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Short communication

Comparison between the macroinfauna of urbanizedand protected beaches in Rio de Janeiro State, Brazil

Valeria G. Veloso*, Elen S. Silva, Carlos H.S. Caetano, Ricardo S. Cardoso

Departamento de Ciencias Naturais, Universidade Federal do Estado do Rio de Janeiro (UNIRIO), Av. Pasteur, 458, Sala 411, Urca,

Rio de Janeiro, RJ, CEP 22290-240, Brazil

A R T I C L E I N F O

Article history:

Received 9 July 2004

Received in revised form 16

September 2005

Accepted 20 September 2005

Available online 17 November 2005

Keywords:

Sandy beaches

Macroinfauna

Trampling

Human impact

0006-3207/$ - see front matter � 2005 Elsevidoi:10.1016/j.biocon.2005.09.027

* Corresponding author: Tel.: +55 21 5877809.E-mail address: chcaetano@zipmail.com.

A B S T R A C T

The intertidal macroinfauna of five sandy beaches with similar morphodynamics condi-

tions was studied to compare composition and structure between beaches near urbanized

centers and protected beaches located in Rio de Janeiro State. The beaches were sampled

in winter 1996 and summer 1997 according to a systematic design with stratification. A

total of nine species were identified with the crustaceans being the most abundant and

frequent animals. Species richness showed little variation between beaches while density

of some species such as Emerita brasiliensis (Crustacea) and Phaleria testacea (Insecta) were

lower at urbanized beaches when compared to protected ones. At the most urbanized

beaches, Pseudorchestoidea brasiliensis was absent during both sampling periods. The neg-

ative relationships between human recreational activities (e.g., trampling) and density of

macroinfaunal species was recently gained support and in Barra da Tijuca beach these

relation is very clear. In the developed and most-visited sector, Barra (Alvorada), the

amphipod Pseudorchestoidea brasiliensis was never collected whereas in the protected por-

tion, Barra (Reserva), the same species occurred in high densities. Thus, our results sug-

gested that the amphipod Pseudorchestoidea brasiliensis is more vulnerable to trampling

than are other species.

� 2005 Elsevier Ltd. All rights reserved.

1. Introduction

In recent decades, studies of beaches have focused on under-

standing the influence of physical factors on communities

(McLachlan, 1983, 1990; Defeo et al., 1992; McArdle and

McLachlan, 1992; Jaramillo et al., 1993; Borzone et al., 1996;

Barros et al., 2001) and populations (Dugan et al., 1994; Gomez

and Defeo, 1999; Defeo et al., 2001; Cardoso et al., 2003). For

most species, seasonal changes in population density are nor-

mal, and are related to many factors such as reproductive

dynamics and changes in abiotic factors (Souza and Gianuca,

er Ltd. All rights reservedbr (C.H.S. Caetano).

1995; Cardoso and Veloso, 1996, 2003; Veloso and Cardoso,

1999; Fonseca et al., 2000). According to the swash exclusion

hypothesis (SEH), swash condition is the main factor control-

ling the intertidal macrofauna communities (McLachlan et al.,

1993, 1995). This hypothesis states that species diversity, total

abundance, biomass increase from reflective towards dissipa-

tive beaches and was corroborated by many subsequent

works (Defeo et al., 1992; McLachlan et al., 1993; Borzone

et al., 1996; Veloso et al., 2003). Despite the increase in rich-

ness, diversity and biomass towards dissipative beaches, as

suggested by the swash exclusion hypothesis, many species

.

B I O L O G I C A L C O N S E R VAT I O N 1 2 7 ( 2 0 0 6 ) 5 1 0 –5 1 5 511

of sandy beaches can tolerate a wide spectrum of morphody-

namic variation, maintaining abundant populations in both

reflective and dissipative beaches.

Hydrodynamics and sediment remobilization continu-

ously alter beach profiles. Because their communities are

well adapted to these dynamics, the beaches do not suffer

significantly when small changes in the beach profiles occur.

Therefore, these ecosystems are not very vulnerable either

to natural changes or to human activities (Jaramillo et al.,

1996).

Pollution, exploitation of natural resources, and mainly

erosion are the most common problems in beach ecosys-

tems (McIntyre, 1995). Although urbanization and tourism

are increasing worldwide, studies on modifications caused

by landfills, recreation and cleaning are still rare. The dam-

age caused by large numbers of people trampling dune veg-

etation is better documented (Hosier and Eaton, 1980;

Brown and McLachlan, 1990; Rickard et al., 1994; Watson

et al., 1996). The first investigations of the effects of recre-

ation and trampling on the intertidal macroinfauna gave

conflicting results. Jaramillo et al. (1996), comparing a free

public access trampled area to a restricted public area on

the Chilean coast over a two-month period, found no influ-

ence of trampling. Contrariwise, trampling has been indi-

cated as the factor responsible for the disappearance of

amphipods from highly frequented beaches in Poland

(Welawski et al., 2000). Experiments done in South Africa

indicate that some species resist the impacts of trampling

better than others (Moffet et al., 1998). Barros (2001) com-

pared the numbers of burrows of ghost crabs, Ocypode cordi-

mana, between urban and non- urban beaches and verifies

that urban beaches presents numbers significantly lower

of burrows than non-urban ones at high shore levels. The

lack of information regarding the long-term effects of tram-

pling on the macroinfauna and the conflicting results so far

have revealed the importance of this subject and have

hinted that the consequences of trampling might be

irreversible.

The present study analyses the changes in the composi-

tion and structure of the macroinfauna in crowded urbanized

beaches in Rio de Janeiro City, Brazil. Comparisons between

beaches near urbanized centers and protected beaches were

made, and potential influence of human recreational activi-

ties such trampling was evaluated.

2. Study area

Five beaches located in the state of Rio de Janeiro, Brazil, were

selected for this study: Copacabana, Ipanema, Sao Conrado,

Barra da Tijuca and Grumari. Barra da Tijuca Beach presents

an area of high visitation frequency and a protected area

(environmental preservation area), respectively, named Barra

(Alvorada) and Barra (Reserva), that was independently

treated.

2.1. Copacabana and Ipanema

These beaches border crowded neighborhoods near the city

center of Rio de Janeiro, where many restaurants, bars and

hotels are located. In these areas, efficient public transporta-

tion provides easy access for visitors from the suburbs; there

are showers and toilets, and many peddlers. The beaches are

frequented year-round. The illumination enables people to go

to those beaches even at night, when they can practice foot-

ball or volleyball.

2.2. Barra (Alvorada) and Sao Conrado

Barra da Tjuca Beach is 18 km long. The urbanized first

6-km stretch is interrupted by a protected area (about

8 km), and the last 4 km are also urbanized. Barra (Alvorada)

constitute the first urbanized portion and together with Sao

Conrado beach are located farther from the urban center,

and have similar facilities to Copacabana and Ipanema.

Although ample infrastructure is provided, Barra (Alvorada)

and Sao Conrado beaches are less crowded in winter, espe-

cially this last beach which is a famous landing site for hang

gliders.

2.3. Barra (Reserva) and Grumari

Barra (Reserva) Beach is located on the same beach arc as Bar-

ra (Alvorada) Beach, but because is part of a protected area,

any kind of construction is forbidden. Also, there is no public

transportation to these beaches. Grumari Beach is 4 km long

and is an environmental preservation area.

3. Materials and methods

3.1. Sampling and laboratory procedures

The beaches were sampled in winter 1996 and summer

1997, during low spring tide. At each beach, two sectors

were established from the lower limit of the swash zone

to above the drift line. Next, five transects were marked

in each sector and 10 equally spaced sampling strata were

marked per transect, according to a systematic design with

stratification, defined: the first below the swash line (N1 –

30 cm water layer), the second last (N9) on the drift line,

and the last (N10) 10 m above the drift line. This systematic

design is necessary to describe the macroinfauna communi-

ties because of conspicuous and intensively recorded

biological zonation (Defeo et al., 1992; Jaramillo et al.,

1993; Borzone et al., 1996; James and Fairweather, 1996).

One sample was taken with a 0.04 m2 quadrat to a depth

of 25 cm from each sampling strata. The collected sediment

was washed through a 0.71 mm sieve, and the retained

material was taken to the laboratory, where the organisms

were sorted by species, counted and fixed in 5%

formaldehyde.

3.2. Physical characterization

Sediment samples for particle size analysis were taken with a

plastic corer of 3.5 cm diameter to a depth of 10 cm at strata

10 (supra), 6 (middle) and 2 (waterline). Samples were oven-

dried at 70 �C and passed through a series of sieves of �2.5,

�2.0, �1.0, 0.0, 0.5, 1.0, 1.5, 2.0, 2.5, 3.0 and 4.0 phi (phi = �log2mm) in order to determine mean particle size (Folk and Ward,

1957). Results were expressed in mm.

512 B I O L O G I C A L C O N S E R VAT I O N 1 2 7 ( 2 0 0 6 ) 5 1 0 –5 1 5

The beach-face slope of each transect was measured by

the height difference (Emery, 1961) between the drift line

and the waterline. Dean�s dimensionless parameter (X) (Short

andWright, 1983) was calculated for each beach as a measure

of its morphodynamic state: X = Hb/Ws. T, where Hb is the

breaker height in cm, Ws the sand settling velocity in cm s�1

(obtained from particle size and Gibbs et al., 1971) and T the

wave period in seconds.

3.3. Statistical analysis

The number of people passing the drift line was counted

every half hour (between 10.00 and 15.00) in 6 h of the peak

of abundance, to verify the trampling intensity. The number

of people was counted in a randomly chosen block of 50 m2.

ANOVA was employed to verify the trampling difference

(mean abundance related to the 6 h peak) between the

beaches.

The two-way ANOVAwas used to verify differences in the

macroinfauna density between beaches and seasons. The Tu-

key–Kramermultiple comparison test was used a posteriori to

assess significant differences. Density values were log(x + 1)

transformed to fulfill the assumptions of normality and hom-

ocedasticity of the data.

4. Results

4.1. Physical characterization

The morphodynamic states of the beaches of Ipanema, Sao

Conrado and Grumari were reflective while Barra (Alvorada)

and Barra (Reserva) were intermediate (Table 1). At Copaca-

bana beach, the morphodynamic state varied according to

the season, being intermediate in winter and reflective in

summer. The beaches classified as intermediate showed val-

ues near the reflective limits (X < 1). During this study, the

granulometry of most beaches varied from 0.37 to 0.50 (med-

ium sand) except at Ipanema beach, in winter, that presented

mean grain size of 0.68 mm (coarse sand).

Table 1 – Characterization of the beaches studied: w = winter;

Beaches Beachextension

(km)

Dean�sparameter

(X)

Intertidalslope (1/m)

M

Copacabana w 6 1.68 23.56

s 0.53 13.79

Ipanema w 5 0.63 9.44

s 0.82 12.69

Sao Conrado w 4 0.84 25.00

s 0.42 13.72

Barra (Alvorada) w 10 2.35 12.90

s 1.36 11.61

Barra (Reserva) w 8 1.15 12.45

s 1.72 11.20

Grumari w 4 0.77 11.38

s 0.72 11.86

Mean visitors = mean number of people in 6 h of the peak of abundance

a Frequency of visitors: 1 = high; 2 = medium; 3 = low.

The beaches of Copacabana and Ipanema are located in

more densely inhabited areas (Table 1) and received the high-

est mean number of visitors. These beaches did not differ sig-

nificantly from the others in regard to the number of visitors,

either in summer or winter (Table 2). The stretches of Barra

(Reserva) and Grumari beaches located within the protected

area were least trampled. Only during summer there was a

significant difference between the number of visitors at Gru-

mari and Barra (Reserva), as indicated by ANOVA. In spite of

being a protected area, Grumari receives more visitors during

summer because of the easy access to facilities and infra-

structure. Although Sao Conrado and Barra (Alvorada) bea-

ches are near a neighborhood area, they have fewer visitors,

mainly during winter, than Copacabana and Ipanema (Table

2).

4.2. Macroinfauna

A total of nine taxa were collected and identified during this

study. Crustaceans were the most abundant and frequent ani-

mals, represented by Emerita brasiliensis, Lepidopa richmondi

(Decapoda), Pseudorchestoidea brasiliensis (Amphipoda) and

Excirolana braziliensis (Isopoda). Molluscs were represented

by the bivalve Donax hanleyanus; and polychaetes by Hemipo-

dus olivieri, Pisionidens indica and unidentified species. At the

most urbanized beaches, Pseudorchestoidea brasiliensis was ab-

sent both in winter and summer. Despite their frequency,

Emerita brasiliensis and Phaleria testacea were present in lower

densities at urbanized beaches when compared to Grumari

and Barra (Reserva). Only one species, Excirolana brasiliensis,

was found at Copacabana, and was absent from the other

urbanized beaches except at Barra (Alvorada) Beach where it

was present in high densities during winter. Donax hanleyanus

and Hemipodus olivieri occurred in almost all the beaches, but

in low densities (Table 3).

The comparison of macroinfauna density between the

beaches and seasons revealed significant differences only be-

tween the beaches (Table 4). The Tukey–Kramer multiple

comparison test showed that more urbanized beaches

s = summer

ean grainsize (mm)

Populationnumber

(per district)

Visitorsabundance(means ± SD)

Frequency ofvisitorsa

0.45 >150,000,000 166.66 ± 20.73 1

0.40 323.50 ± 53.87

0.68 90–100,000,000 154.66 ± 16.68 1

0.45 333.50 ± 44.64

0.41 20–30,000,000 94.33 ± 13.29 1

0.38 203.16 ± 14.03

0.48 120–130,000,000 128.83 ± 20.12 1

0.41 251.33 ± 17.46

0.50 0 6.00 ± 4.00 3

0.43 31.00 ± 10.60

0.37 0 10.50 ± 4.50 2

0.42 93.33 ± 17.96

(persons/50 m2).

Table 2 – ANOVA. Relation between visitors density of beaches studied and results of Tukey-Kramer test analysisundertaken as a posteriori test of multiple comparisons to indicate statistical significance

Summer

F5,30 = 146.91; p < 0.001 Tukey-Kramer multiple comparisons test

Reserva Grumari Sao Conrado Alvorada Copacabana Ipanema

Mean ranks 5.58 9.67 14.28 15.87 17.96 18.25

Non-significance —— —— ——————————————————————————————————

Winter

F5,30 = 216.56; p < 0.001 Tukey-Kramer multiple comparisons test

Reserva Grumari Sao Conrado Alvorada Copacabana Ipanema

Mean ranks 2.54 3.33 9.70 11.36 12.44 13.16

Non-significance ——————————— ——————————————————————————————————

Data transformed to log (x + 1).

Table 3 – Macroinfauna composition and density (ind m–2) at studied beaches

Taxa Ipanema Copacabana Sao Conrado Barra (Alvorada) Barra (Reserva) Grumari

Winter 1996

Emerita brasiliensis 2.5 5.0 22.5 10.0 92.5

Excirolana braziliensis 325.0 100.0 72.5

Pseudorchestoidea brasiliensis 285.0 280.0

Phaleria testacea 60.0

Hemipodus olivieri 2.5 2.5 10.0 35.0 12.5

Lepidopa richmondi

Donax hanleyanus 2.5 2.5 5.0

Polychaeta (unidentified)

Pisionidens indica 7.5 2.5

Species richness 2 4 3 3 5 4

Summer 1997

Emerita brasiliensis 37.5 7.5 52.5 60.0 195.0 1930.0

Excirolana braziliensis 1.0 20.0 180.0 7.5

Pseudorchestoidea brasiliensis 3195.0 260.0

Phaleria testacea 5.0 12.5 62.5 15.0 90.0 17.5

Hemipodus olivieri 5.0 12.5 5.0 30.0

Lepidopa richmondi 5.0

Donax hanleyanus 12.5 2.5

Polychaeta (unidentified) 2.5 7.5

Species richness 6 5 3 4 5 4

B I O L O G I C A L C O N S E R VAT I O N 1 2 7 ( 2 0 0 6 ) 5 1 0 –5 1 5 513

(Copacabana, Ipanema, Sao Conrado) formed a group; Sao

Conrado and Barra (Alvorada) formed a second group, and

the third group was composed by the better-preserved bea-

ches (Barra (Reserva) and Grumari). The two-way ANOVA also

shown a significant interaction between the factors (beach

and season).

5. Discussion

The studied beaches have similar morphodynamic condi-

tions, both granulometric and Dean�s parameter values

showed a low variation (Table 1). Therefore, there is no reason

to suppose that environmental conditions are strong enough

to be responsible for the observed differences in macroinfa-

una composition. Pseudorchestoidea brasiliensis and Excirolana

braziliensis were found over a morphodynamic gradient,

including beaches with granulometric differences, as re-

corded by Veloso et al. (2003). Those results indicate their high

adaptative ability to different environmental conditions. As

shown in Veloso et al. (2003), both species occurs from very

reflective beaches to intermediate ones (Dean�s parameter

0.30–3.12; granulometry 0.19–1.28 mm), supporting the idea

that morphodynamics are not responsible for the changes ob-

served in this study.

The absence of past data on these environments consti-

tuted an insurmountable problem for the evaluation of the

urbanized beaches, and it was not possible to analyse the real

change. On the other hand, the absence of some species and

the drastic decrease in the numbers of individuals of other

species cannot be ignored. At Barra da Tijuca Beach the

changes caused by human activities are very clear. In

the developed and most-visited sector, Barra (Alvorada), the

amphipod Pseudorchestoidea brasiliensis was never collected;

whereas in the protected portion, Barra (Reserva), the same

species occurred in high densities.

The results indicate that the amphipod Pseudorchestoidea

brasiliensis is more vulnerable to trampling than are other spe-

cies. Welawski et al. (2000) suggested that the increasing

Table 4 – Two-way ANOVA. Comparison of macroinfauna density between the beaches and seasons (winter and summer)

Effect df Effect MS Effect df Error MS Error F p

Beach 5 6.56 108 0.33 19.70 0.0000

Season 1 1.30 108 0.33 3.92 0.5033

Interaction 5 0.92 108 0.33 2.77 0.0214

Tukey-Kramer multiple comparisons test

Copacabana Ipanema Sao Conrado Alvorada Grumari Reserva

Mean 0.29 0.41 0.58 0.95 1.51 1.64

Non-significance ———————————————————————————

———————————————

—————————————-

Results of Tukey-Kramer test analysis undertaken as a posteriori test of multiple comparisons to indicate statistical significance are also

showed. Data transformed to log (x + 1).

514 B I O L O G I C A L C O N S E R VAT I O N 1 2 7 ( 2 0 0 6 ) 5 1 0 –5 1 5

tourism on the Polish Baltic coast is one of the causes of the

decrease in the population density of the amphipod Talitrus

saltator. Historical data show a decrease in the occurrence of

this amphipod from 1965 to 1995. The areas where tourism

had most increased showed the greatest declines of this spe-

cies. Along 60% of the coast, the authors recorded 100 people

daily trampling a 1-m area. The method of cleaning was also

indicated as an important limiting factor for the occurrence of

this amphipod. The beaches of Copacabana and Ipanema are

visited year-round. During summer, they are overcrowded

and the space between people is minimal.

The swash zone filter-feeders, such as species of the genus

Emerita and Donax, are able to move to the infralittoral zone,

and their larvae have better power of dispersal and a good

chance to recolonize disturbed areas. For those reasons, they

should be less vulnerable to trampling. Our data showed that

these species occur in all the beaches, although in lower den-

sity in the more urbanized than in the most protected areas.

On the other hand, in most midlittoral species of peracarids

with direct development (such the amphipod Pseudorchestoi-

dea brasiliensis and isopod Excirolana braziliensis), the juveniles

are recruited into the same environment as the adult popula-

tion. Other differences includes: a thinner ‘‘carapace’’ than

Emerita and Donax ; and localization in the most-trampled

central midlittoral zone. Different susceptibility was observed

by Moffet et al. (1998), who attempted experimentally to verify

the effects of different intensities of recreational activity on

the survival rate of four species of the macroinfauna. The re-

sults showed a small change in abundance when trampling

occurred at low intensity. However, there was substantial

damage when trampling increased. The most affected species

were those with the more fragile carapaces, such as mysida-

ceans and juvenile bivalves. Contradictory results were ob-

tained by Jaramillo et al. (1996), who observed no significant

changes between a recreational area and an area restricted

to visitors over a two-month period.

Although several experimental studies have attempted to

show lethal effects of trampling and/or recreational use of

the beaches, the chronic effects on aspects of life cycles such

as fecundity rate, recruitment, growth rate, or the length of

ovigerous females are still unknown. The danger that certain

species may be eliminated from these ecosystems is far from

being understood, considering the sparse knowledge about

energy flow within and between the adjacent ecosystems.

Specific studies should be developed to evaluate the composi-

tion and structure of the macroinfauna related to intense hu-

man activity. Such studies will form a fundamental basis for

developing management and preservation policies.

Acknowledgment

We thank all participants in the field work for their valuable

effort. Thanks also to Dr. Janet W. Reid for English language

revision. This study was supported by CNPq (Conselho Nac-

ional de Desenvolvimento Cientıfico e Tecnologico) and FA-

PERJ (Fundacao de Amparo a Pesquisa do Estado do Rio de

Janeiro).

R E F E R E N C E S

Barros, F., 2001. Ghost crabs as a tool for rapid assessment ofhuman impacts on exposed sandy beaches. BiologicalConservation 97, 399–404.

Barros, F., Borzone, C.A., Rosso, S., 2001. Macroinfauna ofsix beaches near Guaratuba bay, southern Brazil.Brazilian Archives of Biology and Technology 44 (4),351–364.

Borzone, C.A., Souza, J.R.B., Soares, A.G., 1996. Morphodynamicinfluence on the structure of inter and subtidal macrofaunalcommunities of subtropical sandy beaches. Revista Chilena deHistoria Natural 69, 565–577.

Brown, A.C., McLachlan, A., 1990. Ecology of Sandy Shores.Elsevier, Amsterdam.

Cardoso, R.S., Veloso, V.G., 1996. Population biology andsecondary production of the sandhopper Pseudorchesto-idea brasiliensis (Amphipoda: Talitridae) at Prainhabeach, Brazil. Marine Ecology Progress Series 142,111–119.

Cardoso, R.S., Veloso, V.G., 2003. Population dynamics andsecondary production of the wedge clam Donax hanleyanus(Bivalvia: Donacidae) on a high-energy, subtropical beach ofBrazil. Marine Biology 142, 153–162.

Cardoso, R.S., Veloso, V.G., Caetano, C.H.S., 2003. Life history ofEmerita brasiliensis (Decapoda: Hippidae) on two beaches withdifferent morphodynamic characteristics. Journal of CoastalResarch 35, 392–401 (Special issue).

Defeo, O., Gomez, J., Lercari, D., 2001. Testing the swash exclusionhypothesis in sandy beach populations: the mole crab Emerita

B I O L O G I C A L C O N S E R VAT I O N 1 2 7 ( 2 0 0 6 ) 5 1 0 –5 1 5 515

brasiliensis in Uruguay. Marine Ecology Progress Series 212,159–170.

Defeo, O., Jaramillo, E., Lyonnet, A., 1992. Communitystructure and intertidal zonation of the macroinfauna inthe Atlantic coast of Uruguay. Journal of Coastal Research8, 830–839.

Dugan, J.E., Hubbard, D.M., Wenner, A.M., 1994. Geographicalvariation in the life history of the sand crab Emerita analoga(Stimpson) on the California coast: relationships toenvironmental variables. Journal of Experimental MarineBiology and Ecology 118, 255–278.

Emery, K.O., 1961. A simple method of measuring beachesprofiles. Limnology and Oceanography 6, 90–93.

Folk, R.L., Ward, W.C., 1957. Brazos River bar, a study insignificance of grain size parameters. Journal of SedimentaryPetrology 27, 3–26.

Fonseca, D.B., Veloso, V.G., Cardoso, R.S., 2000. Growth, mortalityand reproduction of Excirolana braziliensis Richardson, 1912(Isopoda: Cirolanidae) on the Prainha beach, Rio de Janeiro,Brasil. Crustaceana 73, 535–545.

Gibbs, R.J., Matthews, M.D., Link, D.A., 1971. The relationshipbetween sphere size and settling velocity. Journal ofSedimentary Petrology 41, 7–18.

Gomez, J., Defeo, O., 1999. Life history of the sandhopperPseudorchestoidea brasiliensis (Amphipoda) in sandy beacheswith contrasting morphodynamics. Marine Ecology ProgressSeries 182, 209–220.

James, R.J., Fairweather, P.G., 1996. Spatial variation of intertidalmacrofauna on a sandy ocean beach in Australia. Estuarine,Coastal and Shelf Science 43, 81–107.

Jaramillo, E., Contreras, H., Quijon, P., 1996. Macroinfauna andhuman disturbance in a sandy beach of south-central Chile.Revista Chilena de Historia Natural 69, 655–663.

Jaramillo, E., McLachlan, A., Coetzee, P., 1993. Intertidal zonationpatterns of macrofauna over a range of exposed sandybeaches in south-central Chile. Marine Ecology Progress Series101, 105–118.

Hosier, P.E., Eaton, T.E., 1980. The impact of vehicles ondune and grassland vegetation on a south-eastern NorthCarolina barrier beach. Journal of Applied Ecology 17,173–182.

McArdle, S.B., McLachlan, A., 1992. Sand beach ecology: swashfeatures relevant to the macrofauna. Journal of CoastalResearch 8, 398–407.

McIntyre, A.D., 1995. Human impacts on the oceans: the 1990sand beyond. Marine Pollution Bulletin 31, 147–151.

McLachlan, A., 1983. Sandy beach ecology: a review. In: McLachlan,A., Erasmus, T. (Eds.), Sandy Beaches as Ecosystems. W. JunkPublishers, The Netherlands, pp. 321–380.

McLachlan, A., 1990. Dissipative beaches and macrofauna com-munities on exposed intertidal sands. Journal of CoastalResearch 6, 57–71.

McLachlan, A., Jaramillo, E., Donn, T.E., Wessels, F., 1993. Sandybeach macrofauna communities and their control by thephysical environment: a geographical comparison. Journal ofCoastal Research 15, 27–38 (Special issue).

McLachlan, A., Jaramillo, E., Defeo, O., Dugan, J., De Ruyck, A.,Coetzee, P., 1995. Adaptations of bivalves to different beachtypes. Journal of Experimental Marine Biology and Ecology187, 147–160.

Moffet, M.D., McLachlan, A., Winter, P.E.D., De Ruyck, A.M.C.,1998. Impact of trampling on sandy beach macrofauna.Journal of Coastal Conservation 4, 87–90.

Rickard, C.A., McLachlan, A., Kerley, G.I.H., 1994. The effects ofvehicular and pedestrian traffic on dune vegetation in SouthAfrica. Ocean and Coastal Management 23, 225–247.

Short, A.D., Wright, L.D., 1983. Physical variability of sandybeaches. In: McLachlan, A., Erasmus, T. (Eds.), Sandy Beachesas Ecosystems. W. Junk Publishers, The Netherlands, pp. 133–144.

Souza, J.R.B., Gianuca, N.M., 1995. Zonation and seasonal varia-tion of the intertidal macrofauna on a sandy beach of ParanaState, Brazil. Scientia Marina 59, 103–111.

Veloso, V.G., Cardoso, R.S., 1999. Population biology of the molecrab Emerita brasiliensis (Decapoda: Hippidae) at Fora beach,Brazil. Journal of Crustacean Biology 19, 147–153.

Veloso, V.G., Caetano, C.H.S., Cardoso, R.S., 2003. Composition,structure and zonation of intertidal macroinfauna in relationto physical factors in microtidal sandy beaches at Rio deJaneiro State, Brazil. Scientia Marina 67 (4), 393–402.

Watson, J.J., Kerley, G.I.H., McLachlan, A., 1996. A Humanactivity and potential impacts on dune breeding birds in theAlexandria Coastal Dunefield. Landscape Urban Planning 34,315–322.

Welawski, J.M., Stanek, A., Siewert, A., Beer, N., 2000. Thesandhopper (Talitrus saltator, Montagu 1808) on the PolishBaltic Coast. Is a victim of increased tourism? OceanologicalStudies 29, 77–87.