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(cm)/K (cm/month)
1)
t
50
M
(year) (year
1
)
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
1)
Source: Silva et al. (2007).
TABLE II
Estimates of total mortality (Z), shery mortality (F), exploitation rates (E), and length at rst catch
(L
c
) 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 & Holt
Z F E L
c
Z F E E
MSY
Males
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
Females
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
1)
Source: Silva et al. (2007).
284 FLVIA LUCENA-FRDOU ET AL.
DISCUSSION
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.
POPULATION DYNAMICS OF MACROBRACHIUM AMAZONICUM 285
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
286 FLVIA LUCENA-FRDOU ET AL.
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,
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 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
c
)
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
MSY
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
c
) for both sexes (more evident among males) is
smaller than size at rst maturity (L
50
). This situation accelerates the reduction in
POPULATION DYNAMICS OF MACROBRACHIUM AMAZONICUM 287
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
impossible.
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.
ACKNOWLEDGEMENTS
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).
REFERENCES
ALEKHNOVICH, A. V. & V. F. KULESH, 2001. Variation in the parameters of the life cycle in
prawns 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 Macrobrachium
amazonicum 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
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