Reproductive Output of Uca Vocator (Herbst, 1804) (Brachyura, Ocypodidae) from Three Subtropical Mangroves in Brazil

Reproductive Output of Uca vocator (Herbst, 1804) (Brachyura, Ocypodidae) from ThreeSubtropical Mangroves in BrazilAuthor(s): Karine Delevati Colpo and Maria Lucia Negreiros-FransozoSource: Crustaceana, Vol. 76, No. 1 (Jan., 2003), pp. 1-11Published by: BRILLStable URL: http://www.jstor.org/stable/20105533 .

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REPRODUCTIVE OUTPUT OF UCA VOCATOR (HERBST, 1804)

(BRACHYURA, OCYPODIDAE) FROM THREE SUBTROPICAL

MANGROVES IN BRAZIL

BY

KARINE DELEVATICOLPO1'2) and MARIA LUCIA NEGREIROS-FRANSOZO1'2) 1 ) NEBECC (Group of Studies on Crustacean Biology, Ecology and Culture)

2) Instituto de Bioci?ncias, Departamento de Zoologia, Campus de Botucatu,

Universidade Estadual Paulista (UNESP), C.P. 510, CEP-18618-000, Botucatu, SP, Brazil

ABSTRACT

Crab fecundity is widely known to vary proportionally to female size, but the female's nutritional

state also has an important effect on egg production. This study evaluates intraspecific variability of

reproductive output by monthly sampling Uca vocator populations from the Itapanha?, Indai?, and

Itamambuca mangroves on the southeastern coast of Brazil. The presence of ovigerous crabs, their

carapace width (CW) and their number of eggs were recorded. Additionally, the productivity of the

mangroves and the content of organic matter of the sediments were analysed in order to estimate food

availability in each locality. Size-specific fecundity relationships were obtained for each population

and compared among the three populations. Ovigerous females from Itamambuca are the largest

and their fecundities are also the highest among the populations studied. These results probably are

associated with the favourable environmental conditions in Itamambuca, as this is a young mangrove

with a high productivity level.

RESUMEN

La fecundidad de los cangrejos es muy variable respecto a la talla de la hembra, pero su

estado nutricional tambi?n puede tener importantes efectos en la producci?n de huevos. Este estudio

estudia la variabilidad intraespec?fica del potencial reproductor de Uca vocator mediante mu?streos

mensuales de poblaciones provenientes de los manglares de Itapanha?, Indai? e Itamambuca, en el

litoral sudeste de Brasil. Se registr? la presencia de hembras ov?geras, su anchura del caparaz?n

(CW) y n?mero de huevos. Asimismo, se analiz? la productividad del manglar y el contenido

en materia org?nica de los sedimentos con el fin de evaluar la disponibilidad del alimento en

cada localidad. La relaci?n talla-fecundidad se obtuvo para cada poblaci?n, compar?ndose las tres

poblaciones. Las hembras ov?geras de Itamambuca son las que tienen mayor talla y fecundidades

m?s elevadas de las tres poblaciones estudiadas. Estos resultados est?n probablemente asociados a

las buenas condiciones ambientales existentes en Itamambuca, que es un manglar joven de elevada

productividad.

? Koninklijke Brill NV, Leiden, 2003

Also available online: www.brill.nl

Crustaceana 76 (1): 1-11



2 KARINE D. COLPO & MARIA L. NEGREIROS-FRANSOZO

INTRODUCTION

Mangroves are highly productive zones from which nutrients and organic matter are exported that supply consuming communities in adjacent estuarine

and coastal regions (Ottman et al., 1965; Macintosh, 1988; Robertson, 1991;

Lee, 1999; Schaeffer-Novelli, 1994). According to Schaeffer-Novelli (1994),

mangrove productivity can be evaluated by structure and maturation degree. The

deposit-feeding ocypodid crabs of the genus Uca are typical representatives of

the mangrove invertebrate fauna along the Brazilian coast. Their feeding and

burrowing activities contribute to the turnover of nutrients in those environments

(Aveline, 1980; Genoni, 1985; Macintosh, 1988).

According to Genoni (1985), food resources may constrain fiddler crab popu

lations. The food availability may regulate growth, reproductive output, and ulti

mately settlement rate and survivorship of both young and adult crabs (Christy,

1978). Their main food resource is organic matter, including the endofauna, spec

imens of which are sorted out from the substrate (Murai et al., 1982). Actual food

supply depends on the productivity of the ecosystem, microbial activity, substrate

texture, and tide action (Twilley et al., 1995; Moura et al., 1998).

Quantification of the egg production, from oog?nesis to actual fecundity, is often

used to assess reproductive potential in brachyuran crabs (e.g., Jones & Simons,

1983; Wenner et al., 1985; Hartnoll & Gould, 1988; Hines, 1989). Fecundity is

widely known to vary according to size, but the nutritional state of parental females

also plays an important role (Rabalais, 1991). The behavioural pattern and mating

system, which both vary considerably among species of fiddler crabs, may also

affect reproductive output in this group (Crane, 1975; Christy & Salmon, 1984).

The objective of this study is to assess intraspecific variability of reproductive

output, by analysing Uca vocator (Herbst, 1804) populations from three mangrove areas along the southeastern coast of Brazil, which have different productivity levels.

MATERIAL AND METHODS

Study sites and crab sampling Fiddler crabs were collected in three different mangroves on the coast of

S?o Paulo State, Brazil: Itapanha? (23?49'14"S 46?09/14//W), Indai? (23?24'51"S

45?03/14//W) and Itamambuca (23?24/43//S 45?01/03//W) (fig. 1).

Samples were taken monthly at each site from August 1999 to July 2000. During

fieldwork, crabs were removed from their burrows with the aid of diving knives.

Each month, a catch effort of 15 min. by two collectors was employed at each site.

The presence of ovigerous crabs was recorded and their proportion in the samples calculated.



REPRODUCTIVE OUTPUT OF UCA VOCATOR 3

Fig. 1. Map of the northern coast of the State of S?o Paulo, indicating sampling sites at Bertioga and

Ubatuba.

Habitat analysis Data on the vegetation of each site were obtained according to C?ntr?n &

Schaeffer-Novelli (1984). Height, basal area, and density of trees per hectare

were measured to estimate the productivity of the mangroves. At each site, three

sediment samples were taken each month from which mean grain sizes were




calculated for each season. The organic matter content of sediment samples was

determined by ash-free dry weighing. Since there have been reports of crude oil

contamination over the study region, analyses of oil content and derived substances

were also carried out using N-hexane to separate oil from sediment.

Laboratory work

Maximum carapace width (CW) and the width of the abdomen at the anterior

margin of the 5th somite (AW) were measured with callipers (? 0.01 mm) for all

ovigerous crabs. The egg masses of all ovigerous females were removed and stored

in 70% ethanol until later analysis. Only females carrying early embryos were used

for fecundity estimates. Each clutch was immersed in a 1 : 100 freshwater solution

of 5-6% sodium hypochlorite (NaCIO) to detach the eggs from the pleopods. This

material was agitated for 5 minutes so as to completely dissociate the egg mass.

Subsamples were then obtained using a plankton splitter, from which egg counts

were obtained under a dissecting microscope.

Statistical analyses The characteristics of the mangrove vegetation were compared by the Kruskal

Wallis non-parametric procedure, complemented by Dunn's test. For each season,

the organic matter content in the sediments was compared among the three sites

using a one-way analysis of variance, complemented by the Tukey test (Sokal &

Rohlf, 1979). Median size (CW) of ovigerous crabs was compared among sites using the

Kruskal-Wallis non-parametric procedure, complemented by Dunn's test (Sokal

& Rohlf, 1979). The mean number of eggs (F) was calculated for each site and compared using

a one-way ANOVA, complemented by the Tukey test (Sokal & Rohlf, 1979).

Size-specific fecundity relationships were obtained for each population using the

number of eggs as the dependent variable, and the power function Y = aZb,

in which carapace width (CW) was the reference dimension (Somers, 1991).

The obtained equations were linearized using log10. The equations from each

population were compared using a covariance analysis complemented by the Tukey test. A significance level of 5% was used for rejection of the null hypothesis in all

statistical tests.

RESULTS

The mangrove characterization of each site is shown in table I. The amount of

organic matter found in the sediments from Itapanhau and Itamambuca was higher




TABLE I Plant features in the mangroves studied. The results are represented in median values. A different

letter indicates a significant difference within one column (Kruskal-Wallis test, P < 0.05)

Site Density (trees/ha) Tree height (m) Tree diameter (cm)

Itapanhau Indai? Itamambuca

3750 a

9375 b 1250 a

7.01a

10.0 a

4.8 b

34.2 b 5.2 a

6.0 a

Itapanhau a B Indai?

Itamambuca

Winter Spring Summer Fall

(Jun-Aug) (Sep-Nov) (Dec-Feb) (Mar-May)

Seasons

Fig. 2. Bar graph showing average values and respective standard deviations of organic matter

contents (%) in sediments from each site. Comparisons were performed among sites within the same

season. Columns sharing the same letter in the same season do not differ significantly (P > 0.05).

(P < 0.05) than at Indai? (fig. 2). Sediments from Itapanhau presented remnants

of oil and derived substances, probably due to the contamination by crude oil that

took place in that region in 1983. There is also a significant traffic of ferry-boats, which discharge residual fuel into the river, resulting in a constant waste flow in

the environment.

The percentage of ovigerous females compared to non-ovigerous adult females

was lower than 30% throughout the study period at the three sites. Ovigerous crabs were found year-round, except in August. Breeding intensity was greater

during the months when temperature was higher (November to April) than in

other seasons (late spring and summer months). Unfortunately, a low number of

ovigerous females was found in Indai? (fig. 3).

Size frequency distributions obtained at each site show that ovigerous crabs

reach a larger size at Itamambuca (fig. 4). Their median size was also higher in that

mangrove area (P < 0.05), compared to crabs sampled at Itapanhau and Indai?



KARINE D. COLPO & MARIA L. NEGREIROS-FRANSOZO

30-1

25

20

15

10

5

o-L

Itamambuca

8 30

ro 25

<D 20 LL w 15

2 10 H ?) 5 -

? o

Indai?

Itapanhau

//////////// Months

Fig. 3. Uca vocator (Herbst, 1804): monthly proportion of ovigerous females relative to non

ovigerous adult females at each site.

\*' \V N?> ?jO rfl, rjfc

^ N?>' <$>' <$,'

Carapace Width ( mm)

Fig. 4. Uca vocator (Herbst, 1804): size frequency distributions (CW) of ovigerous females in each

of the populations studied.




Itapanhau Indai? Itamambuca

Sites

I Min-Max

I I 25%-75%

o Median

Fig. 5. Uca vocator (Herbst, 1804): box-plots comparing the size of ovigerous crabs among the three

populations. Median values in sites sharing the same letter do not differ significantly (P > 0.05).

Table II Uca vocator (Herbst): mean and standard deviation values of fecundity in overlapping size classes

of the three populations

Site Size classes

(mm)

Number of

females

Mean number

of eggs

Standard

deviation

Itapanhau

Indai?

Itamambuca

16-18

18-20

16-18

18-20

16-18

18-20

11497 10 353

19 276 20 480

30 515 26 064

2 934 1726

2 022

3 276 1606

(fig. 5). The smallest ovigerous females were found at Itapanhau and Indai?, their

median sizes did not differ significantly (P > 0.05) (figs. 4, 5).

Table II shows the mean number of eggs counted in similar size classes among

sampled Uca vocator populations. Within a given size class, egg production is

highest at Itamambuca, intermediate at Indai?, and lowest at Itapanhau. Fecundity

relationships and covariance analysis results are presented in table III. It can be

verified that the fecundity of the Itamambuca population is much higher than of

that at Itapanhau or Indai?.

DISCUSSION

Growth and reproduction of invertebrates, and also in fiddler crabs, may be

limited by the availability of food (Genoni, 1985). The supply of food accumulated



? KARINE D. COLPO & MARIA L. NEGREIROS-FRANSOZO

Table III Uca vocator (Herbst): regression constants and results of the ANCOVA analyses of size-specific

fecundity (*P < 0.05; ***P < 0.001)

Ho:b = 3.0

Site N Intercept Slope (i-test) r2 value

Itapanhau 20 0.91 2.71 yes 0.75

Indai? 12 0.55 3.00 yes 0.86 Itamambuca 18 3.70 3.70 no 0.39

ANCOVA results F value 4.51* 15.89***

Tukey test Itapanhau = Indai? ^ Itamambuca

in a mangrove bottom depends on the productivity status of the area, the rate of

litter decomposition, tidal action, and the local characteristics of the sediments

(Schaeffer-Novelli, 1994; Twilley et al., 1995; Reitermajer et al, 1998). Detritus

and sediments can also absorb elements, such as heavy metals. Thus, a simple

quantification of organic matter content does not directly indicate the nutritional

value of the food actually contained. It is also necessary to identify its quality.

Mangroves are sensible and vulnerable environments (C?ntr?n et al., 1981).

Contamination by crude oil has a severe impact on communities and their bio

logical processes, thus reducing habitat productivity (Lopez, 1979; Ponte et al.,

1987; Schaeffer-Novelli & C?ntr?n-Molero, 1994) and probably reducing nutri

tional supply. This may be the case at Itapanhau, where a significant portion of the

organic matter associated with the sediments is not a suitable food resource due to

contamination by crude oil. At Indai?, the low amount of organic matter is prob

ably due to specific hydrological and sedimental characteristics, which favour the

exportation of litter. Itamambuca is a young mangrove with a high productivity and

low human disturbance, hence more favourable to sustain fiddler crab populations.

Taking into account the organic matter content in the sediment as well as other

environmental conditions, like oil pollution, food quality or availability probably differ at the studied sites. Such differences may promote different growth rates

between populations within a relative small geographic area (Hines, 1989). This

may also imply size variation at the onset of sexual maturity (Wenner et al.,

1974). At Itamambuca, where the organic matter content supposedly represents

the available food resource, fiddler crabs grow to a larger size at sexual maturity

(Colpo, 2001). The size at the beginning of reproduction and the maximum size

reached by females markedly influence the reproductive potential of individuals,

since the volume of the gonads, and thus of the brood produced, increase with

body size (Hines, 1982). Conde et al. (1989) and Conde & D?az (1992a, b)




reported this relationship between body size and reproductive effort in connection

with the productivity of mangrove forests in Aratus pisonii (H. Milne Edwards,

1837), a grapsid crab feeding on mangrove leaves. However, environmental factors

influence the absolute egg number (Jensen, 1958). During oog?nesis, there is a

great demand for proteins and lipids (Rabalais, 1991). Huner & Romaire (1979)

and Taylor (1983) reported that the freshwater crayfish, Procambarus clarkii

(Girard, 1852) and Procambarus spiculifer (Le Conte, 1856) reach higher size

and fecundity under favourable nutritional conditions as compared to those in an

unfavourable habitat. As mentioned by Momot (1986) and Huner & Lindqvist

(1991), environmental conditions such as food and water quality also potentially influence egg production in the freshwater crayfish, Orconectes virilis (Hagen,

1870). The fecundity of Uca vocator in Itamambuca was higher than in the other

two populations investigated. Such contrasting reproductive output was not only due to size differences. The brood size produced by females of similar size was

also higher at Itamambuca, which indicates a higher reproductive potential in

this population. Thus, it is suggested that favourable nutritional conditions are

promoting optimal reproductive output in that population.

ACKNOWLEDGEMENTS

Thanks are due to NEBECC co-workers for their kind help during fieldwork

and laboratory procedures. To F?lvio Freir? for drawing the map presented here.

To CNPq for a master science fellowship for the first author and to FAPESP for

financing part of the fieldtrips for the co-author (#94/4878-8; #98/03134-6).

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First received 28 February 2002.

Final version accepted 24 October 2002.



Reproductive Output of Uca Vocator (Herbst, 1804) (Brachyura, Ocypodidae) from Three Subtropical Mangroves in Brazil

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