Aquaculture 283 (2008) 1318

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Aquaculture
j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / a q u a - o n l i n e

White shrimp Penaeus vannamei culture in freshwater at three densities: Condition state based on length and weight
Marcelo Araneda a,b, , Eduardo P. Prez c,d, Eucario Gasca-Leyva b
a

Universidad Marista de Mrida, Perifrico norte tablaje catastral 13941 Carretera Mrida Progreso, C.P. 97300 Mrida Yucatn, Mexico Centro de Investigacin y Estudios Avanzados (CINVESTAV) Unidad Mrida, Apdo. Postal 73-CORDEMEX, 97310 Mrida Yucatn, Mexico Universidad Catlica del Norte, Facultad de Ciencias del Mar, Departamento de Biologa Marina, Casilla 117. Coquimbo, IV Regin, Chile d Centro de Estudios Avanzados en Zonas ridas (CEAZA), Casilla 599, Benavente 980, La Serena, Chile
b c

A R T I C L E

I N F O

A B S T R A C T
The commercial culture of shrimp in low water salinities has been successfully achieved. However, there are no long-term studies on the growth response of shrimp culture in freshwater. In the present study white shrimp Penaeus vannamei was cultured in freshwater (0 ppt) at three densities (90, 130 and 180 shrimp m 2) and comparisons made of the resulting growth, lengthweight relationship and condition factor data. Gravimetric data were recorded during a 210-day trial to estimate growth rates, the lengthweight relationship and the condition factor (according to Fulton's equation). Weight gain results showed the freshwater culture system to be viable, with effects from density (P b 0.05). Growth rates decreased as density increased, exhibiting an inverse relationship (P b 0.05, R2 = 0.68). Survival decreased as density increased with differences between treatments (P b 0.05). A potential model was used to analyze the lengthweight relationship of the treatments. The slopes were different between treatments according to an ANCOVA and a Tukey test (P b 0.05). The function exponents showed that the 130 shrimp m 2 treatment produced organisms with isometric growth (b = 2.99) in good condition (t = 0.96; P N 0.05). The 90 shrimp m 2 treatment produced positive allometric growth and the 180 shrimp m 2 treatment negative allometric growth. 2008 Elsevier B.V. All rights reserved.

Article history: Received 30 October 2007 Received in revised form 17 June 2008 Accepted 18 June 2008 Keywords: Penaeus vannamei Freshwater Condition state

1. Introduction Over 90% of shrimp culture in the Western hemisphere is represented by white shrimp (Penaeus vannamei) (Wurmann et al., 2004). This species is generally cultivated in semi-intensive systems located near coasts. Successful culture of white shrimp depends on the quality of seawater used in the system, as well as use of a wastewater treatment plant to prevent pollution of adjacent areas. A number of factors have limited expansion of white shrimp culture, including the high cost of coastal real estate and the constant appearance of viral diseases such as white spot (WSSV), which has brought the industry near collapse (Jory and Dixon, 1999). One proposed solution to white shrimp production problems is the use of water with salinities lower than seawater. Using water with 0.5 ppt salinity, Van Wyk et al. (1999) reported survival and growth rates below those with seawater. In evaluations of postlarvae survival as a function of age in salinities varying from 16.3 to 0 ppt, both McGraw et al. (2002) and Saoud et al. (2003) reported increased survival as the organisms reached 15 days old at salinities greater than 4 ppt, but only 77% survival at 0 ppt. In studies of postlarvae and
Corresponding author. Tel.: +52 999 9410302; fax: +52 999 9410307. E-mail addresses: maraneda@marista.edu.mx (M. Araneda), eperez@ucn.cl (E.P. Prez), eucario@mda.cinvestav.mx (E. Gasca-Leyva). 0044-8486/$ see front matter 2008 Elsevier B.V. All rights reserved. doi:10.1016/j.aquaculture.2008.06.030

juvenile growth and survival in seawater diluted with freshwater and supplemented with a salt mixture, Atwood et al. (2003) and Sowers et al. (2005) reported no osmotic dysfunction although the organisms did not grow as ef ciently compared to those in seawater. Samocha et al. (2004) and Sowers and Tomasso (2006) reported growth higher than in seawater using low salinity (2 ppt) water. No research has yet been done on the long-term growth performance of P. vannamei in freshwater (0 ppt), including the effect of stocking density on weight gain and condition factor. The present study aims to compare the lengthweight relationship and condition state in growth stage white shrimp cultured in freshwater (0 ppt) at three densities (90, 130 and 180 shrimp m 2). The results provide practical data on growth, survival and organism condition indicators (i.e. lengthweight relationship) for these culture conditions. 2. Materials and methods 2.1. Water chemical composition Water used in the experiment came from the subterranean aquifer in the northern Yucatan Peninsula, Mexico, at 18 m depth. Before the trial began, its chemical composition was analyzed in the Marine Chemistry Laboratory, CINVESTAV-IPN, Mrida. The APHA (1989) titration technique was used to determine total hardness (CaCO3) (400 mg l 1),

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alkalinity (325.49 mg l 1) and chloride (Cl) (125.69 mg l 1). The atomic absorption method (using a CBC 932 PLUS spectrophotometer) was used to determine the concentrations of the inorganic ions potassium (K+; 3.37 mg l 1), calcium (Ca2+; 85.49 mg l 1), sodium (Na+; 68.64 mg l 1) and magnesium (Mg2+; 27.52 mg l 1). Concentration of the SO= ion was done with the turbidometric method using a 4 21D UV spectrophotometer (269.50 mg l 1). Salinity (0 ppt) was measured with an induction salinometer (KAHLSICO RS-9), electrical conductivity (1014 s cm 1) with a conductimeter (CORNING CHECKMATE) and pH (7) with a potentiometer (BECKMAN 32). Colorimetric reactions followed by spectrophotometric analysis (SHIMADUZU UV-1201) were used to determine ammonium (0.027 mg l 1), nitrites (0 mg l 1) and nitrates (6.55 mg l 1). 2.2. Experimental organisms and acclimation Postlarvae (PL8; approx. 0.0025 g individual weight according to manufacturer technical data) were acquired from SYAQUA (Mazatlan, Sinaloa, Mxico) and transported by air cargo in plastic bags containing 38 ppt seawater to the experimental unit at the Universidad Marista de Mrida (Mxico). The postlarvae were stocked in three, 600-l circular tanks with constant aeration and 38 ppt salinity for six days. Acclimation was carried out over 10 days and begun once the organisms had reached PL15. During this time salinity was lowered from 38 to 0 ppt at an average rate of 4 ppt day 1, and salinity measured every 2 h with a refractometer (CHASEBRAN). Once acclimated, the organisms were allowed to grow to juvenile stage for 30 days in the 0 ppt water at an average of 26 C, and fed ad libitum. 2.3. Experimental design The study was designed to evaluate the effect of stocking density on weight gain and condition factor in white shrimp P. vannamei cultured in freshwater. The evaluated densities were 90, 130 and 180 shrimp m 2. Juveniles (0.48 0.03 g) were randomly assigned to one of nine tanks, with three tanks per density treatment. The recirculation system consisted of round, polyethylene, 1200-liter tanks (2 m2 surface area and 0.60 m high); two, 5 m-capacity lter cartridges (10 in. Big Blue); a wetdry (or trickling) biological lter with a 405 m2 m 3 effective xation surface; a rectangular concrete sedimentation tank (2.7 m2 surface area) and a 1 hp blower (SWEETWATER) with air diffusers. To maintain good water quality, the water exchange rate was 0.6 renewals h 1, with a daily replacement of approximately 5% total volume and daily removal of unconsumed feed and feces. The organisms were fed a commercial balanced feed (Purina, Camaronina, Agribrands, Mexico) containing 35% protein. Feed ion content was determined with an atomic absorption spectrophotometer (GBC 932 PLUS): K+ = 8.44 g K+ kg 1; Mg2+ = 1.76 g Mg2+ kg 1. Feed rate was four times a day (09:00, 13:00, 17:00 and 21:00 h), as recommended by Van Wyk et al. (1999). Water temperature and oxygen content were measured three times a day (08:00, 13:00 and 18:00 h) in each tank with an oxymeter (YSI 200). Weekly measurements were made of nitrogenated compounds such as nonionized ammonium, nitrite and nitrate with a spectrophotometer (HACH), and of pH with a potentiometer (BECKMAN 32). These were taken in one randomly chosen tank per treatment. 2.4. Growth rates, lengthweight relationship and condition factor Gravimetric data were collected weekly throughout the 210-day (30-weeks) experimental period. An ichthyometer (mm) was used to measure total body length, and rostrum-to-telson distance. A digital scale (0.01 g; EXCELL) was used to record weight after excess moisture was removed from each organism. Sample size (n) for each replicate was estimated following the method of Zar (1999). Growth

rate was estimated for each density treatment as a function of weight and time with the formula: GRi Wfi Woi t 1

Where GRi is the growth rate of treatment i; and Wfi and Woi are average nal and initial weight in time t. The total lengthweight relationship for the different ages was adjusted using the potential model: Wi a TLb i 2

Where Wi is organism weight (g) for treatment i; TLi is total length (mm) for treatment i; a is the proportionality constant; and b the isometric exponent. Fulton's equation was used to determine condition factor (Ricker, 1975; Chow and Sandifer, 1991), which varies according to organism reproductive stage and general culture conditions: CFi Wi TL
3

105
P

Where CFi is the condition factor for treatment i; Wi is average weight (g) and TL i is average total length (mm). 2.5. Statistical analysis Comparison of growth rates and nal survival between treatments was done with a one-way analysis of variance (ANOVA) and a Tukey HSD a posteriori test. Data were expressed as the mean standard error. The effect of density on growth behavior over the 30-week experimental period was estimated with a single factor repeatedmeasures variance analysis. A non linear regression analysis was used to determine the relationship between density and growth rates. The resulting potential curves for the three studied densities were compared with an analysis of covariance (ANCOVA; Zar, 1999) and a Tukey a posteriori test. When parallelism was present between slopes, an intercept analysis was done with a Tukey test (Zar, 1999). To evaluate the relationship between weight and total length in each treatment, the statistical signi cance of the isometric exponent (b) was analyzed with a function proposed by Pauly (1984): t
"

Where t is the t-student statistic; s.d.(x) and (y) are the standard deviation of the log TLi and Wi value of each treatment; ni is the number of recorded organisms; bi is the adjusted b value; and r2 is the potential t determination coef cient. The signi cance level was 0.05. The analyses were conducted using the SPSS for Windows v. 11.0.0 program (SPSS Inc., Chicago, IL). 3. Results The highest growth rate was observed in the 90 shrimp m 2 treatment, with an average of 0.38 g week 1, and the lowest in the 180
Table 1 Average weekly growth rates and nal survival (standard error) of white shrimp grown in freshwater (0 ppt) at three different stocking densities Treatment (shrimp m 2) 90 130 180 Culture time (weeks) 30 30 30 Growth rate (g week 1) 0.38a 0.011 0.34a,b 0.007 0.33b 0.009 Survival (%) 76.1a 3.16 68.9a,b 0.78 65.9b 0.91

s:d:x jbi 3j p p 2 ni 2 s:d: y 2 1r 2

Different letter superscripts in the same column indicate statistically signi cant differences (P b 0.05).

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Fig. 1. Absolute growth rate in white shrimp versus culture density under freshwater (0 ppt) conditions (predicted value 95% con dence interval for the expected dependent variable value).

shrimp m 2 treatment, with a rate 0.33 g week 1 (Table 1). Growth rates were signi cantly different between treatments (F = 8.05; P b 0.05), and there was an inverse relationship (P b 0.05, R2 = 0.68) between higher density and weight gain (g week 1) (Fig. 1). The effect of density on growth performance in terms of weight differed between treatments (F = 2.99; P b 0.05) (Fig. 2). Survival was different (F = 7.18; P b 0.05) between all the treatments with the highest in the 90 shrimp m 2 treatment (76.1%), followed by the 130 and 180 shrimp m 2 (68.9% and 65.9%, respectively) treatments (Table 1). The determination coef cient (r2) values explained the proper t of the model for growth and potential tendency according to the variability observed in each treatment. Slope (b) values were inverse to culture densities, that is, the highest density had the lowest value and the lowest density the highest value (Fig. 3). According to the ANCOVA the potential model slopes were signi cantly different between treatments (F = 33.3; P b 0.05). The highest condition factor value (0.674) was produced in the 180 shrimp m 2 treatment, followed by the 130 (0.670) and 90 shrimp m 2 (0.663) treatments. According to the slope value (b = 2.99), growth in the 130 shrimp m 2 treatment was isometric, and, based on t , was not different from 3 (t = 0.96; P N 0.05). The slopes for the 180 and 90 shrimp m 2 treatments differed from 3 (t = 2.85 and t = 8.25, respectively; P b 0.05) and were therefore allometric. Average water quality parameters between the three treatments during the experimental period were 25.0 to 25.19 C for temperature; 6.40 to 6.56 mg l 1 for dissolved oxygen; 0.049 to 0.06 mg l 1 for non-

Fig. 3. Lengthweight relationship for white shrimp grown under freshwater (0 ppt) conditions at three densities: 180 shrimp m 2 (upper graph); 130 shrimp m 2 (middle graph); and 90 shrimp m 2 (lower graph). Each circle represents an observation.

ionized ammonium; 0.040 to 0.056 mg l 1 for nitrite; 4.16 to 4.56 mg l 1 for nitrate; and 8.08 to 8.12 for pH. 4. Discussion The results demonstrate the feasibility of white shrimp culture in freshwater, at three densities and during 30 weeks of growth. Salinity and speci c conductivity analyses were done (Boyd et al., 2002), and the salinity (0 ppt) and degree of mineralization (1014 s cm 1) values showed it to be within values for freshwater (201500 s cm 1) (Boyd, 1990). Under these conditions, growth rates were lower than reported for hyper-intensive marine, brackish and low salinity culture. However, any comparison should also include factors such as water temperature, initial and nal organism weight, culture density and feed type, among others. Williams et al. (1996) and Davis and Arnold (1998) reported rates of 0.50 to 0.95 g week 1 in seawater. In trials with brackish water and stocking densities of 107 and 100 shrimp m 2, respectively, Samocha et al. (2004) and Sowers and Tomasso (2006) reported very high growth rates (1.17 and 1.23 g week 1, respectively). In a study using low salinity

Fig. 2. Average growth ( SE) of white shrimp P. vannamei cultured in freshwater at three densities during a 210 day experimental period.

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(0.5 ppt) water, Van Wyk et al. (1999) reported growth rates of 0.57 and 0.40 g week 1, which are near those observed here. Survival was similar and growth greater in the 90 shrimp m 2 treatment compared to other trials with white shrimp in freshwater (Laramore et al., 2001; McGraw et al., 2002), although both these studies were done for very short periods (48 h and 35 days, respectively). Lower growth rates and higher mortality are to be expected as culture system salinity decreases; in contrast, survival can be as high as 95% in marine water. Salinity is probably not the sole limiting factor for growth in cultured white shrimp. Van Wyk et al. (1999) reported that salinities less than 0.5 ppt put this species at its physiological limit and cause a large proportion of its energy to be used in osmoregulation, therefore limiting growth and preventing it from reaching commercial sizes. Other studies indicate that growth may depend more on certain required water quality parameters, such as speci c concentrations of the main anions and cations: bicarbonate; sulfates; chlorides; calcium; magnesium; potassium; and sodium (Van Wyk et al., 1999; Boyd et al., 2002; Balbi et al., 2005). The water hardness (CaCO3) and alkalinity levels observed here were above recommended levels (150 mg l 1 and 75 mg l 1, respectively) (Scarpa and Vaughan, 1998; Boyd et al., 2002), and may therefore have provided the physiological conditions for organism growth and survival. Adequate concentrations allow shedding and proper formation of the exoskeleton, promoting growth and survival (McGraw and Scarpa, 2003). Optimum concentrations of SO= (262.3 mg l 1), K+ (40 mg l 1), Na+ 4 (1,114 mg l 1) and Mg2+ (40 mg l 1), as well as the proper Na:K ratio (29:1) in culture system water (4 ppt salinity) produce a growth rate 0.06 g week 1 greater than that reported here for the 90 shrimp m 2 treatment. In the present study, the K+ and Na+ concentrations were lower than the optimum, but the Na:K ratio was 20.4:1, which is near the optimum. This fact leads to the suggestion that the individual K+ and Na+ concentrations may not be as relevant as the ratio between them. Reductions in SO= and Mg2+ are linked to decreased survival. 4 The SO= concentration observed here was slightly higher than the 4 optimum level, while the Mg2+ concentration was slightly lower. Resulting survival was 3.9% and 13.9% lower than under optimum conditions, meaning that the higher SO= concentration may have 4 positively affected survival while the lower Mg2+ concentration negatively affected it (Saoud et al., 2003; Roy et al., 2007a,b). Nutritional studies have shown optimum K+ levels to be 10 g kg 1 and optimum Mg2+ to be 2.63.46 g kg 1, with corresponding growth rates of 0.76 and 0.33 g week 1, respectively, and survival of 78 and 80%, respectively (Cheng et al., 2005; Roy et al., 2007a,b). In the present study, feed K+ content was 1.56 g kg 1 lower than optimum, resulting in a 50% lower growth rate and a 1.9% lower survival rate in the 90 shrimp m 2 treatment. Feed Mg2+ content here was 0.84 g kg 1 below optimum level, producing a 13.2% higher growth rate and 3.9% lower survival rate in the 90 shrimp m 2 treatment. Any further comparison will need to consider the density levels in the respective studies (low in the optimum level studies and high here) and the experimental period (6 vs. 30 weeks). Water quality parameters remained stable throughout the experimental period, although water temperature may have affected growth rates in all three treatments. Water temperature is reported to have a signi cantly greater effect on growth than salinity in penaeid shrimp (Herke et al., 1987; Staples and Heales, 1991; O'Brien, 1994; Tsuzuki et al., 2000). Van Wyk et al. (1999) reported slightly lower growth rates in white shrimp when water temperature is below 26 C, and null rates when less than 22 C. Water temperature was not controlled in the present study and on average was below 26 C (2.60); indeed, from December to February it was below 22 C (minimum = 17 C). Any effect this might have had on growth would have been equal in all three treatments, meaning differences in weight gain can still be attributed to the studied stocking densities. Complementary future

research could be done during the spring and summer to produce comparable data from trials with warmer water temperature. The decreases observed here in growth and survival rates as culture density and biomass increased coincide with other studies of this species at different salinities and densities (Reid and Arnold, 1992; Williams et al., 1996; Davis and Arnold, 1998; Van Wyk et al., 1999; Appelbaum et al., 2002; Samocha et al., 2004). This decrease is probably due to crowding at high densities, leading to stress in the organisms (Foster and Bread, 1974). Large organisms were also seen to dominate smaller ones during feeding and in competition for space, particularly in the higher density treatments. This agrees with Harn et al. (2004) and Arnold et al. (2006), who reported that high densities lead to greater dominance and hierarchy placement of large organisms over small ones in terms of feed, refuge and reproduction. Apparently, territoriality added to social interactions creates an inhibitory effect on growth and survival that is augmented as population density increases. In contrast to other studies (Abdussamad and Thampy, 1994; Arnold et al., 2006), cannibalism was not observed here, although organisms were seen to feed off recently dead shrimp. Reduced population density versus the initial value due to mortality positively affected growth rates in the three treatments since it improved space conditions. For the lengthweight relationship and potential t, the a and b parameters for the three studied densities are similar to those reported for different penaeid species (Dall et al., 1990). Enin (1994) stated that when the b parameter is equal to 3 growth is isometric and when it is less than or greater than 3 it is allometric. Wootton (1992) was more speci c and stated growth to be positive allometric when organism weight increases more than length (b N 3), and negative allometric when length increases more than weight (b b 3). Of the three studied densities, only the 130 shrimp m 2 treatment had a t indicating isometric growth, that is, length and weight increased in the same proportion (Fig. 3). Although close to 3, the t for the 180 shrimp m 2 treatment (b = 2.97) was negative allometric and that for the 90 shrimp m 2 treatment (b = 3.07) was positive allometric. In other words, weight was not symmetrical with length at the highest density and vice versa at the lowest density. Despite being allometric, the 90 shrimp m 2 treatment is apparently the most appropriate under the studied culture conditions because it produced organisms similar in length to the other treatments but with greater weight gain. This assumes a market in which they are sold whole, meaning it would require fewer individuals to reach a kilogram of product. Density conditions clearly affected the lengthweight relationship of individual organisms, which coincides with reports that slopes less than 3 can indicate density and/or feeding problems in a culture system (Murphy et al., 1991). In the present study, the greater competition for feed at the 180 shrimp m 2 density would have affected this parameter, in addition to the stress of crowding. In cases of isometric growth (b = 3), the a (intercept) parameter can be considered an indicator of good physiological condition in a population given the conditions in a certain environment; if it is allometric, however, a cannot be interpreted this way (Pauly, 1984). In the 130 shrimp m 2 treatment, weight gain was isometric, meaning a can be used an indicator of good condition. This parameter was 0.669 in this treatment, similar to the estimated Fulton's function (CF = 0.670). Condition factor and potential relation parameter data can be vital to culture system management because they provide the producer with an evaluation of the speci c conditions under which organisms are developing. Shrimp length and weight were used in the present study, although it should be highlighted that the lengthweight relationship applied must be appropriate for the culture stage and strategy being used since this relationship is known to change in different phases (Dall et al., 1990; Peixoto et al., 2004). For example, Cheng and Chen (1990) and Primavera et al. (1998) reported that spawning Penaeus monodon are heavier than organisms of the same

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length in growth stage. Chow and Sandifer (1991) described mature adult stage white shrimp as thicker, with higher weight in proportion to length, than younger organisms in the growth stage. The present data emphasize that factors such as density can affect isometry in growth stage organisms. The 130 shrimp m 2 density allowed the organisms to remain healthy, which was re ected in a good length weight relationship. The 90 shrimp m 2 density produced positive allometric organisms, which are also healthy and have the added advantage of growing more in weight than in length. The 180 shrimp m 2 density produced negative allometric organisms. 5. Conclusions In conclusion, white shrimp can grow and survive in freshwater (0 ppt) at intensive densities. At the lowest studied density (90 shrimp m 2), it was possible to produce organisms with an average individual weight of 11.72 g and a survival rate of 76.1% at 210 days. The combination and quantities of nutrients in the groundwater of Yucatan is adequate for white shrimp in freshwater, probably due to its CaCO3 concentration, alkalinity and, to a lesser degree, SO= content. 4 The Na:K ratio (20.4:1) was close to optimum and may have played an important role in allowing successful culture in freshwater while highlighting that individual ion concentration may not be as relevant as ion ratio. This said, Mg+2 levels were below optimum water and feed concentrations, and may have had a slight negative effect on survival. The reduction in growth and survival rates as density increased is probably due to competition for space and feed, and negative social interactions creating an inhibitory effect. Signi cant differences between treatments were observed in the lengthweight curves, in which slope values decreased as density increased. This shows the effect of density on organism condition state, con rming the previous result. Only the 130 shrimp m 2 treatment attained isometric growth, indicating that organisms in this treatment had a good lengthweight relationship and were therefore in good condition. The present results can function as a reference for developing shrimp culture in freshwater conditions, particularly in regions where water chemistry is similar to that in the study area. Under these conditions, white shrimp can be cultivated without the need to increase salinity or add essential anions and cations to maintain growth and survival rates. Freshwater white shrimp culture could reduce pressure on coastal areas from this activity, and provide disease incidence below that of marine culture systems. If culture systems were established in rural areas they could increase employment opportunities and aid in diversifying production. The average sizes produced in the present study have a market niche as cocktail shrimp. In this system, greater biomass can be emphasized over individual organism size, making the use of closed systems a good option. Future research will need to include experiments using higher water temperature to determine if it results in higher individual weight and survival, for another market product. Acknowledgments The authors thank Miguel Vela, Leonardo Martinez and Miguel A. Olvera-Novoa for their helpful comments on the manuscript; also to Isabel Real of the Marine Chemistry Laboratory, CINVESTAV-IPN, Mrida, for the assistance with the water chemistry analysis; and the insightful comments of anonymous reviewers greatly improved this manuscript. The SYAQUA Mxico company donated the organisms for this experiment. This research forms part of the Ph.D. dissertation of MAP at CINVESTAV-IPN, Mrida. References
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