. J. Ichthyol. (March 2019), 6(1): 31-40 Received: August 17, 2018 © 2019 Iranian Society of Ichthyology Accepted: November 1, 2018 P-ISSN: 2383-1561; E-ISSN: 2383-0964 doi: 10.22034/iji.v6i1.285 http://www.ijichthyol.org

Research Article

Reproductive biology of the invasive , leucisculus (Basilewsky, 1855), from the southern Caspian Sea basin

Hamed MOUSAVI-SABET*1,2, Adeleh HEIDARI1, Meysam SALEHI3

1Department of Fisheries, Faculty of Natural Resources, University of Guilan, Sowmeh Sara, Guilan, Iran. 2The Caspian Sea Basin Research Center, University of Guilan, Rasht, Iran. 3Abzi-Exir Aquaculture Co., Agriculture Section, Kowsar Economic Organization, Tehran, Iran. *Email: [email protected] Abstract: The sharpbelly, Hemiculter leucisculus, an invasive , has expanded its range throughout much of Asia and into the Middle East. However, little is known of its reproductive information regarding spawning pattern and season that could possibly explain its success as an invasive species. This research is the first presentation of its reproductive characteristics, which was conducted based on 235 individuals collected monthly throughout a year from Sefid River, in the southern Caspian Sea basin. Age, sex ratio, fecundity, oocytes diameter and gonado-somatic index were calculated. Regression analyses were used to find relations among fecundity and size, gonad weight (Wg) and age. The mature males and females were longer than 93.0 and 99.7mm in total length, respectively (+1 in age). The average egg diameter ranged from 0.4mm (April) to 1.1mm (August). Spawning took place in August, when the water temperature was 23 to 26°C. Average gonado-somatic index values for females at the beginning of the reproduction period (March) and in ripe mature spawning females (August) were 2.90% and 16.97%, respectively. The averages of absolute and relative fecundities were 19978.6 and 607.1 eggs, respectively. The absolute fecundity was significantly related to body weight, ovary weight and age. A potentially high reproductive capacity, comparatively small eggs, short time of eggs’ development, fast sexual maturation, short life span, tolerance to a wide range of environmental conditions, all promote the invasion of this invasive sharpbelly in freshwater ecosystems of the temperate climate zone. Keywords: Gonado-somatic index, Fecundity, Freshwater ecosystem, Spawning season. Citation: Mousavi-Sabet, H.; Heidari, A. & Salehi, M. 2019. Reproductive biology of the invasive sharpbelly, Hemiculter leucisculus (Basilewsky, 1855), from the southern Caspian Sea basin. Iranian Journal of Ichthyology 6(1): 31-40.

Introduction of aquaculture fish species, the sharpbelly has Exotic can be caused the greater problem for invaded many other countries including Iran (Holcik the indigenous fish fauna and other organisms (Coad & Razavi 1992; Patimar et al. 2008; Jouladeh- 1980). A variety of exotic fish species have Roudbar et al. 2015), Iraq (Coad & Hussain 2007), introduced in Iranian freshwaters (Coad & Abdoli Kazakhstan (Petr & Mitrofanov 1998), Uzbekistan 1993; Esmaeili et al. 2018). Sharpbelly Hemiculter (Borisova 1972), Turkmenistan (Sal’nikov 1998), leucisculus (Basilewski, 1855) is a small cyprinid fish and Russian Federation (Kolpakov et al. 2010). with an original distribution extending from , Sharpbelly is a non-indigenous fish in the Caspian , Vietnam, to the far-eastern region of Russia Sea basin. In Iran, H. leucisculus is reported from and is the dominant species in many of the water some international wetlands, including the Alma- bodies in those regions (Gao et al. 2010; Tan et al. Gol, Adji-Gol and Ala-Gol (Patimar et al. 2008). 2010). Apparently through the associated movement Esmaeili et al. (2011) reported it from Zarivar Lake,

31

Iran. J. Ichthyol. (March 2019), 6(1): 31-40

Fig.1. Location of the sampling site in the southern Caspian Sea basin, Sefid River, in north of Iran. Abdoli (2000) recorded this species from the lower periods. Knowledge of gonadal development and the Sefid River, Anzali Wetland and the middle Aras spawning season of a species allow the subsequent River and Radkhah et al. (2016) reported it from the studies on spawning frequency of its population, Hamun-e-Jazmourian basin, southern Iran. which is important for its management (Chakraborty Hemiculter leucisculus found in rivers, lakes, small et al. 2007; Eagderi et al. 2013). Within a given ponds and swamps but there is a little information species, fecundity may vary as a result of different available about its habitat requirements (Coad 2017). adaptations to environmental habitats (Witthames et There are many studies on the age and growth of H. al. 1995). Fecundity assessments have been useful in leucisculus in its native ecosystems (e.g., Cao & Wen racial distinction, progeny survival studies, stock 1996; Li et al. 2009). In the invaded ecosystems, evaluation and aquaculture-based induced spawning however, Patimar et al. (2008) did research in three and egg incubation (Bagenal & Braum 1978; Coates lakes of Iran. In order to monitoring this species in its 1988). Sizes of eggs are also important parameters habitat, determination of reproductive biology of the used in reproduction studies to characterize fish fish is an essential factor. Also, descriptions of species and can also be used to predict the spawning reproductive strategies and the assessment of frequency of fish (Wootton 1979). Ecological fecundity are fundamental topics in the study of the conditions such as the water temperature and biology and population dynamics of fish species photoperiod influence the sexual maturity of fish (Cortes 2000; Jamali et al. 2015). greatly (Pawson et al. 2000; Rodríguez et al. 2001). Reproduction has three key components, The aim of the present study was to provide data including sexual maturity, reproductive period and on the reproductive characteristics of H. leucisculus, fecundity, which are vital demographic including gonado-somatic index, fecundity, oocyte characteristics, essential for understanding a species’ diameter, and spawning season/duration. We studied life history (Cortes 2000). Gonado-somatic index has the population inhabiting the down-stream of Sefid been used as an indicator of reproductive activity of River, in the southern Caspian Sea basin. Sefid River fish (Bagenal & Braum 1978; Coates 1988), and in (with 765km in length) is one of the most important the absence of information on eggs and larvae, this rivers in the southern Caspian Sea basin, in north of could be used to give an indication of peak spawning Iran (Fig. 1) (Heidari et al. 2014a, b; Coad 2017). The 32 Mousavi-Sabet et al.- Reproductive biology of Hemiculter leucisculus

results were compared to data available for with eye or by use a binocular. In order to determine populations inhabiting different geographical regions the number of eggs, pieces were removed as well as for other cyprinid fish species to reveal the approximately 0.5-1g, each from the anterior and role of life history traits in promoting the species medial positions of both ovarian lobes. The pieces invasion in the southern Caspian Sea basin rivers were weighed and the eggs counted under a binocular system. microscope. The number of eggs in each female was calculated as the proportion of eggs in the sample to Materials and Methods the weight of the whole ovary. To calculate absolute Study area and Sampling: The research was carried fecundity, ovaries recognized at stages IV or V were out at one site on the down-stream of Sefid River used. The stage of gonad maturity was determined (36°59'31.01"N, 49°34'34.56"E). Samples were visually following the Nikolsky (1963). The relative taken from the shallow littoral pools in Sefid River, fecundity (Fr) was expressed by dividing the absolute clean water, slow water flow, the pools width were fecundity (Fa) to fish body weight (Bagenal & Braum about 5-15 m and maximum depth was up to 1.5m, 1978). To determine the oocyte diameter, the ovaries and the bed was rocky and gravel, somewhere with were preserved in 4% formaldehyde solution. The grassy shores and submergent plants. In total 235 diameters of 30 ova of each specimen were measured specimens were caught by electrofishing device using a binocular microscope M6C–10 which was (200-300V; distance between electrodes: 10-50 fitted with an ocular micrometer. meters; area covered each time: 100 meters; water Statistics: In order to compare significant differences depth covered: 40-120cm) at monthly intervals in the gonado-somatic index among various months throughout the year from March 2014 to February and various size samples, the analysis of variance and 2015. The water temperature of the fish catching site Tukey’s test were applied (P<0.05). The strength and was measured repeatedly. The water analysis in significance of the relationship between the absolute August showed that the water temperature was 23 °C; fecundity (Fa) and selected individual features of the -1 pH 6.5–7.9, O2 5–8 mg. L . All specimens were fixed females included in the study (standard length and and preserved in 4% formaldehyde solution and weight, the gonad weight and the fish’s age) were transferred to the laboratory for other studies. analyzed by determining Pearson’s correlation Measurements and indices used to qualify the coefficient r (P<0.05) and regression equations reproductive biology: For each specimen, total length (Thulasitha & Sivashanthini 2013). The data were (TL), fork length (FL) and standard length (SL) were analyzed by the SPSS version 16.0 software package measured by using measuring board to the nearest and Microsoft Excel 2010 software. 1mm; body weight (W) weighted by a digital balance with 1g accuracy. The fish’s age was assessed based Results on the annual growth of scales taken from the left side During this study, 235 individuals of H. leucisculus of the body, between the end of the pectoral fin and were collected from Sefid River. Fish body beginning of the dorsal fin (Bagenal & Braum 1978; measurements, including total length, fork length, Mousavi-Sabet et al. 2012a, b). Gonado-somatic standard length and total weight ranged 91.0-181.0 index was used to investigate the monthly changes in mm, 82.0-174.0mm, 74.0-162.0mm and from 4.8 to gonads to estimate the spawning season of this 65.9 g, respectively (Table 1). This population of species. In order to determine the Gonado-somatic H. leucisculus had a narrow age range (1+ to 5+ index [= (Wg × Wb–1) ×100] (Biswas 1993) ovaries years). The majority of the fish caught were 3+ and were weighed (Wg) with 0.1g accuracy. Sex was 4+ years old, some were 1+ and 2+ and only a few of determined by examination of the gonad tissue either them were 5+ years old (Table 1). 33 Iran. J. Ichthyol. (March 2019), 6(1): 31-40

Table 1. Total length (TL), fork length (FL) and body weight (W) (mean ± SD) in different ages of Hemiculter leucisculus males and females from Sefid River, the southern Caspian Sea basin Age N FL (mm) TL (mm) W (g) M F M F M F M F 1+ 2 4 84.0±0.28 85.7±0.33 93.0±0.29 99.7±0.29 6.65±0.91 7.89±2.23 2+ 18 20 108.1±0.72 110.7±1.11 119.5±0.87 122.0±1.15 15.88±4.03 16.86±4.27 3+ 48 53 118.6±0.53 126.7±0.49 128.0±0.67 139.0±0.48 21.60±4.78 22.32±4.55 4+ 22 52 138.0±0.53 138.4±0.48 151.7±0.68 156.8±0.86 26.27±4.03 27.33±4.33 5+ 9 7 146.2±0.36 150.2±0.26 163.5±0.56 167.6±0.86 39.02±5.08 39.94±5.69 N = number of specimens, SD = standard deviation, M = male specimens, F = female specimens.

Fig.2. Variation of mean (±SD) gonado-somatic index of female (F) and male (M) Hemiculter leucisculus from Sefid River, in the southern Caspian Sea basin at monthly intervals throughout the year. According to gonad examination, all of the male age of 2+, 3+ or 4+ years. Individual values of the and female specimens with 93.0 and 99.7mm in total absolute fecundity varied in a wide range from length respectively and more than one year old 2749.4 to 58028.1 eggs with an average of 19978.6 (approximately 2 years old) were ripe in both sexes. eggs (SD ± 14761.07) (Table 2). The Fecundity Out of 235 specimens, 136 were females and 99 relative to total weight was 139.8 to 1272.6 with a males. The sex ratio 1:1.37 was significantly deviated mean of 607.1 (SD ± 299.89) per gram body weight from an ideal 1:1. (x2 = 5.582, P<0.05). (Table 2). Absolute fecundity was significantly Significant differences were observed between the related to female body weight, age and also gonad female and male gonado-somatic index (P<0.05). weight the regression coefficient values were 0.419, The female gonado-somatic index increased 0.477 and 0.618 for gonad weight, age and total body gradually in March and reached the maximum value weight, respectively (Table 3). But the relationship in August (at water temperature 23ºC). As seen in between absolute fecundity and standard length was Figure 2, it decreased sharply from August to not statistically significant (P >0.05) (Table 3). September. The lowest and highest mean ovum diameters in Discussion reproductive season were observed in April In this study, the most abundant age group in the (0.42±0.25mm) and August (1.11±0.36mm) (Fig. 3). catches was 3+ (41.5% frequency) with average total The fecundity was estimated from the fishes having length nearly 139.6 mm. Holcik and Razavi (1992) 34 Mousavi-Sabet et al.- Reproductive biology of Hemiculter leucisculus

Table 2. Absolute (Fa) and relative (Fr) fecundity in particular ranges of age, body length (SL) and body weight (W) of Hemiculter leucisculus females from Sefid River, the southern Caspian Sea basin.

Parameters N Fa Fr Range Mean ± SD Range Mean ± SD SL (mm) 100 – 140 21 2739.5–52598.3 18288.6±13353.8 139.8–1203.6 593.2±285.4 141–160 9 9963.8–58028.1 21264.3±2268.7 360.1–1272.6 513.8±137.5 10–25 7 2729.5–40280.0 11950.5±10443.5 139.8–919.6 537.3± 218.2 W (g) 25–45 15 7395.4–52598.0 22471.0±13311.8 310.8–1203.6 658.0±275.6 45–65 8 10416.3–23731.2 17073.2±6656.8 360.09–225.9 293.0±67.1 2+ 5 2739.5–23343.6 11142.3±6240.6 139.8–742.6 509.4±186.6 Age 3+ 21 5140.4–52598.0 24793.6±16925.7 225.9–1203.6 674.2±398.3 4+ 4 21991.2–58028.1 34583.1±20322.6 360.1–1272.5 710.1±491.9 N = number of specimens; SD = standard deviation.

Fig.3. The mean oocyte diameter (mm) of Hemiculter leucisculus in Sefid River from the southern Caspian Sea basin at monthly intervals throughout the year. reported H. leucisculus life span in the Anzali estimated by scale reading the maximum age of Lagoon to reach at least 4 years with 134.5-143.4 mm H. leucisculus in the Anzali Lagoon to be 3+ years standard length, while Tourchi-Moghaddam (2003) with 125mm total length for both sexes. In the Anzali

35 Iran. J. Ichthyol. (March 2019), 6(1): 31-40

Table 3. Correlation coefficients r and regression equations for relationships between absolute fecundity (Fa) and body length (SL), body weight (W), weight of ovary (Wg) and age.

Relationship N Linear Regression r r2 F-value P-value Fa – SL 30 Y = -1261.21x+35517.73 0.178 0.032 0.721 0.405 Fa – W 30 Y = 699.58x – 2155.61 0.618 0.382 13.605 0.001 Fa – Wg 30 Y = 1976.78x – 10a044.00 0.419 0.176 4.696 0.041 Fa – Age 30 Y = 7822.06x – 12287.45 0.477 0.227 6.475 0.018

Lagoon, maximum and minimum of total length for of food are consumed (Wootton 1979; Mousavi- this species reported 200.0 and 148.0mm, Sabet 2012; Heidari et al. 2013; Mansouri-Chorehi et respectively (Moradinasab et al. 2012). Patimar et al. al. 2016; Mohammadi-Darestani et al. 2016; (2008) examined the age structure of H. leucisculus Mousavi-Sabet et al. 2012c, 2017; Pourshabanan et in the Alma-Gol, Adji-Gol and Ala-Gol. They al. 2017; Faghani-Langroudi and Mousavi-Sabet reported the specimens with 3+ and 2+ years old; in 2018). the mentioned region they had the total length of In this study, the average absolute fecundity of 177.15, 163.33 and 165.71mm, respectively. female H. leucisculus from the Sefid River was about Sharpbelly of the wetlands from the southeastern 19978.6 eggs (SD ± 14761.1), ranging from 2749.4 Caspian Sea basin has relatively greater body length to 58028.1 eggs, and the absolute fecundity of H. than those from the same age in the Anzali Lagoon leucisculus caught in Alma-Gol, Adji-Gol and Ala- (Holcik & Razavi 1992; Tourchi-Moghaddam 2003). Gol (Patimar et al. 2008) were comparable to the The monthly gonado-somatic index showed that estimated value for the fish in the Sefid River and the reproductive period of H. leucisculus began from equaled 15058, 12040 and 14142 eggs on average, March and ended in August when the condition respectively. The linear function was found well factor is low in its annual trend (see Mousavi-Sabet enough for expressing the Fa–W (g), Fa–Age and Fa et al. 2013). Condition factor influences the –Wg (g) relationships for this species in Sefid River reproductive cycle in fish (Wootton 1979; Mousavi- (Table 3). Also, Patimar et al. (2008) reported the Sabet et al. 2013). According to obtained results, the existence of a positive significant relationship gonado-somatic index values increases in March and between the absolute fecundity and age, gonad got its highest values in August, then decreased weight and body weight in H. leucisculus, but there sharply from August to September. Also, in the one is not any significant relationship between the Fa– swamp (Hendeh Khaleh) in the Guilan Province, SL. Many factors, such as the size and age of the Iran, spawning of H. leucisculus took place in August females (Thrope et al. 1984), life history strategy (Coad 2017), whereas Patimar et al. (2008) reported (Morita & Takashima 1998), and food supply and a peak spawning for H. leucisculus in March and water temperature (Fleming & Gross 1990) affected April in the international wetlands of Alma-Gol, the fecundity. Ayoade (2011) suggested that Adji-Gol and Ala-Gol. In species which spawn in late variation in fecundity may be due to differential spring and in summer such as H. leucisculus, and its abundance of food. According to Jonsson and sympatric species in the same basin or in adjacent Jonsson (1999), fecundity increases with body size basins in the region (e.g., the genera Alburnoides, because the amount of energy available for egg Alburnus, Cyprinion, Ponticola, Cobitis and production and the body cavity accommodating the Oxynoemacheilus), the index remains low in winter eggs increases with fish size. The length seems to be and then rises sharply just before the spawn then a better indicator of the capacity of oocyte production rapid increase in the weight of ovaries takes place than the weight, because fish dose not decrease when the temperature rises and increasing amounts significantly in size whereas weight may vary along 36 Mousavi-Sabet et al.- Reproductive biology of Hemiculter leucisculus

the year (Bagenal 1967). Usually, the relative 6(5): 571-577. fecundity (number of eggs per body weight) is higher Bagenal, T.B. & Braum, E. 1978. Eggs and early life history. In: T. Bagenal (ed.), Methods for assessment in small females than in large ones (Lobon-Cervia et of fish production in freshwaters. 3rd edn. Blackwell al. 1997). Bagenal & Braum (1978) asserted that fish Scientific, Oxford, UK. pp: 106. species exhibit wide fluctuation in fecundity among Bagenal, T.B. 1967. A short review of fish fecundity. In: fish of the same species, size and age. The effects of S.D. Gerking (Ed.), The Biological basis of fresh fishery management on population dynamics depend water fish production. Blackwell Scientific, Oxford, UK. pp: 89-111. on the biological and environmental characteristics Biswas, S.P. 1993. Manual of methods in fish biology. of the fish (Mousavi-Sabet et al. 2015a, b). South Asian publishery. New Delhi, India, pp: 79–91. As conclusion, the present study on some aspects Borisova, A.T. 1972. Accidental invaders in water of reproductive biology of H. leucisculus has bodies of Uzbekistan. Journal of Ichthyology 12: 49- 53. revealed the followings: the peak of breeding season Cao, Y.P. & Wen, Z.G. 1996. The biology of Hemiculter is in August, the average of absolute fecundity in the leucisculus in Bai Yangdian Lake. Journal of Hebei reproduction period ranged from 2749.4 to 58028.1 University (Natural Science Edition) 16: 27-30. eggs that can be expressed as a function of total Chakraborty, B.K.; Mirza, Z.A.; Miah, M.I.; Habib, length (Ahmed 2010), and ovum diameter is 0.42mm M.A.B. & Chakraborty, A. 2007. Reproductive cycle of the endangered sarpunti, Puntius sarana (Hamilton, and 1.11mm in April and August, respectively. The 1822) in Bangladesh. Asian Fisheries Society 20: mature males and females are longer than 93.0 and 145-164. 99.7mm, respectively in total length (2 year in age). Coad, B.W. & Hussain, N.A. 2007. First record of the The results of this study demonstrate that exotic species Hemiculter leucisculus H. leucisculus is a high fecund and short-lived fish (: ) in Iraq. Zoology in the Middle East 40: 107-109. species found in Sefid River. It is expected that Coad, B.W. 1980. Environmental change and its impact population of this species may increase significantly on the freshwater fishes of Iran. Biological in near future which may lead it an abundant species Conservation 19(1): 51-80. in Sefid River. Therefore, its monitoring is Coad, B.W. 2017. Freshwater Fishes of Iran. unavoidable in the sampled river. http://www.briancoad.com (accessed on 9 August 2017). Coad, W.B. & Abdoli, A. 1993. Exotic Fish Species in Acknowledgements the Freshwaters of Iran. Zoology in the Middle East The authors are grateful to the University of Guilan 9: 65-80. for financial supports. Also, thank to Dr. Saber Coates, D. 1988. Length-dependent changes in egg size, and fecundity in females, and brooded embryo size in Vatandoust for his helpful comments. males of fork tailed catfishes (Pisces: Ariidae) from the Sepik River, Papua, New Guinea, with some References implications for stock assessments. Journal of Fish Abdoli, A. 2000. The Inland Water Fishes of Iran. Biology 33(3): 455-464. Iranian Museum of Nature and Wildlife, Tehran. 378 Cortes, E. 2000. Life history patterns and correlations in pp. [In Persian] sharks. Reviews in Fisheries Science 8(4): 299–344. Ahmed, A.; Sharaf, M.M. & Laban, H.A 2010. Eagderi, S.; Mojazi Amiri, B. & Adriaens, D. 2013. Reproduction of the Egyptian Sole, aegyptiaca Description of the ovarian follicle maturation of the (Actinopterygii: Pleuronectiformes: ), from migratory adult female bulatmai barbel ( Port Said, Egypt, Mediterranean Sea. Acta capito, Güldenstädt 1772) in captivity. Iranian Ichthyologica et Piscatoria 40(2): 161-166. Journal of Fisheries Sciences 12(3): 550-560. Ayoade, A.A. 2011. Population characteristics of Esmaeili, H.R.; Gholamifar, A. & Freyhof, J. 2011. Schilbe mystus (Linne, 1758), from two different Ichthyofauna of Zarivar Lake (Iran) with the first habitats: Asejire and Oyan Lakes Southwestern records of Hemiculter leucisculus and Alburnus Nigeria. Journal of Fisheries and Aquatic Science hohenackeri in the Tigris drainage. Electronic Journal 37 Iran. J. Ichthyol. (March 2019), 6(1): 31-40

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مقاله پژوهشی زیستشناسی تولیدمثل گونه مهاجم تیزکولی،(Hemiculter leucisculus (Basilewsky, 1855 از حوزه آبریز جنوبی دریای خزر

سیدحامد موسوی ثابت*2،1، عادله حیدری1، میثم صالحی3

1گروه شیالت، دانشکده منابع طبیعی، دانشگاه گیالن، صومعهسرا، گیالن، ایران. 2پژوهشکده حوضه آبی دریای خزر، دانشگاه گیالن، رشت، گیالن، ایران. 3شرکت آبزیپروری آبزی اکسیر کوثر، بخش کشاورزی، سازمان اقتصادی کوثر، تهران، ایران.

چکیده: تیزکولی، Hemiculter leucisculus، یک گونه مهاجم است که دامنه پراکنش وسیعی را در سراسر آسیا و خاورمیانه گسترش داده است. با این حال، اطالعات کمی از تولیدمثلی آن بهخصوص در مورد الگو و فصل تخمریزی در دسترس است، که میتواند دلیل موفقیت این ماهی را بهعنوان گونه مهاجم توضیح دهد. این تحقیق اولین ارزیابی ویژگیهای تولیدمثلی این ماهی است که با بررسی 235 نمونه که بهصورت ماهانه در طول یک سال از رودخانه سفیدرود در حوضه آبریز جنوبی دریای خزر صید شدهاند، انجام شده است. سن، نسبت جنسی، هماوری، قطر تخمک و شاخص گنادی-بدنی محاسبه شدند. تجزیه و تحلیل رگرسیون به منظور پیدا کردن رابطه بین هماوری و اندازه ماهی، وزن گناد و سن مورد استفاده قرار گرفت. نرها و مادههای بالغ بهترتیب دارای طولی بیش از 0/93 و 7/99 میلیمتر بودند )1+ ساله از نظر سن(. متوسط قطر تخمک از 0.4 میلی متر )اوایل اردیبهشت( تا 1.1 میلیمتر )اواخر تیر( متغیر بود. تخمریزی در تیرماه، زمانی که درجه حرارت آب 23 تا 26 درجه سانتیگراد بود، انجام شد. میانگین مقادیر شاخص گنادی-بدنی برای مادهها در ابتدای دوره تولیدمثل )اوایل فروردین( و در مادههای بالغ رسیده )تیرماه( به ترتیب 90/2 و 97/16 درصد بودند. متوسط هماوری مطلق و نسبی بهترتیب 6/19978 و 1/607 تخمک بود. هماوری مطلق به طور معنیداری با وزن بدن، وزن تخمدان و سن ارتباط داشت. ظرفیت بالقوه تولید مثلی زیاد، تخمکهای نسبتاً کوچک، توسعه کوتاه مدت تخمکها، بلوغ سریع جنسی، طول عمر کوتاه، تحمل طیف وسیعی از شرایط محیطی، همگی بیانگر قدرت تهاجم این ماهی مهاجم در اکوسیستمهای آبشیرین در منطقه آب و هوای معتدل هستند. کلماتکلیدی: شاخص گنادی-بدنی، هماوری، اکوسیستم آبشیرین، فصل تخمریزی.

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