Population dynamics of Macrobrachium …1)_1-17.pdfMacrobrachium amazonicum is a freshwater shrimp...

Population dynamics of Macrobrachium amazonicum (Heller, 1862)(Decapoda: Palaemonidae) in a Brazilian Amazon Estuary

BIANCA BENTES¹, JUSSARA M. MARTINELLI-LEMOS², ÍTALO A. F. LUTZ¹*, MAYRA S.NASCIMENTO¹ & VICTORIA J. ISAAC²

¹ Federal University of Pará, Campus Bragança. Amazon of crustacean ecology group (GPECA). Leandro Ribeiro Road, s/n Aldeia, Bragança-PA, Brazil. ²Federal University of Pará. Fishery ecology and acquatic resources management Laboratory. Perimetral Avenue, 2651 Guamá, Belém-PA, Brazil.

*Corresponding author: [email protected]

Abstract. Amazonian shrimp Macrobrachium amazonicum is a freshwater species of largeabundance in the rivers of the Amazon basin and widely exploited by small-scale fishing andaquaculture. The objective of this study was to obtain the parameters of growth and mortality ofspecimens collected from May 2006 to August 2007 in the Guajará estuary with the aid ofFISAT II program (FAO-ICLARM Fish Stock Assessment Tools). The asymptotic maximumlength (L∞) estimated was almost always greater for females and individual growth constant(K) greater for males in most cases. L∞ varied from 36.5 to 46.9 mm and 0.2 to 0.44 K peryear-1. The total mortality coefficient (Z) varied from 2.04 to 3.78 per year-1 and (F) from 1.57 to2.75 year-1 due to fishing. Exploitation rates calculated warn about a situation ofoverexploitation of the species. The study of growth and mortality in crustaceans is of extremeimportance, because it provides basic information to subsidize fisheries planning andmanagement.

Keywords: Fishing resource; Amazon river shrimp; Fishing status; Coastal resources; Overexploitation.

Resumo. Dinâmica Populacional de Macrobrachium amazonicum (Heller, 1862)(Decapoda: Palaemonidae) em um Estuário Amazônico Brasileiro. Ocamarão-da-Amazônia, Macrobrachium amazonicum, é uma espécie dulcícola de largaabundância nos rios da bacia amazônica e amplamente explorada pela pesca artesanal e pelaaquicultura. Este estudo teve como objetivo obter os parâmetros de crescimento e mortalidadedos exemplares coletados de maio/2006 a agosto/2007 no estuário Guajará com auxílio doprograma FISAT II (FAO-ICLARM Fish Stock Assessment Tools). O comprimento máximoassintótico (L∞) estimado foi quase sempre maior para as fêmeas e a constante de crescimentoindividual (K) maior para os machos na maioria dos casos. O L∞ variou de 36.5 a 46.9 mm e Kde 0.2 a 0.44 ano-1. O coeficiente de mortalidade total (Z) variou de 2.04 a 3.78 ano -1 e por pesca(F) de 1.57 a 2.75 ano-1 As taxas de explotação calculadas alertam para um estado desobreexplotação da espécie. O estudo do crescimento e mortalidade em crustáceos é de extremaimportância, por fornecer informações básicas para subsidiar o ordenamento e manejopesqueiro.

Palavras-chave: Recurso pesqueiro; Camarão de água doce; Status pesqueiro, Recurso costeiro;Sobrepesca

Pan-American Journal of Aquatic Sciences (2016), 11(1): 1-17

2 B. BENTES ET AL.

IntroductionMacrobrachium amazonicum is a freshwater

shrimp of large distribution in Brazilian continentaland estuarine waters (Holthuis 1952). In theAmazon, it is included among the most abundantspecies and represents an important resource insubsistence and commercial fishing of the Amazonestuary (Bentes et al. 2012). Even so, little is knownabout the volumes produced and/or marketed in theregion (Bentes et al. 2011).

The only data of shrimp fishery production inthe State of Pará are from fishing statistics obtainedby IBAMA (Brazilian Institute of Environment andRenewable Natural Resources), in which the annualfisheries landings in key ports of the coastal zone arerecorded, corresponding to various species together,as marine shrimps (Farfantepenaeus subtilis,Xiphopenaeus kroyeri, and Litopenaeus schmitti,among others) and freshwater shrimps (M.amazonicum and M. surinamicum) (IBAMA 2011).

In recent years they appeared different studieson the biology of M. amazonicum. Highlightingcertain features in the cultivation environment, suchas: the molting cycle (Hayd et al. 2008; Hayd 2007);fecundity and fertility (Lobão et al. 1986); larvaldevelopment (Maciel 2007; Araújo and Valenti2005; Vetorelli 2004; Araújo and Valenti 2003;Rojas et al. 1990; Magalhães 1985); relative growth(Coelho et al. 1982; Roverso et al. 1990; MoraesRiodades and Valenti 2002); density of cultivation(Kimpara 2007; Moraes Riodades et al. 2006;Moraes Riodades 2005; Kimpara 2004; Lobão et al.1994); as well as the economic potential of theseactivities in Brazil (Brown et al. 2010; Kutty andValenti 2010; Moraes Valenti and Valenti 2010; New2005; Moraes-Riodades and Valenti, 2001; New etal. 2000; Odinetz Collart and Moreira 1996; Valenti1993; Valenti 1985; Coelho et al. 1981) and inVenezuela (Romero 1982). In natural environment,known items of the species in different basins are:fecundity and fertility of the species (Silva et al.2004); its oocyte development (Chaves andMagalhães 1993); reproductive strategies (Bentes etal. 2011; Porto 1998; Odinetz Collart 1991a; OdinetzCollart and Rabelo 1996); the population structure inthe Tucuruí Hydroelectric Complex (Odinetz Collart1991b); genetics (Vergamini et al. 2011) andmorphology (Porto 2004); as well as the dynamics inthe population of the Combu Island, Pará (Frédou etal. 2010).

The Amazon estuary is mainly a place of greatproductivity of M. amazonicum, since commercialcatches of this species in this environment supply

the volume of Amazon shrimp consumed in thecapital city of Belém, where the specimens are soldaccording to size (small, medium or large) atdifferent prices. According to the perception ofriverside population, who use M. amazonicum asresource in the Amazon estuary, the average size andabundance of individuals caught have beendecreasing over time, probably due to the increasedeffort in the island region of the Guajará Bay, wherethis resource supports the families that live there.

As there are no studies that evaluate thepopulations of M. amazonicum in the Amazonestuary, except for the work by Frédou et al. (2010),which confirms a situation of overexploitation of thespecies on Combu Island, this study approaches thepopulation dynamics of the species across theestuary region of the Guajará Bay, seeking toestimate growth and mortality parameters for theapplication of evaluation models regarding thesituation of M. amazonicum exploitation. In thisway, we intend to contribute to a greater knowledgeabout the species and subsidize fishing managementinitiatives.

Material and MethodsStudy area: Guajará Bay (Figure 1) is a region of theAmazon estuary submitted to the mixture ofcontinental waters and sediments that drain into the seacarried by the Amazon River (Barthem 1985). It hasirregular bottom composed of sand and mud (fluid andcompact), forming banks of sediments that aredistributed according to the action of the river flowsand tides (Gregório and Mendes 2009).

These characteristics give the estuary a verydynamic environment, including the presence ofmarine and freshwater species, whose catches andmarketing represent important economical profits forthe State of Pará (Bentes et al. 2011).Collection and processing of samples: Specimens ofM. amazonicum were obtained through monthlysurveys from May 2006 to August 2007 in six sites ofthe Guajará Bay (Figure 1), namely: waterfront ofBelém; Combu Island; Icoaraci district; ArapirangaIsland; Mosqueiro Island; and Furo das Marinhas.Sampling was performed with traps –locally calledmatapis, which are made of natural fibers and recycledPET bottles placed in pairs. Three types of ‒ matapiswith different sizes were used: S (small), M (medium)and L (large). Detailed description of this fishing gearand its dimensions can be obtained in Bentes et al.(2011).

Traps were baited with babassu flour (Orbignyaspeciosa) and pieces of fish. The set of traps was


Lt = L 1-e{-k(t-to)-CK sen(2(t-ts)}

2

Population dynamics of M. amazonicum 3

Figure 1. Localization of shrimp collection sites from May 2006 to August 2007 in the study estuary (State of Pará,Brazil). The codes refer to collection sites: MQ = Mosqueiro Island (Port of Pelé); FM = Furo das Marinhas (MosqueiroIsland); AR = Arapiranga Island; IC = Icoaraci District; BL = Waterfront of Belém; CB = Combu Island.

placed at low tide on the day before the new moon. Atthe first low tide the following day, the matapis werecollected, totaling approximately 12 hours of stay inwater.

Caught specimens were properly labeled andtransported to the Laboratory of Fisheries Biology atthe Federal University of Pará, where weremeasured the carapace length CL = measure from‒the back of the orbit to the rear edge of the carapace(cm), without including the rostrum; using a digitalpachymeter (Digimess model, accuracy 0.01mm) -and the total weight (g), with the assistance ofprecision balance (Mettler Toledo model, accuracy0,001g).

Sex was determined by the observation of theendopodite morphology of the second pair ofpleopods, as proposed by Ismael and New (2000).Analysis of the data: From 9,117 individuals ofAmazonian shrimp caught, 9,077 were identifiedregarding sex and they had the CL and the massrecorded. CL and body weight averages were testedthrough ANOVA between sex and months. Analyseswere performed using FISAT II program(FAO/ICLARM Stock Assessment Tools, version

1.1.2, available athttp://www.fao.org.br/fi/statist/fisoft/fisat/, (GayaniloJr. et al., 2000-2004). At the end of the analyses, theresults corresponding to the values of CL wereconverted to total length (TL), according to theequation of linear regression,

TL = 11.158 + 3.8822 CL (Equation 1).

Growth: In order to estimate body growth for each sex, we used the monthly distributions of CL (mm), grouped into class intervals of 2 mm consider length of individuals range (Bentes et al., 2012). For the adjustment of the growth curve, we used von Bertalanffy’s generalized model (1934) (Equation 2),establishing the function between TL and the age of the specimens from the following formula

(Equation 2)In which, L1 = Estimated total length at age t (mm); t= average age at length Lt (years); L = Asymptoticlength (mm); K = Constant growth (year-1) – speed


http://www.fao.org.br/fi/statist/fisoft/fisat/

4 B. BENTES ET AL.

at which the individual tends to reach L; C =Oscillation amplitude of growth rate (ranging from 0to 1); to = Age (year) at length Lt = 0 obtained byPauly’s empirical equation (1979) (Equation 3):Log10(-to) = -0.392 – 0.275Log10L - 1.038Log10K(Equation 3)ts = ‘Summer point’ - varying from 0 to 1 and corresponds to the time of year when the growth rateis higher.

The ts is replaced by WP (winter point) in someroutines, which designates the period of the year inwhich the growth rate is lower. This parameter can alsovary from 0 to 1, in which '0' is equivalent to January1st and '1' to December 31st.

From monthly distributions of CL frequency,we carried out the separation of age groups anddetermined the average length per cohort, viaBhattacharya’s method (1967). For the adjustment ofvon Bertalanffy’s model, from these data of averagelength by age class, we used Appeldoorn’s methods(1987, 1989) and 'Length at age' routine of FISAT IIprogram. Additionally, the same frequencydistribution data were used for another adjustment ofthe growth curve by ELEFAN I method (Pauly andDavid, 1980, 1981), which identifies modes (peaks)and intermodal (valleys), after the calculation ofaverages in a process known as 'restructuring'. In allcases, the adjustments were performed consideringthe recruitment of two distinct cohorts per year,resulting from two different periods of spawning(Frédou et al. 2010).

Longevity or maximum age (tmax) wasestimated from the observation of the growth curvegraphs obtained through ELEFAN I.

Additionally, K and L∞ estimates were usedfor calculation of the growth performance index ('),according to Moureau et al. (1986) (Equation 4) andsubsequently compared with other estimates of otherMacrobrachium species, obtained from growthparameters (L∞ and K) in the literature.’ = log K + 2*logL∞ (Equation 4)

Mortality: Natural mortality rate (M) of M. amazonicum was estimated by five methods using sexes separately:1) Rikhter and Efanov (1976) (Equation 5) thatrelated M with the age of the first maturity (t50);

M = [(1.52 (t50)0.72] 0.16 (Equation 5)‒In which t50 corresponds to the age at which 50% ofthe population is mature, being estimated by vonBertalanffy’s reverse equation, considering L50 =11.5 mm of CL, as obtained by Bentes et al. (2011).

2) Pauly’s formula (1980) (Equation 6), whichassumes that natural mortality is related to vonBertalanffy’s growth parameters, K (year-1) and L

(mm), as well as to the water average surfacetemperature, which for the area of study wasestablished at 28 °C:

(Equation 6)

3) Alverson and Carney (1975) (Equation 7), whichestimates M from K and tmax;

M = 3 K/((e0,38Ktmax) – 1) (Equation 7)

4) Hoenig (1983) (Equation 8), which relates naturalmortality to the maximum age (tmax), indicating thatthe first is inversely proportional to the second;

M = e1.44-0.982*log10(tmax) (Equation 8)

5) Roff (1984) (Equation 9) which relates mortalityto the age of first sexual maturation (t50);

M=3K/(eKt50

-1) (Equation 9)

To obtain the total mortality rate (Z) we used the following methods:1) Ricker’s catch curve method (1975) (Equation10), converted to lengths, through von Bertalanffy’sreverse equation (1934);2) Beverton and Holt’s method (1956) (Equation 11)that calculates Z from the average lengths of catchand K and L parameters; and3) Ault and Ehrhart’s method (Equation 12) that isderived from Beverton and Holt’s formula (1956)adding Lmax factors (higher length class caught).Ln (C (L1, L2) ∆t (L1, L2)) = C – Z*t ((L1 + L2)/2))(Equation 10)In which: C = oscillation amplitude of populationgrowth rate (ranging from 0 to 1);

(Equation 11)

(Equation 12)In which: L’ = length from which 100% ofindividuals are vulnerable to fisheries;

Lmedium= average length of total catch;


Ln (M) = -0,0066 – 0,27Ln(L) + 0,6543Ln(K) + 0,463Ln (T)


From the rates of natural and total mortality,we calculated F (fishing mortality) and E (rate ofexploitation) rates, according to equations 13 and 14.

F = Z – M (Equation 13)

(Equation 14)

All mortality parameters were calculated formales and females separately and later for bothsexes together. In this case, the parameters of growthcurve used were the averages obtained between thetwo sexes.

In order to estimate the pattern of recruitmentin the fishing area, i.e., the intensity of recruitmentpulses, were also used the frequencies of length andthe parameters of von Bertalanffy’s equation.Knowing the growth parameters, it was possible tocalculate how long would have taken a determinedsize group to reach from L = 0 mm up to the currentsize. The accumulation of frequencies (deducted themortality) results in recruitment intensity over thecourse of a year.Yield per recruit: The model of yield per recruit(Beverton and Holt 1956), used assuming 'knifeedge' selectivity, was applied to both sexes together.This method assumes that the selection ofindividuals for fishing occurs at a certain point andthat given a certain pattern of growth (K and L∞)and natural mortality, yields of catches depend onthe value of mortality by fishing (F) and the size orage of the first catch (Lc or tc), respectively. For theuse of this method, it was necessary to estimateLc/L∞ and M/K parameters reasons. Additionally,were used the average size of first catch (Lc),obtained for both sexes together from commercialfishing data by Lucena Fredou et al. (2010). Thistotal length value was converted to carapace lengthby Bentes’ et al. formula (2011). Thus, we obtainedthe average total length (TL) of first catch, Lc = 4.45cm or 8.61 mm of carapace length (CL). In addition,were calculated tc using von Bertalanffy’s growthinverse equation (tc = 0.5) and also average growthparameters by sex.

From the average of the values found forfishing mortality (F) and total mortality (Z), werecalculated the current exploitation rate, which wascompared with the rates provided by FISAT routine:E0.1 = exploitation rate that corresponds to the pointon the curve of yield per recruit, in which the slopeis 10% of the maximum value;Esb50% = rate at which spawning biomass would be at50% of virgin biomass;Emax = value of the exploitation rate obtained whenthe yield is maximum.

ResultsPopulation structure: The smallest individual catchhad 1.55 mm carapace length (CL) and the largestone 44.72 mm. The overall average was 12.88 mm,which corresponds to 61.2 mm total length. Theamplitudes of size and total weight, as well asaverages and deviations for each sex and sexestogether are listed in Table I.

The average length (CL) and mass differedbetween sexes. Females were larger (F = 21.46, p<0.01) and heavier (F = 339.7, p<0.01) than males.Females were more abundant than males, but thelater predominated in the largest size classes(Chi-square: 910.99; p<0.01) (Figure 2). Body growth: Regarding the frequency distributionof normal curves size by Bhattacharya’s method, M.amazonicum showed two modes or one evidentmode (Figures 3 and 4).

From the sequential observation of modalvalues, it was possible to confirm the recruitment oftwo annual cohorts for males (Figures 3 and 5A) andfemales (Figures 4 and 5B), which were probablyborn in April (cohort 1) and August/September (2cohort), in the case of males, and May/June (cohort1) and August/September (cohort 2) in the case offemales. The young shrimps were recruited withcarapace lengths that ranged from 11.44 to 13.64mm.

Table I. Average carapace length variation and mass of Amazonian shrimps collected in the Guajará Bay (Pará). N =number of specimens collected in each category; CL = carapace length; SD = standard deviation; F = female; M = male;IND = indeterminate sex; and T = total of individuals.

SexCL (mm) Mass (g)

N Min Max Mean SD Min Max Mean SDF 4471 2.57 44.72 14.80 2.84 0.04 13.80 2.88 1.37M 4181 1.55 34.23 12.95 1.78 0.02 22.07 1.96 0.91

IND 465 3.09 30.22 11.36 3.39 0.02 12.37 1.27 1.09T 9117 1.55 44.72 12.88 3.68 0.02 22.07 1.97 1.69


E =

FZ

6 B. BENTES ET AL.

Figure 2. Absolute frequency (A.F.) of M. amazonicum carapace length (CL - in mm) collected from April 2006 toAugust 2007. F = females, M = males.

Figure 3. Monthly frequencies of carapace length (CL - mm) by length class of females M. amazonicum, calculated byBhattacharya’s method of mode decomposition. The specimens were collected in the Guajará Bay and on MosqueiroIsland from April 2006 to August 2007. Y axis frequencies calculated are represented as 10¹ and those highlighted with* have Y scale = frequency 10².



Figure 4. Monthly frequencies of carapace length (CL - mm) by males M. amazonicum length class calculated by theBhattacharya’s mode decomposition method. The specimens were collected in the Guajará Bay and on MosqueiroIsland from April 2006 to August 2007. Y axis frequencies calculated are represented as 10 to the first (10¹) and thosehighlighted with * have Y scale = 10² frequency.

The follow-up of the corresponding modes andsubsequent adjustments provided growth parameterssimilar to the two cohorts of each sex recruited peryear. The analyses of CL sequences performed byELEFAN also confirmed this pattern (Figure 5).

Similarly, von Bertalanffy’s curve growthparameters estimated were similar among themethods used. The asymptotic maximum length(L∞) estimated by the methods was almost alwaysgreater for females and the individual growthconstant (K) was higher for males in most cases(Table II). L∞ ranged from 36.5 to 46.9 mm of CLor 152.8 to 193.23 mm pf TL; and K from 0.20 to0.44 year-1 for both the two cohorts as among thedifferent methods used for obtaining estimates.Among all methods used, ELEFAN seemed to be themost adjusted to the data. For this reason, it wasused to obtain estimates of mortality and growthperformance rate.

Growth curve oscillation intensity, representedby C and WP values, were estimated in routineswhere these values could be calculated. Growth wasslower almost always in the second half of the year,except for Appeldoorn’s routine, in which WP wassmaller (Table II).

Values t0 found for females were -0.139 yearsand 0.12 years for males, calculated assuming theuse of growth parameters obtained throughELEFAN. By the observation of the graphs, it isconcluded that longevity was approximately fouryears for males and six years for females. Growthperformance rates ( ) varied between 7.75 and 8.20.ɸMortality and recruitment pattern: Natural mortalityrates (M) estimated ranged from 0.77 (Pauly’smethod) up to 2.12 (Roff’s equation) and wereroughly similar between sexes (Table III).Total mortality (Z) was higher for males in mostcases except when estimated by catch curve, which


8 B. BENTES ET AL.

was higher for females, ranging from 2.37 to 3.78year-1 for males; 2.04 to 3.55 year-1 for females; and2.86 to 3.74 year-1 for sexes together (Table IV;

Figure 6). Fishing mortality rate varied from 0.39 to2.75 for both sexes, and exploitation rate variedfrom 0.19 to 0.77.

Figure 5. Growth curves of two annual cohorts (gray and black lines) estimated by ELEFAN 1 for frequency data ofcarapace length (CL in mm) in M. amazonicum caught in the Guajará Bay between April 2006 and August 2007. Thearrows indicate the recruitments. A- males, B- females.

Table II. Estimation of von Bertalanffy’s growth curve parameters ( ’) in ɸ M. amazonicum collected in the Guajará Bayfrom April 2006 to August 2007 for both cohorts identified and for males (M) and females (F). L∞ = asymptotic length(mm); CL = carapace length (mm) converted to TL = total length (cm) by linear regression, CT = 3.8822*CC+11,158(Bentes et al. 2011); K = constant growth (year-1); C = growth rate oscillation amplitude; WP = winter point.

Method Cohort Sex L∞ (CL – mm) L∞ (TL – cm) K (year-1) C WP (year) ’ɸ

Appeldoorn1

M 42.14 17.47 0.23 1 0.4 7.846F 40.5 16.84 0.2 0.5 0.5 7.754

2M 44.8 18.51 0.21 1 0.34 7.857F 43.0 17.81 0.4 0.5 0.5 8.103

Length at age1

M - - - - - -F 45.0 18.59 0.31 0.5 0.5 8.029

2M - - - - - -F 45.0 18.59 0.3 0.5 0.5 8.015

ELEFAN I1

M 44.8 18.51 0.4 0.8 0.7 8.137F 46.6 19.21 0.31 0.7 0.9 8.058

2M 36.5 15.29 0.44 0.77 0.7 8.012F 46.9 19.32 0.4 1 1 8.174



Table III. Natural mortality rates (M) of Amazon Shrimp M. amazonicum Heller, 1862 – calculated by using indirectmethods of analysis, whose description and parameters used for the estimates are discussed throughout the text.

Method Males Females Grouped sexesRikhter & Efanov 2.03 1.93 1.98

Alverson & Carney (1975) 1.68 1.25 1.44Pauly (1980) 0.85 0.8 0.77

Hoenig (1983) 2.34 1.96 2.12Roff (1984) 2.31 2.07 2.10

Average 1.79 1.52 1.66

Table IV. M. amazonicum total mortality (Z) estimated for males (M), females (F) and sexes together (S.T.) of obtainedthrough different methods. F# = fishing mortality; E = current exploitation rate; L medium = average length ofexperimental catches; L’ (mm) = minimum length at which 100% of individuals are caught.

Method Sex Z year-1 F# year-1 EL medium

(mm)L’(mm)

Catch curveM 2.37 1.57 0.66 12.95 10F 3.55 2.75 0.77 14.8 10

S.T. 2.86 2.06 0.72 12.88 10

Beverton & HoltM 3.78 2.12 0.56 12.95 10F 2.05 0.39 0.19 14.8 10

S.T. 2.92 1.26 0.43 12.88 10

Ault & EhrhartM 3.77 2.11 0.56 12.95 10F 2.04 0.38 0.19 14.8 10

S.T. 3.74 2.08 0.56 12.88 10

Figure 6. Catch curve converted into lengths according to Ricker (1975) for M. amazonicum caught in an Amazonestuary from April 2006 to August 2007. The white dots represent the classes that were not used in the calculation. A-males, B- females, C- both sexes.

The pattern of recruitment for M. amazonicumthrough projection of all length classes’ frequenciesshowed two annual peaks. One of the peaks wasalways more intense than the other, confirming theentry of two groups of young specimens per year inthe adult population, at two different times (Figure7). Yield per recruit: In the current situation, with Lc =8.65 mm, which corresponds to 0.5 years and E =0.50, fishing is at the maximum possible yield(approximately 0.51 g per recruit) for EMax = 0.5 and

E0.10 = 0.41. In this situation, the shrimps’ biomass innature is only 26% of the virgin biomass, i.e., justover 1/4 of the biomass existing before fishingexploitation (Figure 11).

Therefore, to maintain a more sustainablesituation, it would be necessary to increase the sizeof specimens in the first catch, allowing a largeryield without threatening the stock with a growthoverfishing due to excess of little specimens incatches. If the increase were Lc =10 mm, whichcorresponds to tc = 0.62, and keeping the current


10 B. BENTES ET AL.

fishing mortality pattern, Eoptimum will be 0.5,considering that our current rate of exploitation is0.5. In this case, we will have a yield increase ofalmost 9%, with a prediction of 0.54 g per recruit,after we have obtained the new balance. In this newsituation, the biomass exploited will represent 28%of the virgin biomass (Figure 8).

In a situation even more sustainable, Lc couldbe adjusted to 11.5 mm or its equivalent: tc = 0.75years (Table V). This age corresponds to theestimation of first sexual maturation (t50). In thiscase, the maximum exploitation rate is 0.61 and ourcurrent rate (0.50) would coincide with the estimatedvalue for E0.10. Under these conditions, the fishingeffort does not need to be changed and the yieldbalance would be 0.57 g per recruit, representing anincrease of almost 9%, in relation to the currentsituation. Biomass exploited would be 31% of thevirgin biomass. This new situation will allow an

increase in the spawning stock in nature andtherefore in the reproductive potential of the species.

In a situation even more sustainable, Lc couldbe adjusted to 11.5 mm or its equivalent, tc = 0.75years, which corresponds to the estimate of averagesize and age at first sexual maturation (L50 and t50).In this case, the maximum exploitation rate is 0.60and our current rate (0.50) would coincide with theproposed value for E0.10. Considered that E0.10 is amore conservative value and highly suggested forthe regulation of fisheries, this situation is stronglyrecommended. Under these conditions, the fishingeffort does not need to be changed and the yield ofbalance would be 0.69 g per recruit, representing anincrease of 11% in relation to the current situation.Biomass exploited would be 31% of virgin biomass,which will allow an increase in the spawning stockand therefore the reproductive potential of thespecies.

Figure 7. Pattern of M. amazonicum recruitment caught in an Amazon estuary (Guajará Bay) from April 2006 to August2007 showing two pulses of recruits in a year. M = males; F = females; G.S.= grouped sexes.

Figure 8. Yield and biomass per M. amazonicum recruit depending on the fishing mortality (F), for three different agesof first catches, 0.50, 0.62 and 0.75, respectively.



Table V. Parameters of Beverton and Holt’s yield analysis per recruit for sexes together of Amazon shrimp caught in anAmazon estuary from April 2006 to August 2007. Lc = average length of commercial catch (carapace length - mm)according to Lucena Frédou et al., (2010); E0.1 = curve point in which yield per recruit is 10% of virgin biomass; E =current rate of exploitation from tc of commercial fishing.

Parameter tc=0.50 tc=0.62 tc=0.75F year-1 1.6 1.6 1.6E 0.5 0.5 0.5Lc 8.65 10.00 11.50Emax 0.503 0.557 0.617E0,10 0.408 0.455 0.505Yield per recruitment (g) 0.62 0.66 0.69Biomass per recruitment (g) 0.39 0.41 0.43Proportion of virgin biomass 0.26 0.28 0.31Average weigh of caught specimens (g) 1.0 1.3 1.7Average age of caught specimens (years) 0.8 0.9 1.1

DiscussionThe average size of the specimens collected is

greater than most recorded in other studies in theAmazon and in other parts of Brazil (in naturalenvironments). The highest carapace length averagewas recorded on Combu Island (13.75 mm). In thisstudy, we obtained also the largest individual of M.amazonicum reported in the literature, with a femalewith 44.72 mm carapace length or 18.45 cm totallength (Table VI).

The largest specimen collected was a femalenotably, this was the biggest shrimp of the speciesever recorded in the literature.

Generally, the individuals caught in flowingwaters of large rivers have greater lengths thanshrimps collected in calmer waters of floodplainlakes and dams (Odinetz Collart 1993). The largestsize of individuals observed in this study may berelated to increased productivity in the Guajará Bay,with wide food availability and refuge, whereindividuals can optimize growth and fecundity(Paiva et al. 2006).

The population structure varied depending onthe sex of the specimens. Females were alwayslarger and heavier than males. Hartnoll (1982),Mantelatto and Martinelli (2001) state that bodygrowth patterns among crustaceans is similarbetween the sexes until the individual reaches sexualmaturity. From that stage, growth rate becomessmaller in females, which typically reach sizes lowerthan those of males in most species.

To obtain growth parameters of fish or otheraquatic organisms, direct methods can be used, byreading age rings conformed in hard parts such asscales, otoliths and spines; and indirect methods,through length frequency data or capture andrecapture experiments (Etim and Sankare 1998). Theapplicability of the first method is not possible for

shrimps, since they are organisms that do notdevelop calcified structures. In this case, only theanalysis based on length frequencies can be used forthis purpose. Many computational analysis toolshave been developed to work with this type of data.The applicability of their results has been discussed(Isaac 1990; Defeo et al. 1992). However, Tomalin(1995) states that ELEFAN is the most widely used,because this method does not require advancedknowledge of statistics. The method does not requirenormality in the distribution of data to be used,which can be considered as an advantage. Studies onprecision developed by Isaac (1990) show that thismethod is much more accurate for short-livedspecies, as in the case of shrimps.

Von Bertalanffy’s growth curve parameterswere similar among the methods used; however, theasymptotic maximum length (L∞) estimated wasalmost always greater for females, unlike that foundby Silva (2006). From all species ofMacrobrachium, M. fellicinum (Inyang 1984),common in the Western region of the Africancontinent, has the smallest asymptotic maximumsize (total length) already estimated: L∞ = 9.1 cm(Etim and Sankare 1998).

The largest size reached by females favors thesurvival and reproduction through increasedfecundity. In addition, their mortality may bedecreasing by having cryptic habit, due to the factthat they incubate the eggs and seek protection to thelitter.

M. amazonicum growth revealed commonsituations among most crustaceans: rapid growth,low longevity and more than one recruitment peryear, as stated by Frédou et al. (2010). Recruitmentduring two peaks in a year corroborates with Silva(2002) who found recruits in rainy and lessrainy periods, in catches performed in Vigia, State of


12 B. BENTES ET AL.

Table VI. Size variation of M. amazonicum in different sites. TL= Total Length (cm). Empty cells = information notrecorded in the study. Sources: 1: This study; 2: Frédou, 2010; 3: Odinetz Collart, 1987; 4: Odinetz Collart, 1987; 5:Silva, 2002; 6: Borges, 2003; 7Odinetz Collart & Moreira, 1993; 8: Odinetz Collart & Enriconi; 9: Vargas & Paternina,1977; 10: Guest, 1979; 11: Davant, 1963; 12: Coelho et al., 1982; 13: Romero, 1982.

Site Environment TL max TL med AuthorPará River(Pará - Brazil) Natural (lotic)

18.45 6.2 (males)6.8 (females) 1

Guamá River(Pará - Brazil)

Natural (lotic) 14.1 - 2

Tocantins River(Pará - Brazil)

Natural (lotic) 13.2 - 3

Tucuruí Lake(Pará - Brazil)

Reservoir (lentic) 8.0 5.5 4

Guajará-Mirim River(Pará - Brazil)

Swamp 14.4 7 (males)7.6 (females) 5

Rômulo Dam Campos(Bahia - Brazil)

Reservoir (lentic) 17.8 - 6

Careiro Island(Amazonas -Brazil)

Natural (lotic) 10.6 6 7

Central Amazon – Brazil

Natural (lotic) 6,0 - 8

Colombia Culture6,8 (males)

7,7 (females)- 9

Ceará - Brazil Culture 10 6 10Ceará - Brazil - 11 - 11

Pernambuco - BrazilCulture (lentic –culture ponds)

- 8 (commercial) 12

Venezuela Culture (lentic –culture ponds)

- 6 13

Pará, Brazil, and Frédou et al. (2010) on CombuIsland.

At the laboratory, Guest (1979) estimated thetheoretical asymptotic length for this species in102.5 mm total length. At that time, the largestspecimen’s measure recorded was 100 mm. Thisdifferentiation in estimating L∞ in this work andothers with the species is probably related tocatching larger specimens in the Amazon estuary,taking advantage of the fact that it is a nutrient-richenvironment, which justifies a greater growth ofspecimens caught in the area. The influence ofincreased growth density is manifested throughincreased intra-specific competition for food andspace, in addition to other forms of interactionbetween the animals. When food or space becomeslimited, growth will possibly be reduced as a way tosave the accumulated energy (New and Singholka1982; Sandifer and Smith 1985).

The intensity of growth curve oscillation,represented by C and WP values suggest that growthdecrease because of the reproduction, corroboratingthe result of smaller values of catabolic constant forfemales. Seasonality of growth in aquatic animals isa well-known phenomenon, primarily in temperate

environment. In the tropics, growth variations aremore related to the seasonal rain pattern and/orreproduction (Etim and Sankare 1998; Bentes et al.2011).

Growth performance rate ranged between 1.0and 2.6. The value obtained in this study was thelargest among the works with M. amazonicum andother species of the genus that demonstrate vonBertalanffy’s growth curve parameters, which canalso be interpreted as a result of the highproductivity in the Guajarino estuary (Table 7).

Regarding the estimates of mortality rates,they are difficult to obtain, given the complexity ofthe necessary data and yet because the methods areinaccurate, especially in regard to natural mortality(Isaac et al. 1992). This way, the uncertainty in thevalues of natural mortality (M) is recognized. In anattempt to reduce this uncertainty, we chose to useseveral indirect methods, commonly used for crabs.Many empirical models were initially developed forfish and they include few data for invertebrates.However, the use of multiple methods can ensurebetter approximation to the parameter.

Natural mortality (M) coefficients weresimilar between sexes; however, total mortality (Z)



obtained by catch curve was greater for females.Noting that the catch curve showed a convex format,it is assumed that the mortality rate of femalesregarding the lowest classes of length is greater. Onthe other hand, the average size of males’ catcheswas smaller than females’, which can be linked tomales’ vulnerability to catches when they havesmaller sizes than females, since the proportion ofmales is always greater than that of females insmaller length classes.

Similar results were found for males by Silvaet al. (2007). Other works with species of the samegenus show high mortality rates as Z = 10.6 year-1,M = 3.36 year-1 and F = 7.24 year-1 for M.macrobrachium (Enim 1995); Z = 3.4 year-1, M =3.61 year-1, and F = 0.26 year-1 (Gabche and Hockey1995); Z = 3.69 year-1, M = 1.97 year-1 and F = 1.72year-1 (Etim and Sankare 1998) both for M.völlenhovenii. It is discussed that the methods usedto obtain these parameters regard the values of Mconstant for all ages, but this fact is completelyunreal. It is clear that a smaller individual is exposedto greater mortality from predation that a biggerindividual, simply by the fact that smaller animalshave more predators than larger animals. Thisdifference in mortality from predation (which isperhaps the biggest cause of natural mortality insmall aquatic animals) should be relatively high(Sparre and Venema 1992).

Recruitment has two annual peaks, one ofwhich is always more intense than the other. Thenumber of recruitment peaks somewhat compensatesfor the high rates of mortality recorded for thespecies. In any population, there is a certain amountof individuals that reproduce each period and theseare the responsible for producing the sufficientamount of new specimens, so that a sufficientnumber of new individuals enter each time enablingto restock the population.

The optimum sustainable exploitation ratescalculated (Emax) were close to the estimates of thecurrent exploitation of M. amazonicum, i.e., thecurrent rate of exploitation lies in the maximumsustainability. In this situation, any increase of effortmay lead to growth overexploitation or overfishing.Catches, even in small-scale fishing, focus on thejuveniles, whose lengths at first catches were lowerthan those at first maturity. This result is moreevident regarding the males of the species.

It is assumed that larger males were notcaught because they are no longer available in thepopulation. These catches are characterized by highvalues of fishing mortality and by a catch withreduced weight. Because of this apparent reductionof stock, it is suggested to catch specimens with sizegreater than 4.5 cm (tc>0.75 year) total length. Thatis, given this data, by increasing the size of the firstcatch, we would have better fishing yield (increaseof 14% if compared to current yield).

Table VII. Macrobrachium species values of growth performance rate ( ). SX = sex of individuals; M = males; F =ɸfemales; Mn = minimum; Mx = maximum; S.T. sexes together; TL total length; k = growth constant (year -1). Sources:1:Silva et al., 2007 & Frédou et al., 2010; 2: Sampaio & Valenti, 1996; 3: Etim & Sankare, 1998; 4: Alhassan & Armah,2011; 5: Deekae & Abowei, 2010; 6: Enim, 1995; 7: Khan et al., 2004; 8: Mantel & Dudgeon, 2004; 9: RománContreras & Campos Lince, 1993

Specie Site SXL∞

TL (cm) K ɸ SourceMn Mx Mn Mx Mn Mx

M. amazonicumGuajará Bay -

BrazilM 12.4 17.7 0.7 1.4 2.0 2.6

1F 12.7 14.1 0.7 0.9 2.0 2.3

M. rosenbergii Culture - Brazil S.T. 18.4 23.4 0.2 0.4 1.9 2.3 2

M. völlenhovenii Reservoir - Africa S.T. 18.0 0.9 2.5 3

M. völlenhovenii Reservoir - Africa S.T. 14.2 1.0 2.3 4

M.macrobrachium Streams - Nigeria S.T. 8.3 2.0 2.1 5

M. macrobrachium Estuary - Nigeria S.T. 12.9 1.8 2.5 6

M. malcolmsonii Reservoir - India S.T. 20.3 0.6 2.4 7

M. hainanensis Streams - China S.T. 3.6 0.7 1.0 8

M. acanthurus Mexico S.T. 21.2 0.2 2.0 9


14 B. BENTES ET AL.

The decrease in average size of M.amazonicum in catches should be a research target,in view of the complaint by many fishermenregarding exponential increase in the number offishermen and fishing traps used in small-scalefishing. Another discussion is the predation ofAmazon shrimp by the Malaysian prawn (M.rosenbergii) also included in the region of study(Barros and Silva 1997). Additionally, the increasein fishing effort of catches has been occurringgradually. This is a fact observed by the amount oftraps used by fishermen’s families during the time ofcompletion of this work, that is, fishing mortalityseems to be the strongest interference in thecontinuity of stocks. Frédou et al. (2010) havealready warned about this situation on CombuIsland. Co-management initiatives could bediscussed with the coastal communities in order toincrease the selectivity of gear used, thus avoidingthe increasing catch of shrimp that have not yetreproduced.

In terms of management, a priori, we suggestaccurate analyses to assess the sustainability ofAmazon shrimp farming in the State of Pará. Beforebeing established, each enterprise must go through athorough analysis of the likely impacts produced andhold a guarantee of economic and socialsustainability. In Pará, this would possibly be animmediate solution to decrease fishing pressure innatural stocks; however, there should be ensuringaccess to credit for small local producers inparticular, so that the goal can be reached as well asimprovement of the living conditions of theseworkers.

Still, according to Martinelli (2005), referringto marine shrimps of the species Farfantepenaeussubtilis (pink shrimp), Xiphopenaeus kroyeri (seabobshrimp) and Litopenaeus schmitti (white shrimp),preserving estuaries, preventing intensivesmall-scale fisheries of younger shrimps and alsoensuring the health of waters without pollutants ofany nature is essential for the maintenance andsurvival of shrimps.

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Received: June 2015Accepted: December 2015

Published: April 2016


Population dynamics of Macrobrachium …1)_1-17.pdfMacrobrachium amazonicum is a freshwater shrimp...

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Transcript of Population dynamics of Macrobrachium …1)_1-17.pdfMacrobrachium amazonicum is a freshwater shrimp...