Interpopulational morphological analyses and fluctuating asymmetry

in the brackish crab Cardisoma guanhumi Latreille (Decapoda,

Gecarcinidae), on the Brazilian Northeast coastline

MELQUISEDEQUE S. DUARTE1, FRANCISCO A. MAIA-LIMA2 & WAGNER F. MOLINA3

1
Departamento de Biologia Celular e Genética, Pós-Graduação em Genética e Biologia Molecular, Universidade
Federal do Rio Grande do Norte, Centro de Biociências, Campus Universitário, 59078-970, Natal-RN. Brazil.
[email protected]
2
Departamento de Biologia, Faculdade de Ciências, Cultura e Extensão do Rio Grande do Norte – FACEX, 59080-
020, Natal – RN, Brazil. [email protected]
3
Departamento de Biologia Celular e Genética, Laboratório de Genética de Recursos Marinhos, Universidade Federal
do Rio Grande do Norte, Centro de Biociências, Campus Universitário, 59078-970, Natal-RN. Brazil.
[email protected]

Abstract. The brackish crab Cardisoma guanhumi Latreille, 1825 is an economically important species
used for alimentation, commonly sold in open markets throughout Northern and Northeastern Brazil. It
lives mainly in mangrove areas, which have been highly degraded because of misuse and inadequate
settlement. In this study, morphometric data were obtained from segments and dorsal and ventral regions
of 107 individuals from four brackish crab populations on Rio Grande do Norte coastline. The fluctuating
asymmetry indexes and discriminant interpopulation morphological features in measurements at dorsal,
ventral regions and pereiopods were defined. The presence of fluctuating asymmetry was found in the
four first pairs of pereiopods and a correlation between sex and segments from the first pair of pereiopods
and between population and segments in all pairs of pereiopods was observed. As for the discriminating
model, the use of the eye cavity length variable allowed discriminating 82% of individuals. The results
showed a strong correlation between variables and collection site, suggesting a natural grouping of
individuals into two macroregions (Goianinha and Canguaretama, I; and Extremoz and Macau, II). The
data show that this resource should not be exploited homogeneously, inasmuch as it apparently comprises
structured populations of restricted range.

Key words: Guaiamum, blue land crab, morphometry, biological conservation.

Resumo. Análises morfológicas interpopulacionais e assimetria flutuante no caranguejo Cardisoma

guanhumi Latreille (Decapoda, Gecarcinidae), na costa Nordeste do Brasil. O caranguejo
Cardisoma guanhumi Latreille, 1825 é uma espécie economicamente importante, sendo explorado para
alimentação e comercialização, em feiras livres no Norte e Nordeste do Brasil. Habita principalmente os
manguezais, que se encontram em avançado estágio de degradação devido às formas inadequadas de uso
e ocupação. Neste trabalho foram obtidos dados morfométricos da região dorsal, ventral e apêndices de
107 indivíduos de quatro populações do litoral do Rio Grande do Norte. Foram definidos os índices de
assimetria flutuante, bem como padrões morfológicos interpopulacionais discriminantes em medidas da
região dorsal, ventral e pereiópodos dos indivíduos. Foi demonstrada a presença de assimetria flutuante
nos quatro primeiros pares de pereiópodos e correlação dos fatores sexo e segmentos no 1º par de
pereiópodos e de população e segmento em todos os pares de pereiópodos. Quanto ao modelo
discriminante, a utilização da variável comprimento da cavidade ocular, permitiu discriminar 82% dos
indivíduos. Os resultados evidenciaram uma forte correlação das variáveis em relação à região de origem,
sugerindo um agrupamento natural dos indivíduos em duas macrorregiões (Goianinha e Canguaretama, I;
e Extremoz e Macau, II). Os dados indicam que este recurso não deve ser explorado de forma homogênea
tendo em vista que representam, aparentemente, populações estruturadas e de âmbito restrito.

Palavras-chave: Guaiamum, caranguejo, morfometria, conservação biológica.

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Interpopulational morphological analyses and fluctuating asymmetry in Cardisoma guanhumi. 295

Introduction
Brazilian mangroves are distributed from (3º45’47’’S 38º31’23’’W), until Santa Catarina
Oiapoque (04°30’N; Amapá) to Laguna (28°30’S; (27º35’36’’S 48º35’56’’W).
Santa Catarina), comprising 6,800 Km along the It lives mainly in mangroves, where they
coast and 25,000 Km² (Novelli 1989). build galleries open to receive sea water (Melo
The State of Rio Grande do Norte is located 1996). This species is economically important and
in the Northeastern coast, encompassing about 400 commonly sold at large scale in open markets in
km of seashore, with mangroves along its northern northeastern Brazil. Although there are no specific
and northeast coastal zones. The northern seashore - populational records, the uncontrolled extractivism
estuarine zone of Mossoró river, Piranhas-Açu river of this species incur the ecologic unbalance in some
- and the estuarine system Guamaré-Galinhos make mangrove zones. Thereby, strategies of stock
up an area of 3,034 ha. The northeastern seashore - protection must be defined.
estuarine zone of Maxaranguape river, Ceará-Mirim Morphological analyses were able to inform
river, Potengi-Jundiaí river, Piranhas river and the about distinct features of crustacean population and
estuary–lagoon complex of Nísia Floresta, Papeba, species (Sullivan 1998, Brian 2005). Some studies
Guaraíras and the estuaries Jacu and Curimataú– demonstrate that the expression of inter-population
Cunhaú rivers – cover an area larger than 8,898 ha. variability in the crab morphology has both
The mangrove total area in Rio Grande do Norte is environmental and genetic components that need to
about 11,992 ha. On the northeastern seashore, more be accounted for in population-level research (Brian
dense and larger mangrove areas can be found, such et al 2006) or product of ecogenesis (Daniels et al
as the ones from Curimataú–Cunhaú river, with 1998, 2001).
3,100 ha and Potengi–Jundiaí river, with 1,530 ha. Most individuals present bilateral symmetry
On the northern seashore, the biggest mangrove is and their random deviation are called fluctuating
located in the estuary of Piranhas–Açu river, with asymmetry (FA) (Møller 1998). The fluctuating
1,230 ha, characterizing the Macau region (Souza asymmetry level provides relevant information about
1996). individual performance in relation to its
Special attention has been given to development (Pravosudov & Kitaysky 2006)
ecological studies in estuaries, and efforts have been working as a useful tool to evaluate the quality or
developed for a better and a reasonable use of the developmental instability of individuals in
natural resources found in such environment association with environment (Zakharov &
(Nascimento 1980). These surveys are justified since Yablokov 1990, Li 2001, Petavy et al. 2006) and
estuaries are among the most affected areas by both genetic stress (Leary & Allendorf 1989, Palmer
natural changes and human activities (IDEC 1993). 1996, Møller 1998). Some results suggest that traits
Along northeastern Brazil, particularly in with relaxation of selection for functionality exhibit
Rio Grande do Norte, the mangroves are usually higher FA (Crespi & Vanderkist 1997).
deforested for implantation of salt mines, shrimp In the present work, we carried out
farms and oil extraction. On the northern and morphological analyses within and among
northeastern shores, salt mines, sugar cane Cardisoma guanhumi populations from mangroves
industries, fishing and in natura dumping of on the Brazilian Northeast coastline, in order to
domestic and industrial sewage also contribute to evaluate the presence of populational differences
their degradation status. Currently, the urban based on morphological analyses of locomotion and
expansion and the growth of shrimp farming body structures and fluctuating asymmetry.
represent the most important causes of negative
impacts on mangroves (Freire 1993). Nonetheless, Materials and Methods
they are considered environments of high diversity, Samples of C. guanhumi (brackish crab)
with a rich and commercially exploited fauna, were collected alive in four mangrove areas – Macau
chiefly related to collection of mollusks and (5º 06’ 56" S - 36º 38’ 08" W), Extremoz (5º 42’ 20”
crustaceans. S - 35º 18’ 26” W), Goianinha (6º 15’ 53’’ S - 35º
The brackish crab Cardisoma guanhumi 12’ 45’’ W), and Canguaretama (06° 13’ 13.4’’ S -
(Latreille 1825) has a semi-terrestrial, nocturnal and 35° 08’ 32.7’’ W), in Rio Grande do Norte State
gregarious life mode and presents a wide geographic (Figure 1). After manual collecting, the animals
distribution (i.e. Florida, Bermudas, Gulf of Mexico, were placed in tanks, separated by collection site,
Antilles, Colombia, and Venezuela up to Brazil). In numbered according to the place they were collected
Brazil, this species can be found from Ceará state and sex, and then preserved in plastic bags at -20º C

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296 M. S. DUARTE ET AL.

for further measurements. The samples were rando- The fluctuating asymmetry (AF) was
mly taken, comprising 107 adult individuals of both calculated as proposed by Palmer & Strobeck
sexes, divided into four populations: Goianinha (G), (1986), disregarding the influence of individual
with 27 individuals (14 ♂ and 13 ♀); Extremoz (E) segment size, in which the mean inverse ponderate
with 29 individuals (13 ♂ and 16 ♀); Macau (M) asymmetry was used: AFi=Ri-Li/(Ri+Li)/2. Where Li
with 31 individuals (21 ♂ and 10 ♀) and Can- corresponds to the mean left segment and Ri the
guaretama (C), with 20 individuals (16 ♂ and 4 ♀). mean right segment.
Dorsal and ventral regions, as well as This method was chosen because of the
segments, were measured in centimeters using a possibility to correct the measurements of each
digital caliper (0.01mm precision). Absolute values individual, once the sample is composed of
for each individual were calculated based on the heterogeneous elements. Initially, the mean of two
mean of two repeated measurements. The data were measures was calculated (left and right side) for each
ordered in tables and further analyzed through sampled individual and then a Multivariate Analysis
statistical packages. All the measures (Figure 2) of Variance (MANOVA) test was applied on
were obtained with replicates by the same person repeated measures about the response vector
and the following nomenclature was used to identify comparing different populations. The response
the measures in this study: vector comprised the width and length
measurements (RCW, LCW, RCJW, LCJW, RCDL,
Dorsal Region LCDL, RMC, LMC, RMW, LMW, RML, LML)
CTL Carapace total length taken from the pair of pereiopods (1st, 2nd, 3rd and
CTW Carapace total width 4th). The first letters used in the measurements
CFW Carapace final width abbreviations above (R and L) correspond to right
ECL Eye cavity length and left side, respectively.
IOD Inter orbital distance The effects of population (G, E, MA and C),
MPW Medial peduncle width sex (male and female) and segment (CW, CJW,
CDL, MC, MW and ML) and side (R and L) as well
Ventral Region
as the mean profile of measurements per population,
ATL Abdomen total length
ASW Abdomen first suture width
sex and side were also analyzed. Complementary
tests were incorporated in order to identify the
Pereiopods morphometrical differences among the several
CW First movable chela width groups studied. Variance analyses and the use of
CJW Width of the joining between the Pearson’s correlation coefficient, besides
propodus and the movable chela thigh discriminating function and main component
CDL Chela dactylus length analyses were employed to detect variation in
RMC/LMC Right and left major movable chela quantitative characters and possible distinctive
MW 2nd, 3rd and 4th pereiopod merus width features among populations by using Statistica V7
ML 2nd, 3rd and 4th pereiopod merus length (Statsoft).

Figure 1. Collection sites of Cardisoma guanhumi in Rio Grande do Norte State, Brazil.

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Interpopulational morphological analyses and fluctuating asymmetry in Cardisoma guanhumi. 297

Figure 2. Corporal measures analyzed in C. guanhumi populations. Dorsal region. (A-I) Eye cavity length; (A-II) Inter orbital
distance; (A-III) Medial peduncle width; (B-IV) Carapace total length; (B-V) Carapace total width; (B-VI) Carapace final width.
Major chela. (C-VII) Width of the joining between the propodus and the movable chela thigh; (C-VIII) Chela dactylus length; (C-IX)
First movable chela width. Pereiopod. (D-X) 2nd, 3rd and 4th pereiopod merus width; (D-XI) Merus length. Ventral region. (E-XII)
Abdomen total length; (E-XIII) Abdomen first suture width. Bars = 2 cm.

Results
Fluctuating asymmetry analysis was on the variables regarding individual’s morphology.
performed in a sample of 107 individuals of brackish Information about dorsal and ventral regions was
crab, distributed in four populations, from which two taken as demonstrated in Tables II and III.
measurements were taken in each studied segment Based on the results, a significant effect of
and side, resulting in a mean value. Aiming to population over all measurements can be suggested,
measure the effects of population, sex, segment and revealing that populations of C. guanhumi present
side, besides estimate the mean measurements for distinctive morphological characteristics.
each population and sex, a MANOVA model with The frequencies of the major movable chela
repeated measurements was adjusted to the sample at in each body side and sex were established, as
issue, which results can be observed in Table I. shown in Table IV.
A significant effect of population and Table IV shows that right-sided major
segment and their interaction was observed chelas were present in 51.40% of crabs. In females,
concerning the four first pairs of pereiopods. In this structure occurred in a frequency of 45.7% on
addition, sex and its interaction with segment and the left side. There was a regular distribution of this
population were also significant in the 1st pair of feature in females, giving rise to a proportion of 1:1
pereiopods. In all cases, side presented no significant related to the body side, whereas in males this
effects, thus characterizing fluctuating asymmetry in relation was 1:1.52. Some individuals presented
the pairs of pereiopods. symmetrical chelas, more frequent in females
Effect and factor-interaction tests by (10.8%), but this condition was not observed in
MANOVA, with repeated measures, were performed Canguaretama sample.

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298 M. S. DUARTE ET AL.

Table I. Effect tests and interaction between factors, through MANOVA, with repeated measures, regarding
1st, 2nd, 3rd and 4th pereiopod pair.
Factor Effect Experimental Error
Pereiopods Factor F test P value *
DF MS DF MS
Population 3 5.446 96 0.241 22.535 0.000
Sex 1 2.211 96 0.241 9.151 0.003
1st pair Segment 2 158.292 192 0.036 4343.611 0.000
Population x Segment 6 0.737 192 0.036 20.231 0.000
Sex x Segment 2 0.709 192 0.036 19.476 0.000
Population 3 1.583 95 0.136 11.631 0.000
2nd pair Segment 1 474.191 95 0.116 4058.323 0.000
Population x Segment 3 1.132 95 0.116 9.696 0.000
Population 3 2.124 94 0.128 16.496 0.000
Segment 1 657.078 94 0.064 10126.206 0.000
3rd pair
Population x Segment 3 0.989 94 0.064 15.242 0.000
Sex x Segment 1 0.263 94 0.064 4.064 0.046
Population 3 2.039 85 0.136 14.912 0.000
th Segment 1 518.278 85 0.077 6678.164 0.000
4 pair
Population x Segment 3 1.071 85 0.077 13.800 0.000
DF = Degrees of freedom; MS = Mean square; (*) significant if P <0.05.

Table II. Effect and interaction test between factors by MANOVA, with repeated measures, regarding the
dorsal region.
Factor Effect Experimental Error
Measure Factors F test P value*
DF MS DF MS
Population 3 4.234 99 0.212 19.931 0.000
CTL Sex 2 0.101 99 0.212 0.476 0.491
Population x Sex 2 0.014 99 0.212 0.067 0.976
Population 3 4.642 99 0.251 18.478 0.000
CTW Sex 1 0.198 99 0.251 0.789 0.376
Population x Sex 3 0.042 99 0.251 0.169 0.916
Population 3 0.607 99 0.035 17.221 0.000
CFW Sex 1 0.202 99 0.035 5.742 0.018
Population x Sex 3 0.013 99 0.035 0.378 0.768
Population 3 2.854 99 0.134 21.200 0.000
ECL Sex 1 0.019 99 0.134 0.147 0.702
Population x Sex 3 0.020 99 0.134 0.152 0.928
Population 3 0.242 99 0.017 13.844 0.000
IOD Sex 1 0.004 99 0.017 0.248 0.619
Population x Sex 3 0.002 99 0.017 0.169 0.916
Population 3 0.005 99 0.000 15.423 0.000
MPW Sex 1 0.000 99 0.000 0.080 0.776
Population x Sex 3 0.000 99 0.000 1.264 0.290
DF = Degrees of freedom; MS = Mean square; (*) significant if P <0.05. CTL = Carapace total length; CTW = Carapace total
width; CFW = Carapace final width; ECL = Eye cavity length; IOD = inter orbital distance; MPW = Medial peduncle width.

Table III. Interaction tests between population and sex by MANOVA regarding body ventral region.
Factor Effect Experimental Error F test P value*
Measure Factor
DF MS DF MS
Population 3 1.489 99 0.100 14.827 0.000
ATL Sex 1 0.876 99 0.100 8.728 0.003
Population x Sex 3 0.006 99 0.100 0.068 0.976
Population 3 1.985 99 0.171 11.582 0.000
ASW Sex 1 41.378 99 0.171 241.427 0.000
Population x Sex 3 0.424 99 0.171 2.475 0.065
DF = Degrees of freedom; MS = Mean square; (*) significant if P<0.05. ATL = Abdomen total length; ASW =Abdomen first suture
width.

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Interpopulational morphological analyses and fluctuating asymmetry in Cardisoma guanhumi. 299

The correlation coefficient between the lower, it would be related to macroregion I

measures taken from dorsal (CTL, CTW, CFW, (Goianinha and Canguaretama).
ECL, IOD and MPW) and ventral (ATL and ASW)
regions of C. guanhumi was very high, indicating a
correlation among all the variables. Table V shows
significant correlation values among all the
variables, except for first suture abdomen width,
which presented a lower correlation than the others,
however also significant.
This result suggests that it is possible to
work with a single factor to infer the whole variance
pattern. To test this hypothesis, a factorial analysis
was performed, showing that the first seven features
can properly represent the variables. In this case, the
variable abdomen first suture should be excluded,
because it is influenced by the sex of individual. Figure 3. Morphological interpopulation analysis of the
The communalities elicited from the carapace total length.
variables showed the amount of the original variable
behavior that is shared as a resulting factor behavior,
named size. 97.4% of the eye cavity length variable
behavior is shared with the elicited factor. Thus, the
behavior of the elicited single factor represents
90.6% of the behavior of the seven original variables
(Table VI).

Main component analyses

The largest individuals were observed in
Extremoz sample and the smallest ones in
Goianinha. Figures 3 and 4 show the pattern of
variables separated according to the collection site.
There was a wider variation range in individuals
from Canguaretama, virtually in all variables, except
for abdomen first suture width. The results suggest Figure 4. Morphological interpopulation analysis of the eye
that individuals from Goianinha and Canguaretama cavity length.
compose a single group (macroregion I), without
significant differences among them. Individuals Discussion
from Macau and Extremoz would comprise another The exoskeleton of crabs provides a rich
group (macroregion II). source of biological information. Recent studies
Such differentiation allowed us to define a have detected extensive phenotypic variability in
discriminating pattern between the macroregions. shore crabs within relatively restricted geographical
The specimens from macroregion I were smaller areas (Bentley et al. 2002, Brian 2005, Lye et al.
than those in the macroregion II. The discriminating 2005). For example, specimens collected from
model, using all the eight variables, classified locations around the coast of the UK have been
correctly 83.2% of the original cases and 78.5% of found to differ in terms of their morphology (Brian
cases in crossed validation. Using the stepwise et al. 2006).
procedure, we observed that it is not necessary to The male crab uses the dominant chela as a
use all the original variables. A model using just the sexual ornament and as a weapon, in addition to
eye cavity length variable is able to classify correctly being a feeding structure (Mariappan et al. 2000).
82.2% of the original cases and 82.2% of cases in The lack of a predisposition to the occurrence of
crossed validation. Thus, the origin of an individual, major chela in a specific body side in C. guanhumi,
related to its macroregion, can be predicted by for both sexes, differs from previous reports on the
measurements of the eye cavity length in order to crab Carcinus maenas, which showed a preferential
calculate the discriminant function values. If this occurrence of right-sided chelas (Sneddon &
value is higher than -0.0925, the specimen would Swaddle 1999). These data show that the presence of
belong to macroregion II (Extremoz and Macau); if a highly developed chela in C. guanhumi constitutes

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300 M. S. DUARTE ET AL.

Table IV. Distribution of major chela frequency in relation to sex in Cardisoma guanhumi.
Sex
Males % Females % Total %
Majorchela
Right side 35 57.4 20 43.5 55 51.3
Left side 23 37.7 21 45.7 44 41.2
Symmetrical 3 4.9 5 10.8 8 7.5
Total 61 100.0 46 100.0 107 100.0

Table V. Pearson’s correlation coefficient for all sampled elements.

Carapace Carapace Carapace Eye Inter Medial Abdomen Abdomen
total total final cavity orbital peduncle total first suture
length width width length distance width length width
Carapace total Coef 1 0.972 0.926 0.987 0.861 0.893 0.916 0.365
length Sig. . 0.000 0.000 0.000 0.000 0.000 0.000 0.000
Carapace total Coef 0.972 1 0.902 0.975 0.859 0.877 0.883 0.340
width Sig. 0.000 . 0.000 0.000 0.000 0.000 0.000 0.000
Carapace final Coef 0.926 0.902 1 0.925 0.809 0.838 0.96 0.605
width Sig. 0.000 0.000 . 0.000 0.000 0.000 0.000 0.000
Eye cavity Coef 0.987 0.975 0.925 1 0.877 0.89 0.915 0.391
length Sig. 0.000 0.000 0.000 . 0.000 0.000 0.000 0.000
Inter orbital Coef 0.861 0.859 0.809 0.877 1 0.783 0.808 0.327
distance Sig. 0.000 0.000 0.000 0.000 . 0.000 0.000 0.000
Medial Coef 0.893 0.877 0.838 0.89 0.783 1 0.821 0.372
peduncle width Sig. 0.000 0.000 0.000 0.000 0.000 . 0.000 0.000
Abdomen total Coef 0.916 0.883 0.96 0.915 0.808 0.821 1 0.625
length Sig. 0.000 0.000 0.000 0.000 0.000 0.000 . 0.000
Abdomen first Coef 0.365 0.34 0.605 0.391 0.327 0.372 0.625 1
suture width Sig. 0.000 0.000 0.000 0.000 0.000 0.000 0.000 .

Table VI. Communality values elicited from C. guanhumi interpopulational samples.

Baselines Initial Elicited communalities Factorial charge
Carapace total length 1.000 0.970 0.985
Carapace total width 1.000 0.944 0.972
Carapace final width 1.000 0.913 0.955
Eye cavity length 1.000 0.974 0.987
Inter orbital distance 1.000 0.808 0.899
Medium peduncle width 1.000 0.838 0.915
Abdomen total length 1.000 0.896 0.947

a sexual dimorphism between males and females, methods to investigate the environmental quality.
but its localization (right or left body side) appears The developmental instability of an organism can be
to neither influence the adaptive coefficient nor the measured according to fluctuating asymmetry,
sexual selection in this species. defined as a random deviation from perfect bilateral
On the other hand, the presence of symmetry of morphological traits, caused by genetic
symmetric chelas in 4.9% of males and in 10.8% and environment disturbances (Møller 1998). This
of females, representing 7.47% of all individuals, procedure has been successfully applied to detect
could be an effect of the genetic factors associated stress in natural populations (Palmer & Strobeck
to development of this structure or to influence of 1986, Leary & Allendorf 1989, Zakharov 1992,
sex-related genes. The lower frequency of this Palmer 1996, Møller 1999).
feature in males might represent a less adaptive The asymmetry observed in the first pair
condition, caused by a lower copulation, feeding or of pereiopods showed a significant sex-related
defense performance (intra and interspecific effect, without any significance of this factor
interactions). over other pereiopods. The evidences of asymmetry
Over the last two decades there has been a observed in the 2nd, 3rd and 4th pairs of pereiopods
growing increase in the use of biological monitoring reveal the interaction between asymmetry

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Interpopulational morphological analyses and fluctuating asymmetry in Cardisoma guanhumi. 301

and population and segment as well as between completely discriminated by free-size independent
both factors, with no significance related to sex. components.
The association between asymmetry magnitude and In the present study, the variation
side was not detected. observed among C. guanhumi populations, allowed
Studies on marine shrimp species have not us to differentiate them into two groups
been concordant when it comes to the association represented by the samples from Goianinha and
between fluctuating asymmetry and sex. Thus, while Canguaretama (macroregion I) and Extremoz and
Maia (2002) evaluating fluctuating asymmetry levels Macau (macroregion II). Different causes can be
in L. schmitti, observed a similar fluctuating involved in the definition of a smaller body
asymmetry index among sexes, Silva (2001) verified size in macroregion I, such as particular
that females of L. vannamei were more asymmetric environmental conditions, pollution or a greater
than males. selective pressure (exploitation) on larger
Under a genetic point of view, high individuals.
inbreeding rates have been associated to The correlation between the differences
developmental instability increases in different in the level of morphological and genetic
organisms (Lacy 1996, Eldridge 1999). In similarity in geographical distinct populations
crustaceans, the relationship between fluctuating of crabs suggests that the phenotypic characteristics
asymmetry and endogamy remains poorly can be related to patterns of genetic variability
understood. Silva (2001) observed that highly (although this relationship is not necessarily
endogamic broodstocks of the captive-reared marine causal) (Brian at al. 2006). So, the present result
shrimp, Litopenaeus vannamei, presented high in C. guanhumi populations could also indicate
asymmetry levels and a higher number of the presence of independent and structured
locomotion appendages. Fluctuating asymmetry was populational stocks, with a possible restricted gene
also found in locomotion appendages within wild flow, despite the absence of visible physical barriers
populations of Litopenaeus schmitti from among them.
Northeastern Brazil (Maia 2002), however in a As for the group classification, two
significantly lower level than that observed for the factors have significantly contributed to the
congeneric species L. vannamei under intense inclusion of an individual in one of the groups,
captive breeding. eye cavity and carapace total length. These
Besides providing information about features can, therefore, be used as practical tools
fluctuating asymmetry, population morphological in the environmental monitoring and analyses
studies like those carried out in crabs of the genus of collected animals concerning biological
Uca (Filho 1990) or fish species (Smith 1973, Kerby conservation programs.
1979) can be helpful to define stocks as well as The populations that presented the higher
taxonomic status. size variation were those from macroregion I
According to Reis (1988), the discriminating (Goianinha and Canguaretama) and, based on these
pattern among populations and species can be data, it is possible to infer that the variation observed
ascertained through discriminating functions using might be related to either overfishing (selective
canonical variables. Another multivariate technique, collection) or intrinsic biological characteristics of
the main component analysis, is more suitable to these populations.
study evolutionary biology issues. Main component Genetic analyses must be carried out in
method basically analyses the relationship between a C. guanhumi populations as a complementary
set of correlated variables, transforming them into a way to identify a possible phenotypic
new set of noncorrelated variables, so-called the correlation with interpopulational genetic differ-
main components. rentiation, contributing to the establishment of
A morphometric discrimination study in exploitation and management policies of this natural
two fish species of the genus Leporinus found a resource.
very high vectorial correlation coefficient among
the elements (Reis et al. 1987). In this study, Acknowledgements
the two samples were regarded as a homogeneous We would like to thank CNPq, CAPES and
group according to main component analysis, Universidade Federal do Rio Grande do Norte for
but the results indicated the presence of two the financial support and for providing the facilities
distinct groups corresponding to L. trifasciatus and proper conditions for the accomplishment of the
and L. macrocephalus. Both species were present survey.

Pan-American Journal of Aquatic Sciences (2008), 3(3): 294-303

302 M. S. DUARTE ET AL.

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Received June 2008

Accepted September 2008
Published online September 2008

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