Revista Chilena de Historia Natural

69: 565-577, 1996

Morphodynamic influence on the structure

of inter and subtidal macrofaunal communities
of subtropical sandy beaches

Influencia de la morfodinamica sobre la estructura de !as comunidades

intermareales y submareales de la macrofauna de playas arenosas subtropicales

CARLOS A. BORZONE 1, JOSE R.B. SOUZA 2 and ALEXANDRE G. SOARES 1

1 Centro de Estudos do Mar. Universidade Federal do PDUDQi.

Av.Beira Mar s/n. Pontal do SuO. Paranagui. 3255-000. Brazil.

2 Curso de 3yV*UDGXDFDRem Zoo!ogia. E-mail: [email protected].

ABSTRACT

%Hnthic macrofauna communities of ten beaches were studied during a survey in February 1993, in Parani State, southern
Brazil. Sampling stations were distributed along a transect from the limit of vegetation on the upper sub-aerial beach, to nearly
2 m deep on the sub-aqueous beach. Biological and sedimentological samples, slope of the beach and wave height and period
were recorded for each beach. Subtidal samples were collected by scuba diving. Dimensionless fall velocity () and surf scal-
ing (I) parameter modal values ranged from retlecti\'e to dissipative extremes. Sedimentological parameters varied little
among beaches. with average grain size from 2.57 to 2.88 ij Only one intermediate beach had an average grain size of medium
sand (  ij). Intertidal species richness and total abundance had significant correlations with  values. increasing from
reflective to dissipativc beaches. The trend for the subtidal was the reverse, with species richness negatively correlated with ‫گ‬.
Species richness of each station increased along the transect towards offshore stations. This increase was more accentuated at
the reIlective extreme. lnte1tidal species composition and dominance were similar on all beaches, varying little throughout the
morphodynamic spectrum. The subtidal communities had more variable species composition and dominance than the intertidal
cotlllnunities. Zonation patterns obtained from classification and ordination analyses were similar for all beaches, with at least
four faunistic zones recognized. Biological zonation in the subaerial portion of dissipative beaches was fitted to Salvat's
physical zones. with saturation and suhtida! zones sharing species. The homogeneity of sand composition buffered the possible
morphodynamic influence on the intertidal dominance and density patterns, suggesting that these parameters are controlled
by sedimentological characteristics. Subtidal communities seem to be controlled by water and sediment movement, which is
higher in dissipative beaches.
Key words: sandy beaches. subtropical macrofauna. morphodynamic influence.

RESUMEN

Las comunidadcs bentynicas macrofaunales de diez playas fucron estudiadas a partir de un muestreo realizado en febrero de
99. en el estado de Parani. sur del Brasil. Las estaciones de muestreo estuvieron distribuidas a lo largo de un transecto desde
cl limite de la vegetaciyn, en la parte superior de la playa subaerea. hasta aproximadamente 2 m de profundidad, en la porciyn
subacuosa de la playa. Para cada playa se obtuvieron muestras biolygicas y sedimentolygicas, perfil topogrifico, altura y
pHUtodo Jc !as olas. Las muestras infralitorales se colectaron por medio de buceo autynomo. Valores modales de Ios pDUime-
WURVde "dimensionlcss fall velocity") y ‫"( گ‬surf scaling'') variaron del extremo retlectivo a! disipativo. Los pararnetros
sHdimcntolygicos mostraron una pHquexa variaciyn entre playas, con valores medios del tamafio de particulas entre 2,57 y
 ij SRlamHntc una play a intermedia presenty un valor medio correspondiente a arena media ( 1.84 ij). La riqueza de espe-
ciHs y la abundancia total del intennareal estuvo correlacionada significativamente con Ios valores de  aumentando de pla-
yas reflectivas a disipativas. La tendencia en el infralitoral fue inversa, estando la riqueza negativamente correlacionada
con cl parimctro de ‫گ‬La riqueza de especies de cada estacityn aumentya lo largo del transecto en direcciyQ a !as estaciones
infralitoralcs. Estc aumento fue PiV acentuado en el extremo rcflectivo. La composicitin y la dominancia de especies del
intennarcal fue similar en todas !as playas. variando poco a traves del espectro morfodinamico. El patryn de zonaci6n,
obtenido de Ios anilisis de clasificaci6n y de ordenaciyn. resulty similar en todas ODV playas, con el reconocimiento de por lo
WQaQRVcuatro zonas IDXQtVWLFDV En la porciyn subaerea de playas disipativas. la zonacityn biol6gica sigue el esquema de zonas
fisicas de Sal vat. con la zona de saturaciyn comparticndo especies con la zona infralitoral. La homogeneidad de Ios sedimentos
ocultyalguna influencia morfodinimica en Ios patroncs de dominancia y densidad, sugiriendo que !as caracterfsticas sedimen-
tolygicas cstarian controlando estos parametros. Las comunidades infralitorales estarian controladas por el movimiento de agua
y scdimento. que seria mayor en !as playas disipativas.
Palabras clavc: playas arenosas. macrofauna subtropical. intluencia morfodinamica.

(Receivcd I January 1995: accepted 28 July 1996)

566 BORZONE ET AL.

INTRODUCTION beaches spread over the entire morpho-

dynamic spectrum.
Most of the studies dealing with beach com-
munities have been restricted to the subaerial
portion of the beach. Generalizations about MATERIAL AND METHODS
trends and patterns of beach community
structure have been inferred from surveying The Parami Coast stretches for 100 km in a
that narrow portion, in spite of the fact that a NE-SW direction and include several Atlantic
beach is that area which can be actively re- open beaches. Mel Island separates the Para-
worked by wave action and extended, across nagua Bay estuarine system access to the
the nearshore, from depths at the wave base open sea. It has many beaches exposed to the
to the upper limit of swash action (Swart sea, delimited by rocky shore headlands. Les-
1983, Short & Wright 1983). In fact, sampl- te Coastal Plain is 30 km long and comprises
ing the benthos in the subaquous portion only one beach. This beach presents a hetero-
of a beach is not a simple task, requiring in- geneous morphology and sediment charac-
tensive and special sampling effort. The first teristics throughout its extent. The tides are
attempts were undertaken by Day et al. characterized by diurnal inequality and attain
( 1971) and Field ( 1971 ), who used van Veen maximum and minimum amplitudes of ap-
grab in an extensive sampling throughout proximately 2 and 0.5 m respectively (Knop-
the entire continental shelf. More accurate pers et al. 1987). Salinity and temperature of
subtidal sampling was done with by scuba the surf zone water varied from 26 to 33%c
diving and the development of suction and 26 to 30 oc, respectively. These differ-
samplers (for a brief review of the different ences were more related to climatic variation
suction samplers used in benthic studies, during the sampling survey than to the dif-
see also Borzone et al. 1990). Masse (1972), ferent beach locations.
Oliver et al. ( 1980), Christie ( 1976), McLa- Ten beaches of Parana State, southern
chlan et al. (1984), Morin et al. (1985), Fleis- Brazil (Lat. 25° 30-50' S; Long. 48° 15-30'
chack & Freitas ( 1989) and Soares (1992) W) were studied during a survey in February
sampled the subtidal of different sandy and March 1993. Six beaches were located
beaches using this new approach. However, on Me! Island, namely Ponta do Bicho
few studies included the intertidal and the (PPB), Fortaleza (PFZ), Faro! (PFL), Fora
subtidal of a beach. Hill & Hunter (1976), Norte (PFN), Grande (PG) and Fora Sui
Leber ( 1982) and Knott et al. (1983) did an (PFS). Four other beaches were located south
extensive sampling extending from inter- of the island, on the Leste Coastal Plain,
tidal to subtidal. Some general trends were namely Centro (CBM), Atami (ATA), Leste
evident from these studies: species composi- (PLST) and Gaivotas (GAl) (Fig. 1). Nine to
tion of the intertidal was different from the twelve sampling stations were distributed
subtidal, and species richness increased along a transect, from the limit of the vegeta-
offshore with diminishing water move- tion to nearly 2 m depth, in the subidal. Tri-
ment. plicate macrofaunal samples were collected
Over the last ten years, the new morpho- at each station with an iron core of 0.05 m 2
dynamic concept of beach classification de- surface area, taken to a depth of 20 cm. Sand
veloped by Short & Wright (1983) was ex- was sieved through a 0.5 mm mesh and
tensively used by beach ecologists for the de- organisms fixed in 10% formalin, these were
scription of the sandy beach environment. identified to the lowest taxonomic level pos-
Studies searching for patterns of macro- sible. One sand sample was collected at each
benthic communities structure along the re- station for standard mechanical-sieving grain
flective-dissipative beach spectrum are nu- size analysis. Mean and standard deviation
merous, but restricted to the intertidal envi- were computed according to Folk & Ward
ronment. The present contribution is the first (1957) and results expressed as <jl values (<jl =
attempt to analyze the influence of morpho- - log 2 diameter in mm). Slope of the beach,
dynamics on the structure of inter and sub- water table profile, water salinity and tem-
tidal macrobenthic communities of different perature in the surf zone, wave height and
 SUBTROPICAL SANDY BEACH MACROFAUNA 567

~~
1-PPB
2-PFZ
were pooled and a cluster analysis of the
3-PFL
root-root transformed matrix performed,
4-PFN using the Bray-Curtis coefficient and the
5-PG Unweigthed Mean Arithmetic clustering
6-PFS method (Clifford & Stephenson 1975). A
7-CBM Nonmetric Multidimensional Scaling ordina-
8-ATA tion technique was carried out with the same
9-PLST matrix, in order to verify the relationship be-
10-GAI
tween groups. A "mega" cluster analysis was
finally performed with all ten beaches ( 106
48°20'
8
samples), in order to test the robustness of
the biological zonation over the morpho-
dynamic spectrum.
PLC

RESULTS

10
The beaches

The profiles, sediment and morphodynamics

...............
0 5km characteristic of PFN, PG, and PPS represent-
ed the dissipative extreme. These beaches had
Fig. 1: Parami coast, Brazil, showing the location gentle slopes, 1140 - I /59, fine sand, 2.80-
of the beaches. GB: Guaratuba Bay, MI: Me! 2.75 <j>, good sorting, 0.23-0.31, intertidal
Island, PB: Paranagua Bay, PLC: Praia de Leste beach widths from 90 to 116 m, and high
Coastal Plain. PS: Pontal do Sui. values of Q and I parameters (Table 1). PG
Costa de Parana. Brasil. mostrando la Iocalizaci6n de las presented a ridge and runnel morphology on
play as. GB: Bahia de Guaratuba. MI: Ilha do Me!. PB: Bahia the lower shore during the sampling period.
A value of Q = 3.8, corresponding to an in-
de Paranagua. PLC: Planicie costera de Praia de Leste. PS:
Pontal dn Sui.
termediate state, was presented by AT A, but
in combination with a high value of I = 46,
period were measured for each beach. Sub- corresponding to a dissipative surf zone.
tidal samples were collected by scuba diving. This beach, with similar sedimentological
Modal morphodynamic states were com- and morphological features to the former
puted employing the dimensionless fall beaches, streched to the south in an unin-
velocity parameter Q = Hb/W 5 T (Dean terrupted typical bar-trough system that
1973 ), where H 0 is the breaker height, W s is characterizes PLST and GAL On these
the mean fall velocity of the intertidal sand beaches the intertidal sand grain size in-
and T the wave period (see also Short & creased, reaching a medium sand value of
Wright 1983); and the surf-scaling parameter 1.89 <1> on GAl. The intertidal slope ranged
I = a0 2/g tan 2 (Guza & Inman 1975), where from 1/30 to 1/25, and n values from 2.9 to
ab is the breaker amplitude, is incident wave 1.5. The subtidal bar of these beaches was
radian frequency (2n/T), g is acceleration well developed, and the inclusion of these
of gravity and is the intertidal I surfzone features in the profile resulted in gentle
gradient (see also Wright et al. 1982). Mean slopes.
values of H 0 and T were obtained from field As a result, I reached values greater than
observations over one year. 20, allowing some dissipativeness. CBM and
A series of linear regression analyses were PFL beaches had very similar sedimento-
performed between biological and physical logical and morphological characteristics,
variables to assess the relationships between e.g. fine and well sorted sands and subaerial
community structure and morphodynamics. slopes ranging from 1/30 to 1/39. Because of
For the analysis of species zonation patterns, their geographical position, these beaches
abundance data of samples at each station are impacted by low energy refracted waves.
568 BORZONE ET AL.

TABLE I

Physical characteristic of the beaches. IT: intertidal, ST: subtidal

Caracterfsticas fisicas de las playas. IT: intermareal, ST: submareal

Sediment grain characteristic Beach profile Morphodynamics

Beach Width Mean IT Mean ST Mean IT Mean ST .SIT 8 ST Slope Slope Hb w s

(m) (<j>) (<j>) (mm) (mm) (<j>) (<j>) IT ST (cm)

PG 116 2.80 2.77 0.144 0.147 0.24 0.31 49.8 59.2 Ill 5.6 55
PFS 105 2.78 2.75 0.146 0.149 0.27 0.31 40.9 43.4 108 5.3 28
PFN 90 2.80 2.78 0.144 0.145 0.24 0.23 40.3 47.7 134 6.7 50
ATA 105 2.88 2.94 0.136 0.130 0.34 0.37 49.4 69 69 3.8 46
PLST 75 2.57 2.49 0.168 0.178 0.40 0.60 30 50.3 72.5 2.9 26
GAl 54 1.89 2.19 0.270 0.219 0.72 0 68 25.1 49.8 70 1.5 24
PFZ 50 2.81 2.66 0.143 0.158 0 28 0.40 22.5 20 80 4.1 4
CBM 60 2.79 2.85 0.145 0.139 0.24 0.32 32.5 20.9 15 0.7
PFL 60 2.79 2.76 0.145 0.147 0.23 0.28 39.2 30 25 1.2 3
PPB 54 2.65 2.68 0.159 0.156 0.41 0.56 25 13.7 39 1.7 3

PFZ and PPB beaches had a steep intertidal brasiliensis and Excirolana braziliensis
and subtidal profile without a bar system. occurred occasionally, the former only on
The former, with finer sediments, received GAl and the latter on the beaches PLST,
stronger wave action, resulting in an inter- GAl, and CBM.
mediate value of .Q. On this beach longshore The subtidal stations had a more variable
currents associated with the north tidal chan- species composition. However, the clype-
nel determined a very steep subtidal profile asteroid Mellita quinquiesperforata and
(I /20). PPB beach, at the reflective extreme, again the bivalve Donax gemmula dominated
showed a small increase in average particle in numbers and were constant on all of the
size and standard deviation, a steep profile beaches. D. gemmula is one of several spe-
and low values of .Q and L due to weak wave cies that extended its distribution from
action. subtidal to intertidal stations. The mysids
Metamysidopsis neritica and Bowmaniella
Species composition, richness and brasiliensis, and the polychaete Dispio
abundance remanei, occurred on most of the beaches,
with lower abundance (Table 2). CBM beach
Species composition, richness and abundance presented two exclusive species with highest
were analyzed separately for intertidal and abundance, one unidentified tanaidacean and
subtidal portions of the beach, in order to the polychaete Goniada multidens.
allow comparison with the previous litera- Significant correlations between species
ture. richness, mean beach density (number of
Species composition and dominance of individuals m- 2) and total abundance (num-
the intertidal stations were similar on all the ber of individuals m- 1) with physical para-
beaches, varying little throughout the mor- meters were only obtained when dealing
phodynamic spectrum (Table 2). Eight separately with the intertidal and subtidal
beaches were dominated by the polychaete (Table 3). Intertidal species richness had the
Scolelepis squamata. Two isopods, Excirolana best correlation with dimensionless fall
armata and Tholozodiwn rombofrontalis, and velocity (r = 0.75, p < 0.0 12) and surf scaling
the bivalve Donax gemmula, showed high (r = 0.79, p < 0.007), increasing from reflec-
abundances on at least six beaches. The in- tive to dissipative beaches. In contrast, sub-
sects Bledius bonaerensis and B. microcephalus tidal richness was negatively correlated with
appeared on the dissipative beaches. Other the slope and surf scaling (r = -0.67, p < 0.03
common components of the Atlantic beach and r = -0.78, p < 0.01, respectively): spe-
macrofaunal communities, such as Emerita cies richness decreased with increasing surf
 SUBTROPICAL SANDY BEACH MACROFAUNA 569
TABLE 2

Species composition and abundances (individual m·2 ). A: amphipod, B: bivalve, C: coleopteran,

D: decapocl. E: echinoid, G: gastropod, 1: isopod, M: mysidacean, P: polychaete,
Composi~il\n cspecffica y abundancias (individuos m·2 ). A: anffpodo, B: bivalvo. C: coleptero. D: decapodo.
E: cquinoidco. G: gaster6podo. 1: is6podo. M: miscidaceo. P: poliqueto.

INTERTlDAL BEACHES

Species PG PFS PFN ATA PLST GAl PFZ CBM PFL PPB

ScoleleJ'is .Hflliiiii<IW (P) 877 1372 1426 865 1574 79 2146 4212 291 839
Puelche sp.G tAl 4'.16 -15 282 10 () () 265 0 7 139
Emeriw hmsiliensis (DJ () () () () 0 41 0 0 0 0
L.rciro!ww ill'liiill/1 (l) 231 2-1-1 376 358 37 14 107 140 371 170
L\cimftlll/1 /Jm:iliensis (l) 0 () () 0 27 102 0 34 () 0
Tlw!o:odium ro/li/)()fi"imlufis (I) 100 7 428 194 0 () Ill 20 598 336
Donux f!CIIIIiillla tB) 66 !54 10-1 250 71 I 92 37 27 54
Donax hanlcyanus 1. B) 7 0 0 4 7 I 41 0 () 0
Eutonus furciferus ( Pl 27 51 0 36 48 0 24 41 44 0
Hcmipouus olivieri ( PJ 14 7 7 20 () 0 16 0 14 32
Macrochindothea giamhiagi (!) () 0 27 () 0 0 20 0 0 7
Le pi dopa richmondi (D) 20 !I 7 7 7 0 20 20 7 0
l\1cllita quinquiesperforata (E) I-I () 0 0 () 0 () 0 0 0
Dispio remanci tP) () 10 54 17 0 () 20 () 0 0
Orbiniuac ( Pl 0 0 0 0 0 0 0 () 0 54
Bathyporeiapus ruffoi (A) 14 () 41 7 0 0 20 0 7 14
Lioll·lllilllidlu /Jmsi!icnsis (1\1) 7 12 7 7 0 0 7 0 14 7
Bledius /Jonuerensis (C) 78 61 20 41 7 0 0 7 14 ()
Sipuncula () () () 0 0 () 0 () 14 0
Bkdius microcephalus (C) 14 285 () () 0 () 0 0 0 0
Puelche sp.P tA) I-I 0 () 23 27 () 0 0 0 ()
Tivela sp. (8) 0 0 0 0 20 0 0 () 0 ()
Crasinella sp. tB) () 0 7 7 7 () () 0 0 ()
Pinnixa patagoniemis (D) 7 () 0 31 0 0 0 0 0 0

Richness (no. spp.J 21 13 16 15 13 7 14 10 13 10

1\lean Density (x 1000 m·'l 1.16 U8 1.46 1.05 1.33 0.09 2.16 2.26 0.57 13
Ahundance (~ 1000 nY 1 ) 119.3 140.'1 150.7 121.2 100.7 3.56 92.4 125.7 32.2 37.3

Dona.\ gellllllllfa (B) 0 34 186 265 276 239 439 56 48 120

Puelche sp.P tA) I-I 16 54 27 0 () 139 0 36 17
Melliln\'sidofJSis nerilica (!'vi) 7 () 7 34 7 () 109 14 7 14
Macrochiriuotea giambiagi tl) () 7 14 () 7 () 42 () 0 ()
Mcllita quinquiesperfnrata tE) !56 240 34 113 66 24 75 51 10 54
Di,pto remanei (Pi 0 () 31 20 18 27 27 19 16 27
Orhiniidae I Pl () 7 10 () 7 7 43 14 25 41
Sco!~lef'is Sifl/Ciilililll lP) 24 0 () () 17 () 102 14 68 0
Cumacea 0 0 7 7 () () 31 14 58 20
Sipuncula () () 0 0 () () 20 34 7 ()
ParanoiJac ( P) () 41 () 20 () () 0 88 29 0
Lirllt'IIIUIIiella lnasiliensis (l\1) 7 50 7 0 !I 17 14 12 27 10
Hemipouu' olivieri ( P) () 0 0 () 27 32 7 10 7 ')
Ncmertinca () () 0 () () 0 7 70 9 14
Bathyporciapus rulloi (A l 10 () 0 () 0 0 14 0 7 14
Spionidac inuet. ( Pl () () () () 7 () 0 68 14 27
Nepbtys simoni (Pl 7 0 7 7 () 0 17 25 41 10
Olivancillaria vesica tG) 0 () 7 0 0 0 0 () 7
An~ in us hraziliensis ( [) () 0 7 () 17 7 () 0 0 0
Tanaidac.:a () 0 0 () () () 0 2427 14 0
Ogyriues hayi 1DJ 7 7 0 0 () 0 !I 7 14 10
Armandia sp. (P) 0 () 0 () 0 7 0 81 10 14
Synidothea marplatcnsis (I) 7 7 () () 0 0 () 0 0 0
Phoxocepbalopsis sp. (A) 7 () () 0 0 () () 0 14 0
Pinnixa patagoniensis (D) 7 () () 0 () 7 10 0 7 0
Dissnuactylus crinitichelis (D) 10 10 () () 7 0 0 0 0 0
Psioniuens indica (Pl () 7 () 0 29 7 0 0 7 0
Ophiuroiuea 0 34 () 0 7 34 7 () 20 0
Onuphis emercti (P) () () () () 0 () 0 0 20 0
570 BORZONE ET AL.

TABLA 2 (CONTINUED)

INTERTIDAL BEACHES

Species PG PFS PFN ATA PLST GAl PFZ CBM PFL PPB

Caridca 0 14 0 0 0 0 () () () ()
Felaniclla vilardehoana (B) () () () 0 17 () () () 0 ()
Crasinella sp. tBJ 0 0 0 0 37 () 14 0 0 0
Goniada multidens tP) 0 0 () 0 0 0 0 229 0 ()
Strigilla carnaria (B) () () 0 14 0 0 7 44 7 0
TiYela ventricosa (B) 0 0 () 0 27 0 7 7 0 0
Lcpiuopa richmondi (D) 7 7 7 0 7 7 14 7 0 0
Sthcnelais limicola (P) 0 () 0 14 () 0 17 7 0 7
OliYella minuta (G) () 0 0 () () () () 17 7 0
Sigaliurn cirriferum (PJ () 0 0 7 7 7 0 7 7 0

Richness (no. sppJ 13 16 14 15 22 14 30 33 30 17

l\1ean Density (x 1000 m·2 ) 0.22 0.4 0.32 0.44 0.43 0.32 0.84 2.58 0.34 0.29
Abundance (x 1000 m·l) 8.71 2251 14.1 44.47 49.35 35.4 46.62 147.37 8.76 5.72

INTER+ SUB TIDAL

Richness (no. spp) 26 25 25 25 25 20 35 38 36 22

1\lcan Density (x I 000 m 2 ) 0.88 109 112 0.84 0.88 0.19 1.61 2.39 0.49 0.88
Abundance (x I 000 m· 1 ) 128.0 163.4 164.8 165.7 150.1 38.9 139.0 273.1 40.9 43.1

TABLE 3

Correlations between physical and community parameters (n =I 0).

RIT: intertidal species richness, RST: subtidal species richness, RT: total species
richness, D: density , A: linear abundances. Underlined values = p < 0.05
Correlacwncs entre parametros fisicos y de la comunidad (n = 10). RIT: riqueza especffica del intennareal, RST: riqueza
especifica del submarcal. RT: riqueza espccffica total, D: densiuau. A: abunuancia lineal. Valores subrayados = p < 0.05

RIT RST RT DIT DST DT AIT AST AT

Width 0.7-1 tO 01-11 -0.61 (0 063 I -0.32 (Q 365 I -0 02 (0 948) -0.32 (0.372) -0.11 (0.758) 0.68 (0.031) -0.28 (0.419) 0.31 (0 375)
Slope IT 0.72 10019) -0.39 tO 270J -0.05 t0.887) -0.12 (0.7-18) -0.20 (0.571) -0.12 t0.739) 0.54 (0 10-l) -0.23 (0.521) 0.25 t0.472)
Slope ST 0.-16 (0 180) -0.66 (Q.tJ:17 I -0.50 10.139) -040 tO 2-151 -0.44 (0 206) -046 (0 185) 0.31 (0.383) -0.26 (0.459) 0.08 (0.828)
w 0.75(0012) -O.S.l (() I 05 J -0.29 (0.421 ) 0.20 (0 579) -0.42 (0 123) -0 14 (0 969) 0.70 (() 025) -0.44 (0.207) 0.23 (0.516)
s 0.66 ({) OYil -O.?H (() 007) -0.54 (0.104) -0.21 (0557) -0.-17 (() 170) -0.31 (0.380) 0.51 (0.135) -0.38 (0 275) 0.14 (0.695)

zone dissipativeness. Density was not PFN, PG and ATA) had a smaller rate of
correlated with any of the physical para- increase offshore (Fig. 2c). Variation of
meters, and only intertidal abundance (m- 1) density was very similar on these beaches
was significantly correlated with the beach and in PPB and PLST, with a maximum
width (r = 0.68, p < 0.031) and dimension- value of 3,000 individuals m·2 in the inter-
less fall velocity (r = 0.69, p < 0.025). No tidal, around station 4-2 above low water
significant relationship was obtained with level. Seaward from that point there was a
sedimentological parameters, as they varied very pronounced decrease, down to 200
little over the morphodynamic spectrum. individuals m·2 , that was maintained offshore
Species richness and density of each sta- along the transect (Fig. 3a, b). PFZ and CBM
tion along the transect showed characteristic showed higher intertidal values (5,000 and
trends on all beaches. Species richness in- 7,000 individuals m·2 respectively), decreas-
creased towards offshore stations (Fig. 2). ing around low water level and increasing
This increase was more accentuated at the subtidally again (Fig. 3c). The lowest density
reflective extreme (PFZ, CBM and PFL), occurred on GAl and PFL (Fig. 3d). The first
with an abrupt change around low water had a maximum of I ,700 individuals m·2 in-
level (Fig. 2a). Dissipati ve beaches (PFS, tertidally, decreasing around low water level
 SUBTROPICAL SANDY BEACH MACROFAUNA 571

a b
20 20
-PFZ -GAl
+-CBM -+- PLST
16 -o-PFL 16 ·0.· PPB

12 12 *
8

-7 -6 -5 -4 -3 -2 -1 0 1 2 3 4 5 -7 -6 -5 -4 -3 -2 -1 0 1 2 3 4 5

c
Richness 20
-ATA
-+--PG
16 ~--PFS
·<>·-PFN
12

-7 -6 -5 -4 -3 -2 -1 0 1 2 3 4 5
Stations from l W
Fig. 2: Variation of the species richness along the transect. Stations arc numbered from the low tide
water level (0).
Variaci6n de la riqueza de especies a lo largo del transecto. Las estaciones estan enumeradas desde el nivel de marea baja (0).

and the second had lower intertidal density bonaerensis and B. microcephalus, charac-
values. increasing up to 500 individuals m- 2 terized that zone. At CBM, GAl and PLST
subtidally (Fig. 3d). the isopod Excirolana brasiliensis was
restricted to this upper zone, showing more
adaptation to drying than Excirolana armata.
Zonation The latter species characterized the retention
zone, where the sand remains wet during
Zonation patterns obtained from classifica- low tide, on all the beaches. The isopod
tion and ordination analyses were similar for Tholozodium rhombofrontalis occurred in
all beaches. with at least four faunistic zones this zone on all Me! Island beaches and on
discriminated along the whole transect (Fig AT A, showing no relationship between its
4). These biological zones were named (in distribution and morphodynamic character-
part) according to Sal vat's physical scheme istics. The deposit feeder polychaete Euzonus
(Salvat 1964). An upper drying zone was furciferus had an occasional presence in the
present on all the beaches of the coastal plain retention zone, but extended its distribution
and on two beaches of Mel Island, PG and across other zones on AT A. The resurgence
PFS. The presence of two staphylinids, Bledius zone had the greatest abundance of Scolelepis
572 BORZONE ET AL.

(X1000)
a (X1000)
b
4 5
--PG -PPB
··o-PFS
~·ATA 4 -O··PLST 9.
3
-t--PFN
3
f\
2
2

-7 -6 -5 -4 -3 -2 -1 0 1 2 3 4 5 4 ~ ~ ~ -3 ~ ~ 0 1 2 3 4 5

(x1000) r--_ _ _ _ _ _ _ c________, d

8
-PFZ 1800 -GAl o

.. 6
-o·-CBM l 1500 ·-<>··PFL !\
f \
(1.1 '
c:
Gl
0 4
? j

b~
900

2
I
.... o--6
0~~~-~~~~~~~~~~~ 0~~~~~._--~~~~._._~~
7 6 5 4 3 2 1 0 1 2 3 4 5 -7 -6 -5 -4 -3 -2 -1 0 1 2 3 4 5
Stations from LW

Fig. 3: Variation in density (individuals m·2 ) along the transect. Stations are numbered from the low
tide water level (0).
Variaci6n de la densidad (individuos m·2 ) a lo largo del transecto. Las estaciones estan numeradas desde el nivel de marea
haja (()).

squamata on all beaches. This suspension Macrochiridothe giambiagi, Puelche sp. P,

feeder spread its distribution towards off- Hemipodus olivieri, and Lepidopa richmondi.
shore zones, especially at the dissipative ex- The other group of stations, named infra-
treme. Only on PLST did it spread onshore, litoral, corresponded to the surfzone on
overlapping with E. armata in the retention these beaches. The scutellid Mellita quin-
zone. Together with S. squamata, the am- quiesperforata and the polychaete Dispio
phipod Puelche sp. G characterized the remanei are the most representative species
resurgence zone of all Mel Island beaches of this zone.
and AT A. Except for PFN, stations included A similar zonation pattern was formed
in the retention and resurgence zones cluster- when all the stations of the ten beaches were
ed together on all the beaches. Two other analyzed together (Fig. 5). Drying, retention
groups of stations on beaches PFZ, PFS, PG and resurgence zones were very robust, with
and AT A were formed. The first group, satu- 90 to 100% of the stations clustering toge-
ration zone, included stations at the low ther. Saturation zone stations were similar to
water level This zone represented a transition those of the resurgence zone (28%) and
between the two main environments (inter- especially with the infralitoral zone (72% ).
tidal and subtidal). It was characterized by Finally, all the infralitoral stations clustered
the overlapping distributions of S. squamata together (I 00% ), although with the highest
with subtidal species, e.g. Donax gemmula, dissimilarity values.
 SUBTROPICAL SANDY BEACH MACROFAUNA 573
Stations composition
100" DY

~ 90" RT
~

100" RS
I ..----l 10"RT
...--- 28" ST
~

.--
~ 100" ll
72" ST
Lr--C
'----

1.00 0.75 0.50 0.25 0.00

Fip,. -1: Cluster analysis and ordination of stations (MDS) of the beaches. Groups identifi-
ed in the dendrogram are encircled in the MDS. Scale represents dissimilarity.
Analisis de agrupamientos y ordenaci6n de las estaciones (MDS) de las playas. Los grupos identificados en el
dendrograma estan circunscriptos en el MDS. La escala representa disimilitudes.

DISCUSSION This great sedimentary homogeneity is

responsible for the similar species composi-
The definition of the morphodynamic state tion and abundance of the intertidal macro-
of any beach using the "dimensioness fall fauna on all the beaches, with the exception
velocity" formulation is a function of three of GAl, with a coarser intertidal grain size,
parameters: grain size, wave height and wave resulting in a significant reduction of the spe-
period. The combination of different values cies abundance. In spite of this homogeneity,
of these parameters will determine the posi- the intertidal species richness and the linear
tion of a beach on the reflective-dissipative abundance were positively correlated with
morphodynamic spectrum. When the geo- morphodynamic parameters and increased
logical history of a region determines the from reflective to dissipative conditions,
existence of a sediment source of great showing the independence of this relation-
volumes of fine sands, as is the case with the ship with sediment grain characteristics. A
Paranagua Bay area, the morphodynamic correlation between beach face slope and
evolution of the beaches of this area will be linear abundance and diversity of the macro-
an exclusive function of wave action. The fauna was first found by McLachlan et al.
particular geographic positions of the Me! ( 1981 ). Subsequent studies (McLachlan
Island beaches resulted in beaches with the 1990, McLachlan et al. 1993), included in
same sedimentological characteristics, but these positive correlations "omega" values
receiving varied wave action and dispersed and grain size and found this correlation for
throughout the morphodynamic spectrum. different zoogeographical areas.
Guaratuba Bay is the sediment source of Studying the intertidal of Uruguay's
medium sand that occurs at GAl, the only beaches (South Atlantic), Defeo et al. (1992)
medium sand intermediate beach. stated that grain size and slope are among the
574 BORZONE ET AL.
PG
PLST

lO O.'IS 0.5 i;IZ5 LO 0.1 0.1 0.4 0.2

GAl PFS

LO O.'IS 0

ATA
PFI.

LO 0.1 0.4 o.z lO 0.1 o.e 0 ..4 o.z

CBM PFN

lO 0.1 0.1 0.4 o.z

PFZ
PPB

0.1 0.4 0.2 1.0 Q'IS 0.5 0.25 0

Fig. 5: Results of cluster analysis of all the stations sampled. Scale represents dissimilarity.
Resultados del analisis de agrupamiento de todas las estaciones muestreadas. La escala representa disimilitudes.

dominant factors controlling sandy beach with decreasing dissipativeness of a beach,

community structure. They found an increase but also with increasing beach-surf zone
in number of species and total abundance profile. No significant correlation was found
(density) from reflective to dissipative between physical parameters and macro-
beaches, and a negative and exponential fauna! community parameters when the inter-
correlation between grain size and slope with tidal and subtidal data were combined.
density. The trend of richness along whole transects
No correlation was found in the present exhibited an offshore increase, as has been
study between intertidal density and any shown by numerous authors (e.g. McLachlan
physical parameters. However, only one et al. 1984). This increase was different with-
beach had a larger grain size sediment in in the morphodynamic spectrum, being gentle
comparison to the others, with a remarkable on dissipative beaches, and very accentuated
reduction in macrofauna density. This result at the reflective extreme. This is a direct con-
could suggest that density is independent of sequence of a morphodynamic difference in
the morphodynamic state of the beach but wave energy dissipation, that is limited at the
closely related to the sediment particle size. reflective extreme to a narrow zone of strong
Significant correlations in subtidal species breaker action on the beach face. Density
richness were found with the surf scaling patterns were very similar at the dissipative
parameter and the subtidal slope, showing extreme, as a result of the distribution of the
that the number of subtidal species increases dominant species Scolelepis squamata. Simi-
 SUBTROPICAL SANDY BEACH MACROFAUNA 575

Jar high density values were subtidally found tidal species in the intertidal environment.
on CBM, probably due to a more estuarine The list of species (Table 2), that was "a
influence on the macrofauna. GAl was the priori" divided in intertidal and subtidal,
only beach that showed higher subtidal than shows this pattern.
intertidal density values. It is generally accepted that sandy beaches
Zonation patterns of the ten beaches were constitute a unique environment where sand
very similar, with three intertidal zones that and water are in continuous movement, and
maintained their identity throughout the mor- that macrofauna is controlled primarily by
phodynamic spectrum. Each of these zones, physical conditions. The physical environ-
named as drying, retention and resurgence ment has discontinuities and zones, not only
zone, have at least one species with the peak in the intertidal (Salvat 1964, Pollock &
of its distribution centered in one of them. Hummon 1971), but also in the subtidal
This scheme of biological zonation, that fits (Riedl 1964, Riedl & McMaham 1974,
Sal vat's physical one, was extensively studi- Hiscok 1983, Short 1983). The existence of
ed over a year by Souza & Gianuca (1995) physical zones indicates that a beach may be
near AT A beach, showing its persistence divided into different physical subenviron-
over any seasonal fluctuation. The saturation ments that will vary in width and general
zone, identified by several authors as a characteristics throughout the morphody-
unique biological zone (Bally 1983, Wendt namic spectrum. If macrobenthic communi-
& McLachlan 1985) presented great varia- ties are dominated by the physical environ-
tion between the beaches, and lost its identity ment, then their structure may be influenced
when analyzed by "mega" clustering of all by physical zonation. The variation of the
stations. This confirms previous observations width of subtidal biological zones in relation
(Day et al. 1971, Field 1971, Christie 1976, to change in morphodynamic state was in-
McLachlan et al. 1984, McLachlan 1990) directly shown by Knott et al. (1984 ),
that species occupying the saturation zone Fleischack & Freitas ( 1989) and Borzone &
also extend into the sublittoral and may more Gianuca ( 1990), working on beaches in the
appropriately be called surfzone species proximity of jetties. In the present work, the
whose upper limits of distribution are on the name of infralitoral zone included two differ-
lower shore. Therefore, the saturation zone ent subenvironments, surfzone and nearshore.
represents an area that links intertidal and On dissipative beaches, the subtidal sampling
subtidal compartments, allowing biological included only the surfzone, well developed
interaction amongst their members. For on these beaches. On reflective beaches, a
instance, in this zone occur the polychaetes narrow surfzone was represented by only one
Hemipodus olivieri and Nephtys simoni station, and the rest of the subtidal stations
which are predators of juvenile and adult represented the nearshore zone.
intertidal macrofauna (McDermott 1987). The three intertidal biological zones found
McLachlan et al. (1993) remarked that it is on these beaches were persistent throughout
not the type of beach state that is important the morphodynamic spectrum. Nevertheless,
to the fauna, but the swash climate associated some authors have recently remarked on the
with it. Swash climate is highly correlated difficulties in establishing major intertidal
with morphodynamics, an increase of swash biological zones (Raffaelli et al. 1991, Defeo
periods and swash lengths existing from et al. 1992), suggesting that the only valid
reflective to dissipative beaches (McArdle & zonation scheme might be division into a
McLachlan 1992). If the existence and width high shore assemblage of air-breather and a
of the saturation zone are related to swash lower shore zone of water-breathers (Brown
climate, this zone will be better developed at & McLachlan 1990). This is an oversimplifi-
the dissipative extreme, allowing more wide- cation of both the physical and ecological
spread occurrence of subtidal species in in- environment, and not in agreement with
tertidal stations. In this case, the increase in many studies that have been showing
species richness from reflective to dissipa- complex adaptations of species to this
tive on beaches with the same granulometric dynamic environment (Ansell 1983, Brown
characteristics, reflects the inclusion of sub- 1983).
 576 BORZONE ET AL.

It is obvious that beach zonation, con- BROWN C & A McLACHLAN ( 1990) Ecology of sandy
shores. Elsevier, Amsterdam, 327 pp.
sidering both the intertidal and the subtidal CHRISTIE NO (I '!76) A numerical analysis of the
environment, is highly influenced by mor- distribution of a shallow sublittoral sand macrofauna
phodynamic state and fauna! composition. along a transect at Lamberts Bay, South Africa. Tran-
sactions of the Royal Society of South Africa 42:
Biological zones are more evident on dissipa- 149-172.
tive than on reflective beaches, since some CLIFFORD H & W STEPHENSON (1975) An introduction
physical zones (e.g. resurgence and satura- to numerical classification. Academic Press, London.
229 pp.
tion) tend to diminish in width or disappear DAY JH. JG FIELD & MP MONTGOMERY (1971) The
towards the reflective extreme. Biological use of numerical methods to determine the distri-
zones will also be more evident in the bution of the benthic fauna across the continental
shelf of North Carolina. Journal of Animal Ecology
presence of less motile and sedentary spe- 40: 93-126.
cies, e.g. polychaetes or tube builder thalas- DEAN RG 1973 Heuristic models of sand transport in the
sinid crustaceans, than in the presence of surf zone. Proceeding of the Conferences on engi-
neering dynamics in the surf zone, Sydney. NSW,:
tidal migratory species. 208-214.
In this study we have shown the impor- DEFEO 0, E JARAMILLO & A LYONNET (1992) Com-
tance, when analyzing the structure of beach munity structure and inte11idai zonation of the macro-
infauna on the Atlantic Coast of Uruguay. Journal of
communities, of considering the entire beach Coastal Research 8: 830-839.
ecosystem boundaries and morphodynamic FIELD JG ( 1971) A numerical analysis of changes in the
spectrum. The homogeneity of sand compo- soft-bottom fauna along a transect across False Bay,
South Africa. Journal of Experimental Marine
sition buffered the possible morphodynamic Biology and Ecology 7: 215-253.
influence on the intertidal dominance and FLEISCHACK PC & AJ FREIT AS ( 1989) Physical para-
density patterns, suggesting that these para- meters influencing the zonation of surf zone benthos.
Estuarine Coastal and Shelf Science 28: 517-530.
meters are not controlled only by morpho- FOLK RL & WC WARD ( 1957) Brazos River Bar: a study
dynamics but also by sedimentological in the significance of grain size parameters. Journal
characteristics. This alternative hypothesis of Sedimentary Petrology 27: 3-26.
GUZA RT & DL IN MAN ( 1975) Edge waves and beach
awaits further testing. cusps. Journal of Geophysical Research 80: 2997-
3012.
HILL GW & RE HUNTER (1976) Interaction of biological
and geological process in the beach and nearshore,
ACKNOWLEDGMENTS
northern Padre Island, Texas. In: Davis JC & E
Ethington (eds) Beach and nearshores sedimentation:
We wish to thank Yara A. Garcia Tavares, 169-187. Tulsa, Okla.
HISCOCK F (1983) Water movement. In: Earll J & OH
Vicente Prata Jr. and Fernanda Gemael for Erwin (eds) Sublittoral ecology. The ecology of sub-
their cheerful help with the field work. littoral benthos: 58-96. Oxford.
Fernanda Gemael did the sand grain size KNOPPERS BA. FP BRANDINI & CA THAMM (1987)
Ecological studies in the bay of Paranagua !I: some
analysis and An De Ruyck the English physical and chemical characteristics. Neritica 2: 1-
corrections. 36.
KNOTT DM. DR CALDER & RF V AN DOLAH ( 1983)
Macrobenlhos of sandy beach and nearshore en-
vironments at Murrells Inlet, South Carolina, U.S.A.
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