Nauplius
e-ISSN 2358-2936
www.scielo.br/nau
www.crustacea.org.br
ORIGINAL ARTICLE
Morphometric aspects of two coexisting
amphidromous shrimps, Atya gabonensis
Giebel, 1875 and Atya scabra (Leach, 1816),
in the Paraíba do Sul River, Brazil
Abner Carvalho-Batista1,2 orcid.org/0000-0001-8193-1269
Caio M.C.A. Oliveira1 orcid.org/0000-0002-4662-1422
Guilherme Souza3 orcid.org/0000-0002-0834-6415
Fabrício Lopes Carvalho4 orcid.org/0000-0001-7851-0578
Fernando Luis Mantelatto1 orcid.org/0000-0002-8497-187X
1
Universidade de São Paulo (USP), Faculdade de Filosofia, Ciências e Letras de
Ribeirão Preto (FFCLRP), Departamento de Biologia, Laboratório de Bioecologia
e Sistemática de Crustáceos (LBSC). Av. Bandeirantes, 3900, 14040-901, Ribeirão
Preto, São Paulo, Brazil.
AC-B E-mail: carvbatis@gmail.com
FLM E-mail: flmantel@usp.br
2
Universidade Paulista (UNIP), Instituto de Ciências da Saúde, Avenida Marquês
de São Vicente, 3001, 05037-040, São Paulo, São Paulo, Brazil.
3
ONG Projeto Piabanha. Campo de Sementes, 28570-000, Caixa Postal 123546,
Zona Rural / Itaocara, Rio de Janeiro, Brazil.
GS E-mail: guilhermesouza.bio@gmail.com
4
Universidade Federal do Sul da Bahia (UFSB), Centro de Formação em Ciências
Agroflorestais (CFCAf), Grupo de Pesquisa em Carcinologia e Biodiversidade
Aquática (GPCBio). Rodovia Ilhéus-Itabuna, km 22, CEPEC/CEPLAC,
45662-200, Ilhéus, Bahia, Brazil.
FLC E-mail: flcarvalho@ufsb.edu.br
ZOOBANK: http://zoobank.org/urn:lsid:zoobank.org:pub:6B13D127-5657-47C7-
9087-C6FDCFD31969
ABSTRACT
Corresponding Author
Abner Carvalho-Batista
carvbatis@gmail.com
SUBMITTED 21 March 2020
ACCEPTED 03 December 2020
PUBLISHED 16 April 2021
DOI 10.1590/2358-2936e2021018
All content of the journal, except where
identified, is licensed under a Creative
Commons attribution-type BY.
Nauplius, 29: e2021018
Atya gabonensis Giebel, 1875 and Atya scabra (Leach, 1816) are amphidromous
shrimps. In some areas, these species populations are vulnerable due to
the anthropogenic impact on their habitats and commercial exploitation.
However, basic morphometric data is still lacking. This study provides
morphometric data on both species in the Fluvial Island Domain of Paraíba
do Sul River basin, Brazil. Sampling was performed bimonthly from January
2013 to March 2014. Individuals were analysed according to sex, weight,
carapace length and width, abdomen length, second abdominal pleura height
and width, length of third pereopod articles, and length and width of the
male appendix interna. We sampled 42 individuals of A. gabonensis and 16
individuals of A. scabra. In both species, females showed abdomen and second
abdominal pleura larger than males, while males showed third pereopod
1
Carvalho-Batista et al.
Morphometric aspects of Atya gabonensis and A. scabra
articles larger than females, both related to sexual dimorphism. We detected differences in the carapace length
× carapace width relationship and in the growth pattern of the male appendix interna between species, with A.
gabonensis presenting the carapace and the male appendix interna wider than A. scabra. Morphometric aspects
of both species are unpublished for this region, and this data is valuable for A. gabonensis in the western Atlantic
and for A. scabra from Rio de Janeiro state. Such information is essential for future systematic assessment and
establishment of conservation management policies.
KEYWORDS
Atyidae, biogeography, Caridea, Fluvial Island Domain, systematics.
INTRODUCTION
Atya gabonensis Giebel, 1875 and Atya scabra
(Leach, 1816) are phylogenetically close species of
the family Atyidae De Haan, 1849 (Page et al., 2008;
von Rintelen et al., 2012; Oliveira et al., 2019) with an
amphidromous life cycle: their larvae are dependent
on estuarine or marine environments (McDowall,
2007; Bauer, 2013). The adults feed mainly on
planktonic algae, organic debris and insect parts
(Chace, 1972; Fryer, 1977; Obande and Kusemiju,
2008) and are distributed in the countries of South,
Central and Caribbean America as well as in western
Africa (Hobbs and Hart, 1982; Melo, 2003). Atya
gabonensis and A. scabra are target species of artisanal
fishing at different intensities, both in Brazil and Africa
(Oliveira, 1945; Motoh and Murai, 2006; Kadjo et al.,
2016), and present significant interest in the aquarium trade (Werner, 2003; Lipták and Vitázková,
2015; Weiperth et al., 2019).
Despite the wide geographic distribution and
commercial interest, there are still few studies on
the population biology of these two species. For A.
gabonensis, the scarcity of information is evident,
and studies are restricted to the populations of the
African continent. In Brazil, so far, there are mainly
studies focusing on the diagnosis of the species and
reporting its occurrence (Hobbs, 1980; Fonseca et al.,
1994; Ramos-Porto and Coelho, 1998; Melo, 2003).
Galvão and Bueno (2000) studied the population
structure of A. scabra and its reproductive biology
in the Guaecá River, in the city of São Sebastião,
while Herrera-Correal et al. (2013) studied the
fecundity and reproductive output at São Sebastião
Nauplius, 29: e2021018
Island, both located on the northern coast of São
Paulo state. Almeida et al. (2010) and Barros et al.
(2020), as well, investigated several aspects of the
reproductive biology of this species in southern Bahia.
However, there is a clear lack of information about
populations of A. scabra in other locations throughout
Brazil, and information on A. gabonensis in American
regions remains scarce. This has led to A. scabra and
A. gabonensis being considered as Near Threatened NT and Data Deficient - DD, respectively, in the latest
evaluation made for species inhabiting the Brazilian
territory (Mantelatto et al., 2016).
The presence of these species was recorded
during the monitoring of the aquatic fauna of the
Paraíba do Sul River islands (Carvalho et al., 2019).
This is the last stretch of the lower middle course of
the Paraíba do Sul River, between the cities of São
Sebastião do Paraíba and São Fidélis, covering the
municipalities of Aperibé, Cambuci, Itaocara and
Santo Antonio de Padua, Rio de Janeiro state (Souza
et al., 2007). It is also found in the Pomba and Grande
rivers. Environmental diversification in the area is
high, represented by a relatively conserved riparian
forest mosaic with various depths and different
hydrodynamic conditions caused by the presence of
islands (Bizerril, 1998).
The capture of shrimps of the genus Atya has
been reported among fishermen of the Fluvial Island
Domain (Souza et al., 2007), indicating that both
species are under fishing pressure in this region,
especially for human consumption by local people.
In addition, the Paraíba do Sul River basin is located
in one of the most industrialised regions of Brazil
(between the states of Minas Gerais, São Paulo and
2
Carvalho-Batista et al.
Rio de Janeiro) (Marengo and Alves, 2005; Devide
et al., 2014), with a history of intense deforestation of
riparian forests (Dantas et al., 2000), water pollution
in most of the municipalities (Cavadas-Barcellos et
al., 2011), installation of dams and hydroelectric
plants, numerous environmental accidents and the
introduction of exotic species (Polaz et al., 2011;
Moraes et al., 2017).
In this context, the present study aims to provide
unprecedented information about these populations
for A. gabonensis on the American continent and
for A. scabra in the state of Rio de Janeiro, Brazil.
Such information can provide a scientific basis for
future reassessment of the genetic diversity, for the
monitoring of these species, as well as for developing
sustainable fishing strategies and management policies
for this region.
Morphometric aspects of Atya gabonensis and A. scabra
MATERIAL
AND
METHODS
Sampling
Specimens of A. gabonensis and A. scabra were
collected bimonthly from manual collections
performed by a 30-minute active search via free diving
by a local fisherman. Sampling was performed from
January 2013 to November 2014 at six sites in the
Fluvial Island Domain of the Paraíba do Sul River.
Three sites were located in the course of the Paraíba
do Sul River and three in the tributaries: Pomba
River and Grande River (Fig. 1). This region can
be considered a priority area for the conservation of
aquatic endangered species (Polaz et al., 2011) since
there are large parts of the river in continuity with
the estuary.
Figure 1. Sampling sites for shrimp of the genus Atya Leach, 1816 in the Fluvial Island Domain in the Paraíba do Sul River basin
(modified from Carvalho-Batista et al., 2019). NOTE: Crustacean sampling sites/Paraíba do Sul River Basin/Main Hydrography/
State Limits.
Nauplius, 29: e2021018
3
Carvalho-Batista et al.
After capture, individuals were placed in labelled
plastic bags and frozen. At the laboratory, individuals
were thawed and stored in 80 % ethanol in the Decapod
Crustacean Collection of the Department of Biology
(CCDB), at the Faculty of Philosophy, Sciences and
Letters at Ribeirão Preto, University of São Paulo,
Brazil. All individuals were identified to the species
level, following Melo (2003), and sexed, based on the
presence of the male appendix interna on the second
pair of pleopods.
Carapace length (CL) was measured in millimetres
and standardised as an independent variable. We used
the Student’s t-test to compare the carapace length of
males and females. All individuals were weighed on a
0.1 g precision digital scale. The relationship between
carapace length and individual weight was represented
by the equation y = axb, where “y” is the individual’s
weight, “x” is the length of the carapace, “b” is the
allometric coefficient of the structure studied and “a”
is the line intercept on the ordinate axis. The other
measurements were as follows: total length (TL),
carapace width (CW), abdomen length (AL), width
Morphometric aspects of Atya gabonensis and A. scabra
of abdomen at the second segment (AW), height and
length of the second abdominal pleura (PH and PL,
respectively), length of the merus (MerL), carpus
(CarL), propodus (ProL), dactylus (DacL) of the third
pereopod, length and width of the male appendix
interna (AmL and AmW) and ratio L/W of Am (Fig. 2),
which were used as dependent and CL-related
variables.
The parameters were estimated by linear regression
of logarithmic data, i.e., with a linear version of the
model (log y = log a + bx log x). Comparisons between
males and females in terms of the measured ratios were
performed by covariance analysis (ANCOVA). The
allometric state of each structure was analysed (b >
1 - positive allometry; b < 1 - negative allometry; b =
1 - isometry), and the null hypothesis H0 (b = 1) was
tested by the Student’s t-test for linear regression (α =
0.05) (Zar, 1999). All analyses assumed a significance
level of 0.05, and all data were tested for normality
and homoscedasticity; when data did not meet these
assumptions, the corresponding nonparametric tests
were used.
Figure 2. Dimensions used in the morphometric analyses of Atya Leach, 1816. A, (CL) carapace length, (PL) length of second
abdominal pleuron, (PH) height of second abdominal pleuron; B, (CW) carapace width, (TL) total length, (AW) abdomen width,
(AL) abdomen length; C, (MerL) merus length, (CarL) carpus length, (ProL) propodus length, (DacL) dactylus length; D, (AmL)
length of male appendix interna, (AmW) width of male appendix interna.
Nauplius, 29: e2021018
4
Carvalho-Batista et al.
Morphometric aspects of Atya gabonensis and A. scabra
RESULTS
Atya gabonensis
In total, 42 individuals were sampled: 17 (40.5 %)
males and 25 (59.5 %) females. The carapace length of
males ranged from 19.54 to 58.49 mm, with an average
of 41.26 ± 11.01 mm, while that of females ranged
from 25.74 to 46.40 mm, with an average of 39.18 ±
5.28 mm. The overall average was 40.02 ± 7.92 mm.
The weight of males ranged from 3 to 85 g (mean
= 35.6 ± 26.1 g), while that of females ranged from
8 to 44 g (mean = 26.7 ± 18.7 g); the overall average
was 30.3 ± 18.7 g. The relationship between carapace
length and individual weight was described by the
potential equation y = 0.0002CL3.1895, where y is the
weight and CL is the carapace length in mm, with a
determination coefficient (R 2) of 97 %. There was
no statistically significant difference between sexes
for the CL × weight relationship (p <0.05) (Fig. 3).
The total length of males ranged from 51.4 to 137.6
mm, with an average of 103.0 ± 24.1 mm; females
Figure 3. Relationship between carapace length and weight of
Atya gabonensis Giebel, 1875 in the Fluvial Island Domain in the
Paraíba do Sul River basin.
ranged from 75.6 mm to 124.1 mm, with an average of
102.5 ± 12.0 mm. The overall average was 102.7 ± 17.7
mm. The CL × CW relationship was the only one that
showed no statistically significant difference between
sexes (Tab. 1). The equations that describe each of
these relationships, as well as each R², are shown in
Figure 4. Both sexes presented negative allometry for
Table 1. Results of the Covariance Analysis (ANCOVA) of morphometric data for Atya gabonensis Giebel, 1875 at the Fluvial
Island Domain in the Paraíba do Sul River basin. (CL) carapace length, (PL) length of the second abdominal pleuron, (PH) height
of the second abdominal pleuron, (CW) carapace width, (TL) total length, (AW) abdomen width, (AL) abdomen length, (MerL)
merus length, (CarL) carpus length, (ProL) propodus length, (DacL) dactylus length. Significant difference: * P <0.05; ** P <0.01.
Relationship
Factor
(Group)
CL vs. TL
Male vs. Female
CL vs. CW
Male vs. Female
CL vs. PL
Male vs. Female
CL vs. PH
Male vs. Female
CL vs. AW
Male vs. Female
CL vs. AL
Male vs. Female
CL vs. MerL
Male vs. Female
CL vs. CarL
Male vs. Female
CL vs. ProL
CL vs. DacL
Nauplius, 29: e2021018
Male vs. Female
Male vs. Female
Par. (Log)
F
P
0.0012**
a
12.256
b
0.4330
0.5145
a
0.0868
0.7730
b
1.5027
0.2438
a
---------
-------
b
7.7304
0.0084*
a
137.8092
0.0000**
b
0.3777
0.5425
0.0000**
a
124.0965
b
0.4630
0.5003
a
17.8022
0.0001**
b
0.0013
0.9715
a
---------
--------
b
6.3221
0.0165*
a
20.2880
0.0011**
b
2.4464
0.1259
a
----------
---------
b
15.8218
0.0007**
a
13.6782
0.0007**
b
3.6156
0.0657
5
Carvalho-Batista et al.
the CL × TL relationships and CL × CW and isometry
for CL × AL. In other relationships, there were different
growth patterns between males and females. Males
presented positive allometry for most of relationships
with third pereopod articles (CL × MerL, CL × CarL,
CL × DacL), while females showed isometry. On the
Morphometric aspects of Atya gabonensis and A. scabra
other hand, females presented isometry for CL × PL
and CL × PH, and positive allometry for CL × AW,
while male presented negative allometry for all these
relationships. Males presented isometry for the CL ×
AmL and CL × AmW relationships and decreasing
ratio Am L/W. (Tab. 2, Fig. 4).
Figure 4. Comparison between morphometric relationships of males and females of Atya gabonensis Giebel, 1875 at the Fluvial
Island Domain in the Paraíba do Sul River basin. (CL) carapace length as independent variable and the dependent variables: A, (CW)
carapace width; B, (TL) total length; C, (AL) abdomen length; D, (AW) abdomen width; E, (PL) length of the second abdominal
pleuron; F, (PH) height of the second abdominal pleuron; G, (MerL) merus length; H, (CarL) carpus length; I, (ProL) propodus
length; J, (DacL) dactylus length; K, (AmL) length of male appendix interna; L, (AmW) width of male appendix interna; M, (Am
L/W) male appendix interna length/width.
Nauplius, 29: e2021018
6
Carvalho-Batista et al.
Morphometric aspects of Atya gabonensis and A. scabra
Table 2. Regression analyses of morphometric data of Atya gabonensis Giebel, 1875 at the Fluvial Island Domain in the Paraíba
do Sul River basin. Carapace length (CL) is the independent variable. (PL) length of second abdominal pleuron; (PH) height of
second abdominal pleuron; (CW) carapace width; (TL) total length; (AW) abdomen width; (AL) abdomen length; (MerL) merus
length; (CarL) carpus length; (ProL) propodus length; (DacL) dactylus length; (AmL) length of male appendix interna; (AmW)
width of male appendix interna.
Relationship
CL vs. TL
CL vs. CW
CL vs. PL
CL vs. PH
CL vs. AW
CL vs. AL
CL vs. MerL
CL vs. CarL
CL vs. ProL
Sex
N
Intercept
Inclination
R²
T (b-1)
P
Allometry
Females
25
0.6289
0.8654
0.9073
2.3375
<0.05
-
Males
17
0.5519
0.902
0.9891
4.0065
<0.05
-
Both sexes
41
0.3863
1.0518
0.9766
1.9871
<0.05
+
0
Females
25
0.1062
0.9028
0.6501
0.7038
<0.05
Males
17
0.0522
0.8003
0.9868
8.3612
<0.05
-
Females
25
0.1632
0.9794
0.8258
0.2192
<0.05
0
Males
17
0.1383
0.9114
0.9844
2.9903
<0.05
-
Females
25
0.6322
1.214
0.8385
1.9262
<0.05
+
Males
17
0.2706
0.9401
0.9852
1.9468
<0.05
-
Females
25
0.4377
0.837
0.7777
1.7470
<0.05
-
Males
17
0.4057
0.8375
0.973
4.5110
<0.05
-
Females
23
0.0608
0.9294
0.8026
0.7019
<0.05
0
Males
17
0.4304
1.1802
0.9796
1.7917
<0.05
+
Females
25
0.6193
1.0992
0.9808
0.9637
<0.05
0
+
Males
17
0.864
1.2743
0.8383
5.9639
<0.05
Females
25
0.7025
1.0016
0.7387
0.0126
<0.05
0
Males
17
0.6049
0.9585
0.9393
0.6597
<0.05
0
Females
25
0.568
0.8286
0.4318
0.8458
<0.05
0
Male
17
1.0394
1.1569
0.9401
2.0812
<0.05
+
CL vs. AmL
Males
17
0.8531
0.989
0.9477
0.1640
<0.05
0
CL vs. AmW
Males
17
1.1037
1.0089
0.9622
0.1542
<0.05
0
CL vs. DacL
Atya scabra
In total, 16 individuals were sampled: 8 (50 %)
males and 8 (50 %) females. Carapace length of males
ranging from 29.23 to 43.75 mm, with an average of
38.34 ± 10.61 mm, while that of females ranged from
8.37 to 30.09 mm, with an average of 22.06 ± 10.04
mm. The overall average was 29.19 ± 10.27 mm.
Male weight ranged from 11 to 39 g (mean = 26
± 10.27 g), while female weight ranged from 1 to 11
g (average = 6.62 ± 3.92 g); the overall average was
16.31 ± 12.5 g. The relationships between carapace
length and weight of males and females were described
by the potential equations y = 0.0123CL2.0247 and y =
0.005CL2.29748, respectively, where y is the weight and
CL is the carapace length in mm (Fig. 5).
The total length of males ranged from 78.62 to
112.64 mm, with an average of 100.40 ± 24.26 mm.
Female length ranged from 31.13 to 81.12 mm, with
an average of 64.95 ± 23.15 mm. The overall average
Nauplius, 29: e2021018
Figure 5. Relationship between carapace length and weight of
Atya scabra (Leach, 1816) in the Fluvial Islands Domain in the
Paraíba do Sul River basin.
was 80.46 ± 23.44 mm. The CL × TL, CL × CW,
and CL × DacL relationships showed no statistically
significant differences between sexes (Tab. 3).
The equations that describe each of these relationships,
as well as each R², are shown in Figure 6. Both sexes
presented negative allometry for the CL × TL and
CL × CW relationships and isometry for CL × ProL,
7
Carvalho-Batista et al.
Morphometric aspects of Atya gabonensis and A. scabra
Table 3. Results of the Covariance Analysis (ANCOVA) of morphometric data for Atya scabra (Leach, 1816) at the Fluvial Island
Domain in the Paraíba do Sul River basin. (CL) carapace length, (PL) length of the second abdominal pleuron, (PH) height of the
second abdominal pleuron, (CW) carapace width, (TL) total length, (AW) abdomen width, (AL) abdomen length, (MerL) merus
length, (CarL) carpus length, (ProL) propodus length, (DacL) dactylus length. Significant difference: * P <0.05; ** P <0.01.
Relationship
Factor (Group)
CL vs. TL
Male vs. Female
CL vs. CW
Male vs. Female
CL vs. PL
Male vs. Female
CL vs. PH
Male vs. Female
CL vs. AW
CL vs. AL
Male vs. Female
Male vs. Female
CL vs. MerL
Male vs. Female
CL vs. CarL
Male vs. Female
CL vs. ProL
Male vs. Female
CL vs. DacL
Par. (Log)
Male vs. Female
CL × AL and CL × DacL. In other relationships, there
were different growth patterns between males and
females. Males presented positive allometry for CL
× MerL and CL × CarL, while females presented
negative allometry for the former and isometry for the
second relationship. Females presented isometry for
CL × PL, while males presented negative allometry for
this relationship. Males presented negative allometry
for the CL × AmL and CL × AmW relationships, and
constant ratio Am L/W (Tab. 4, Fig. 6).
DISCUSSION
As far as we know, the present study provides
unique biological information about A. gabonensis on
the American continent, allowing comparisons with
population data from the African continent for the first
time. The individuals sampled in the present study
reached high values, especially of weight (a maximum
of 13.7 cm in total length, with a weight of 85 g), when
Nauplius, 29: e2021018
F
P
A
0.5977
0.4533
B
1.3101
0.2747
A
0.0868
0.7730
B
1.5027
0.2438
A
22.2460
0.0004**
B
0.3457
0.5675
A
14.9071
0.0020**
B
0.7469
0.4043
A
23.9904
0.0003**
B
1.7851
0.2063
A
2.0988
0.1711
B
0.0025
0.9612
A
6.7336
0.0249*
B
10.4102
0.0090**
A
6.4206
0.0278*
B
20.2880
0.0011**
A
1.6998
0.2189
0.0032**
B
14.8744
A
0.5149
0.4868
B
0.7740
0.3978
compared to other populations previously studied
(Tab. 5). Such differences can be due to intrinsic
(genetics), as well as extrinsic reasons (environmental
conditions, such as food availability, temperature,
etc.) (Staples and Heales, 1991; Alford and Jackson,
1993, Okayi et al., 2012). The relationships between
the weight and length of a species are important in
determining its survival condition, providing the
basis for growth and production estimates, and
describing the structural characteristics of individuals
in populations. In this way, our results indicate that,
despite the potential anthropogenic impact, the
population of A. gabonensis from the Fluvial Island
Domain in the Paraíba do Sul River basin had a growth
potential equal to or greater than other populations in
Africa, showing an area of important occurrence for
the maintenance of the species. Systematic analysis
based on molecular data performed by two of us
(CMCAO and FLM, unpublished) maintains the
Amphi-Atlantic A. gabonensis hypothesis as well as
the morphological revision of Hobbs and Hart (1982).
8
Carvalho-Batista et al.
In the literature, there is no information on the
body weight for A. scabra. The individuals of this
species collected in the Fluvial Island Domain
reached larger sizes than previously observed in
other locations in Brazil. Even with considerably
larger sample numbers, previous studies conducted
in southern Bahia state (Almeida et al., 2010; Barros
Morphometric aspects of Atya gabonensis and A. scabra
et al., 2020) and northern São Paulo state (Galvão
and Bueno, 2000) did not find individuals with a
maximum size similar to that found in the present
study. Individuals with a maximum size larger than
that observed in the Paraíba do Sul River were
reported in Mexico by Lorán-Núñez et al. (2009)
and reached 117 mm in total length.
Figure 6. Comparison between morphometric relationships of males and females of Atya scabra (Leach, 1816) in the Fluvial Island
Domain in the Paraíba do Sul River basin. (CL) carapace length as independent variable and the dependent variables: A, (CW)
carapace width; B, (TL) total length; C, (AL) abdomen length; D, (AW) abdomen width; E, (PL) length of the second abdominal
pleuron; F, (PH) height of the second abdominal pleuron; G, (MerL) merus length; H, (CarL) carpus length; I, (ProL) propodus
length; J, (DacL) dactylus length; K, (AmL) length of male appendix interna; L, (AmW) width of male appendix interna; M, (Am
L/W) male appendix interna length/width.
Nauplius, 29: e2021018
9
Carvalho-Batista et al.
Morphometric aspects of Atya gabonensis and A. scabra
Table 4. Regression analyses of morphometric data of Atya scabra (Leach, 1816) in the Fluvial Island Domain in the Paraíba do Sul
River basin. Carapace length (CL) is the independent variable. (PL) length of second abdominal pleuron; (PH) height of second
abdominal pleuron; (CW) carapace width; (TL) total length; (AW) abdomen width; (AL) abdomen length; (MerL) merus length;
(CarL) carpus length; (ProL) propodus length; (DacL) dactylus length; (AmL) length of male appendix interna; (AmW) width of
male appendix interna.
Relationship
Sex
N
Intercept
Inclination
R²
T (b-1)
P
Allometry
CL vs. TL
Both sexes
16
0.7045
0.8178
0.9908
8.6273
<0.05
-
CL vs. CW
Both sexes
16
0.0573
0.843
0.986
5.8462
<0.05
-
Females
8
0.1204
0.9708
0.996
1.1654
<0.05
0
CL vs. PL
CL vs. PH
CL vs. AW
CL vs. AL
CL vs. MerL
CL vs. CarL
CL vs. ProL
Males
8
0.4864
0.4681
0.6809
4.0657
<0.05
-
Females
8
0.0458
0.9379
0.99
1.6163
<0.05
0
Males
8
0.2087
0.7078
0.877
2.7007
<0.05
-
Females
8
0.1398
0.9179
0.9962
3.5348
<0.05
-
Males
8
0,0131
0.7741
0.9199
2.4226
<0.05
-
Both sexes
16
0.6268
0.723
0.9685
7.9474
<0.05
0
Females
7
0.0159
0.8678
0.9766
2.2015
<0.05
-
Males
8
0.7356
1.4186
0.8761
1.9216
<0.05
+
Females
7
0.4186
0.9365
0.9738
0.9243
<0.05
0
Males
8
1.8167
1.9344
0.8854
3.2892
<0.05
+
Females
7
0.3999
0.8807
0.8834
0.8335
<0.05
0
Males
8
1.281
1.5068
0.7376
1.3814
<0.05
0
CL vs. DacL
Both sexes
15
0.8705
1.0202
0.9098
0.2266
<0.05
0
CL vs. AmL
Males
8
0.3128
0.6562
0.8958
3.7620
<0.05
-
CL vs. AmW
Males
8
0.4533
0.6129
0.8109
3.2031
<0.05
-
Males of both species had relatively larger articles
of the third pereopod when compared to females.
Almeida et al. (2010) also detected a difference in
the growth of the merus of the third pereopod in A.
scabra. Sexual dimorphism in chelipeds has been
widely reported in caridean shrimp, as such structures
are involved in agonistic disputes between males,
territories, and females for mating (Bauer, 2004).
However, there is no information whether the third
pereopod can be used for such purposes in Atya. The
third pair of pereopods is used by shrimps of this genus
to attach themselves to the substrate (usually rocky)
in strong water f low environments or waterfalls,
preventing the animal from being carried away by
the stream while performing particle filtration (Fryer,
1977; Villalobos and Álvarez, 1997).
Females, on the other hand, showed greater
abdominal growth than males, with greater abdomen
width and length as well as greater length and height
of the second abdominal pleura. These characteristics
were observed for the two studied species, being
adaptations to increase egg hatching area and,
consequently, fecundity, and useful for the detection
Nauplius, 29: e2021018
of morphological sexual maturity size in females in
future studies (Mantelatto and Barbosa, 2005; Pralon
and Negreiros-Fransozo, 2006; Pantaleão et al., 2012).
For both species, there was no difference between
males and females in the relationship between carapace
length and width, although A. gabonensis presented
positive allometry for this relationship and A. scabra
showed negative allometry. A similar allometric
pattern for this species was observed by Barros et al.
(2020), in Contas River, Bahia state, although the
authors have detected differences between sexes in
the slope and in the intercept for this relationship.
Such difference gives A. gabonensis a more robust
appearance when compared to A. scabra, the former
being considered the largest species of the genus
and more attractive for aquarists. For both sexes, of
both species, negative allometry was observed in the
correlations between the carapace and total length and
abdomen length. Both species suffer fishing pressure
in certain localities, and larger individuals reach higher
prices, although our results indicate that in larger sizes,
these animals have lower meat yields in relation to
their total size. Lima et al. (2018) observed a reduction
10
Carvalho-Batista et al.
Morphometric aspects of Atya gabonensis and A. scabra
Table 5. Minimum, maximum and mean values ± standard deviation of total length (mm) and weight (g) of Atya gabonensis Giebel,
1875 and Atya scabra (Leach, 1816), obtained in different locations, available in previous studies and the present study.
Atya scabra
Atya gabonensis
Locality
Total length
range (mm)
Sex
Min
Max
Mean ± S.D.
Weight range (g)
Min
Max
Weight range
± S.D.
Reference
Cross River, Itigidi,
Nigeria
Not
determined
65
140
-
8.1
60.3
-
Nwosu (2009)
Mu River, Makurdi,
Nigeria
Not
determined
50
122
79.9 ± 21.2
3.3
51.6
13.98 ± 10.36
Okayi and Iorkyaa
(2004)
Benue River, Nigeria
Females
32
105
-
3.2
34
-
Obande et al.
(2009)
Bandama River,
N´Douci, Ivory Coast
Males
81
138
-
11.68
86.8
40.89 ± 15.47
Females
73
129
-
11.35
47.5
23.79 ± 5.62
Paraíba do Sul River,
Brazil
Males
51.4
137.6
103.0 ± 24.1
3
85
35.6 ± 26.1
Females
75.6
124.1
102.5 ± 12.0
8
44
26.7 ± 18.7
Santana River, Ilhéus,
Brazil
Males
14.98
88.77
-
-
-
-
Females
29.03
64.4
-
-
-
-
Guaecá River, São
Sebastião, Brazil
Males
22.7
89.3
54.5 ± 13.5
-
-
-
Females
25.3
61.1
46.9 ± 6.8
-
-
-
Males
48
117
90.75 ± 11.26
-
-
-
Females
32
103
68.67 ± 7.22
-
-
-
Males
54
105
78.93 ± 10.23
-
-
-
Females
38
88
65.28 ± 7.12
-
-
-
Females
42.8
77.3
-
-
-
-
Los Pescados River,
Mexico
Actopan River, Mexico
São Sebastião Island,
Brazil
Contas River, Brazil
Paraíba do Sul River,
Brazil
Males
37.45
96.6
73.04 ± 10.83
-
-
-
Females
38.0
87.4
57.65 ± 8.65
-
-
-
Males
78.62
112.64
100.4 ± 24.26
11
39
26.0 ± 10.27
Females
31.13
81.12
64.95 ± 23.15
1
11
6.62 ± 3.92
in meat yield with increasing size of individuals from
the Amazon River in Macrobrachium amazonicum
(Heller, 1862). Atya species are used as a protein source
by riverine human populations and, although informal
trade for food is reported, this is at a smaller scale when
compared to other species, such as those of the genus
Macrobrachium, with a focus on subsistence artisanal
fishing (Oliveira, 1945; Holthuis, 1980; Hobbs and
Hart, 1982; Almeida et al., 2010).
The male appendix interna showed differences
in the growth pattern between the two species. Atya
gabonensis presented isometry for both width and
length, while A. scabra presented negative allometry
for both relationships. For A. gabonensis, the male
appendix interna length × width ratio decreases along
with animal growth, i.e., such structures becomes
proportionally wider in larger individuals. In A.
scabra, on the other hand, such relationships remained
constant in all sizes analysed. Thus, this presents
another distinctive characteristic for the two species.
Nauplius, 29: e2021018
Madelaine et al.
(2015)
Present study
Almeida et al.
(2010)
Galvão and Bueno
(2000)
Lorán-Núñez et al.
(2009)
Herrera-Correal et
al. (2013)
Barros et al.
(2020)
Present study
This study presents previously unpublished
morphometric aspects of both species for the Paraiba
do Sul River Basin, as well as the first information on
the biology of A. gabonensis on the American continent
and A. scabra for the state of Rio de Janeiro, Brazil.
The maximum sizes reached by both species and its
relationship with weight indicate the Fluvial Island
Domain as an important area for the maintenance of
population stocks of these species, which may act as a
source of individuals for other regions. Furthermore,
such populations are still viable in this area due to the
fact that the estuary is not fragmented by dams, which
enables the juvenile recruitment of these populations.
In both species, males showed a larger relative growth
of the third pereopod articles while females had a larger
growth of the abdomen. Moreover, morphometric
differences between the species were found, even for
CL × CW and for male appendix interna growth. Such
knowledge about the basic biology and morphology
of these species may serve as a basis for future genetic
11
Carvalho-Batista et al.
population assessments and the establishment of
conservation management policies for the region,
and as evaluations of the conservation status of these
species, whose habitats suffer from anthropogenic
pressures.
ACKNOWLEDGMENTS
We thank the members of LBSC (Alvaro Costa,
Edvanda Carvalho, Elis Mesquita, Natalia Rossi
and Rafael Robles), the Piabanha Project team
(Thiago Berriel, Christian Natalia, Evodio Peçanha,
Adam Carvalho, Andreza Pacheco, Dominguinhos
Afonso, João Lucas and Jacson Gomes), CEPTA/
ICMBIO (Luis Gaspar, Noel Martins and Sandoval
Santos Jr.), PESAGRO-RIO and MZUSP for their
assistance in specimen collection. FLM expresses
sincere thanks to the National Council for Scientific
and Technological Development – CNPq (Proc.
473050/2007-2, 302748/2010-5, 304968/2014-5,
302253/2019-0 and 471011/2011-8) and the São
Paulo Research Foundation - FAPESP (Temáticos
BIOTA Proc. 2010/50188-8 and INTERCRUSTA
2018/13685-5) for financial support during collection
and analysis activities. FLC thanks CEPTA/ICMBIO
for their assistance in the collections carried out under
Project UNDP - BRA/08/023 and to CNPq (Proc.
425769/2016-0). We are thankful to the anonymous
reviewers for constructive and useful comments
during the reviewing process.
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