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Faculdade de Oceanograa, Instituto de Geocincias, Universidade Federal do Par,
Av. Augusto Corra 01, Guam, Belm, Par, CEP 66075-110, Brazil
This study estimates reproductive parameters (length at rst maturity, reproduction period, and
fecundity), mortality, and maximum sustainable yield for Macrobrachium amazonicum on Combu
Island (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 the
ood 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 maximum
sustainable yield (E
) for males was lower than that for females. For both sexes, E
below the exploitation rates (E). The present study demonstrates the need for management of the
prawn shery on the islands in the Amazon estuary, such as establishing a minimum commercial
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 coletadas
mensalmente, entre maro de 2003 e fevereiro de 2004. Fmeas ovgeras foram registradas ao longo
de todo ano, com picos reprodutivos no meio da enchente (maro), vazante (setembro) e pico de
mnima vazo (dezembro) no Rio Guam. O tamanho na qual 50% das fmeas esto maduras de
60,8 mm de comprimento total. Fecundidade absoluta variou entre 40 e 3375 ovos por fmea. A taxa
de explorao para a obteno do rendimento mximo sustentvel (E
) para machos foi inferior
ao obtido para as fmeas. Para ambos sexos E
foi abaixo das taxas de explotao atuais (E).
O presente estudo demonstra a necessidade do manejo da pesca de camaro nas ilhas do esturio
Amaznico, como por exemplo o estabelecimento de tamanho mnimo de comercializao.
) Fax: +55.9132017609; e-mail: ucena@ufpa.br
Koninklijke Brill NV, Leiden, 2010 Crustaceana 83 (3): 277-290
Also available online: www.brill.nl/cr DOI:10.1163/001121609X12596543952298
The river prawn, Macrobrachium amazonicum (Heller, 1862) (Decapoda, Palae-
monidae) is an endemic species in South America and occurs throughout the basins
of the rivers Orinoco, Amazon, and Paraguay (Holthuis, 1952). This species is also
found 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 has
free-swimming larvae, which, under natural conditions, may develop in both fresh
and estuarine waters (Alekhnovich & Kulesh, 2001).
M. amazonicum is the main species of freshwater shrimp exploited by the
commercial artisanal eet in the Amazon (Odinetz-Collart, 1987; Odinetz-Collart
& Moreira, 1993). In the rivers and estuaries of the Amazon, this prawn is mainly
caught 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 for
aquaculture (Kutty et al., 2000).
Dynamic pool models are widely used in population dynamics and have become
a 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 et
al., 2007), Pandalus (cf. Hvingel & Kingsleu, 2006), Aristeus (cf. Maynou et al.,
2006), and Melicertus (cf. Conides et al., 2006). Amazon stocks of M. amazonicum
have never been assessed, despite its enormous socio-economic importance in the
region, 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 other
regions in the richness of the exploited species, the quantity of the catch, and the
dependence 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 aquatic
environments of the Amazon remain poorly known. In many areas, river and
estuarine communities have a low quality of life, even through they live within
an ecosystem with a high potential for providing goods and services.
In order to provide the knowledge and baseline information required for the
management of M. amazonicum stocks in the Amazon estuaries, the aim of the
present study was to estimate the reproductive parameters (length at rst maturity,
reproduction period, and fecundity), mortality, and maximum sustainable yield for
this species on Combu island (Amazon estuary).
Combu (Par, Brazil) is an island of approximately 15 km
, located on the left
margin of the Guam River, 1.5 km from the city of Belm (Par) (g. 1). Annual
air temperature ranges from 23 to 32

C. Total monthly rainfall ranges from 200 to

350 mm in the rainy season (December to May) and from 90 to 170 mm in the dry
season (June to November) (Moraes et al., 2005).
Each month, approximately 500 g (ca 100 individuals) of M. amazonicum were
obtained from commercial catches caught using matapi traps between March
2003 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, separated
into ve categories according to criteria established by Carvalho & Pereira (1998):
(a) stage I, underdeveloped (ovary transparent); (b) stage II, developing (ovary
cream or pale green); (c) stage III, ripe or almost ripe (ovary at full size, dark green,
oocytes visible); (d) stage IV, ovigerous (enlarged brood chambers bearing eggs);
and (e) stage V, spent (large ovary with whitish aspect). The reproductive period
was dened as those months with a maximum percentage of ovigerous females.
Size at sexual maturity (TL
) was determined by the proportion of ovigerous
or 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.
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 coefcients
of 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
), corresponding to a proportion of 50% sexually mature individuals, was
estimated as the negative ratio of the two coefcients TL
= (a/b).
Eggs from 118 individuals in maturity stage V were used to determine species
fecundity. 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 obtained
and extrapolated to the total volume of the egg mass. The correlation between
total length and number of eggs was tested by the non-parametric Spearman R
The instantaneous coefcient of total mortality (Z) was estimated using the
length-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 rst
maturity (t
). Age at rst maturity was obtained using the inverse Von Bertalanffy
equation considering the length at rst maturity obtained in this study. As length
at rst maturity was only obtained for females, M was calculated by grouping
sexes. The instantaneous rate of shing mortality (F) was computed from the
difference of the instantaneous rates of total mortality and natural mortality. The
exploitation 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 expected
numbers to those that were actually caught. From this curve, the length at rst
capture (L
) 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 E
(the exploitation rate at which maximum
relative yield-per-recruit is obtained).
Fishery mortality, length at rst capture, and maximum sustainable yield were
obtained for males and females separately as well as for pooled sexes. Growth
parameters used as input for the estimates were based on various methodologies
described in Silva et al. (2007). The effects of the different values of growth
parameters in estimating the population parameters were assessed.
Fig. 2. Frequency histogram for ovigerous females (dark bars) of Macrobrachium amazonicum
(Heller, 1862).
Ovigerous females were recorded throughout the year, except in July. There
were three reproductive periods established, i.e., September, December, and
March, when, respectively, 35%, 45%, and 21% of the females were ovigerous
(g. 2). Among the 1338 females analysed, 239 (18%) carried eggs on their
abdomen and 1099 (82%) did not. Total length for females ranged from 52 to
106 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 3375
eggs/female, with a mean value of 905 590. The correlation between fecundity
and 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 method
varied little, considering the various growth parameters L

and K (Silva et al.,

2007) used to obtain age at rst maturity (t
). The exception occurred for the
estimate derived from the growth parameter obtained from Electronic Length
Frequency Analysis (ELEFAN) (table I).
Regarding length at rst capture (L
), the various methods resulted in similar
values, and females showed larger sizes than males. Total mortality (Z) estimated
from the catch curve method was similar for each set of growth parameters, with
the exception of the ELEFAN method. The estimate for males (6.39 year
) was
higher than that for females (3.86 year
). Similar results were obtained from the
Beverton & Holt (1956) method. The exploitation rate for maximum sustainable
yield (E
) for males was lower than that for females when considering the
various methods (table II). In most cases, E
for both males and females was
Fig. 3. Size at rst maturity (TL
in mm) for females of Macrobrachium amazonicum
(Heller, 1862).
below the exploitation rates (E) found using the Ricker catch curve method and
the Beverton & Holt method.
Fig. 4. Fecundity and total length relationship for females of Macrobrachium amazonicum
(Heller, 1862).
Estimates of age of rst capture (t
) and natural mortality under different growth parameters for
Macrobrachium amazonicum (Heller, 1862)
Growth method L

(cm)/K (cm/month)
(year) (year
Gulland & Holt 15.75/0.98 1.60 0.93
Fabens 17.66/0.78 1.64 0.91
Appeldoorn 15.84/0.94 1.52 0.96
Seasonal length-at-age 12.38/1.35 1.56 0.95
Non-seasonal length-at-age 12.37/1.36 1.61 0.92
ELEFAN 14.00/0.70 0.87 1.53
Source: Silva et al. (2007).
Estimates of total mortality (Z), shery mortality (F), exploitation rates (E), and length at rst catch
) through both the catch curve and the Beverton & Holt method obtained for Macrobrachium
amazonicum (Heller, 1862)
Growth method L

Catch curve Beverton & Holt
Gulland & Holt 15.75/0.98 6.04 5.11 0.85 4.530 9.31 8.38 0.89 0.51
Fabens 17.66/0.78 6.13 5.22 0.85 4.666 8.90 7.99 0.89 0.49
Appeldoorn 15.84/0.94 5.87 4.91 0.84 4.574 9.03 8.07 0.89 0.50
Seasonal 12.38/1.35 5.04 4.09 0.81 4.373 8.24 7.29 0.90 0.56
Non-seasonal 12.37/1.36 5.06 4.14 0.82 4.342 8.30 7.38 0.90 0.56
ELEFAN 14.00/0.70 3.15 1.62 0.51 4.248 5.43 3.90 0.72 0.54
Gulland & Holt 12.79/0.80 4.25 3.32 0.78 6.112 4.43 3.50 0.79 0.65
Fabens 13.16/0.75 4.27 3.36 0.79 6.015 4.43 3.52 0.79 0.64
Appeldoorn 12.66/0.92 3.86 2.90 0.75 5.878 4.92 3.96 0.81 0.64
Seasonal 14.14/0.66 4.32 3.37 0.78 6.051 4.55 3.60 0.79 0.62
Non-seasonal 14.03/0.67 4.41 3.49 0.79 6.058 4.55 3.63 0.80 0.62
ELEFAN 14.00/0.70 6.89 5.36 0.78 6.279 4.73 3.20 0.68 0.73
Pooled sexes
Gulland & Holt 15.70/0.37 4.06 3.13 0.77 4.621 3.66 2.73 0.75 0.54
Fabens 15.11/0.38 4.06 3.15 0.78 4.621 3.54 2.63 0.74 0.55
Appeldoorn 12.17/0.56 3.66 2.70 0.74 4.413 3.57 2.61 0.73 0.58
Seasonal 10.50/0.71 3.63 2.68 0.74 4.404 3.34 2.39 0.72 0.61
Non-seasonal 10.51/0.68 3.28 2.36 0.72 4.384 3.20 2.28 0.71 0.61
ELEFAN 14.00/0.80 6.89 5.36 0.78 4.551 6.56 5.03 0.77 0.55
Source: Silva et al. (2007).
The freshwater prawn, Macrobrachium amazonicum occupies a special position
among the resources of economic and social importance in the Amazonian region.
The species is widely caught and commercialized the Amazon estuary. This
resource serves as the main and often only source of animal protein for human
river and estuarine communities.
The identication of eggs adhering to the pleopods of females throughout the
year 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. amazonicum
has two reproduction strategies in the Amazon basin regarding the duration of the
reproductive period: populations with well-dened seasonal reproduction, which
occur in coastal rivers; and those with continual reproduction, which occur in
inland rivers. Prawns on Combu Island belong to the second group (continual
reproduction), but with periods in which reproduction is more intense.
Reproductive peaks were observed in March, September, and December, with
the highest percentage of females with eggs adhered to the pleopods recorded in
December. These months mark the middle of the ood period (March), the low
water period (September), and the peak dry period (December) of the Guam River.
Odinetz-Collart (1991) reports the same reproductive pattern in lakes and rivers of
the Central Amazon and Silva et al. (2002) report this pattern along the coast of
the state of Par. In the Tocantins River, however, reproduction of the species is
concentrated during the peak ood season (Silva et al., 2005).
Gonad maturation in crustaceans depends upon internal factors (endogenous
control) as well as environmental characteristics (Odinetz-Collart, 1991). Pinheiro
& Hebbing (1998) report that the reproductive period of freshwater decapods is
closely 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 rivers
must be the determinant factor in the reproduction of M. amazonicum.
On Combu Island, the main stimulus for M. amazonicum reproduction appears
to be the increased velocity of the currents and increased food supply. In the Guam
River, the strongest currents occur in the middle of the ood and the low water
periods (Pinheiro, 1987), which is when prawns take advantage of the greater
dispersive power of the water to release their larvae and thereby occupy a greater
number 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 et
al., 2006) and, consequently, zooplankton in the Guam River. As larvae of M.
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 40
to 3345 (mean 850 eggs per female). A signicant positive correlation was
found 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 individuals
of the same species, increasing in quantity with the size of the female. Studying
the 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 within
the range of the variables considered.
According to Coelho et al. (1982), the river prawn can produce as many as
6000 eggs per spawning. However, studies carried out in Venezuela found lower
numbers (Gmba, 1984). Females with 148 to 1554 eggs have been recorded in
northeastern 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 fecundity
in comparison to other species of the same genus, the absolute fecundity of which
is less than 200 eggs per female. However, this species fecundity is considered
low in comparison with species that reach larger sizes and have greater economic
interest, 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 fecundity
ranges from 12 800 to 194 350 eggs (Coelho et al., 1982; Valenti, 1984; Lobo
et al., 1985; Scaico, 1992). According to Lobo et al. (1986) and Scaico (1992),
M. amazonicum spawns monthly, which may determine the low fecundity, while
compensating for the lower number of eggs per spawn in comparison to species of
greater 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 sizes
larger than those recorded for populations of the same species inhabiting fresh
water (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 lower
values for freshwater populations in comparison to estuarine populations. Mashiko
(1983) found that freshwater M. nipponense (De Haan, 1849) females mature
at signicantly lower sizes than for those in estuaries, attributing this difference
to 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 prawn
stocks. Management advice for most prawn stocks in the North Atlantic is basically
formulated by a qualitative assessment of trends in stock condition in response
to catch history (Koeller et al., 2000). For the relatively slow growing prawns of
northern waters, the use of standard techniques such as VPA and length-based
methods has not been successful (Quinn et al., 1998). This may be due to the fact
that 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 shery
models, which assume natural mortality to be constant (Hvingel & Kingsley,
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 with
constant parameters, they are important in determining whether an area is either
heavily shed or under-exploited in order to establish appropriate management
measures (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 also
tested 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 of
the methods employed, the results of the stock assessment models converge and
are consistent with knowledge on the shery and biology of the species.
Instantaneous coefcients of total mortality (Z) were higher for males. This
probably stems from the fact that males have a smaller length at rst capture (L
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 sex
proportion favouring females on Combu Island (Silva et al., 2007), which is a
common situation among prawns.
Consequently, the recommendable exploitation rate for maximum sustainable
yield is lower for males than for females. For both sexes, E
is below current
exploitation rates (E) (considering the different methods and scenarios), thereby
demonstrating that this activity may not be sustainable in the near future. This
information is corroborated by other evidence regarding the species on Combu
Island. Size at rst capture (L
) for both sexes (more evident among males) is
smaller than size at rst maturity (L
). This situation accelerates the reduction in
numbers of individuals in the environment, thereby negatively altering population
fecundity, 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 parameters
available in the literature for the species or even other species of the genus, which
renders comparisons of the values obtained here with those from other locations
The present study demonstrates the need for management of the prawn shery
activity on islands of the Amazon estuary, given the importance of and local human
dependence on the exploitation of natural resources. There is no information on the
volume of M. amazonicum catches, as ofcial statistics have failed to consider
this resource. This places serious limitations to the application of analytical
models and, consequently, the application of specic management measures.
It also restricts knowledge regarding the impact of this shery activity on the
Amazon estuary. It is likely that the species is not renewing itself adequately
due to the capture of immature individuals. Controlling shery efforts is not a
very viable measure, considering the large number of people involved in the
prawn shery activity and the dispersal of boats over the large area of the
Amazon estuary. However, based on the results of the present study, an alternative
management measure would be the establishment of a minimum commercial
size (not considering what is used for sustenance) to be obtained through the
determination of a xed trap opening size or minimum commercialization size.
The authors acknowledge Dr. Victoria Isaac Nahum for comments on earlier
drafts of the manuscript. This study was partially nanced by National Council
for Scientic and Technological Development (CNPq) through a grant to the third
author and a research grant to the rst author (302280/2007-3).
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First received 31 October 2008.
Final version accepted 11 May 2009.