Iran. J. Ichthyol. (March 2021), 8(1): 1-13 Received: August 21, 2019 © 2021 Iranian Society of Ichthyology Accepted: July 28, 2020 P-ISSN: 2383-1561; E-ISSN: 2383-0964 doi: 10.22034/iji.v8i1.434 http://www.ijichthyol.org

Research Article

Reproductive biology of Indian halibut, Psettodes erumei from the northern Persian Gulf and Oman Sea (Teleoststei: Psettodidae)

Mehrnaz GHANBARZADEH*1, Ehsan KAMRANI2, Mohammad Sharif RANJBAR1, Ali SALARPOURI3, Carl WALTERS4

1Department of Marine biology, Faculty of Marine Sciences and Technologies, University of Hormozgan, Bandar Abbas, Iran. 2Fisheries Department, Faculty of Marine Sciences and Technologies, University of Hormozgan, Bandar Abbas, Iran. 3Persian Gulf and Oman Sea Ecological Center, Iranian Fisheries Science Research Institute (IFSRI), Agricultural Research Education and Extension Organization (AREEO), Bandar Abbas, Iran. 4Fisheries Centre, The University of British Columbia, 2204 Main Mall, Vancouver, BC V6T 1Z4, Canada. *Email: [email protected] Abstract: The reproductive biology of Indian halibut, Psettodes erumei, a commercially valuable flatfish , in the northern Persian Gulf and Oman Sea was studied. A total of 433 specimens were collected monthly from October 2016 to November 2017, and their length, weight, sex, gonad and liver weight, and maturity status were recorded. Monthly variations in Gonado-somatic index showed two obvious peaks in the spawning period in both sexes. Observations from the seasonal maturation curve, the frequency distribution of the female gonadal development stages and the condition factor confirmed recent findings that the spawning periods have two peaks, a higher peak in April-May (spring) and a lower peak in October (autumn). The size at first maturity of females was estimated to be 38.20 cm in total length. The absolute fecundity ranged from 18072 to 332171 eggs while the relative fecundity ranged from 22 to 161 eggs/g. Absolute fecundity was positively correlated with body related parameters (total length/weight) and with gonad weight. Combined, the information provided in this study could be useful for resource managers to set seasonal closure during peak spawning periods and implement minimum size limits above the size at first maturity to let the fish spawn at least once before capture. Keywords: Reproduction, Maturity stages, Sex ratio, Flatfish, Iran. Citation: Ghanbarzadeh, M.; Kamrani, E.; Ranjbar, M.Sh.; Salarpouri, A. & Walters, C. 2020. Reproductive biology of Indian halibut, Psettodes erumei from the northern Persian Gulf and Oman Sea (Teleoststei: Psettodidae). Iranian Journal of Ichthyology 8(1): 1-13.

Introduction usually finds in the shallower parts of the continental The Indian halibut, Psettodes erumei (Bloch and shelf at depths between 20 and 50m (Hensley 1997), Schneider, 1801), is a large widespread commercial where it feeds on nektons and benthos flatfish of the order Pleuronectiformes/ (Devadoss et al. 1977). Psettodes erumei is the only Carangiformes that occurs in the Indian Ocean and species belonging to the Psettodidae family that lives the southwestern part of the Pacific Ocean, from the in the Persian Gulf and Oman Sea (Hensley 1997; Red Sea and eastern Africa to the coasts of Papua Yasemi et al. 2008; Eagderi et al. 2019). In the north New Guinea and Australia (Nelson 2006; Fricke et of the Persian Gulf and Oman Sea, P. erumei is a al. 2020). It inhabits sandy and muddy bottoms highly valued commercial species mainly caught (Hensley 1997; Kuiter & Tonozuka 2001) and with bottom trawl and composes a high percentage

1

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Fig.1. The annual yield of Psettodes erumei in the north of the Persian Gulf and Oman Sea (Hormozgan Province, Iran) (IFO 2017).

(0.21% of total catch) of the commercial shrimp Persian Gulf and Oman. This information includes fishery catch (Kazemi et al. 2013; Eighani & reproduction period, the size at first sexual maturity, Paighambari 2014). In the northern Persian Gulf and sex ratio, and fecundity; such information is critical Sea of Oman, the official catch statistics do not for the sustainability of P. erumei stock. separate between P. erumei and other flatfishes. In recent years, the catch trend of halibuts, including Materials and Methods P. erumei, has increased in the study area (Fig. 1) A total of 433 specimens of P. erumei were collected (IFO 2017). This increase in catch requires basic monthly during the period between October 2016 and knowledge on the reproductive biology of P. erumei, November 2017 by gill net and bottom trawl (at to avoid overexploitation. depths between 20 and 50m) from three sampling The biology of P. erumei has been studied in sites along the northern parts of the Persian Gulf and different regions such as the Bombay waters Oman Sea (Hormozgan Province, Iran; Fig. 2). The (northwest Indian coast) (Pradhan 1965), the vicinity samples were collected randomly and transferred in of Dar es Salaam and Kunduchi in western Indian iceboxes to the laboratory. Ocean (Darracott 1977), the Porto Novo coast For each fish, the total length (TL) was measured (southeast Indian Ocean) (Devadoss et al. 1977; to the nearest cm and the total weight (TW) to the Ramanathan & Natarajan 1979), the coast of Orissa nearest g. Then a ventro-lateral section was made for (northeast Indian Ocean) (Das and Mishra 1990) and each specimen and the individuals were sexed by the Kota Kinabalu, Sabah, Malaysia (Yoshikawa et macroscopic examination of the gonads as females, al. 2012). Despite the commercial importance of males and undetermined (for specimens which could P. erumei in the Persian Gulf and Oman Sea (Iran), not be sexed). very little is known about its reproductive biology; To investigate the reproductive biology, gonadal the only data on its reproductive strategy were development (the maturity stages of females) was reported by Azh et al. (2014). The purpose of the recognized macroscopically according to the ovarian present study is to provide detailed information on maturity scale proposed by (Biswas 1993). Five the breeding biology of P. erumei in the north of the maturity stages were identified: I-immature, II- 2 Ghanbarzadeh et al- Reproductive biology of Indian halibut

Fig.2. Location of the sampling sites in different regions of Hormozgan harbor, Iran; *- sampling sites. developing, III-maturing, IV-mature gonads and To estimate the first maturation length (the length spawning, V-spent. at which 50% of individuals are mature (Lm50)), the The gonads and liver of all specimens were following equation was applied (King 2007): = 1 removed and weighed to the nearest 0.1g and for 1 + ( ) determination of the spawning period, the monthly Where Pl is 𝑃𝑃the𝑙𝑙 proportion−𝑟𝑟 𝑚𝑚of𝑙𝑙− 𝑙𝑙reproductive/adult𝑚𝑚50 mean Gonado-somatic index (GSI = (gonad weight/ � 𝑒𝑒 females for each size-class; rm is the rate of change total weight) × 100 and Hepatosomatic index (HIS = from non-reproductive to reproductive status; l is the liver weight/total weight) × 100 were estimated total length (cm) and Lm50 is the size at first sexual (Mazlan and Rohaya 2008). The spawning period maturity (cm). For parameter estimation, the was also confirmed by the determination of a proportion of mature individuals (with gonads in monthly percentage occurrence of different gonadal stages IV and V) was calculated in each body size- development stages. Because the gonads of females class at an interval of 2cm. Then Lm50 was estimated show greater temporal weigh changes than males by using the mentioned logistic equation by (Chellappa et al. 2003), we described the spawning maximizing the likelihood of binomial distribution period of the species according to the female’s by using the Excel add-in tool “Solverˮ (Tokai & gonads (West 1990). Mitsuhashi 1998). In order to link resource allocation to the The sex ratio (male to the female) was calculated reproductive process, the mean of the total condition according to the proportion of the two sexes relative factor (K), for both sexes, and the mean of the to one another and it was statistically tested for any somatic condition factor (K’), only for females, were significant deviation from the expected 1:1 ratio. calculated monthly using the following formula: K = For the estimation of fecundity (the number of ripe (total weight)/ (total length)b, K’ = (total weight- b eggs in the female fish before the next spawning gonad weight)/(total length) , b = angular coefficient period) the “sub-sampling dry gravimetric methodˮ determined from the length-weight relationship. modified after Simpson, was applied (Simpson 3

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Fig.3. Monthly variations of the mean Gonado somatic index (GSI) of (A): females and (B): males of Psettodes erumei in the north of the Persian Gulf and Oman Sea. The error bars correspond to the standard deviation.

1959a, b). A sum of 30 mature ovaries (ovarian and absolute fecundity- gonad weight (Ekanem developmental stage IV) was used for the assessment 2000). of fecundity. After removing and weighing of Statistical analyses were performed using ovaries, three sub-samples were taken from the front, Microsoft Excel 2016 and SPSS software (Version mid and rear sections of each ovary, weighed (all 22). All data were tested to check normality using the were approximately 5g) and preserved in Gilson’s Kolmogorov-Smirnov and Shapiro Wilk tests. Where solution for 3 months. Preserved gonads were then the test for normality assumption was not met, the washed gently, eggs were placed in a Petri dish and non-parametric tests were used to determine air-dried for 72 hours at room temperature. Then differences. To test for any significant deviation of these eggs were weighed (for each fish separately) the sex ratio from the expected 1:1 ratio, the Chi- and three subsamples of 0.02g eggs from each fish Square test was used. All statistical effects were were taken and the number of eggs in each subsample considered significant at α=0.05. was counted under the binocular microscope and the mean number of eggs was calculated. Then the total Results number of eggs in each ovary sub-sample was Of the specimens examined, 231 (53.35%) were proportionally estimated using the following females and 177 (40.88%) males. The sex of the formula: remaining 25 (5.77%) specimens could not be = determined. Females ranged from 20.00 to 64.00cm TL (44.16±7.34cm) and from 147.80 to 4068.00g Where n is the number1 𝑛𝑛𝑛𝑛𝑛𝑛of eggs in the sub-sample, 𝐹𝐹 �𝑔𝑔 GW is the total weight of gonad, g is the weight of TW (1370.17±671.99g). Males ranged from 21.80 to sub-sample (here=0.02g). Then by taking the mean 57.50cm TL (36.60±5.21cm) and from 167.00 to 3305.00g TW (731.14±393.40g). number of three sub-sample fecundities (F1, F2, F3), the absolute fecundity for each female fish was For females, different gonadal development stages estimated as: were easily identified. The monthly values of GSI FA = (F1 + F2 + F3) /3 ranged from 0.74 to 7.67 (2.85±2.27) in females and For the estimation of relative fecundity (FR), from 0.30 to 1.39 (0.66±0.32) in males. Investigation individual fecundity was divided by the weight of the of monthly GSI mean values showed two peaks in respective fish. Regression analyses and linear both sexes; one in spring and the other in autumn. functions were used to describe the relationship GSI mean values were high from February to May between absolute fecundity- fish total length/weight (both sexes), with peaks in April and May for females

4 Ghanbarzadeh et al- Reproductive biology of Indian halibut

Fig.4. Monthly percentage occurrence of different gonadal development stages of females of Psettodes erumei in the north of the Persian Gulf and Oman Sea. and males, respectively (spring peak; Fig. 3). The GSI then decreased to a minimum value in August followed by an increase from September to October with a peak in October for both sexes (autumn peak; Fig. 3). Statistical analysis of this index for both females and males showed significant differences throughout the study period (P<0.05). The reproduction period of P. erumei in spring and autumn was confirmed by the monthly percentage occurrence of female gonadal development stages (Fig. 4). Monthly variations of HSI in females showed the same trend as the GSI. Monthly values varied from 1.09 to 1.78 (1.37±0.20) and the maximum values occurred in April and October. In contrast, in males, monthly values varied from 0.80 to 1.53 (1.14±0.23) and HSI had the maximum values in post spawning periods (Fig. 5). Statistical analysis showed significant differences in both sexes throughout the study period (P<0.05). Fig.5. Monthly variations of the mean Hepatosomatic index In females, the mean values of K were high from (HSI) of (A): females and (B): males of Psettodes erumei in the north of the Persian Gulf and Oman Sea. The error bars March to May, with a peak in April-May. In males, correspond to the standard deviation. these values were high from January to May, with a peak in May. Then, the mean values of K decreased throughout the study period (P<0.05). The mean to a minimum value in June and again followed by an values of K showed significant differences between increase from July to October with a peak in October the two sexes throughout the study period and the for both sexes. Statistical analysis showed significant mean values were significantly higher in females differences in the mean values of K in both sexes 5

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Table 1. The sex ratio of Psettodes erumei during different months in the north of the Persian Gulf and Oman Sea.

Chi Months Male Female Total Male : Female square P value (χ2) Number % Number % Oct-16 24 72.73 9 27.27 33 1 : 0.38 6.82 P<0.05 Nov-16 8 21.62 20 54.05 37 1 : 2.50 5.14 P<0.05 Dec-16 22 73.33 8 26.67 30 1 : 0.36 6.53 P<0.05 Jan-17 9 30.00 18 60.00 30 1 : 2.00 3.00 P>0.05 Feb-17 15 50.00 15 50.00 30 1 : 1.00 0.00 P>0.05 Mar-17 12 40.00 18 60.00 30 1 : 1.50 1.20 P>0.05 Apr-17 21 58.33 15 41.67 36 1 : 0.71 1.00 P>0.05 May-17 7 23.33 16 53.33 30 1 : 2.29 3.52 P>0.05 Jun-17 14 46.67 10 33.33 30 1 : 0.71 0.67 P>0.05 Jul-17 3 10.00 27 90.00 30 1 : 9.00 19.20 P<0.05 Aug-17 8 26.67 22 73.33 30 1 : 2.75 6.53 P<0.05 Sep-17 17 56.67 13 43.33 30 1 : 0.76 0.53 P>0.05 Oct-17 11 36.67 19 63.33 30 1 : 1.73 2.13 P>0.05 Nov-17 6 22.22 21 77.78 27 1 : 3.50 8.33 P<0.05 Total 177 40.88 231 53.35 433† 1 : 1.31 7.15 P<0.05 † 433= 177 males, 231 females, 25 Undetermined than that of males (P<0.05). The greatest differences between the values of K and K’ (females) occurred from March to May and from September to November (highest GSI mean values). The sexual maturity logistic curve for females of P. erumei is shown in Figure 6. The total length of 38.20cm corresponded to 50% of the sexually mature female ovary samples. Also, the lengths at which 25, 75 and 100% of individuals were able to actively participate in the reproductive process, were 34.10, 41.90 and 58.00cm, respectively. A total of 231 females and 177 males were observed with a ratio of 1:1.31 (M:F) over the study period which was significantly different from the expected ratio of 1:1 (P<0.05). The monthly distribution of the sex ratio showed that the sexes did not occur in the same proportion throughout the year Fig.6. Maturity logistic curve for females of Psettodes and this difference was statistically significant in erumei in the north of the Persian Gulf and Oman Sea. some months (Table 1). The sex ratio per length class showed significant differences, except in the class ranged from 18072 eggs (in a fish with 37.6cm TL that varied from 25.1-30.0cm and in the length and 806g TW) to 332171 eggs (in a fish with 56.4cm classes of ≤25 and >60cm. TL and 2920g TW). Mean fecundity of The number of eggs in each mature ovary (FA) 157833±7476.4 eggs per female was estimated. The 6 Ghanbarzadeh et al- Reproductive biology of Indian halibut

number of eggs per gram body weight (FR) and per (1990) reported that the highest reproductive activity gram gonad weight varied from 22 to 161 of P. erumei occurred from September to December (101±29.24) and from 1021 to 2695 (1796±362.81), with a peak in October in the coast of Orissa. respectively. There was a strong positive correlation Ramanathan & Natarajan (1979) noted that the between absolute fecundity-total length (r = 0.87, spawning periods of this flatfish extended from May P<0.01), fecundity-weight (r = 0.86, P<0.01) and to September with a peak from May to August in the between absolute fecundity-gonad weight (r=0.91, Porto Novo coast. Spawning activity of P. erumei P<0.01). The regressions between absolute occurred from September to February in the Western fecundity-total length/weight were F=14529 TL- Indian Ocean (Darracott 1977). Pradhan (1965) 510287 (R2=0.7626) and F=177.63TW-100673 (R2= reported a short spawning period (during September 0.7368), respectively. The regression between and October) for this species in the Bombay coast. absolute fecundity-gonad weight was F = 1289.2 The spawning of P. erumei occurred from September GW+39044 (R2=0.8261). to January with peaks from October to November along the Trivandrum coast (Abraham & Nair 1976). Discussion In general, the existence of some differences in the In the presentation above, we have studied the reproductive period of this species in different reproductive biology of P. erumei a widely regions is not surprising, because geographic distributed commercial species in the Persian Gulf variations in the timing of reproduction have been and Oman Sea (Valinassab et al. 2006). Monthly observed in many fish species (Caputo et al. 2001). variations of GSI in P. erumei showed two obvious Indeed, these differences are likely due to differences peaks, one in spring and the other in autumn (Fig. 3). in environment features of the habitats, such as This finding is corroborated by some other temperature and photoperiod (Bye 1984; Jiménez et researchers who investigated the reproductive cycle al. 1998). of fish species in the Persian Gulf and Oman Sea, In females, HSI and GSI had the same trend. HSI where they reported that spring and autumn to be the is associated with the liver energetic reserves and periods of the highest reproductive activity for a large metabolic activity and its variations imply energy number of resident fish species of these areas (Dadzie storage for reproduction (Garcia-Diaz et al. 2006). et al. 2000; Kaymaram et al. 2010; Abaszadeh et al. Our results imply that P. erumei stores its energetic 2013; Alizadeh et al. 2014; Ghaffari et al. 2015). In reserves in the liver during the gonadal maturation the Porto Novo coast, Devadoss et al. (1977) reported periods and it may use these reserves for the energetic a prolonged spawning season-extending between requirements of the spawning activity. November and May for P. erumei, which is Some variations were observed in K and K’ over consistent with the findings of this study. On the the study period. Before the gonadal maturation peak, other hand, a different finding was obtained by Azh the mean values of K were approximately high due to et al. (2014), where they observed a short spawning the active feeding of individuals (both sexes) season (October- January) for the same species in the (Ghanbarzadeh et al. 2020). During the gonadal northern parts of the Persian Gulf. Different results maturation peak, there was a sharp increase in the were also recorded for P. erumei in other regions; for mean values of K and the highest values were example, P. erumei exhibited few changes in GSI in observed in April-May and October for females and Kota Kinabalu, Malaysia, throughout the year and so in May and October for males. Some researchers the study concluded that this species might spawn stated that condition factor does not merely reflect year-round (Yoshikawa et al. 2012). Das & Mishra the feeding condition of the adult stage, but includes 7

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the state of gonadal development, based on the reported that the spawning stage in females of consumption of fat reserves during the spawning P. erumei in the Kota Kinabalu, Malaysia, begins at period (Vazzoler & Vazzoler 1965; Martins-Juras lengths larger than 33.4cm (standard length). Size at 1980). So, in the present study, one reason for first maturity of P. erumei in the Porto Novo coast increasing the mean values of K in April, May, and was between 37.1 and 39 cm TL (Ramanathan & October could be related to the development of Natarajan 1979). Abraham & Nair (1976) reported a gonads, because the growth of gonads alters the body significantly small size at first maturity for specimens weight of the fish significantly. After gonadal from the Trivandrum coast. In the Bombay coast, maturation peak, which coincides with spawning, the Lm50 of P. erumei was higher than that observed in mean values of K decreased, because in these periods the north of the Persian Gulf and Oman Sea (especially in June) individuals did not have much (Lm50=41.1cm TL) (Pradhan 1965). It seems that active feeding (Ghanbarzadeh et al. 2020), and the geographical distribution and consequently different weight of their gonads had decreased due to the environmental conditions (such as food productivity, spawning process (both sexes). After these periods water salinity, rainfall, etc.) could be important and from July to September, there was a gradual factors, which generally influence the maturity of increase in the mean values of K due to the active fishes (Smida & Hadhri 2014). Another important feeding, and simultaneous with the second gonadal factor that could influence Lm50 from one area to maturation peak in October, a sharp increase another is the increase in fishing pressure observed in the mean values of K in both sexes. (Lappalainen et al. 2016), which can lead to According to the results, the nature of monthly moderation in growth and a decrease in maturation variations of K values in females and males is almost size (Mcphail et al. 2001). similar, but the mean values of K in females were Sex ratio is an important indicator of the significantly higher than those of males. One reason population state and the sustainability of species could be related to this fact that the testes do not grow (Kartas & Quignard 1984) and is generally close to as large as ovaries and so they do not change the body 1:1 for the majority of species (Nikolski 1963). weight of the fish to a large extent. On the other hand, However, the sex ratio of P. erumei showed a and based on the results of Gastro-somatic index, significant difference between the sexes over the feeding intensity in females was higher than males entire study period and there were more females than (Ghanbarzadeh et al. 2020) which could impress the males. This pattern is consistent with almost all fish values of K. species that do not form schools (e.g., anchovies, Knowledge of the first maturation size is useful sardines)–and similar sex ratio pattern was observed for regulating fisheries activity and fish resource for fishes inhabiting the Persian Gulf and Oman Sea management (for example for regulating mesh size (Valinassab et al. 2008; Abaszadeh et al. 2013; and avoiding recruitment overfishing) and also for Hamzeh et al. 2017). The departure of sex ratio from implementation of aquaculture activities (Fontoura et the expected 1:1 could be the result of some factors, al. 2009; Bindu & Padmakumar 2014). In the present such as partial segregation of mature forms through study, the size at first maturity was found to be habitat preference (Ahmed et al. 2010) or due to 38.2cm total length for females. This value is near to differential migration and/or other behavioral that found for the same species in the north of the differences between sexes, rendering one sex more Persian Gulf and Oman Sea (Lm50=37.3cm TL) likely to be caught than the other (Mehanna 2014). (Ashghaly Farahani et al. 2005; Kamali et al. 2006). The works carried out by Pradhan (1965), Some regional variations were found in size at first Ramanathan & Natarajan (1979), Yoshikawa et al. maturity of this species; Yoshikawa et al. (2012) (2012) and Azh et al. (2014) with the same species, 8 Ghanbarzadeh et al- Reproductive biology of Indian halibut

were four of the works that corroborated the would only deteriorate the problem by affecting prediction above. Investigation of sex ratio per length mainly the larger ones (Kamler 2005). Nevertheless, class showed significant differences in most length different results on the fecundity of a given species classes, where females considerably outnumbered could be related to some factors such as the age of males in the larger size-classes (>40cm TL). This is samples, different environments and conditions in a common pattern in fish studies, because, generally, which fish live under them (Zijlstra 1973). Since body size in females is favored by the increase in fecundity is greatly dependent upon nutrition, some fecundity; while in males, size is affected by sexual differences in the fecundity of a given species from selection (Shine 1990). various geographical areas may be as a result of The absolute fecundity (FA) of P. erumei females differences in food supply (Rahman et al. 2016). fluctuated from 18072 to 332171 eggs for lengths During the present investigation a straight line between 37.6 and 56.4cm, respectively. Das & relationship was found between fecundity and body- Mishra (1990) reported the fecundity of P. erumei related parameters (total length/weight) and between ranged from 59875 to 187414 eggs (for lengths fecundity and gonad weight; and it has been observed between 35.5 and 47.5cm) from the coast of Orissa. that the correlation coefficient (r) between absolute On the other hand, Abraham & Nair (1976), reported fecundity and gonad weight was higher than those a fecundity range of 33000 to 82079 eggs for lengths observed between absolute fecundity and total of 36-46cm from Trivandrum coast. Both the weight. These results show that absolute fecundity previous findings are nearly in accordance with the was more dependent on gonad weight than body present results. The value of fecundity of P. erumei related parameters. So, gonad weight is a better index in the Porto novo coast ranged from 303840 to of fecundity than body related parameters. On the 1316735 eggs for lengths between 38 and 55.8cm other hand, absolute fecundity was more closely (Ramanathan & Natarajan 1979). Pradhan (1965) related to total length than total weight, however the reported that fecundity of the same species in the differences were not so great. The results of earlier Bombay coast ranged from 313803 to 1070354 eggs investigations on P. erumei in different parts of India (for the lengths of 43.5-52.4cm). In the two aforesaid also support this finding (Ramanathan & Natarajan studies, the observed fecundity was very high when 1979; Das & Mishra 1990). In summary, this study compared to the results of fecundity obtained in this has provided some basic and valuable information on study. The relative fecundity (FR) of P. erumei in the the reproductive biology of P. erumei in the north of different locations of India ranged from 425 to 1095 the Persian Gulf and Oman Sea. This information is (Porto Novo coast) (Ramanathan & Natarajan 1979) useful for the proper fisheries management, and from 123 to 349 (Trivandrum coast) (Abraham conservation, and aquaculture of this species in the & Nair 1976). Whereas, the relative fecundity in this region. Also, the information of this study may study, was found to vary from 22 to 161 with a mean provide a basis for future studies on life-history value of 101.0±29.24 which is considerably lower parameters of other flatfish species in the region. than any populations of India. As reported by many researchers, relative fecundity may change over time Acknowledgements within species, although the changes seem to be The authors would like to thank the marine biology relatively modest (Yoneda & Wright 2004; Thorsen & stock assessment department of the Persian Gulf et al. 2006). Another factor that may affect the and Oman Sea Ecological Research Institute for the relative fecundity is the size of species and it may provision of laboratory facilities used in this study increase with size within species, but in this case, this and to A. Ben-Hasan for his technical assistance in 9

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some stages of this study. This research was funded Pleuronectiformes) in the Adriatic Sea. Italian Journal by the University of Hormozgan. of Zoology 68: 107-113. Chellappa, S.; Câmara, M.R.; Chellappa, N.T.; Beveridge, References M.C.M. & Huntingford, F.A. 2003. Reproductive Abaszadeh, A.; Keivany, Y.; Soofiani, N.M. & ecology of a Neotropical cichlid fish Cichla monoculus Falahatimarvast, A. 2013. Reproductive biology of the (Osteichthyes, Cichlidae). Brazilian Journal of greater lizardfish, Saurida tumbil (Bloch, 1795), in Biology 63: 17-26. Bushehr coastal waters of Iran. Turkish Journal of Dadzie, S.; Abou‐Seedo, F. & Al‐Shallal, T. 2000. Zoology 37: 717-722. Reproductive biology of the silver pomfret, Pampus Abraham, M. & Nair, N.B. 1976. Food and feeding habits argenteus (Euphrasen), in Kuwait waters. Journal of and breeding biology of the Indian halibut Psettodes Applied Ichthyology 16: 247-253. erumei (Bloch and Schncider). Aquatic Biology 1: 43- Darracott, A. 1977. Availability, morphometrics, feeding 63. and breeding activity in a multi-species, demersal fish Ahmed, A.I.; Sharaf, M.M. & Laban, H.A. 2010. stock of the Western Indian Ocean. Journal of Fish Reproduction of the Egyptian sole Solea aegyptiaca Biology 10: 1-16. (: Pleuronectiformes: Soleidae), from Das, M. & Mishra, B. 1990. On the biology of Psettodes Port Said, Egypt, Mediterranean Sea. Acta erumei (Bloch & Schn.), an Indian Halibut. Indian Ichthyologica et Piscatoria 40: 161-166. Journal of Fisheries 37: 79-92. Alizadeh, R.; Kamrani, E.; Safaei, M. & Momeni, M. Devadoss, P.; Mahadevan Pillai, P.K.; Natarajan, P. & 2014. Reproductive biology of Sillago sihama in the Muniyandi, K. 1977. Observations on some aspects of Persian Gulf (Hormozgan Province). Journal of the biology and fishery of Psettodes erumei (Bloch) at Oceanography 5: 41-47. (In Farsi with an English Porto Novo. Indian Journal of Fisheries 24: 62-68. abstract) Eagderi, S.; Fricke, R.; Esmaeili, H.R. & Jalili, P. 2019. Ashghaly Farahani, S.; Valinasab, T. & Keyvan, A. 2005. Annotated checklist of the fishes of the Persian Gulf: Length frequency, length-weight relationship and Diversity and conservation status. Iranian Journal of distribution of Indian halibut in the Oman Sea. Iranian Ichthyology 6(Suppl. 1): 1-171. Scientific Fisheries Journal 14: 169-174. Eighani, M. & Paighambari, S.Y. 2014. Performance of a Azh, Z.; Sourinezhad, I.; Kamrani, E. & Ghodrati shojaei, Square Mesh Panel in Capturing Immature Indian M. 2014. Ovary maturation process of Psettodes Halibut in Shrimp Trawlers in the Persian Gulf. erumei in the coastal waters of the northern Persian Journal of the Persian Gulf (Marine Science) 5: 1-8. Gulf (in Persian). Journal of Oceanography 10(19): 61- Ekanem, S.B. 2000. Some reproductive aspects of 70. (In Farsi with an English abstract). Chrysichthys nigrodigitatus (Lacepede) from Cross Bindu, L. & Padmakumar, K.G. 2014. Reproductive River Nigeria. Naga, the ICLARM Quarterly 23: 24- biology of Etroplus suratensis (Bloch) from the 28. Vembanad wetland system, Kerala. Indian Journal of Fricke, R., Eschmeyer, W.N. & Van der Laan, R. Geo-Marine Sciences 43: 646-654. 2020. Eschmeyer's catalog of fishes: Genera, Species, Biswas, S.P. 1993. Manual of methods in fish biology. 3 References. (http://researcharchive.calacademy.org/re Nejati subhoshmary. the South Asian publishers Pty search/ichthyology/catalog/fishcatmain.asp). Electron ltd, Daryagam, New Dehli, India. ic version accessed 28.07.2020. Bye, V. 1984. The role of environmental factors in the Fontoura, N.F.; Braun, A.S. & Milani, P.C.C. 2009. timing of reproductive cycles. In: Potts, G.W. Estimating size at first maturity (L50) from &Wootton R.J. (eds.), Fish reproduction, strategies Gonadossomatic Index (GSI) data. Neotropical and tactics. London: Academic Press. Ichthyology 7: 217-222. Caputo, V.; Candi, G.; Colella, S. & Arneri, E. 2001. Garcia-Diaz, M.; González, J.A.; Lorente, M.J. & Tuset, Reproductive biology of turbot (Psetta maxima) and V.M. 2006. Spawning season, maturity sizes and brill (Scophthalmus rhombus) (Teleostei, fecundity in Hacktail Comber (Serranus atricauda)

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12 Iran. J. Ichthyol. (March 2021), 8(1): 1–13 Received: August 21, 2019 © 2021 Iranian Society of Ichthyology Accepted: July 28, 2020 P-ISSN: 2383-1561; E-ISSN: 2383-0964 doi: 10.22034/iji.v8i1.434 http://www.ijichthyol.org

ﻣﻘﺎﻟﻪ ﭘﮋوﻫﺸﯽ ﺑﯿﻮﻟﻮژي ﺗﻮﻟﯿﺪﻣﺜﻞ ﻣﺎﻫﯽ ﮐﻔﺸﮏ ﺗﯿﺰدﻧﺪان (Psettodes erumei) در ﺷﻤﺎل ﺧﻠﯿﺞ ﻓﺎرس و درﯾﺎي ﻋﻤﺎن(ﻣﺎﻫﯿﺎن اﺳﺘﺨﻮاﻧﯽ ﻋﺎﻟﯽ: ﮐﻔﺸﮏ ﻣﺎﻫﯿﺎن)

ﻣﻬﺮﻧﺎز ﻗﻨﺒﺮزاده*1، اﺣﺴﺎن ﮐﺎﻣﺮاﻧﯽ2، ﻣﺤﻤﺪﺷﺮﯾﻒ رﻧﺠﺒﺮ1، ﻋﻠﯽ ﺳﺎﻻرﭘﻮري3، ﮐﺎرل واﻟﺘﺮز4 1ﮔﺮوه زﯾﺴﺖﺷﻨﺎﺳﯽ درﯾﺎ، داﻧﺸﮑﺪه ﻋﻠﻮم و ﻓﻨﻮن درﯾﺎﯾﯽ، داﻧﺸﮕﺎه ﻫﺮﻣﺰﮔﺎن، ﺑﻨﺪرﻋﺒﺎس، اﯾﺮان. 2ﮔﺮوه ﺷﯿﻼت، داﻧﺸﮑﺪه ﻋﻠﻮم و ﻓﻨﻮن درﯾﺎﯾﯽ، داﻧﺸﮕﺎه ﻫﺮﻣﺰﮔﺎن، ﺑﻨﺪرﻋﺒﺎس، اﯾﺮان. 3ﭘﮋوﻫﺸﮑﺪه اﮐﻮﻟﻮژي ﺧﻠﯿﺞ ﻓﺎرس و درﯾﺎي ﻋﻤﺎن، ﻣﺆﺳﺴﻪ ﺗﺤﻘﯿﻘﺎت ﻋﻠﻮم ﺷﯿﻼﺗﯽ ﮐﺸﻮر، ﺳﺎزﻣﺎن ﺗﺤﻘﯿﻘﺎت و آﻣﻮزش و ﺗﺮوﯾﺞ ﮐﺸﺎورزي، ﺑﻨﺪرﻋﺒﺎس، اﯾﺮان. 4ﮔﺮوه ﺷﯿﻼت، داﻧﺸﮕﺎه ﺑﺮﯾﺘﯿﺶ ﮐﻠﻤﺒﯿﺎ، وﻧﮑﻮور، ﮐﺎﻧﺎدا.

ﭼﮑﯿﺪه: ﻫﺪف از اﻧﺠﺎم اﯾﻦ ﻣﻄﺎﻟﻌﻪ، ﺗﻌﯿﯿﻦ ﭼﺮﺧﻪ ﺗﻮﻟﯿﺪﻣﺜﻞ ﻣﺎﻫﯽ ﮐﻔﺸﮏ ﺗﯿﺰدﻧﺪان (Psettodes erumei) ﺑﻪ ﻋﻨﻮان ﯾﮏ ﻣﺎﻫﯽ ﭘﻬﻦ ﺑﺎ ارزش اﻗﺘﺼﺎدي در ﺷﻤﺎل ﺧﻠﯿﺞ ﻓﺎرس و درﯾﺎي ﻋﻤﺎن ﺑﻮد. ﺟﻬﺖ اﻧﺠﺎم اﯾﻦ ﺗﺤﻘﯿﻖ، 433 ﻧﻤﻮﻧﻪ از ﻣﻬﺮﻣﺎه 1395 ﺗﺎ آﺑﺎنﻣﺎه 1396ﺑﻪ ﺻﻮرت ﻣﺎﻫﺎﻧﻪ ﺟﻤﻊآوري ﺷﺪﻧﺪ و ﻃﻮل ﮐﻞ، وزن، ﺟﻨﺴﯿﺖ، وزن ﮔﻨﺎد و ﮐﺒﺪ و وﺿﻌﯿﺖ رﺳﯿﺪﮔﯽ ﺟﻨﺴﯽ ﺑﺮاي ﻫﺮ ﻧﻤﻮﻧﻪ ﺛﺒﺖ ﺷﺪ. ﺗﻐﯿﯿﺮات ﻣﺎﻫﺎﻧﻪ ﺷﺎﺧﺺ ﮔﻨﺎدي دو ﭘﯿﮏ ﻣﺸﺨﺺ ﺗﻮﻟﯿﺪﻣﺜﻠﯽ را در ﻫﺮ دو ﺟﻨﺲ ﻧﺸﺎن داد. ﻧﺘﺎﯾﺞ ﻣﻨﺤﻨﯽ رﺳﯿﺪﮔﯽ ﻓﺼﻠﯽ ﻧﻤﻮﻧﻪﻫﺎ، ﺗﻮزﯾﻊ ﻓﺮاواﻧﯽ ﻣﺮاﺣﻞ ﺗﻮﺳﻌﻪ ﮔﻨﺎدي ﻣﺎدهﻫﺎ و ﻓﺎﮐﺘﻮر وﺿﻌﯿﺖ، ﯾﺎﻓﺘﻪﻫﺎي اﺧﯿﺮ در ﻣﻮرد وﺟﻮد دو ﭘﯿﮏ ﺗﻮﻟﯿﺪ ﻣﺜﻠﯽ در ﮔﻮﻧﻪ ﮐﻔﺸﮏ ﺗﯿﺰدﻧﺪان، ﯾﮏ ﭘﯿﮏ اﺻﻠﯽ در ﻣﺎهﻫﺎي ﻓﺮوردﯾﻦ-اردﯾﺒﻬﺸﺖ (ﺑﻬﺎر) و ﯾﮏ ﭘﯿﮏ ﻓﺮﻋﯽ در ﻣﺎه ﻣﻬﺮ (ﭘﺎﯾﯿﺰ) را ﺗﺄﯾﯿﺪ ﻧﻤﻮد. اﻧﺪازه در اوﻟﯿﻦ ﺑﻠﻮغ ﺟﻨﺴﯽ ﻣﺎدهﻫﺎ، 20/38 ﺳﺎﻧﺘﯽﻣﺘﺮ ﻃﻮل ﮐﻞ ﺑﻪ دﺳﺖ آﻣﺪ. ﻫﻤﺂوري ﻣﻄﻠﻖ ﺑﯿﻦ 18072 و 332171 ﺗﺨﻤﮏ و ﻫﻤﺎوري ﻧﺴﺒﯽ ﺑﯿﻦ 22 و 161 ﺗﺨﻤﮏ ﺑﻪ ازاي ﻫﺮ ﮔﺮم وزن ﺑﺪن ﺗﻐﯿﯿﺮ ﮐﺮد. ﻫﻤﺎوري ﻣﻄﻠﻖ داراي ارﺗﺒﺎط ﻣﺜﺒﺖ ﺑﺎ ﻃﻮل ﮐﻞ و وزن ﮐﻞ ﺑﺪن و ﻧﯿﺰ وزن ﮔﻨﺎد ﺑﻮد. ﺑﻪ ﻃﻮر ﮐﻠﯽ، اﻃﻼﻋﺎت در اﯾﻦ ﻣﻄﺎﻟﻌﻪ، ﻣﯽﺗﻮاﻧﺪ ﺑﻪ ﻣﺪﯾﺮان ﺷﯿﻼﺗﯽ در ﺗﻌﯿﯿﻦ ﻣﺤﺪودﯾﺖﻫﺎي ﻓﺼﻠﯽ در ﻃﻮل دورهﻫﺎي ﺗﺨﻢرﯾﺰي ﻣﺎﻫﯽ و ﻧﯿﺰ اﻋﻤﺎل ﻣﺤﺪودﯾﺖﻫﺎي ﺣﺪاﻗﻞ اﻧﺪازه ﺻﯿﺪ ﮐﻤﮏ ﮐﻨﺪ و از اﯾﻦ ﻃﺮﯾﻖ ﺣﺪاﻗﻞ ﯾﮑﺒﺎر ﺑﻪ ﻣﺎﻫﯽ اﺟﺎزه ﺗﺨﻢرﯾﺰي ﻗﺒﻞ از ﺻﯿﺪ را ﺑﺪﻫﺪ. ﮐﻠﻤﺎتﮐﻠﯿﺪي: ﺗﻮﻟﯿﺪﻣﺜﻞ، ﻫﻤﺎوري، ﻧﺴﺒﺖ ﺟﻨﺴﯽ، ﭘﻬﻦ ﻣﺎﻫﯽ، اﯾﺮان.

13