygen and etc.), internal factors (egg size, parent's effects and etc.) or a complex of both (Kamler 2002).
Eyes pigments in Bunni were observed sooner than other species of Barbus. However, this phenomenon has been observed in some other endemic species like Hamri (Penaz 2001), Gatan and Shirbot (Pyka et al. 2001). Pigments in Bunni larvae are concentrated like big brown patches in certain parts of the body (head, along notochord and lateral line). Pigment appearance in fish larvae has a close relationship with metabolism, some hormones and growth factors (Christensen & Korsgaard 1999; Solbakken et al. 1999), nutrients and diet (Bolker & Hill 2000; Diler & Dilek 2002), genetic and environmental factors (Urho 2002).
Compared to all life periods, swimming ability in the larval phase has a fast development because of fast morphological changes in the phase (Fuiman & Webb 1988). Swimming ability in Bunni larvae is determined by the growth and development process. An increase in mechanical power of muscles and swimming organs function make the fish swim like Common Carp Cyprinus carpio (Wakeling et al. 1999).
Fast growth and development have been reported in many species of Barbus species (Kohno et al. 1986). In Bunni, larval growth and development can be studied in two phases. The first phase is recognizable by fast growth and development rates in newly hatched larva. Also, according to the small size of yolk sac in Bunni, after hatching food should be available immediately to begin exogenous feeding, a similar characteristic is reported for Pagrus pagrus (Conides & Glamuzina 2001).
After the fast growth (first phase), we observed a slow growth phase (second phase). The second phase begins a little after yolk sac depletion. In this phase, growth rates in body length and other organs are slow. This growth rate can demonstrate larval development and interior organs function (Klaoudates et al. 1990).
Barbus sp. larvae are significantly larger than many marine species and smaller than Salmon because of egg size differences (Calta 1996). Bunni adults are short like Hamri because they are small in the beginning of their larval development compared to other cyprinids (Szypula et al. 2001). Similar to other endemic fishes like Gatan, Shirbot and Hamri (Pyka et al. 2001), in the present study, total length of Bunni is short at hatching time and compared to other related species no Barbus sp. are as short as Bunni larvae in average length. In the present study body length of newly hatched Bunni larvae was observed to be 6.26 mm, while it was reported as 5mm by Mukhaysin and Jawad (2012). Other reports on total length in newly hatched larvae are available for Hamri larvae, 6.81 mm (Al-Hazza & Hussein 2007), Pagrus pagrus, 3.02 mm (Conides & Glamuzina 2001) and Chelon labrosus, 4 mm (Klaoudates et al. 1990). Although Mulchaysin and Jawad (2012) reported eye diameter as small in newly hatched Bunni larvae and rapidly increased with growth in scaled juveniles, our results showed that the eye diameter decreases in proportion to head length with growth.
In Bunni fish larvae, growth of head length and tail length, are positive and growth of eye diameter is almost isometric. In a similar study on Hamri, growth of eye diameter and head length were positive (2.72 and 2.5, respectively) and growth of tail length was almost isometric 1.25 (Al-Hazza & Hussein 2007).
Development of digestive system in many fishes has significant development and progress in their structure and function during the larval period observed with an increase in their gut twist (Cuvier-Perez & Kestmont 2002). Growth rate of the digestive system of Bunni fish is slow, like Hamri (Al-Hazza & Hussein 2007). Of course the slow growth process doesn't mean that the digestive system isn't important in the development path.
There is a close relationship between mouth opening of larvae and yolk sac absorption and co-occurs with complete absorption of yolk sac (Lasker et al. 1970). When larvae mouth opens, shows larvae readiness and its need for exogenous food (Sato et al. 2003). This period happens simultaneously with completion of digestive system and internal organs (Nakatani et al. 2001).
The size of the yolk sac is one of the important strategies of fish survival and reproduction. So, if the size of yolk sac is large, requirement of exogenous food will be postponed (Blaxter 1988). Yolk sac depletion may be related to significant developments in respiratory system, swimming ability and swimming activity (Al-Hazza 2007). The volume of the yolk sac in newly hatched Bunni was 0.264 [+ or -] 0.034 [mm.sup.3]. By comparison, the volume of the yolk sac in Danio rerio is 0.1497 [mm.sup.3] (Schmidt & Starck 2004), in Pagrus pagrus, 0.3178 [+ or -] 0.085 m[m.sup.3] (Mihelakakis et al. 2001) in Acanthopagrus latus, 0.1856 m[m.sup.3] (Sarvi-Ghiasabadi 2008) and in Dentex dentex it has been reported as 0.2476 [+ or -] 0.1020 [mm.sup.3] (Firat et al. 2003).
Yolk sac absorption in Bunni fish is done until 3 days after hatching and they begin exogenous feeding from the fourth day. This process is faster compare to some other species of Barbus sp. for example yolk sac absorption in Hamri fish lasts one week (Al-Hazza & Hussein 2007). Leuciscus cephalus in laboratory conditions consumes the yolk sac in 8 days, but only 30% of larvae population can begin exogenous feeding (Calta 2000). Compare to other cyprinids the short period of yolk sac absorption may be caused by the small size of yolk sac in Bunni fish.
In conclusion, study of the larval development of Bunni and other fishes in controlled conditions can lead to an understanding of biological and environmental requirements of the development process, which can be important in endemic fish stock management.
The authors are grateful to Dr. Frank Pezold and Heiko Bleher for their helpful comments, Dr. Lak, Ms. Maktabi and Mr. Savari for their cooperation and Zeynab Moradkhani for her help in English revision.
Received: 3 September 2012--Accepted: 15 April 2013
ABDOLI, A. 2000. The Inland Water Fishes of Iran, Iranian Museum of Nature and Wildlife, Tehran: 378 pp. [In Persian]
AL-HAZZA, R. & HUSSEIN, A. 2007. Larval development of Himri, Barbus luteus (Cyprinidae: Cypriniformes) Reared in the Laboratory Turkish Journal of Zoology 31: 27-33.
AL-NASIH, M. H. 1992. Preliminary observation related to the culture of Barbus sharpeyi (Bunni). Journal of Aquaculture in the Tropics 7: 69-78.
BEHZADI, S. 1991. A study on embryonic development stages of Kutum (Rutilus frisii kutum). MSc thesis. Azad University. Tehran Shomal Branch: 15-22.
BLAXTER, J. H. S. & HEMPEL, G. 1966. Utilization of the yolk sac by herring larvae. Journal of the Marine Biological Association of the United Kingdom 46: 219-234.
BAXTER, J. H. S. 1988. Eggs and larvae. In: Fish physiology. (Eds. Hoar, W. S. & Randall, D. J): 17-48. Academic press, New York.
BOLKER, J. A. & HILL, C. R. 2000. Pigmentation development in hatchery-reared flatfishes. Journal of Fish Biology 56: 1029-1052.
BUKE, E., AYEKIN, B. & IMAM, H. 2003. (unpublished data). The research project on the culture potential of new fish species as alternative for sea bass and sea bream. Ministry of Agriculture and Rural Affairs, General Directorate of Agricultural Research.
CALTA, M. 1996. Early development and gill function in freshwater fish. Ph. D. Thesis. The University of Nottingham. Nottingham. UK.
CALTA, M. 2000. Morphological development and growth of chub, Leuciscus cephalus (L.), larvae. Applied Ichthyology 16: 83-85.
CETTA, C. M. & CAPUZZO, J. M. 1982. Physiological and biochemical aspects of embryonic and larval development of the winter flounder, Pseudopleuronectes americanus. Marine Biology 71: 327-337.
CHRISTENSEN, M. N. & KORSGAARD, B. 1999. Protein metabolism, growth and pigmentation patterns during metamorphosis of plaice (Pleuronectes platessa) larvae. Journal of Experimental Marine Biology and Ecology 237: 225-241.
CONIDES, A. J. & GLAMUZINA, B. 2001. Study on the early larval development and growth of red porgy, Pagrus pagrus with emphasis on the mass mortalities observed during this phase. Scientia Marina 63 (3): 193-200.
CUVIER-PEREZ, A. & KESTMONT, P. 2002. Development of some digestive enzymes in Eurasian perch larvae Perca fluviatilis. Fish Physiology and Biochemistry 24: 279-285.
DILER, I. & DILEK, K. 2002. Significance of pigmentation and use in aquaculture. Turkish Journal of Fishery and Aquatatic Science 2: 97-99.
FIRAT, K. S. & COBAN, D. 2003. The effect of light intensity on early life development of the common dentex (Dentex dentex) larvae. Aquaculture Research 34: 1-6.
FIRAT, K., SAKA, S. & COBAN, D. 2005. The cleavage and embryonic phase of gilthead sea bream (Sparus aurata) eggs. Journal of Fisheries & Aquatic Sciences 22: 205-207.
FUIMAN, L. A. & WEBB, P. W. 1988. Ontogeny of routine swimming activity and performance in Zebra danio (Teleostei: Cyprinidae). Animal Behaviour 36: 250-261.
HUXLEY, J. S. 1932. Problems of relative growth. Methuen & Co., London.
KAMLER, E. 2002. Ontogeny of yolk-feeding fish: An Ecological Perspective. Reviews in Fish Biology and Fisheries 12: 79-103
KIMMEL, C., BALLARD, W, ULLMANN, B. & SCHMILLNG, T. 1995. Zebra fish (Danio rerio) embryonic development. Developmental Dynamics 203: 205-310.
KLAOUDATES, S., TSEVIS, N. & CONIDES, A. 1990. Energy sources during early larval development of the Eurropean sea bass, Dicentrarchus labrax (L.). Aquaculture 87: 361-372.
KOHNO, H., HARA, S. & TAKI, Y. 1986. Early larval development of the sea bass Lates calcarifer with emphasis on the transition of energy sources. Bulletin of the Japanese Society for the Science of Fish 52 (10): 1719-1725.
LASKER, R., FEDER, H. M., THEILACHER, G. H. & MAY, R. C. 1970. Feeding, growth and survival of Engraulis mordax larvae reared in the laboratory. Marine Biology 5: 345-353.
LIEW, H. J., AMBAK, M. A. & ABOL-MUNAFI, A. B. 2006. Embryonic development of clownfish Amphiprion ocellaris under laboratory conditions. Journal of Sustainability Science and Management 1: 64-73.
MARAMMAZI, Gh. 1995. Ichthyology report on Shadegan Marsh. Southern Iran Aquaculture Fishery Research Centre, Ahvaz. 63 pp. [In Persian]
MIHELAKAKIS, A., YOSHIMATSU, T & TSOLKAS, C. 2001. Spawning in captivity and early life history of cultured red porgy, Pagrus pagrus. Aquaculture 199: 333-352.
MUKHAYSIN, A. A. & JAWAD, L. A. 2012. Larval development of the cyprinid fish Barbus sharpeyi (Gunther, 1874). Journal of Fisheries and Aquatic Science 7: 307319.
NAJAFPUR, N. 1996. Final report of project of identification of some fresh water fishes of Khuzestan. Iranian fisheries research organization 1-96.
NAKATANI, K. 2001. Ovos e larvas de peixes de agues doce: desenvolvimento e manual de identificacao. Maringa: Eduem, 378 p.
NIKPEI, P. M. 1996. Final report of project of biologic study of Barbus grupus and B. sharpeyi. Iranian Fisheries Research Organization: pages 1-10, 52-64.
PARANDAVAR, H. 2004. A Study on embryonic development stages of South Caspian stellate sturgeon (Acipenser stellatus). International Research Institute of Sturgeons. IFRO publications. pp. 111.
PENAZ, M. 2001. A general framework of fish ontogeny: A review of the ongoing debate. Folia Zoologica 50: 241-256.
PYKA, J., BARTEL, R., SZCZERBOWSKI, J. A. & EPLER, P. 2001. Reproduction of gattan (Barbus xanthopterus), shabbout (Barbus grypus) and Bunni (Barbus sharpeyi) and rearing stocking material of these species. Archives of Polish Fisheries 9 (Suppl. 1): 235-246.
REYNALTE-TATAJE, D., ZANIBONI-FIHOL, E. & MUELBERT, B. 2001. Stages of the embryonic development of the Leporinus macrocephalus. Department of Aquaculture, University of Federal of Santa Catarina, Santa Catarina Brasil. pp. 824-826.
SADO, T. & KIMURA, S. 2002. Developmental morphology of the cyprinid fish, Candidia barbatus. Ichthyological Research 49: 350-354.
SADO, T. & KIMURA, S. 2005. Developmental morphology of the cyprinid fish Tanichthys albonubes. Ichthyological Research 52: 386-391.
SADO, T & KIMURA, S. 2006. Developmental morphology of the cyprinid fish, Inlecypris auropurpureus. Ichthyological Research 53: 34-40.
SARVI-GHISABADI, A. 2008. A study on embryonic development stages of Acanthopterus latus. MSc thesis. Azad University. Sciences and Research Branch: 4-14.
SATO, Y. N., FENERICH-VERANI, A. P. O., NUNER, H. P. & GODINHO, J. R. 2003. Padores reprodutivevos de peixes da bacia do sao Francisco. In: Aguas, peixes e pescadores do Sao Francisco das Minas Gerais. (Eds. H. P. Godinho & A. L. Godinho): 224-268. Belo Horizonte.
SCHMIDT, K. & STARCK, M. 2004. Developmental variability during early embryonic development of Zebra fish (Dania rerio). Journal of Experimental Zoology Part B: Molecular and Developmental Evolution 302B (5): 446-457.
SHAFIZADEH, S. 1993. A study on embryonic development stages of Persian sturgeon (Acipenser persicus). International Research Institute of Sturgeons. 1-2 pp.
SOLBAKKEN, J. S., NORBERG, B., WATANAB K. & PITTMAN, K. 1999. Thyroxine as a mediator of metamorphosis of Atlantic halibut, Hippoglossus hippoglossus. Environmental Biology of Fishes 56: 53-65.
SZYPULA, J., EPLER, P., BARTEL, R. & SZCZERBOWKI, J. A. 2001. Age and growth of fish in lakes Tharthar, Razzazah, and Habbaniya. Archives of Polish Fisheries 9 (Supll. 1): 185-197.
TACHIHARA, K., & KAWAGUCHI, K. 2003. Morphological development of eggs, larvae and juveniles of laboratory-reared Ryukyu-ayu Plecoglossus altivelis ryukyuensis. Fisheries Science 69: 323-330.
URHO, L. 2002. Characters of larvae-what are they? Folia Zoolica 51: 161-186.
WAKELING, J. M., KEMP, K. M. & JOHNSTON, I. A. 1999. The biomechanics of fast-starts during ontogeny in the common carp (Cyprinus carpio). Journal of Experimental Biology 202: 3057-3067.
Sara Ahmadi (l), Mojgan Khodadadi (2), Ammar Salehi Farsani (3), Bahareh Samadi Kuchaksaraei (4) and Hamed Mousavi-Sabet (5) *
(1.) Department of Fisheries, Science and Research Branch, Islamic Azad University, Khuzestan, Iran
(2.) Deepartment of Fisheries, Ahvaz Branch, Islamic Azad University, Ahvaz, Iran
(3.) Department of Fisheries, Science and Research Branch, Islamic Azad University, Tehran, Iran
(4.) Shilat, Ensan, Salamati Company, Tehran, Iran
(5.) Department of Fisheries Sciences, Faculty of Natural Resources, University of Guilan, Sowmeh Sara, Iran
* Corresponding author: H. Mousavi-Sabet, Department of Fisheries Sciences, Faculty of Natural Resources, University of Guilan, P.O. Box 1144, Sowmeh Sara, Guilan, Tel: +98 182 322 3599--Fax: +98 182 322 3600. Email: Mousavifirstname.lastname@example.org
|Printer friendly Cite/link Email Feedback|
|Author:||Ahmadi, Sara; Khodadadi, Mojgan; Farsani, Ammar Salehi; Kuchaksaraei, Bahareh Samadi; Mousavi-Sabet,|
|Publication:||aqua: International Journal of Ichthyology|
|Date:||Apr 26, 2013|
|Previous Article:||Leptophilypnion, a new genus with two new species of tiny central Amazonian gobioid fishes (Teleostei, Eleotridae).|
|Next Article:||Description of a new species of annual fish of the genus Neofundulus (Cyprinodontiformes: Rivulidae) from the upper rio Mamore basin, Bolivia.|