ORI GIN AL PA PER

Subtropical Brazilian mangroves as a refugeof crab (Decapoda: Brachyura) diversity

Karine Delevati Colpo • Monica Mungai Chacur •

Fernanda Jordao Guimaraes • Maria Lucia Negreiros-Fransozo

Received: 17 November 2010 / Accepted: 18 July 2011 / Published online: 6 August 2011� Springer Science+Business Media B.V. 2011

Abstract This study characterized seven mangrove areas of different sizes, located on

the southeastern Brazilian coast, with respect to their sediment and vegetation features. We

also estimated the richness and composition of the brachyuran assemblages in these

mangroves. Crab, vegetation and sediment data were collected during low tide. The larger

mangroves showed richer and complex forests, and may be more ecologically stable than

smaller mangroves. Twenty-eight species of crabs were recorded, members of nine fam-

ilies: Panopeidae, Pinnotheridae, Gecarcinidae, Grapsidae, Sesarmidae, Varunidae,

Ocypodidae, Ucididae, and Portunidae. Ocypodoidea predominated over Grapsoidea in the

smaller mangrove areas in pioneer stages of forest establishment, whereas the opposite was

recorded for larger and more stable mangroves, where the forest can reach advanced stage

of development. Comprido is a small mangrove, but its crab assemblage was similar to the

larger ones, probably because of other factors not investigated here (e.g., megalopa settling

Electronic supplementary material The online version of this article (doi:10.1007/s10531-011-0125-x)contains supplementary material, which is available to authorized users.

K. D. Colpo (&)Universidade Estadual Paulista, Campus Experimental do Litoral Paulista—UNESP,Sao Vicente, SP 11330-900, Brazile-mail: kacolpo@clp.unesp.br

M. M. ChacurUniversidade Estadual de Mato Grosso do Sul—PB 351—UEMS, Dourados, MS, Brazile-mail: mmchacur@uems.br

F. J. GuimaraesUniversidade Estadual de Santa Cruz—UESC, DCB/Sistemas Aquaticos TropicaisRodovia Ilheus/Itabuna, km 16, Ilheus, BA CEP 45650-000, Brazile-mail: fernandajguimaraes@yahoo.com.br

M. L. Negreiros-FransozoUniversidade Estadual Paulista, Departamento de Zoologia, IBB—UNESP, Botucatu,SP 18618-000, Brazile-mail: mlnf@ibb.unesp.br

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Biodivers Conserv (2011) 20:3239–3250DOI 10.1007/s10531-011-0125-x

rate). This study provided basic knowledge for developing conservation strategies for

vulnerable mangrove ecosystems.

Keywords Brazil � Crab assemblages � Forest features � Sediment �Subtropical mangroves

Introduction

In coastal regions, the mouth of the drainage system (estuary) is surrounded by a coastal

plain with a low slope and tidal influence. In tropical and subtropical regions, mangrove

forests grow on these plains. The halophyte vegetation tolerates the constant variations of

salinity, temperature, moisture, and luminosity (Schaeffer-Novelli et al. 1990).

The water flow of the estuarine stretch of a river and the extent of its coastal plain are

the most important factors that determine the size that a mangrove forest can reach. In the

northern sector of the southeastern coast of Brazil, the coastline shows several bays, which

are mostly limited by bedrock outcrops of the Serra do Mar coastal mountain range

(Villwock et al. 2005). In this sector, the coastal plains and the rivers are smaller, and

consequently the mangrove areas are less extensive. However, in the southern sector, the

bedrock of the Serra do Mar lies about 10–15 km from the coastline. Larger rivers flow

through these wider plains, allowing the establishment of vast mangrove forests (Martin

and Suguio 1976)

The mangrove vegetation contributes to habitat complexity and the diversity of the

mangrove ecosystem (Macintosh et al. 2002). According to Schaeffer-Novelli et al. (1990),

the typical species of the Brazilian mangrove flora are Laguncularia racemosa, Avicenniashaueriana and Rhizophora mangle. Rhizophora is the most vulnerable genus, and its

species grow in mangrove forests in advanced stages of development (Fromard et al. 1998),

which are allowed by less dynamic and frequent disturbances (Semeniuk 1980; Cintron-

Molero and Schaeffer-Novelli 1992).

Mangroves are productive ecosystems that support highly diverse faunas (Twilley et al.

1995), because they offer food, refuges and areas for the reproduction and growth of many

species (Rodrıguez 1987; Bradford et al. 2002; Nordhaus et al. 2006). Brachyuran crabs

and mollusks are the most numerous components of the benthic macrofauna of mangroves

(Jones 1984; Nagelkerken et al. 2008). The majority of mangrove crabs belong to the

Grapsoidea and Ocypodoidea (Schubart et al. 2002). Mangrove crabs have important

impacts on energy flow by converting litter into debris, which can be exported to supply

heterotrophic organisms of the adjacent aquatic communities (Macintosh 1988; Robertson

1991; Micheli 1993; Twilley et al. 1997; Lee 1999; Werry and Lee 2005; Cannicci et al.

2008). Crabs also affect the sediment chemistry and structure, and therefore are considered

ecosystem engineers (Kristensen 2008).

The diversity and richness of macrofauna can be related to the structure of the mangrove

forest, indicating the conservation status of this ecosystem (Macintosh et al. 2002). These

parameters are little known for Brazilian subtropical mangroves, which makes it difficult to

take management decisions or to formulate laws in order to protect this ecosystem. The

aims of this investigation were to: (1) characterize seven mangrove areas of different sizes

located on the southeastern Brazilian coast, with respect to their sediment and vegetation

features. (2) estimate the richness of the brachyuran crab faunas in these mangroves.

The information provided in this study can be used by Brazilian governmental agencies for

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environmental conservation purposes, since mangroves are permanently protected

ecosystems.

Materials and methods

Study area

Seven mangroves in the state of Sao Paulo, Brazil (Fig. 1) were selected for this study. They

are within the same range of latitude, in order to avoid confounding latitudinal effects on the

biological data. Three of these mangroves were Itapanhau (23�4900700S 46�0900700W),

Itaguare (23�4603000S 45�5800800W), and Guaratuba (23�4501300S 45�5304200W). They are

situated in the southern sector of the Sao Paulo coast, in the municipality of Bertioga, where

the coastal plain is 7–8 km wide (Martin and Suguio 1976). The other four mangroves were

Comprido (23�2902200S 45�0905500W), Indaia (23�2405700S 45�0301000W), Itamambuca

(23�2402500S 45�0004700W), and Ubatumirim (23�2001800S 44�5300200W), in Ubatuba

municipality in the northern sector of the Sao Paulo coast, where the coastal plain is narrow.

The software GE Path was used to estimate the area of each mangrove, in order to confirm

the size differences between the southern and northern sectors.

Sampling

The study period extended from January through December 1999. Physical factors such as

temperature and relative humidity were monitored monthly, using a psychrometer.

In each mangrove, an area of about 0.1 ha, a rectangle of 24 m 9 48 m (1,152 m2) was

delimited perpendicular to the river margin and subdivided into 12 contiguous parallel

bands of 6 quadrants (4 m 9 4 m). The size of each quadrant (16 m2) was established

based on the previously measured size of the trees, or a median basal area less than 0.05 of

the total area of the quadrant (c.f. Green 1979). This method was proposed by Schaeffer-

Novelli and Cintron (1990) to evaluate mangrove vegetation structure. The number of

quadrants effectively sampled for vegetation was determined based on the density obtained

Fig. 1 Sao Paulo coast: locations of the seven mangroves studied. Itapanhau, Itaguare, and Guaratuba arelocated in the southern sector of the coast; and Comprido, Indaia, Itamambuca, and Ubatumirim in thenorthern sector

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in previous sampling (Conde and Dıaz 1985). Definitive forest sampling was carried out

once during the investigation period, in each locality. The mangrove species were iden-

tified, and the total density of stems and the relative density of tree species (%) were

estimated for each mangrove. The diameter at breast height (DBH, measured with a

caliper) of each tree was recorded in each quadrant, and the height (measured with a

clinometer) of at least ten of the tallest trees in each site (inside the rectangle) was

recorded, as suggested by Cintron and Schaeffer-Novelli (1984).

In each rectangle, three samples of sediment were collected monthly throughout the

year, and grain size composition, organic-matter and nitrogen contents were analyzed. The

samples were oven-dried at 60�C for 24 h or to constant weight. Next, they were sieved to

determine the percentage of each particle size, to calculate the central tendency (/) of the

grain sizes in the entire sample, according to the Wentworth (1922) scale, using the sieving

method (Bale and Kenny 2005). The proportion of silt and clay was estimated for each

mangrove. The organic-matter and nitrogen contents of the sediment were assessed by the

ash-free dry weight and the Kjeldahl technique (Pompeo 1996), respectively.

The data for sediment and vegetation features were evaluated for similarity using the

Levene test. Data with equal variances were compared among mangroves by one-way

ANOVA, followed by a Student–Newman–Keuls pairwise comparison procedure. Data

with unequal variances were compared by a Kruskal–Wallis analysis, followed by Dunns’

test (Zar 1999).

These aspects of mangrove forests were chosen because according to Fromard et al.

(1998), the total density of trees and relative density of species (trees ha-1), diameter at

breast height (cm), and tree height (m) are important vegetation aspects that provide

information about the structure of mangrove. Also, the sediment characteristics (central

tendency, silt and clay proportion, organic-matter and nitrogen contents) seem to show a

relationship to the mangrove vegetation (Rossi and Mattos 2002). These variables were

used in a non-metric multidimensional scaling (nmMDS), which was carried out with a

Bray-Curtis similarity matrix, to compare the seven mangrove forests (Gotelli and Ellison

2004).

Crabs were sampled monthly at low tide by five people, for 30 min at each site.

Mangroves are complex ecosystems, and the brachyuran species are associated with sev-

eral microhabitats (i.e., different plants and their parts, tide level, sediment types).

Therefore, in order to determine the actual richness of crabs in the mangroves, the col-

lectors used several capture and sampling techniques (e.g., visual counts, manual capture,

digging and sieving of sediment). Specimens were identified based on Melo (1996) and

classified according to Ng et al. (2008). The specimens are deposited in a reference

collection at the Department of Zoology of the Instituto de Biociencias of the Universidade

Estadual Paulista, Campus de Botucatu, state of Sao Paulo, Brazil. The composition of the

crab assemblages was compared among the mangroves by cluster analysis, using the

Jaccard coefficient of similarity that describes how similar the communities are in terms of

shared species (presence/absence) (Gotelli and Ellison 2004). All statistical analyses were

performed at the 5% significance level.

Results

The mangroves are located near each other, and the physical factors affecting them were

similar. Air temperatures ranged from 15.1�C in winter to 27.6�C in summer at all sites.

The relative humidity was always high, over 78% in all mangroves throughout the year.

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Itapanhau was the largest mangrove (853 ha), followed by Guaratuba (299 ha) and

Itaguare (204 ha). The mangroves from Ubatuba were smaller: Comprido about 28.5 ha,

Ubatumirim 18.8 ha, and Indaia 11.5 ha; Itamambuca was the smallest, with 3.5 ha.

The mangroves were composed of three species of trees, L. racemosa, A. shaueriana,and R. mangle. L. racemosa was the predominant species in the Ubatuba mangroves

(Fig. 2a), while in the Bertioga mangroves A. shaueriana and R. mangle were also con-

sistently present. The mangroves of the southern sector (Itapanhau, Itaguare, and Guara-

tuba) showed lower densities of larger-diameter trees, indicating an advanced state of

development. However, in the northern sector, the forest density was higher and the trees

had smaller diameters, except Itamambuca which had the lowest density and the shortest

trees (Fig. 2b, c, d).

Table 1 presents the mean values of the central tendency of grain size (/), sediment

type, median proportions of silt and clay, median values of organic matter (%), and mean

nitrogen content (%) of the sediment in all localities. Very fine sand comprised the sedi-

ments in most mangroves (Itaguare, Guaratuba, Comprido, Indaia, and Ubatumirim). The

sediments from Itapanhau and Itamambuca contained the highest proportions of silt and

clay, so were muddier than the other mangroves. Organic-matter and nitrogen contents in

the mud of Itapanhau and Itamambuca were higher than in the other mangroves.

The nmMDS analysis clustered, at 85% similarity, Itapanhau, Itaguare, and Guaratuba,

which are the largest mangroves in the southern sector. The analysis also grouped the

Comprido, Indaia, and Ubatumirim mangroves of the northern sector. Itamambuca, the

smallest (3.5 ha) and monospecific mangrove was isolated from the others (Fig. 3).

Fig. 2 Vegetation features of seven mangroves on the coast of Sao Paulo: a relative density (%) of eachtree species recorded in the mangroves. b Median values of total tree density per hectare in each mangrove.c Median values of diameter at breast height—DBH (cm) of trees in the mangroves. d Median height of treesin each mangrove (m). Abbreviations for mangroves as in Table 2

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We collected 28 species of brachyuran crabs, members of 15 genera, 9 families, and 5

superfamilies. The superfamilies Xanthoidea, Pinotheroidea, Grapsoidea, Ocypodoidea,

and Portunoidea were recorded, of which Grapsoidea and Ocypodoidea were the most

important. In Xanthoidea, the family Panopeidae (3 species) and in Pinnotheroidea, the

Table 1 Comparison of sediment features (mean ± standard error) among mangroves on the northerncoast of Sao Paulo

Mangroves

Itu Ite Gua Com Ind Ita Uba

Central tendency

(/)

2.7 ± 0.9 3.5 ± 0.1 3.3 ± 0.1 3.5 ± 0.1 3.2 ± 0.5 2.1 ± 0.1 3.2 ± 0.5

ab a a a a bc a

F = 4.23, df = 6,

P = 0.0123

Silt and clay

proportion (%)

67.8 ± 31.9 12.6 ± 3.1 11.1 ± 6.5 10.5 ± 2.5 11.9 ± 21.3 19.4 ± 17.6 10.7 ± 5.6

a b b b b a b

H = 56.8, df = 6,

P \ 0.0001

Organic matter

content (%)

10.5 ± 5.0 2.7 ± 1.6 1.8 ± 1.9 1.5 ± 0.6 2.1 ± 3.0 3.1 ± 3.2 2.3 ± 1.6

a b cd cd bd b b

H = 98.1, df = 6,

P \ 0.0001

Nitrogen content

(%)

6.9 ± 0.9 1.4 ± 0.6 1.0 ± 0.3 0.8 ± 0.2 1.5 ± 0.8 2.9 ± 0.5 1.3 ± 0.4

a c c c c b c

F = 54.5, df = 6,

P \ 0.0001

For data with equal variances, means were compared by one-way Anova (F), and for data with unequal variances, means

were compared by Kruskal–Wallis (H). df degrees of freedom. Values with at least one letter the same did not differ

statistically (P [ 0.05)

Itu Itapanhau, Ite Itaguare, Gua Guaratuba, Com Comprido, Ind Indaia, Ita Itamambuca, Uba ubatumirim

Fig. 3 Non-metric multidimensional scaling (nmMDS) based on aspects of mangrove forest and sedimentcharacteristics. Three sets were determined at 85% similarity (Bray–Curtis coefficient). The first set groupedthe larger mangrove areas of the southern sector of Sao Paulo (Itapanhau, Itaguare, and Guaratuba). Thesecond set linked the small mangroves Comprido, Indaia, and Ubatumirim, in the north sector. Itamambuca,the smallest mangrove, was isolated by the analysis

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family Pinnotheridae (2 species) were recorded. Grapsoidea was represented by 4 families:

Gecarcinidae (1 species), Grapsidae (3 species), Sesarmidae (4 species), and Varunidae

(2 species). In superfamily Ocypodoidea, 2 families were recorded: Ocypodidae (8 species)

and Ucididae (1 species). In Portunoidea, 4 species of Callinectes (Portunidae) were found

(Table 2). More information about geographical distribution, habitat, behavior, and life

cycle of each species in the Appendix 1.

In Itaguare and Guaratuba mangroves, species of all 5 superfamilies were found. In

Indaia and Itamambuca only members of Grapsoidea, Ocypodoidea, and Portunoidea were

recorded. The proportions of species in each superfamily for the seven mangroves are

shown in Table 3. Guaratuba, Comprido, and Ubatumirim had the richest crab faunas, with

a total of 19 brachyuran species (67.8% of the total sampled) from 8 families in each

mangrove. In Itapanhau and Itamambuca, 16 brachyuran species (57.1%) were recorded,

from 8 and 7 families, respectively. The mangroves with the lowest richness of

brachyurans were Itaguare and Indaia, where 15 species (53.6% of the total sampled),

belonging to 8 and 6 families, respectively, were found.

At 60% similarity (Jaccard coefficient), the cluster analysis showed two groups of

mangroves, according to the compositions of their crab assemblages. The first group was

composed of the smallest mangroves in the northern sector (Indaia, Itamambuca and

Ubatumirim). The second group was composed of Itapanhau, Itaguare, Guaratuba

(southern sector) and Comprido (northern) (Fig. 4).

Discussion

Mangrove forests grow on coastal plains associated with estuaries throughout the equa-

torial or humid tropical and subtropical coasts (Por and Dor 1984). The three mangrove

areas studied in the southern sector of the Sao Paulo coast (Itapanhau, Itaguare, and

Guaratuba) are larger than the four mangroves of the northern sector (Comprido, Indaia,

Itamambuca, and Ubatumirim). The areas of these mangroves are constrained by the width

of the coastal plain, which is wide in the south and narrow in the north.

The mangrove ecosystem is considered to be ‘‘land builders’’ (Davis 1940), because

of the slow tidal water movements that allow sediment to accumulate (Alongi 2008). The

sediment type and the local hydrology affect the maintenance and retention of organic

material (Twilley 1985). Muddy sediments with high silt-clay concentrations have a

greater capacity to retain elements in the interstitial spaces than do coarser sandy sed-

iments (Gray 1974; Watling 1988; Snelgrove and Butman 1994; Raffaelli and Hawkins

1996). Therefore, water, chemical elements, organic debris, algae, bacteria, and infaunal

organisms are naturally more abundant in mud substrates. The muddy sediment from

Itapanhau and Itamambuca, which showed the highest proportions of silt and clay, also

showed the highest concentrations of organic matter and nitrogen. In contrast, the sand

sediments from Guaratuba and Comprido, which had smaller proportions of silt and clay,

contained less of these elements. Nevertheless, the differences observed in these sedi-

ment characteristics did not prove useful to discriminate the mangroves of the southern

and northern sectors.

Knowledge of the forest structure of mangroves could provide information about their

age and stages of development (Silva et al. 1991; Fromard et al. 1998; Chen et al. 2009),

degree of ecological stability (Alongi 2008), or post-disturbance recuperation ability

(Menghini 2008). The mangrove forests in the southern sector of Sao Paulo were richer and

the composition of trees showed more equity, since R. mangle and A. shaueriana occurred

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Table 2 List of crab species found in each mangrove studied on the coast of Sao Paulo

Mangroves

Species Itu Ite Gua Com Ind Ita Uba

Xanthoidea

Panopeidae

Eurytium limosum X X X X

Panopeus austrobesus X

Panopeus occidentalis X X X

Pinnotheroidea

Pinnotheridae

Austinixia patagonensis X

Zaops ostreus X X

Grapsoidea

Gecarcinidae

Cardisoma guanhumi X X X X X X X

Grapsidae

Goniopsis cruentata X X X X X X X

Pachygrapsus gracilis X X X

Pachygrapsus transversus X X

Sesarmidae

Aratus pisonii X X X X X X

Armases angustipes X X X X

Armases rubripes X X X X X X X

Sesarma rectum X X X X X X X

Varunidae

Neohelice granulatus X X

Cyclograpsus integer X

Ocypodoidea

Ocypodidae

Uca cumulanta* X X

Uca leptodactylus X X X X X X X

Uca uruguayensis X X X X

Uca burgersi X X X X X

Uca mordax X X X

Uca rapax X X X X X

Uca thayeri X X X X X X X

Uca vocator X X X X X X

Ucididae

Ucides cordatus X X X X X X X

Portunoidea

Portunidae

Callinectes bocourti X X

Callinectes danae X X X X X X X

Callinectes exasperatus X X

Callinectes sapidus X X X X X

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in high proportions together with L. racemosa. These mangroves also showed a low density

of larger-diameter stems. These characteristics suggest forests with greater stability, which

allows them to reach an advanced stage of development (Cintron-Molero and Schaeffer-

Novelli 1992). However, the mangroves of the northern sector showed a dominance of

L. racemosa, a pioneer species (Fromard et al. 1998). The density of trees was higher in

Comprido, Indaia, and Ubatumirim. This feature, together with the slender stems, indicates

young stands or an unstable environment that prevents the establishment of mature forests.

The Itamambuca mangrove was composed by sparse, short and slender individuals of

L. racemosa. The structure of the Itamambuca mangrove indicates a younger stand or less

Table 3 Percentage of brachyuran species in each superfamily, in the seven mangroves

Superfamilies Ita Ite Gua Com Ind Ita Uba

Xanthoidea 12.5 13.3 10.5 10.5 0 0 0

Pinnotheroidea 0 6.6 5.3 0 0 0 5.3

Grapsoidea 37.5 33.5 42.2 42.2 33.3 37.5 42.1

Ocypodoidea 25 33.3 21 36.8 53.4 56.3 47.4

Portunoidea 25 13.3 21.1 10.5 13.3 6.2 5.2

Itu Itapanhau, Ite Itaguare, Gua Guaratuba, Com Comprido, Ind Indaia, Ita Itamambuca, Uba Ubatumirim

Fig. 4 Cluster analyses, using the Jaccard coefficient of similarity (presence/absence of species), showedthat Indaia, Itamambuca, and Ubatumirim shared similar crab assemblages, while Itapanhau, Itaguare,Guaratuba, and Comprido composed a second group (60% similarity, indicated by the vertical line)

Table 2 continued

Mangroves

Species Itu Ite Gua Com Ind Ita Uba

Total of species 16 15 19 19 15 16 19

Itu Itapanhau, Ite Itaguare, Gua Guaratuba, Com Comprido, Ind Indaia, Ita Itamambuca, Uba ubatumirim

* The first record of this fiddler crab species for the coast of Sao Paulo

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stable area. The nmMDS analysis grouped, at 85% similarity, the stable and advanced

mangrove areas of the southern sector (Itapanhau, Itaguare, and Guaratuba). A second

group was constituted by pioneer mangroves in the early stages of establishment

(Comprido, Indaia and Ubatumirim). Finally, the smallest, youngest and probably least

stable Itamambuca mangrove was isolated from the others in the analysis, because of its

distinct features.

Mangroves damp the effects of waves, tides, and shoreline erosion (Mazda et al. 2007).

These ecosystems are constantly influenced by these environmental stressors (Alongi

2008). Our results support the presumption that extensive mangrove areas mitigate more

efficiently the effect of these stressors and may exhibit a high degree of ecological stability,

since the larger Bertioga mangroves (southern sector) showed richer and more complex

forest structures than the smaller Ubatuba mangroves (northern sector).

The availability of energy resources in mangrove ecosystems varies locally according to

the production, consumption, decomposition, and stock of detritus (Rodrıguez 1987;

Robertson 1991; Wafar et al. 1997). Benthic macrofaunal organisms initiate the litter

breakdown process in mangrove systems (Johnstone 1981; Twilley 1985; Robertson and

Daniel 1989; Robertson 1991; Micheli 1993; Silva et al. 1991; Ashton et al. 1999; Lee

1999; Gribsholt et al. 2003; Cannicci et al. 2008). In tropical mangroves, brachyuran crabs

comprise most of the macrofauna (Macintosh 1988), and contribute to litter processing

during their feeding activities (Robertson and Daniel 1989; Lee 1999; Micheli 1993; Werry

and Lee 2005; Cannicci et al. 2008).

Although the distributional ranges of brachyuran species along the Brazilian coast are

known (Melo 1996), no study has examined the richness and diversity of crabs in Brazilian

subtropical estuaries. Richer and more-diverse assemblages of crabs are associated with

mature and preserved mangroves (Macintosh et al. 2002). In the seven mangroves, we

recorded 28 species of brachyuran decapods, indicating a diverse and abundant assemblage

in these systems. The resident biota in these systems shows morphological, physiological,

and behavioral adaptations to tolerate the wide fluctuations in environmental parameters

(e.g., salinity and tides) (Macintosh 1988; Macintosh et al. 2002). Members of five

superfamilies were collected; Grapsoidea (35.7%) and Ocypodoidea (32.2%) crabs were

the dominant component, as in other mangroves worldwide (Por and Dor 1984; Jones

1984; Macintosh 1988; Schubart et al. 2002; Macintosh et al. 2002). Another 32.1% of the

species belonged to Xanthoidea, Pinnotheroidea, and Portunoidea. We also report a range

extension for Uca cumulanta, which was found in the Ubatumirim and Itamambuca

mangroves. This is the first record of this species for the Sao Paulo coast.

The cluster analysis, using a Jaccard coefficient of similarity, indicated that Indaia,

Itamambuca, and Ubatumirim shared similar crab assemblages (more than 60% similarity).

In these mangroves, members of Xanthoidea were not found, and species of Ocypodoidea

were more predominant than Grapsoidea (Table 3). According to Macintosh et al. (2002),

ocypodoids were more abundant than grapsoids in degraded forest areas. Although these

mangroves were not degraded, their degree of instability and pioneer stages of establish-

ment may be similar to that seen in post-disturbance recovering forests (Menghini 2008).

The mangroves of the southern sector (Itapanhau, Itaguare, and Guaratuba) together with

Comprido showed similar brachyuran faunas (60% similarity). Xanthoidea comprised

between 10 and 14% of the crabs, and Grapsoidea predominated over Ocypodoidea.

Although Comprido is a northern mangrove, its crab assemblage was similar to the

Bertioga areas; perhaps some characteristics not investigated in this study may be affecting

the Comprido crab fauna (e.g., local currents and conditions that foster megalopa settling

in this estuary).

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This study suggested that the size of the coastal plain and the mangroves influence the

stability of the forest structure, since large mangroves more efficiently dampen the effects

of waves, tides and shoreline erosion than small ones. This study also provides the first

information about the composition of estuarine brachyuran assemblages on the subtropical

Brazilian coast. These observations provided basic knowledge to develop strategies for

conservation of these vulnerable ecosystems.

Acknowledgments To FAPESP (Fundacao de Amparo a Pesquisa do Estado de Sao Paulo) for financingthis project (#94/4878-8; #95/8520-3; #98/3134-6; #98/15292-5), and to the members of Nebecc for theirhelp during field activities (Alvaro L. D. Reigada, Jelly M. Nakagaki, and Cecilia M. Guerrero Ocampo). Allthe collections of crabs were carried out in compliance with current federal and state laws (IBAMA andICMBio). The authors are grateful to Dr. Janet W. Reid for English improvement in the draft of themanuscript, and to the anonymous reviewers for their helpful comments.

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