Artigo Macrobrachium (Crustaceana)

8/11/2019 Artigo Macrobrachium (Crustaceana)

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POPULATION DYNAMICS OF THE RIVER PRAWN, MACROBRACHIUM AMAZONICUM (HELLER, 1862) (DECAPODA, PALAEMONIDAE) ON

COMBU ISLAND (AMAZON ESTUARY)

BY

FLVIA LUCENA-FRDOU1), JOS S. ROSA FILHO, MRCIA C. N. SILVA

and ELPDIO F. AZEVEDOFaculdade de Oceanograa, Instituto de Geocincias, Universidade Federal do Par,Av. Augusto Corra 01, Guam, Belm, Par, CEP 66075-110, Brazil

ABSTRACT

This study estimates reproductive parameters (length at rst maturity, reproduction period, andfecundity), mortality, and maximum sustainable yield for Macrobrachium amazonicum on CombuIsland (Amazon estuary). Samples were taken monthly between March 2003 and February 2004.Ovigerous females were recorded throughout the year, with reproductive peaks in the middle of theood period (March), in the low water period (September), and the peak dry period (December)of the Guam River. The size at which 50% of the females were mature was 60.8 mm TL.Absolute fecundity varied between 40 and 3375 eggs per female. The exploitation rate for maximumsustainable yield (EMSY) for males was lower than that for females. For both sexes, EMSY wasbelow the exploitation rates (E). The present study demonstrates the need for management of theprawn shery on the islands in the Amazon estuary, such as establishing a minimum commercialsize.

RESUMO

Este estudo tem o objetivo de estimar os parmetros reprodutivos (tamanho de primeira maturi-dade, perodo reprodutivo e fecundidade), mortalidade e rendimento mximo sustentvel do Mac-robrachium amazonicum da Ilha do Combu (Esturio Amaznico). As amostras foram coletadasmensalmente, entre maro de 2003 e fevereiro de 2004. Fmeas ovgeras foram registradas ao longode todo ano, com picos reprodutivos no meio da enchente (maro), vazante (setembro) e pico demnima vazo (dezembro) no Rio Guam. O tamanho na qual 50% das fmeas esto maduras de60,8 mm de comprimento total. Fecundidade absoluta variou entre 40 e 3375 ovos por fmea. A taxade explorao para a obteno do rendimento mximo sustentvel (ERMY) para machos foi inferior

ao obtido para as fmeas. Para ambos sexos ERMY foi abaixo das taxas de explotao atuais (E).O presente estudo demonstra a necessidade do manejo da pesca de camaro nas ilhas do esturioAmaznico, como por exemplo o estabelecimento de tamanho mnimo de comercializao.

1) Fax: +55.9132017609; e-mail: [email protected] Koninklijke Brill NV, Leiden, 2010 Crustaceana 83 (3): 277-290Also available online: www.brill.nl/cr DOI:10.1163/001121609X12596543952298


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278 FLVIA LUCENA-FRDOU ET AL.

INTRODUCTION

The river prawn, Macrobrachium amazonicum (Heller, 1862) (Decapoda, Palae-monidae) is an endemic species in South America and occurs throughout the basinsof the rivers Orinoco, Amazon, and Paraguay (Holthuis, 1952). This species is alsofound in the northeastern and eastern states of Brazil (Coelho & Ramos Porto,1985; Barros & Braun, 1997). In the Amazon basin, the species inhabits estuar-ies, lakes, and rivers of Central Amazon (Odinetz-Collart, 1987). This species hasfree-swimming larvae, which, under natural conditions, may develop in both freshand estuarine waters (Alekhnovich & Kulesh, 2001).

M. amazonicum is the main species of freshwater shrimp exploited by thecommercial artisanal eet in the Amazon (Odinetz-Collart, 1987; Odinetz-Collart& Moreira, 1993). In the rivers and estuaries of the Amazon, this prawn is mainlycaught with traps, locally known as matapi, made from palms ( Astrocaryum spp.and Atrix spp.) or jupaty ( Raphia spp.). This species also has a great potential foraquaculture (Kutty et al., 2000).

Dynamic pool models are widely used in population dynamics and have becomea major technique for shrimp management. A large variety of models have been

applied to species from the genera Penaeus (cf. Cha et al., 2002; Niamaimandi etal., 2007), Pandalus (cf. Hvingel & Kingsleu, 2006), Aristeus (cf. Maynou et al.,2006), and Melicertus (cf. Conides et al., 2006). Amazon stocks of M. amazonicumhave never been assessed, despite its enormous socio-economic importance in theregion, and the many studies on aspects of reproduction, abundance, distribution,diet, aquaculture, and growth of the species (Guest & Durocher, 1979; Odinetz-Collart, 1991; Alekhnovich & Kulesh, 2001; Morales-Riodades & Valenti, 2004;Silva et al., 2007).

Both coastal and inland sheries in the Amazon differ from those of otherregions in the richness of the exploited species, the quantity of the catch, and thedependence of traditional communities on this activity (Barthem & Fabr, 2004).Despite its importance as a source of income and food for river communities,the functioning of the biological communities and shery activities in aquaticenvironments of the Amazon remain poorly known. In many areas, river andestuarine communities have a low quality of life, even through they live withinan ecosystem with a high potential for providing goods and services.

In order to provide the knowledge and baseline information required for themanagement of M. amazonicum stocks in the Amazon estuaries, the aim of thepresent study was to estimate the reproductive parameters (length at rst maturity,reproduction period, and fecundity), mortality, and maximum sustainable yield forthis species on Combu island (Amazon estuary).


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POPULATION DYNAMICS OF MACROBRACHIUM AMAZONICUM 279

MATERIAL AND METHODS

Combu (Par, Brazil) is an island of approximately 15 km2

, located on the leftmargin of the Guam River, 1.5 km from the city of Belm (Par) (g. 1). Annualair temperature ranges from 23 to 32 C. Total monthly rainfall ranges from 200 to350 mm in the rainy season (December to May) and from 90 to 170 mm in the dryseason (June to November) (Moraes et al., 2005).

Each month, approximately 500 g (ca 100 individuals) of M. amazonicum wereobtained from commercial catches caught using matapi traps between March2003 and February 2004. In the laboratory, the specimens were sexed, measured

(total length: TL in mm) and weighed (total wet weight: TW in grams).For each female, the sexual maturity stage was determined visually, separatedinto ve categories according to criteria established by Carvalho & Pereira (1998):(a) stage I, underdeveloped (ovary transparent); (b) stage II, developing (ovarycream or pale green); (c) stage III, ripe or almost ripe (ovary at full size, dark greenoocytes visible); (d) stage IV, ovigerous (enlarged brood chambers bearing eggs);and (e) stage V, spent (large ovary with whitish aspect). The reproductive periodwas dened as those months with a maximum percentage of ovigerous females.

Size at sexual maturity (TL50) was determined by the proportion of ovigerousor spent females. The proportion of these females by size was tted to a logistic

Fig. 1. Study area: the Island of Combu in the Guam River, Amazon estuary, Brazil.


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equation as described by King (1995):

P =

1/ 1+

exp(a+

bT L),

where P is the predicted mature proportion, a and b are the estimated coefcientsof the logistic equation, and T L is total length. Parameters were estimated by non-linear regression analysis using the quasi-Newton method. Size at sexual maturity(TL50), corresponding to a proportion of 50% sexually mature individuals, wasestimated as the negative ratio of the two coefcients TL50 = (a/b) .

Eggs from 118 individuals in maturity stage V were used to determine speciesfecundity. For each female, eggs from four samples of diluted eggs (5 ml of a 100 ml solution) were counted. The mean number of eggs was obtainedand extrapolated to the total volume of the egg mass. The correlation betweentotal length and number of eggs was tested by the non-parametric Spearman Rtest.

The instantaneous coefcient of total mortality (Z) was estimated using thelength-converted Ricker (1975) catch curve and by the Beverton & Holt (1956)equation, which estimates the correlation between total mortality and mean length.

The instantaneous rate of natural mortality (M) was estimated using the Rikhter& Efanov (1976) formula, which associates natural mortality (M) and age at rstmaturity (t50). Age at rst maturity was obtained using the inverse Von Bertalanffyequation considering the length at rst maturity obtained in this study. As lengthat rst maturity was only obtained for females, M was calculated by groupingsexes. The instantaneous rate of shing mortality (F) was computed from thedifference of the instantaneous rates of total mortality and natural mortality. Theexploitation rate (E) was the ratio of the instantaneous rate of shing mortality

to the instantaneous rate of total mortality. The probability of capture for sigmoid-type selection was estimated for different size classes from the ratio of the expectednumbers to those that were actually caught. From this curve, the length at rstcapture (Lc) was estimated as the length corresponding to a 50% probability of capture. To estimate the levels of exploitation that would give optimum yields,the relative yield-per-recruit (Y/R) model was applied (Beverton & Holt, 1966).This analysis provides estimates of Emax (the exploitation rate at which maximumrelative yield-per-recruit is obtained).

Fishery mortality, length at rst capture, and maximum sustainable yield wereobtained for males and females separately as well as for pooled sexes. Growthparameters used as input for the estimates were based on various methodologiesdescribed in Silva et al. (2007). The effects of the different values of growthparameters in estimating the population parameters were assessed.


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Fig. 2. Frequency histogram for ovigerous females (dark bars) of Macrobrachium amazonicum(Heller, 1862).

RESULTS

Ovigerous females were recorded throughout the year, except in July. Therewere three reproductive periods established, i.e., September, December, andMarch, when, respectively, 35%, 45%, and 21% of the females were ovigerous(g. 2). Among the 1338 females analysed, 239 (18%) carried eggs on theirabdomen and 1099 (82%) did not. Total length for females ranged from 52 to106 mm. The size at which 50% of the females were mature was 60.8 mm TL(g. 3). The smallest females with a maturing ovary (stage II), mature ovary(stage III), and carrying eggs on their abdomen (stage IV) measured 31 mm,52 mm, and 52 mm TL, respectively. Absolute fecundity ranged from 40 to 3375eggs/female, with a mean value of 905 590. The correlation between fecundityand the length of the female was positive and signicant (p < 0.05). However,variance in fecundity per length class was high (g. 4).

Estimates of natural mortality (M) obtained from the Rikhter-Efanov methodvaried little, considering the various growth parameters L and K (Silva et al.,2007) used to obtain age at rst maturity (t50). The exception occurred for theestimate derived from the growth parameter obtained from Electronic LengthFrequency Analysis (ELEFAN) (table I).

Regarding length at rst capture (Lc), the various methods resulted in similarvalues, and females showed larger sizes than males. Total mortality (Z) estimatedfrom the catch curve method was similar for each set of growth parameters, withthe exception of the ELEFAN method. The estimate for males (6.39 year 1) washigher than that for females (3.86 year 1). Similar results were obtained from theBeverton & Holt (1956) method. The exploitation rate for maximum sustainableyield (EMSY) for males was lower than that for females when considering thevarious methods (table II). In most cases, EMSY for both males and females was


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Fig. 3. Size at rst maturity (TL50 in mm) for females of Macrobrachium amazonicum(Heller, 1862).

below the exploitation rates (E) found using the Ricker catch curve method andthe Beverton & Holt method.

Fig. 4. Fecundity and total length relationship for females of Macrobrachium amazonicum(Heller, 1862).


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TABLE IEstimates of age of rst capture (t50) and natural mortality under different growth parameters for Macrobrachium amazonicum (Heller, 1862)

Growth method L (cm)/K (cm/month)1) t50 M(year) (year 1)

Gulland & Holt 15.75/0.98 1.60 0.93Fabens 17.66/0.78 1.64 0.91Appeldoorn 15.84/0.94 1.52 0.96Seasonal length-at-age 12.38/1.35 1.56 0.95Non-seasonal length-at-age 12.37/1.36 1.61 0.92

ELEFAN 14.00/0.70 0.87 1.531) Source: Silva et al. (2007).

TABLE IIEstimates of total mortality (Z), shery mortality (F), exploitation rates (E), and length at rst catch(Lc) through both the catch curve and the Beverton & Holt method obtained for Macrobrachium

amazonicum (Heller, 1862)

Growth method L (cm)/ K(cm/month)1)

Catch curve Beverton & HoltZ F E Lc Z F E EMSY

MalesGulland & Holt 15.75/0.98 6.04 5.11 0.85 4.530 9.31 8.38 0.89 0.51Fabens 17.66/0.78 6.13 5.22 0.85 4.666 8.90 7.99 0.89 0.49Appeldoorn 15.84/0.94 5.87 4.91 0.84 4.574 9.03 8.07 0.89 0.50Seasonal 12.38/1.35 5.04 4.09 0.81 4.373 8.24 7.29 0.90 0.56Non-seasonal 12.37/1.36 5.06 4.14 0.82 4.342 8.30 7.38 0.90 0.56ELEFAN 14.00/0.70 3.15 1.62 0.51 4.248 5.43 3.90 0.72 0.54

FemalesGulland & Holt 12.79/0.80 4.25 3.32 0.78 6.112 4.43 3.50 0.79 0.65Fabens 13.16/0.75 4.27 3.36 0.79 6.015 4.43 3.52 0.79 0.64Appeldoorn 12.66/0.92 3.86 2.90 0.75 5.878 4.92 3.96 0.81 0.64Seasonal 14.14/0.66 4.32 3.37 0.78 6.051 4.55 3.60 0.79 0.62Non-seasonal 14.03/0.67 4.41 3.49 0.79 6.058 4.55 3.63 0.80 0.62ELEFAN 14.00/0.70 6.89 5.36 0.78 6.279 4.73 3.20 0.68 0.73

Pooled sexesGulland & Holt 15.70/0.37 4.06 3.13 0.77 4.621 3.66 2.73 0.75 0.54Fabens 15.11/0.38 4.06 3.15 0.78 4.621 3.54 2.63 0.74 0.55Appeldoorn 12.17/0.56 3.66 2.70 0.74 4.413 3.57 2.61 0.73 0.58Seasonal 10.50/0.71 3.63 2.68 0.74 4.404 3.34 2.39 0.72 0.61Non-seasonal 10.51/0.68 3.28 2.36 0.72 4.384 3.20 2.28 0.71 0.61ELEFAN 14.00/0.80 6.89 5.36 0.78 4.551 6.56 5.03 0.77 0.55

1) Source: Silva et al. (2007).


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DISCUSSION

The freshwater prawn, Macrobrachium amazonicum occupies a special positionamong the resources of economic and social importance in the Amazonian region.The species is widely caught and commercialized the Amazon estuary. Thisresource serves as the main and often only source of animal protein for humanriver and estuarine communities.

The identication of eggs adhering to the pleopods of females throughout theyear indicates that the species exhibits continual reproduction on Combu Island,as already reported in other studies (Chaves & Magalhes, 1993; Bialetzki et al.,1997; Silva et al., 2005). According to Odinetz-Collart (1991), M. amazonicumhas two reproduction strategies in the Amazon basin regarding the duration of thereproductive period: populations with well-dened seasonal reproduction, whichoccur in coastal rivers; and those with continual reproduction, which occur ininland rivers. Prawns on Combu Island belong to the second group (continualreproduction), but with periods in which reproduction is more intense.

Reproductive peaks were observed in March, September, and December, withthe highest percentage of females with eggs adhered to the pleopods recorded inDecember. These months mark the middle of the ood period (March), the lowwater period (September), and the peak dry period (December) of the Guam River.Odinetz-Collart (1991) reports the same reproductive pattern in lakes and rivers ofthe Central Amazon and Silva et al. (2002) report this pattern along the coast ofthe state of Par. In the Tocantins River, however, reproduction of the species isconcentrated during the peak ood season (Silva et al., 2005).

Gonad maturation in crustaceans depends upon internal factors (endogenouscontrol) as well as environmental characteristics (Odinetz-Collart, 1991). Pinheiro& Hebbing (1998) report that the reproductive period of freshwater decapods isclosely associated with variations in precipitation, temperature, and photoperiod.In the Amazon region, where the photoperiod and temperature vary little through-out the year, variation in rainfall and, consequently, the characteristics of the riversmust be the determinant factor in the reproduction of M. amazonicum .

On Combu Island, the main stimulus for M. amazonicum reproduction appearsto be the increased velocity of the currents and increased food supply. In the GuamRiver, the strongest currents occur in the middle of the ood and the low waterperiods (Pinheiro, 1987), which is when prawns take advantage of the greaterdispersive power of the water to release their larvae and thereby occupy a greaternumber of habitats. This reproductive behaviour is described by Odinetz-Collart(1987) as being typical of Palaemonidae populations in coastal rivers. In turn,December is the month with the highest production of phytoplankton (Paiva etal., 2006) and, consequently, zooplankton in the Guam River. As larvae of M.


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amazonicum are planktotrophic in their early stages (Arajo & Valenti, 2007),reproduction in the period of greater plankton abundance ensures the survival of

the larvae during the most critical period of the life cycle.The number of eggs recorded for M. amazonicum females ranged from 40to 3345 (mean 850 eggs per female). A signicant positive correlation wasfound between size and fecundity. According to Valenti (1984) and Silva et al.(2004), there is a direct correlation between size and fecundity in the genus Macrobrachium ; and the number of eggs laid is quite variable among individualsof the same species, increasing in quantity with the size of the female. Studyingthe fecundity of M. amazonicum under laboratory conditions, Lobo et al. (1986)

found considerable variability among females in the same length or weight class.The same was found in ovigerous females during the present study. Scaico (1992)also stated that fertility and hatching rate increase with length and weight withinthe range of the variables considered.

According to Coelho et al. (1982), the river prawn can produce as many as6000 eggs per spawning. However, studies carried out in Venezuela found lowernumbers (Gmba, 1984). Females with 148 to 1554 eggs have been recorded innortheastern Brazil (Scaico, 1992; Silva et al., 2004). Under laboratory conditions,

Lobo et al. (1986) reported between 178 and 1344 eggs/female.Lobo et al. (1986) mentioned that M. amazonicum has relatively high fecundityin comparison to other species of the same genus, the absolute fecundity of whichis less than 200 eggs per female. However, this species fecundity is consideredlow in comparison with species that reach larger sizes and have greater economicinterest, such as M. acanthurus (Wiegmann, 1836), M. carcinus (Linnaeus, 1758), M. malcolmsonii (H. Milne Edwards, 1844), M. rosenbergii (De Man, 1879),and M. vollenhovenii (Herklots, 1857), in which maximum absolute fecundityranges from 12 800 to 194350 eggs (Coelho et al., 1982; Valenti, 1984; Loboet al., 1985; Scaico, 1992). According to Lobo et al. (1986) and Scaico (1992), M. amazonicum spawns monthly, which may determine the low fecundity, whilecompensating for the lower number of eggs per spawn in comparison to species ofgreater economic interest.

Size at rst maturity was around 60 mm (TL). Although euryhaline Macro-brachium species exhibit considerable variation in length at rst maturity (Mantel& Dugeon, 2005), the prawns on Combu Island only begin reproducing at sizeslarger than those recorded for populations of the same species inhabiting freshwater (Mashiko, 1983; Mossoli & Bueno, 2002).

According to Alekhnovich & Kulesh (2001), females from most species of Macrobrachium begin to reproduce between 30 and 40 mm (TL), with lowervalues for freshwater populations in comparison to estuarine populations. Mashiko(1983) found that freshwater M. nipponense (De Haan, 1849) females mature


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at signicantly lower sizes than for those in estuaries, attributing this differenceto the fact that the food supply for omnivorous species, such as the majority of Macrobrachium , is greater in estuaries than for those in rivers or lakes.There is no consensus in shery science regarding the management of prawnstocks. Management advice for most prawn stocks in the North Atlantic is basicallyformulated by a qualitative assessment of trends in stock condition in responseto catch history (Koeller et al., 2000). For the relatively slow growing prawns ofnorthern waters, the use of standard techniques such as VPA and length-basedmethods has not been successful (Quinn et al., 1998). This may be due to the factthat natural mortality is thought to be at least as important as shing mortality in

the dynamics of shrimp stocks and this limits the application of traditional sherymodels, which assume natural mortality to be constant (Hvingel & Kingsley,2006).

Despite limitations, traditional stock assessment models have been used sat-isfactorily for prawn stocks in tropical and temperate waters (Isaac et al., 1992;Conides et al., 2006; Niamaimandi et al., 2007). These models require few para-meters (Sparre & Venema, 1997) and, despite the presupposition of a system withconstant parameters, they are important in determining whether an area is either

heavily shed or under-exploited in order to establish appropriate managementmeasures (Niamaimandi et al., 2007).Models based on different presuppositions are used to estimate total mortality

and natural mortality. These models often result in similar values (Oh et al., 1999).Moreover, different inputs (growth parameters, natural mortality, etc.) are alsotested by assessing the sensitivity of the resulting estimates (Gallagher et al., 2004).This was the case in the present study, which found that, despite the limitations ofthe methods employed, the results of the stock assessment models converge andare consistent with knowledge on the shery and biology of the species.

Instantaneous coefcients of total mortality (Z) were higher for males. Thisprobably stems from the fact that males have a smaller length at rst capture (Lc)than females, resulting from the smaller size of recruitment to shery activity.This high rate of exploitation leads to a lesser availability of males and a sexproportion favouring females on Combu Island (Silva et al., 2007), which is acommon situation among prawns.

Consequently, the recommendable exploitation rate for maximum sustainableyield is lower for males than for females. For both sexes, EMSY is below currentexploitation rates (E) (considering the different methods and scenarios), therebydemonstrating that this activity may not be sustainable in the near future. Thisinformation is corroborated by other evidence regarding the species on CombuIsland. Size at rst capture (Lc) for both sexes (more evident among males) issmaller than size at rst maturity (L50). This situation accelerates the reduction in


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numbers of individuals in the environment, thereby negatively altering populationfecundity, as larger females have a greater number of eggs and, consequently, a

greater likelihood for providing a larger number of recruits to the population.There are no estimates on mortality and maximum sustainable yield parametersavailable in the literature for the species or even other species of the genus, whichrenders comparisons of the values obtained here with those from other locationsimpossible.

The present study demonstrates the need for management of the prawn sheryactivity on islands of the Amazon estuary, given the importance of and local humandependence on the exploitation of natural resources. There is no information on the

volume of M. amazonicum catches, as ofcial statistics have failed to considerthis resource. This places serious limitations to the application of analyticalmodels and, consequently, the application of specic management measures.It also restricts knowledge regarding the impact of this shery activity on theAmazon estuary. It is likely that the species is not renewing itself adequatelydue to the capture of immature individuals. Controlling shery efforts is not avery viable measure, considering the large number of people involved in theprawn shery activity and the dispersal of boats over the large area of theAmazon estuary. However, based on the results of the present study, an alternativemanagement measure would be the establishment of a minimum commercialsize (not considering what is used for sustenance) to be obtained through thedetermination of a xed trap opening size or minimum commercialization size.

ACKNOWLEDGEMENTS

The authors acknowledge Dr. Victoria Isaac Nahum for comments on earlierdrafts of the manuscript. This study was partially nanced by National Councilfor Scientic and Technological Development (CNPq) through a grant to the thirdauthor and a research grant to the rst author (302280/2007-3).

REFERENCES

ALEKHNOVICH, A. V. & V. F. KULESH, 2001. Variation in the parameters of the life cycle inprawns of the genus Macrobrachium Bate (Crustacea, Palaemonidae). Russian Journ. Ecol.,32 : 420-424.

ARAJO

, M. C. & W. C. VALENTI

, 2007. Feeding habit of the Amazon river prawn Macrobrachiumamazonicum larvae. Aquaculture, 265 : 187-193.BARROS, M. P. & A. S. BRAUN, 1997. Contribuio ao estudo dos Atyidae e Palaemonidae

(Crustacea, Decapoda) do leste brasileiro 14 21 e 20 55 de latitude sul. Biotemas, 10 : 7-26.BARTHEM, R. B. & N. N. FABR, 2004. Biologia e diversidade dos recursos pesqueiros na

Amaznia brasileira. In: M. L. RUFFINO (ed.), A pesca e os recursos pesqueiros na Amazniabrasileira: 17-62. (IBAMA/PROVRZEA, Brasilia).


12/14


13/14


LOBO, V. L., N. E. T. ROJAS & W. C. VALENTI, 1986. Fecundidade e fertilidade de Macro-brachium amazonicum (Heller, 1862) (Crustacea, Decapoda) em laboratrio. Bol. Inst. PescaSo Paulo, 18 : 15-20.

MANTEL, S. K. & D. DUDGEON, 2005. Reproduction and sexual dimorphism of the palaemonidshrimp Macrobrachium hainanense in Hong Kong streams. Journ. Crust. Biol., 25 : 450-459.

MASHIKO, K., 1983. Comparison of growth pattern until sexual maturity between the estuarine andupper freshwater populations of the prawn Macrobrachium nipponense (De Haan) within ariver. Japanese Journ. Zool., 33 : 207-212.

MAYNOU, F., F. SARD, S. TUDELA & M. DEMESTRE, 2006. Management strategies for redshrimp ( Aristeus antennatus ) sheries in the Catalan Sea (NW Mediterranean) based onbioeconomic simulation analysis. Aquat. Living Resour., 19 : 161-171.

MORAES, B. C., J. M. N. COSTA, A. C. L. COSTA & M. H. COSTA, 2005. Variao espacial etemporal da precipitao no estado do Par. Acta Amazonica, 35 : 207-214.

MORAES-RIODADES, P. M. C. & W. C. VALENTI, 2004. Morphotypes in male Amazon Riverprawns Macrobrachium amazonicum . Aquaculture, 236 : 297-307.

MOSSOLIN, E. C. & S. L. S. BUENO, 2002. Reproductive biology of Macrobrachium olfersi(Decapoda, Palaemonidae) in So Sebastio, Brazil. Journ. Crust. Biol., 22 : 367-376.

NIAMAIMANDI, N., A. B. ARSHD, S. K. DAUD, R. C. SAED & B. KIABI, 2007. Populationdynamic of green tiger prawn, Pennaeus semisulcatus (De Haan) in Bushehr coastal waters,Persian Gulf. Fish. Res., 86 : 105-112.

ODINETZ-COLLART, O., 1987. La pche crevettire de Macrobrachium amazonicum (Palae-monidae) dans le Bas Tocantins, aprs la fermeture de barrage de Tucurui (Brsil). Rev. Hy-drobiol. trop., 20 : 131-144.

, 1991. Stratgie de reproduction de Macrobrachium amazonicum en amazonie centrale(Decapoda, Caridea, Palaemonidae). Crustaceana, 61 : 253-270.ODINETZ-COLLART, O. & L. C. MOREIRA, 1993. Migrao vertical nictemeral das larvas de

Macrobrachium amazonicum num lago de vrzea na Amaznia Central, Ilha do Careiro, Brasil.Amazoniana, 3 : 385-389.

OH, C.-W., R. G. HARTNOLL & R. D. M. NASH, 1999. Population dynamic of the commonshrimp, Crangon crangon (L.), in Port Erin Bay, Isle of Man, Irish Sea. Journ. mar. Sci., 56 :718-733.

PAIVA, R. S., E. ESKINAZI-LEA, J. Z. O. PASSAVANTE, M. G. G. SILVA-CUNHA & N. F. A.MELO, 2006. Consideraes ecolgicas sobre o toplncton da baa do Guajar e foz do rio

Guam (Par, Brasil). Bol. Mus. Paraense Emlio Goeldi, 1

: 133-146.PINHEIRO, M. A. A. & N. J. HEBLING, 1998. Biologia de Macrobrachium rosenbergii (De Man,1879). In: W. C. VALENTI (ed.), Carcinicultura de gua doce. Tecnologia para produo decamares: 21-46. (IBAMA, Brasilia).

PINHEIRO, R. V. L., 1987. Estudo hidrodinmico e sedimentolgico do esturio Guajar Belm(PA): 1-164. (M.Sc. Thesis, Universidade Federal do Par, Belm).

QUINN, T. J., C. T. TURNBULL & D. FU, 1998. A length-based population model for hard-to-age invertebrate populations. In: F. FUNK, T. J. QUINN II, J. HEIFETZ, J . N. IANELLI,J. E. POWERS, J. F. SCHWEIGERT, P. J. SULLIVAN & C.-I. ZHANG (eds.), Fishery stockassessment models: 531-556. (Alaska Sea Grant College Program, University of AlaskaFairbanks, AK-SG-98-01, Fairbanks).

RICKER, W. E., 1975. Computation and interpretation of biological statistics of sh populations.Bull. Fish. Res. Board Canada, 191 : 1-382.

RIKHTER, V. A. & V. N. EVANOV, 1976. On one of the approaches to estimation of naturalmortality of sh population. ICNAF Res. Doc., 76 : 1-12.

SCAICO, M. A., 1992. Fecundidade e fertilidade de Macrobrachium amazonicum (Crustacea,Decapoda) de um aude no nordeste brasileiro. Bol. Inst. Pesca So Paulo, 19 : 89-96.


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SILVA, K. C., R. A. L. SOUZA & I. H. A. CINTRA, 2002. Camaro-cascudo Macrobrachiumamazonicum (Heller, 1862) (Crustacea, Decapoda, Palaemonidae) no Municpio de Vigia-Par-Brasil. Bol. tec. cient. CEPNOR, 2 : 41-74.

SILVA, K. C. A., I . H. A. CINTRA & A. P. M. MUNIZ, 2005. Aspectos bioecolgicos de Macrobrachium amazonicum (Heller, 1862) a jusante do reservatrio da hidreltrica deTucuru-Par. Bol. tec. cient. CEPNOR, 5 : 55-71.

SILVA, M. C. N., F. LUCENA FRDOU & J. S. ROSA FILHO, 2007. Estudo do crescimento docamaro Macrobrachium amazonicum (Heller, 1862) da ilha de Comb, Belm, Estado doPar. Amaznia: Cienc. & Desenv., 2 : 85-104.

SILVA, R. R., C. M. S. SAMPAIO & J. A. SANTOS, 2004. Fecundity and fertility of Macrobrachiumamazonicum (Crustacea, Palaemonidae). Brazilian Journ. Biol., 64 : 489-500.

SPARRE, P. & S. VENEMA, 1997. Introduction to tropical sh stock assessment. FAO Fish. tech.Pap., 306 (1): 1-376.

VALENTI, W. C., 1984. Estudo populacional dos camares de gua doce Macrobrachium acan-thurus (Wiegmann, 1836) e Macrobrachiun carcinus (Linnaeus, 1758) do rio Ribeira Iguape(Crustacea, Palaemonidae): 1-156. (M.Sc. Thesis, Universidade de So Paulo, So Paulo).

First received 31 October 2008.Final version accepted 11 May 2009.

Documents

Artigo Macrobrachium (Crustaceana)