Indian Journal of Geo-Marine Sciences

Vol. 43 (12), December, 2014,pp Reproductive biology of lar (fabricius, 1798) in Andaman

Islands

S. N. Sethi*, Nagesh Ram & V.Venkatesan

Madras Research Centre of Central Marine Fisheries Research Institute (CMFRI), Chennai-28, India.

Central Agricultural Research Institute (CARI), Port Blair-744103, Andaman and Nicobar Islands, India

Central Marine Fisheries Research Institute (CMFRI), Cochin -682018, India.

* [E-Mail: [email protected] ]

Received; revised

Reproductive biology of the Monkey river prawn (Macrobrachium lar) was studied in the Rangat river, Andaman Islands, during 2007 to 2009, with the aim of determining the male: female ratio, maturity cycle, spawning seasons and the fecundity in those ecosystems. During 18 months period, 2078 individuals were collected, with a male: female ratio (M/F) of 1: 2.84. Total number of individuals caught per month ranged 65 -172. Males were generally larger than females. Percentage occurrence of reproducing and non-reproducing females, ovigerous females, seasonal occurrence of maturity stages, and maturity indices (GSI and HSI) exhibited that M. lar breeds twice in a year with two breeding peaks (June and November) every year. Highest reproduction indices (GSI) in female were observed in November (4.0 ± 0.07), December (3.88 ± 0.28), May (3.63 ± 0.08), and June (3.88 ± 0.16). In both the sexes, HSI showed inversely proportional relationships with GSI. With respect to average fecundity (F) by length classes, the lowest and highest number of eggs observed was 3090 and 8177, respectively. As for fecundity by weight classes, the lowest number of eggs observed was 4069 and the highest, 8543. Overall mean fecundity irrespective of the length and weight classes was 5402 (± 772.35) eggs per female. Diameter of non-eyed and eyed eggs ranged between 0.32 mm to 0.50 mm and 0.52 mm to 0.68 mm respectively.

[Key Words: Fresh water prawn, Macrobrachium lar, Andaman Islands, Sex ratio, Reproductive biology]

Introduction sex ratio, spawning period, its fecundity and egg size etc. Most of the prawn species of commercial interest Materials and Methods belong to the Macrobrachium genus, distributed in the tropical and subtropical regions of the world1. Currently, there are nearly 210 species of the Study area is a stretch of Rangat River, Andaman Macrobrachium genus known in the world2. The Islands, flowing west to east is located in south east understanding of the reproductive biology of region of India. The river in its upper reaches flows palaemonid prawns is imperative for development through a rainforest zone, while its terrain in the of management and culture programs as well as vicinity of Rangat, small town, Andaman Islands helping to describe sound strategies to preserve where the study was carried out (Fig.1). A total of biodiversity3-4. Studies on the reproductive biology 18 collections of M.lar were collected with the of other palaemonids have been reported by several assistance of fishermen using cast net from Rangat workers5-6 mainly M. rosenbergii7-8, M acanthurus (lat.12° 38' 17.196" N and long. 92° 45' 55.635"E) (Wiegmann, 1936)9,7, M. carcinus (Linnaeus, during December 2007 to May 2009. Collected 1758)10,11 , M. amazonicum (Heller, 1862)12-13 , , M. specimens were placed immediately in plastic jars olfersii (Wiegmann, 1836) 3, M. iheringi (Ortmann, containing clove oil solution. In the laboratory, 1897) 4,Palaemon paucidens (De Haan, 1844)14, P. specimens were segregated according to the size longirostris (Milne-Edwards, 1837)15 and Atya and sex and counted. Total length, TL of the prawn scabra (Leach, 1815)16. However, information was measured with a meter rule to the nearest 0.1 concerning the reproductive biology of M. lar is not mm, while the body weight was measured, to the available in any scientific literature. Present study nearest 0.01 g with an electronic balance. Prawns is to investigate some aspects of the reproductive were grouped into male, female, ovigerous female biology of the fresh water prawn M. lar such as the and undifferentiated prawns. Determination of the maturity stages of M.lar was done by observing “INDIAN J. MAR. SCI .VOL. 45, NO.10, OCTOBER 2014” macroscopic characters as suggested by Singh and measured in micrometer division using ocular Roy (1994)17 and the annual reproductive cycle was micrometer. Ova with diameter 0.02-0.08 mm were determined on the basis of characteristics such as considered as immature and were ignored. percentage of sex ratio in sampling catch Fecundity was determined by counting the number composition, preponderance of reproducing and non of eggs of 30 berried females by following the - reproducing females, percentage of occurrence of methods described by Sarojini20. Specimens were ovigerous females, percentage of maturity of female randomly selected that carried eggs in the initial gonads and Gonado-somatic and Hepato- somatic developmental stages only because egg loss may indices etc. Sexual differentiation was assessed by occur during the incubation period in caridean the observation of secondary sexual characteristics prawns 21. The egg mass was completely removed such as the masculine appendix in the second from the pleopods with a small brush, and the eggs pleopods of the males and its absence in females were placed in Petri dishes for counting under a and in juveniles. Juveniles were not included in the stereoscopic microscope. Egg size measurement is study. To know the homogeneity of the distribution done with the aid of a microscope using miotic of males and females, Chi square test18 was applied. software and eggs were classified by the absence The females undergo series of ovarian and presence of eye spots in the embryo. developmental stages, and the ovary exhibits Results variation in its coloration, which can be visualized Out of 2079 numbers of individuals examined through the carapace. Proportion of reproducing during 18 months of sampling period, 550 were and non-reproducing females was distinguished; by males and 1529 were females. Sex composition of the presence of dark brick red coloured ovaries, M. lar in the examined samples varied monthly, But fertilised eggs in the brood pouch, before with an overall average (M: F) sex ratio of 1: 2.84 undergoing post-pertural moulting while all the with a chi square value of 26.58 for both sexes. immature females having white translucent ovaries, This indicates that the ratio of males to females was with light pink coloured ovaries are considered as significantly different. Females predominated in all non- reproducing females. Reproductive period the months; the ratio varied between 1.90 (Jan. was determined by the presence of ovigerous 2009) and 4.06 (Mar. 2008). High percentage of female during the sampling period. occurrence of females was observed in the month of Gonads and livers undergo regular seasonal and March, and October whereas the same for males cyclical changes in weight in relation to the total was observed in the month of December to weight of the prawn. Such changes are indicative of February and again in the month of September and the spawning season of the prawn and the November. Total length of M. lar male and female relationship is respectively termed as ‘Gonado ranged between 82.0 - 123.18 mm and 73.15-118.21 somatic index’ (GSI) and Hepato somatic index mm respectively. Males are relatively larger than (HSI). Both indices were calculated according to the female (Plate 1). Observations on percentage method of Giese and Pearse 19. occurrence revealed that both the sexes were found The ‘GSI’ and ‘HSI’ were calculated as: throughout the year in natural water bodies of Andaman water. GSI = (weight of gonad X 100)/ Total body weight.

HSI = (weight of liver X 100)/ Total body weight. In order to determine the spawning periodicity of M. lar, matured female prawns were caught: their total length and total weight were measured and preserved in 5% formalin for 3-4 days to harden it. Ovaries were dissected out from mature female to study the size distribution of the ova. Ova were spread on a glass slide and were observed under Fig.1.Map showing the location of sampling site (arrow mark) microscope and the diameter of about 140 ripe ova in Andaman Island, India from each berried female (total six berried) were

2 SETHI et al.: REPRODUCTIVE BIOLOGY OF MACROBRACHIUM LAR (FABRICIUS, 1798) IN ANDAMAN ISLANDS

Plate 2.Ovigerous female of M.lar collected from Rangat River,

Andaman.

Plate1. Male and female individual of M. lar.

Plate 3. Measurement of ova diameter of M. lar using miotic soft ware. During the study period, the maximum percentage of ovigerous females was observed in the months of December 2007, June and November 2008. The ovigerous females were found totally absent in the months from February - April 2008, September, 2008 and again in the months of February - April, Fig.2. Percentage occurrences of reproducing and non 2009 (Fig. 3). These observations revealed that M. reproducing females in Andaman waters. lar breeds twice in a year with two breeding peaks

occurred in the months of June and November every During the present study, maximum percentage of year. Photograph of few specimens of ovigerous reproducing females was observed in the months of female is shown in the Plate 2. June and November with two peaks. Preponderance of non-reproducing females was highest in the months of March and September (Fig. 2). These observations on preponderance of reproducing and non-reproducing females revealed that M. lar breeds twice in a year with two breeding peaks.

Fig.3. Percentage occurrence of Ovigerous female in Andaman waters.

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mean values of GSI and HSI in M. lar. Monthly mean GSI values for females and males were estimated and graphical representation is given in the Fig. 5 and 6.

Fig. 4. Monthly stages of maturity of female of M. lar from Andaman waters.

During the study period, regular seasonal changes in the maturity occur in female gonads. The matured females were found all the months except January to March. The annual reproductive cycle showed two peaks periods. The first peak was observed in May followed by second peak in the month of

October. Maximum percentage of immature females was found in the month of February, March and August. Maximum percentages of maturing females were found in the month of April and September. Similarly maximum numbers of spent females were found in the month of January, July and December respectively (Fig. 4).

Fig.7. Frequency Polygons of Oocytes in 78 -105 mm female M. lar.

In males, maximum GSI was observed during the months of Nov. (2.76 ± 0.31), Dec. (3.61 ± 0.78), May (3.60 ± 0.12) and June (2.91 ± 0.15). Range of GSI during the whole year varied between 1.54 ± 0.24 to 3.61 ± 0.78. Minimum GSI was noticed Fig. 5. Monthly variation in mean GSI and HSI for female M. during the month of Aug. (1.54 ± 0.24) and Feb. lar from Dec ‘07 - May ’09. (1.63 ± 0.26). In females, maximum ovarian index was recorded during the month of Nov. (4.0 ± 0.07), Dec. (3.88 ± 0.28), May (3.63 ± 0.08), and June (3.88 ± 0.16). Range of GSI during the whole year in females varied between 1.68 ± 0.02 to 4.00 ± 0.07.This fluctuations in the mean values in both sexes exhibit two peak breeding periods in the annual reproductive cycle of M. lar. HSI showed inversely proportional relationships with the GSI in both sexes. In males, maximum HSI was observed Fig. 6. Monthly variation in mean GSI and HSI for male M. lar in the months of Feb. (3.93 ± 0.71) and Aug. (3.48 ± 0.18) and minimum in the months of June (1.54 ± from Dec ‘07 – May ’09. 0.12) and Nov. (1.48 ± 0.19) and similar trend was noticed in females also. It was also found that the Maturity indices negative correlation between the GSI and HSI in Overall fluctuations in the reproductive cycle could male is not significantly different (P < 0.05) be demonstrated through the cyclic changes in the

4 SETHI et al.: REPRODUCTIVE BIOLOGY OF MACROBRACHIUM LAR (FABRICIUS, 1798) IN ANDAMAN ISLANDS

whereas it was significantly different in female (P > Discussion 0.05) (Fig.5 and 6). The sex ratio in the natural population of M. Spawning periodicity lar collected in the Rangat River, Andaman was quite similar to that observed by Porto22 and In the present study, 15 matured females of Sampaio23 for the species M. amazonicum M. lar with matured ovaries were used for respectively collected in the Meir Ponte River, determination of ova diameter. Immature eggs in Goisa, and in the Jaguaribe River, Ceará with the the size range of 0.02 – 0.08 mm, which formed the female predominating over the males. Similarly ground stock, were ignored. Distributions of ova in Mantelatto and Barbosa24 studied the species M. six ripe females in the size range of 78 mm to 105 brasiliense in Serra Azul, Sao Paulo discovered the mm were measured (Fig. 7). It is seen from the same, that is, and females predominated over males figure that in the case of all the six matured females, in a proportion of 1: 2.6. The sex ratio varied majority of ova were in the size range of 0.21 mm between the Macrobrachium species. Barros25, to 0.35mm with maximum numbers in 0.26 mm to Ammar13 and Mossolin and Bueno3 studied the M. 0.30 mm in size and they showed a single peak. olfersii population found a predominance of males Appearance of a single peak in the ovary clearly over females at a proportion of 1: 0.53; 1: 0.71 and indicates that all the matured ova are shed at one 1: 0.23, respectively. Antunes and Oshiro 26 studied instance. samples of the species M. potiuna collected in the coastal waters of the State of Rio de Janeiro, Brazil Average fecundity found a ratio of 1: 0.94, that is, a predominance of males over females higher than M. olfersii. Silva27 The berried females carrying fertilized eggs in the studied the species M. amazonicum collected in the periopods were used for the estimation of fecundity. State of Pará discovered the same, that is, males Present observations revealed that mean fecundity predominating over females. Male attained the (F) varied from 3090 to 8177 in the individuals of largest size among all sexual categorical stages in length class of 6.0 to 11.5 cm, whereas by weight M. lar. The studied sexual, size dimorphism could class of 5.1 to 21.0 g, it was ranged between 4069 be resulted from differential growth among the and 8543. The estimated mean fecundity of M. lar sexes because females need to invest more energy was 5402 (± 772.35). Average fecundity/total length into gonadal development. This change has been and average fecundity/total weight may be reported in several other palaemonid species13, 28. expressed by a linear relationship represented in the Mossolin & Bueno29, Fransozo 4, and Mantelatto following equations: &Barbosa 24 studied M. olfersi , M. iheringi and M. brasiliense respectively, also observed the similar F = - 1697.47 + 811.34 L (P > 0.01) differential growth rates between sexes. Mantelatto 24 F = 5632.12 + 4. 439 W (P < 0.01) and Barbosa also reported the additional fact for dimorphism that the larger size reached by males Where F = average fecundity; L = total length (cm); may be related to domination over females, as well and W = total weight (g). pre-adult males, during the copulation process. In the present study, low numbers of adult prawns The obtained result infers that there is no significant were collected. This could be probably related to relationship between the fecundity (F) and the large quantity of shelters available in the river edge, length classes (L) however, significant relationship together with several aquatic macrophyte species, exists between the fecundity (F) and the weight reducing their chance of capture. This truth was also classes (W). Diameter of non-eyed eggs ranged observed by Galvao and Bueno 30 in Atya scabra between 0.32 mm to 0.50 mm whereas those of (Leach, 1815). The four different stages of ovarian eyed eggs measured between 0.52 mm to 0.68 mm maturity for M. lar classified in this paper were very (Plate 3). similar to the ones described by Singh and Roy 17. Breeding period is assessed by the presence of

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“INDIAN J. MAR. SCI .VOL. 45, NO.10, OCTOBER 2014” berried female15. The studies related to reproductive between 4069 and 8543 with estimated mean characteristics such as percentage occurrence of fecundity of 5402 (± 772.35). Fecundity of reproducing and non-reproducing females, Macrobrachium sp prawns is highly unpredictable. percentage of occurrence of ovigerous females, The female of M. rosenbergii and M. carcinus can seasonal occurrence of maturity stages, and lay eggs between 80,000 and 100,000 numbers in maturity indices (GSI and HSI) exhibited that M. each spawning during the fully mature condition. lar breeds twice in a year with two breeding peaks During the first spawning, they can lay the eggs occurred in the months of June and November every between 5,000 and 20,000 numbers 36. Third highest year. Presence of ovigerous females only during fecundity in species of this genus is observed in M. those monsoon months indicate that this species acanthurus females, which can hatch about 18, 000 breeds seasonally. Walker and Ferreira 31 studied eggs 7-8, whereas Shokita, cited by Scaico 39 hatched the samples of M. nattereri (Heller, 1862) and M. bout 13, 600 eggs. Although fecundity of M. lar inpa (Kensley and Walker, 1982) species collected collected in lower stream of river is smaller in the Taruma-Mirim River in central Amazon compared to those above species, it shows higher recorded the pattern of seasonal reproduction. fecundity than M. amazonicum whose fecundity is Presence of ovigerous females in those species were lower than 2193 eggs 40 and M. iheringi, M. Borelli, found only during rainy season or monsoon months. M. australiense, M. potiuna, and M. jelski, whose Chaves and Magalhães 32, Porto 22 and Silva 27 absolute fecundity is lower than 200 eggs 39. observed that ovigerous M. amazonicum females Fecundity of M. lar was found higher than those of are more abundant in the rainy season than at other M. equidens 41 but it was quite related to that times of the year. Sastry 33 reported that there are observed by Krishna-Murthy 42 for the same two main strategies for reproduction in decapods, species. Egg size of decapods that reproduce in that is, continuous strategy, when spawning occurs fresh water is frequently enlarged compared with at roughly constant levels throughout the year and marine relatives 43. Egg size measured in the present seasonal strategy, when spawning is seasonal or study was lower than those of M. potiuna and M. restricted in a certain period of the year. In the olfersi whose mean size was 1.79 ± 0.13 (long axis) latter one, there is a close link between and 2.17 ± 0.17 (long axis) respectively 44 but reproduction, environmental conditions and food fecundity was found more than those species. supply, which are fundamental for the survival of Nazari 44 studied in M. olfersi and M. potiuna larvae and juveniles. M. lar is coming under the reported that the strategy of carrying a large number latter case. However, in general, it is accepted that of small eggs by M. olfersi allows a greater egg loss the reproduction is continuous in the tropical without affecting the reproductive success of the regions whereas it is seasonal and more and more species whereas in M. potiuna has a reduced egg restricted in species inhabiting subtropical to loss because this species spawns smaller number of temperate 34-35. Occurrence of non-ovigerous voluminous eggs and a larger loss could females during most of the month of collection compromise its reproductive investment. could be related to a difficulty in catching ovigerous Conclusions females as they migrate to lower regions during the In M. lar, Male attained the largest size than female reproductive phase where egg incubation take but female predominant over male throughout the places or they try to find shelter in protected areas, entire year. Occurrence of ovigerous females with that is, in marginal plants to keep away from mature ovaries twice in a year, that is, June and predation pressure 36. Fecundity is an important November, reinforces the fact that this species breed parameter to know the population ecology and life seasonally during rainy season with low realized history of any species 37. Different fecundity fecundity. indexes are differentiated: potential fecundity

(number of oocytes in the ovary), realized fecundity Acknowledgement (number of eggs attached under the pleopods) and Authors wish to express their sincere gratitude to actual fecundity (number of larvae hatched) 38. the Director, Central Agriculture Research Institute Generally realized fecundity is illustrated in studies (CARI) for their keen interest and encouragement of reproductive biology 16. In the present during the course of study. study, realized fecundity was found to be ranged

6 SETHI et al.: REPRODUCTIVE BIOLOGY OF MACROBRACHIUM LAR (FABRICIUS, 1798) IN ANDAMAN ISLANDS

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