//

~}'\
t \ "

in 'Sar\dv(\ec; k

SeaL h ~~
") ~ ~~

~q c: ~,

[\11-1
'~

C?

"ch

13;-\

~-,

~COS~ y/s.tems' &, T r r :~smu.s,

~ ~. . .~ \ ,~)~):~
-

-',

~- ~-

""'/

321

!~'

J ~~~; k

~ ~

H 8 Q U ~

SANDY

BEACH

ECOLOGY

REVIEH

ANTON McLACHLAN (Zoology Department, University of Port Elizabeth, P.O. Box 1600, Port Elizabeth 6000, South Africa)

1. INTRODUCTION Sandy beaches dominate

/

open to the sea. most

temperate

This does not include

sand lagoons

flats out

and assets
-;ea.

~ropical both and In

coastlines
impor~ant

where

they

re?resent

recreational

Nevertheless,

in estuaries CJr closed only open marine LJeaches. sucb beaches may differ in their Jegree of exposure

.:-ocJuctive

..::ommercially. regarc1ed

zones against the buffer they areas are very some exploited are and However, they have been marine and T.1!ere deserts largely

considerably

to wave action. On a 20 point exposure scale, developed studies for on (McLachlan, intertidal fauna 19800.) , scoring 11 to 18 are considered by are characterised and continuous, often heavy, 'v-lave action, the absence of silt, a mobile fauna and a high degree of oxygenation of the sand. (scores Sheltered beaches have 5-10) less,
fa Ulia

as

uy

many

biologists

neglected

beaches exposed

until Remane(1933) coasts of Germany. et 0.1 (1942) was

:epreser~~ed attempt
Sjster\':.

began studies on the Tbe vl0 r k of Pearse also pioneering and first a
sandy
b 1 y,

the

qualitative whole
b,::ach

to
Since
c e d

evaluate the n
sid era

often
wh i c b

intermittent
rnay cons t rue t

i.4aVe
sern i -pe

C3. c

t i
'8

r-~

beach ecology
tho ugh i t has

rrnanen

[-, s a
a 1 v7 a

\; a c1 an
y s

con

GurroVls

and

reduced

layers to

present

and

behind other aspects of coastal sarine ecology in the attention During this period it has enjoyed. research on beaches has biological early from taxonomic spread and
~alitative ol09:l
.species

lagged

close sometimes Beacbes scoriny rating and review.

develo]?ed

the surface. 5 points on this scale are not open ocean beaches
Ge 1 0\-1

fall

studies through qua~titative physiology important and of a more bolistic systems

beacbes

Lave a six ]?oint scale classifying from dissipative to reflective

outside Hright,

the scope cf (1982) Short

this

tOVlards today.

extremes based on morphodynamics. As the sandy beach occupies the a dynamic adjacent

approach

The

term

sandy

beach

has

been

loosely

interface its

position between sea and land, with

in the literature to cover a range '-'sed of environments from high energy open to extremely sheltered ocean beaches
est ua r i ne sand

boundaries

flats. a

of

this

review

For the purposes sandy beach is area

terrestrial and marine environments are not always clear and the functional extent of the beach itself bas seldom sand beach discussed. The Dee" considered here to be part a system

considered

to be a sandy littoral

322

comprising highest boundary of the envelope limit they which ecosystem, tropical not sandy interactions landvlard seavlard situations, driven inshore features in Fig. 1. by water. of include and of exist. I drift out waves of

i)

the line to and the

sand near beyond ii)

oody the the the to

from eJune/beach break

the point water outer wbere

been and

written relationships

on beach less on the water

sediments, between slope and

wave wave has

action action, IJeen

seeJimentary

parameters, considerably focused relating zone.

moving the

attention important movement in

surf circulation

zone

processes the of surf surf systems in IJeach

surf

cells

to The

This call is the are boundary boundary surf breaking The of the

eJynamic great dunes although important. is clearer

environment, zone in It of both does many extent

significance cells, rip to be current realized

beach/surf areas. typical

circulation etc., ecological Biological has

still stueJies. work

temperate

coasts,

dune/beacb Its than in its

to the aspects surf

eJate

has

concentrated sand lJeach lJolly

on and

the most

fauna other the neglected. of

of

intertidal of

ecology, have sand is and plants. two

where, circulation waves, basic

exposed cells, mix with and

including lJeen body

zone The sandy a of mobile

as a whole,
intertidal lJeaches substratum provides

open lJy

topography

characterized the Normally absence the

this

system

are

illustrated

attached rat UIll

subst

..

DUNES
DUNES ACTIVE DUNES

BEACH
II
I J

NEARSHORE
INNER TURBULENT ZONE
A
'\ (

'STABLE I ~

BACK SHORE '~)r~,

FORE SHORE

OUTER TURBULENT

ZONE

OUTER LIMIT OF SURF CIRCULATION CELLS 1

FIGURE 1. Profile the text. The will other physical not be papers

of

a typical

sandy beach environment,

showing areas referred to in

features examined in this

of

this

system Besides

here. volume,

habitats population. habitat

suitalJle faunal for Tbese are i) the wacrofauna

descript-

consisting

of

the

sand

surface

ions of physical features of beaches importan to ecol og i sts may be f our.d in t

many publications, including Bascom (1964), King (1972), Davies (1972) and Brush (1973). Though much has

and upper layer of sediment and ii) the interstitial habitat consisting of the
porous
former, is a

system
un ifo rm

of
lJu t

the

sand
and

lJody.

The

particularly

on exposed

lJeaches,
unstable,

Inman,

dynam ic

323 essentially tyJO dimensional latter is a more stable system. The and complex environment. macrofauna (Hummon these et are Thus although meiofaunainteractions may occur a1. , Reise, 1979) 1976; generally of negligible flow on

habitat with clear vertical gradients and is essentially a three dimensional system (1)1cLachlan,1977a). It has great vertical extent on exposed beaches or high energy 'windows' where strong hydrodynamic deep into the forces introduce oxygen sediment. Elsewhere its layers reduced

significance in terms of energy exposed sandy beaches.

The only review on sand beach ecology as a whole is some that of Hedgpeth of veach (1957), ecology Both although aspects

base consists of (Fenchel, Riedl, 1970).

are covered

lJy Eltringbam

(1971).

':2;;e macrofauna of sandy beaches and the (and microfauna) of meiofauna their entirely comprise two interstices components with separate faunal virtually no overlap or exchanges of energy. This is because the meiofauna forms small interstitial are extremely 'while the macrofauna are several orcJers of
mag:iitude
1

these works are outdated and there is a a synthesis distinct need for of information on sandy lJeaches. The aIm of this review is to outline our knowledge of the ecology of individual components structure ecosystems through of sandy beach biotas and the and as the a function whole. It
interstitial

of beach progresses

larger

(McIntyre, 1973; (1981)

1971 ;
disting-

~.'1cI:1tyre

Murison, Schwinghamer

McL&chlan,

macrofauna of the phytoplankton, zooplankton

fauna and body, sand and fishes of

:977a) .

the surf zone and birds of the beach and dune nlargin to functional aspects within energy the f r ame'v';o r k of flow and nutrient cycling.
2. The THE INTERSTITIAL SYSTEM

uished three biomass peaks in intertidal corresponding to Lacteria, sediments

interstitial

meiofauna size,

and

rnacrof

auna.

As of gap

sediments

mean

grain

become finer below 200 urn large lJur roy/ing forms increasingly become meiofauna and
'v"lee n

important
bet

tend

to

bridge

the

size

and meiofauna. macrofauna separation into two makes Thi s components less marked in finer and facilitates better energy sediments
101,0.'

between the sand averages about 40% of Most the total sediment volume. physical properties of this system are grains generally directly c1etermined by the sediment properties which in turn are related to the wave and current regime the geological history of
as vlell

porous

system

Energy between these components. components also between these meiofauna because the are increases surface the and concentrated near
f f 1 01,01

an

as area.

Grain size, shape and sorting are most important in fixing porosity and permeability Drainage which influence drainage. the organic layers. coarser and is critical in determining

macrofauna
abundance

deposit in sediments

feeders such probably

increase

in In

finer

~ediments.

coarser

intermedi",te

because they forms are absent, represent a size range selected against by the physical force s of the beach

moisture content, oxygen and input and the depth of reduced Permeability substrate increases and better with

sorting

324 also increases on steeper drainage beaches. The interactions between these and covered factors are well documented in several papers (Hebb, 1958; Hulings, Gray, 1971; Crisp, Hilliams, 1971). The major interstitial process involving the system on open beaches is

the filtration of sea water. ':Chis water the sediment either is introduced into by flushing by waves and tides in the intertidal (Riedl, 1971; Riedl, Machan, 1972; r'1cLachlan,1979a) or the pumping effects of waves in the subtidal (Steel et al., 1970a; Riedl et al., 1972). In exposed situations filtered volumes may strip of average S-lD m3 per metre intertidal times on beaches. In the subtidal surf zone off open beaches wave pumping may filter and particulate organics in the sand. Under conditions, however, the high energy input of oxygen and the high drainage rates maintain and
oxygenated

. .
0

Yellow layer: Gray layer Black layer

oxygenaled RPD zone reduced

sand sand

beach per day and be several grained very steep, coarse

FIGURE 2. vertical sheltered

Fenchel, 19790.)

Schematic chemical and Riedl, an

representation in
et

of

gradients exposed beach.

a
0.1.,

( P.fter

1970; McLachlan

D.OS-S.DOm3 m-2.d-l. This flushing dissolved or pumping concentrates

moisture content and surface temperature (Pollock, Hummon, 1971; r'lcLachlan et al. , 1977a) while in very sheltered situations it sharp can result in changes gradients, in chemical for example pH changes coupled to temperature IS78). a1. , vii th reduced layers three vertical zones (Fig. 2) may be distinguished, namely In sheltered beaches clean (oxygenated) sand, a gray (or RPD) transition zone and black (reduced) conditions (Fenchel, Riedl, 1970) . Horizontal zones may also be distinguished down the relatively constant beach from conditions of tbe the changes (Gnaiger et

this there

system are no

fully steep

vertical gradients in oxygen and oxidation states of sulphur or nitrogen et (Riedl, MacMahan, McLachlan 1969; al., 1979a). Permeability and flushing generally decrease with increasing shelter the until is drainage is so poor that sand constantly saturated.

vertical chemical gradients are then steep and reduced layer s occu r close to the surface (Fig. 2). The interstitial system cyclic changes related is to subject to tidal and

sublittoral temperature (Salvat, ~1ielke, because

to the backshore 'VIithlarge and salinity fluctuations 1967; On greater Schmidt, exposed vertical 1970; beaches, extent

diurnal cycles and the seasons. In high energy beaches this mainly results in fluctuations in the water table, pore

1964, 1976). of its

325 the interstitial and better drainage, system lends itself to subdivision into strata. recognized
g ra ined (M d

Pollock, five strata

660

Hummon in
t-1 ass

(1971 ) a coarse
tts

CRANES BEACH, MASSACHUSETTS 2 S: ..J W >

A TYPICAL

EAST

CAPE

BEACH

urn)

a c h use

beach which they could relate Sa Ivat's (1964) horizontal HcLachlan (1980b) strata on East Cape beaches urn) which closely parallel Pollock

back to zones. four 250 of

g ]
Z 0 <

distinguished (Me] those

~
~
10 15 HORIZONTAL
---WATER TABLE

10 DISTANCE

20
(m)

30

40

50

60

and Hummon. In both cases the strata range from surface dry sand at the top of the beach down through moist
belo,,! tbe

layers to the permanently saturated zone water table (Fig. 3 and Table

of intertidal FIGURE 3. stratification based on interstit'ial sand moisture content during low tide for a beach in f-1assachusetts and the East Cape, South Africa. (After Pollock, Hummon, 1971; McLachlan, 1980). See also Table 1.

1).

Table 1.

Summary of stratification

of exposed sandy beaches based on interstitial

See also Fig.3.

water content (Modified from pollock, Hummon 1971; McLachlan, 1980). Moisture Content & Water
Zone!StratuFI

Degree of
Oxyger,ation 'Temperature

Circulation Low, loses capillary water

Zone of dry sandi :8ry sand stratum Zone of drying/ Dry sand stratum Zone of retention/ Moist sand stratum Zone of resurgence/ Water table stratum

High

Highly variable

Low, loses capillary water

High

Variable

Loses gravitational

water but

High

Little variation

retains capillary water Gravitational water from zone Stable

r.1

d era

t e

of retention drains through here during ebb tide

to low

Zone of saturation/ Low oxygen stratum

Permanently

saturated, little

Low to
very 10V!

Very stable

water circulation

326 Large numbers of microscopic organisms occupy the interstices: fungi, algae, protozoans metazoans. and Amongst the animals, metazoans that pass
bacteria,

cells per gram dry sand and increase with finer sediment and greater surface area (Dale, 1974; et al., t'ieyer-Reil
1978; ~1azure, Branch, 1979). Khiyama, Makemson (1973) recorded 362 bacterial strains in beach sand, many associated symbiotically wi th metazoans. Those on

through a lmm screen are meiofauna and the rest Individual dry weights are microfauna. 10-5_l0-8g 10-6 meiofauna, for 10-10 -IO-llg protozoans and for -1012g for bacteria (Fenchel, 1978; Fungi in sancJy Warwick, pers. comm. ) beaches studied are generally et concentrated a1., 1978). at A higher tide levels and have been little (Upadhyay flora of diatoms or flagellates only develops in sheltered beaches and is absent on very exposed beaches (Steel et This also al. , 197 Oa ; Brown, 1971a) . appears to be more characteristic of temperate than tropical beaches (Munro et ale , 1978) . In sheltered sands the diatom flora may be locally important and, due to mixing by wave action, may occur to some depth in the sand (Steele, Baird, 1968;
MeadoHs, recorded diatoms McIntrye,
1'1unro,

undamaged considered

sand grains made up 54-78% of the total numbers of 107_108 per gram of wet sand. Hicks (1974) demonstrated the presence of Azotobacter in Hla r in e sand beaches while Rheinheimer (1977) 31 strains from beach sand, of the majority showed salinity at 30-500/ gram and were 00 negative rods that cJecompose could proteins, carbohydrates and fats. which optima Andrews et al., (1976) studied microbial development in a model sandy beach, recording greatest abundance near the high and low tide marks. Organic have carbon and nitrogen been in the sand isolated

1977;
Brock

Anderson, (1968) and

1978). numerous attached

living to sand

bacteria grains

found to correlate with bacterial biomass and abundance (Dale, 1974; An d e r son eta 1 ., 1981; Bolter et 0.1., 1981) which can be up tc four times the 0.1. Liomass (1978) of (~1eyer-Reil, Faubel, procJuction a.t 15gc.m-2 the 1980). for a meiofauna Munro et microbial temperate

up

to

10cm below the sand surface and Round assemblage (1979) described a diatom living below the sand surface. Amspoker described the intertidal (1977) distribution of epipsammic diatoms on a
Californian permanently

estimated

beach. saturated

Hhere diatoms

sand is generally

beach and 72gc.m-2 for a tropical beach, the higher latter value due to greater water percolation. Mazure, Branch (1979) estimated an average bacterial biomass of weight in the sands Lagoon. disturbance
bacterial

Pearse et occur closer to the surface. 0.1., (1942) recorded Chrysophyta, Chlorophyta, Pyrrophyta and Cyanophyta on sandy beaches. Bacteria beach to the 1966, abundant and important in mostly being attached sediments, are sand 1968). grains (Meadows, range Anderson, 108_1010 Numbers

l4.2gm-2 dry of Langebaan showed that increased

Dye

(19790.)

caused

greatly

McLachlan bacterial dry sand sediment

activity beach sand. in et 0.1., (19790.) recorded numbers around 108 per gram down to more than 1m in the of an exposed beach. Dye

327 (1980a) was able to show fluctuations in oxygen uptake by tidal beach 1977; Small Fenchel, 1978; Hartwig, protozoans l1ave also 1982) . been

sand. Maximum uptake coincided with maximum water percolation through the sand and maximum fluctuation occurred at Desiccation of the higher tide levels. sand caused uptake or a severe bacterial drop in oxygen activity. Dye total

enumerated

on

exposed beaches in the

McLachlan et Their numbers

Eastern Cape (Dye, 1979b; al., 1979a; Dye, 1981).

ranged 101_103 per gram ury sand and they were found to more than 1m depth in a well oxygenated beach. Their contribution to total benthic oxygen consumption was significant, accounting for 15-25% of interstitial oxygen uptake.
r'1eiofauna

(1981) also attempted

to partition

benthic oxygen uptake on exposed East Cape beaches and concluded that bacteria were responsible for most of the oxygen co~sumption except where there was a very t-1eye r -Re i 1 et rich macrofauna. microbial turna1. , (1980) estimated over time of 100h with daily microbial carbon production of 43I11g.m-2 in sheltered beaches in the Baltic. They concl uded that 50% of microphytobenthos primary production was fixed by microorganisms. Koop, Griffiths (1982) found a high bacterial biomass on an exposed beach receiving high kelp input on the
\,'lest

bave

received

considerable

attention reviews ecology

on sandy beaches and several cover aspects of meiofauna

(Delamare-Deboutteville, 1960; Swedmark, 1964; Fenchel, Riedl, 1970; Hieser, 1975; Giere, 1975; Fenchel,

1978).
Small beaches
meiofauna!

metazoans may be i.e. ,

occurring either larval

on forms

sandy

temporary

being Koop

coast of South Africa, found to 1.2 m in the

bacteria
sediment.

that et a1. , (1982a, b) estimated bacteria were responsible for over 90% of these k e-l p s . of carbon utilization

of macrofauna, As or permanent meiofauna. we are mainly cJealing with interstitial meiofauna here, the temporary component is generally not of importance on open beaches. components The dominant sandy Leach meiofauna are nematodes of and

Annual turnover

(P/B)

of bacteria

in

this system was estimated (Koop, Griffiths, 1982; Griffiths, information the ecology 1983). we still of

to be 30 times Stenton-Dozey, this about of

harpacticoid taxa

copepods with several other

Despite know nothing species

importance. These of variable oligochaetes, include turbellarians, ostracods, mystacocarids, gastrotichs,

individual

micro-organisms All groups

on sandy beaches. may be present

halacarid mites, tardigrades, gnathostomulids, hydrozoans and bryozoans. There is a well established relationship between the relative p roport ions of nematodes und l1arpaticoids and grain size. finer coarser mean Nematodes tend to dominate in sediments, harpacticoids in sediments and in sediments wi th size around et al., 300-350 urn they (Gray, 1971; Raffaelli,

of protozoans

in the interstitial and foraminiferans studied as they are

system but ciliates have been best usually relatively

la rge and qu ite abundant, espec ially on sheltered beaches with fine sands (Swedmark, 1964; Fenchel, Jansson, 1966; Panikkar, Rajan, 1970; Hartwig, parker,

grain

are about equally important McLachlan 1981b;

328 Mason, 1981; Hockin, 1982). Fricke, related reducing to differing tolerances to

Fleming (1983) experimentally the for finer sediments.

have demonstrated neamtode preference

1970).
generally

condi tions (Fenchel, Riedl, vertical distribution is shallower in the surf zone

is In sands above 200 urn the meiofauna usually entirely interstitial while
below 200 urn burrowing forms become increasingly important due to pore size restrictions (vlieser, 1959). Interstitial harpacticoids may occur down to t-1oore, 160 urn (McLachlan et a1., 1977b; can pursue an 1979a) while nematodes
interstitial

than on the beach but high numbers may st i11 occu r 10-30 crn into the sediment (McLachlan et a1., 1977b) . Dean (1981) looked at the effects of wave action on meiofauna abundance.
Horizontal cJistribution

may and

take

form

of

layers,

horizontal Riedl,

the zones,

125-100
Murison,

mode of life down to McIntyre, (Wieser, 1959; urn Fenchel, 1978) . 1973; In sediments nematodes As most open sizes in the nematodes always
(1973) optimum

bathymetrical steps regions (Fenchel,

Horizontal distribution

Sjeograpbical 1970).

zones have been

meiofauna of described from

noncapillary

dominate (Fenchel, 1978). sandy beaches have grain and harpacticoids

Hclntyre,

some beaches, particularly relatively sheltered conditions 1970; Schmidt,

~1 0 0 r e ,

under (Coull, Harris,

1972a,

b;

range 200-500 urn, interstitial are

dominant. considered

almost
l-1 r i son u

Itlhe r e 1979b) the 197 2a ; meiofauna is close to the surface and the distribution tends to take on a more

sands

of

230 urn to be

two-dimensional Seasonality

character

(Fig. 4).

for
faur;.a

the

development Gray, ',..'hi Ie that

of

interstitial Rieger (1971) diversity In South

1981b).

suggested

meiofaunal

increases
of 250-350

as sands get finer.

urn (McLachlan et al.,

Africa richest meiofauna occurs in sands

bas Deen documented in several cases in temperate areas with the occurring lower meiofauna in abundance and moving deeper into the winter (Renaud-Debyser, in Harris, 1972b; Schmidt, 1972a, 1963; t.]i elk b,; Hulings, 1976; 1974; e, Faubel, Nodot, Feder, 1976; 1976; paul, areas 1980). warmer In seasonality is less clear (McLachlan,
sediment

On a vertical basis meiofauna distrito the degree of bution is related drainage and oxygenation of the sediment (see Fig. 3 and Table 1 ). Meiofauna abundance 1970; drops off drastically 1978) and in a the deep RPD and reduced layers (Fenchel, Riedl,

1977b) and be more vertical

in sublittoral
cOIilplex

areas

(Emberton,

it may 1981).

McLachlan,

vertical distribution therefore only occur s on exposed beaches where reduced layers are absent (Renaud-Debyser, 1963; et

other than seasonal response Deen recorded as to have factors such as heavy rains (Bush, 1966), \-lavedisturbance (Boaden, 1968), tidal
r-1 i n eke, e

migrations

Fenchel, Riedl, 1970;

ale , 1979a) (see species and vertical also taxa

Different

McLachlan Fig. 2). show well

patterns

factors
~Jest

(Rieger,
1 9 7 9)

ott,
and

1971;

he ide,

c h a n 9 e sin

moisture and oxygen over the tidal cycle (t-1cLachlan al., 1977a). et

defined

distribution

329

(a) SHERT ALLAI

BEACH, INDIA

~
illillJ

Arenopontia oriental is Arenopontia acantha

Psammopsyllus operculatus

All

5]
(b) A TYPICAL

INTERSTITIAL
FORMS

MANX BEACH
INTERSTITIAL) 3 EULITTORAl SPECIES (1 EPIBENTHIC 2 INTERSTITIAL)

3 EURYHALINE UPPER SHORE SPECIES (All

MTl

6 SUBLITTORAL FRINGE SPECIES ( EPIBENTHIC) lWN

SAND GETS FINER

FIGURE 4.

Comparison of vertical and horizontal distribution patterns of

harpacticoid copepods in a well drained, exposed beach (a) and a sheltered beach (b) where distribution patterns tend to be very three-dimensional and rather two(After (a) Munro et al., 1978 and (b) Moore, 1979b). dimensional respectively. all cases the copepods are concentrated above the low tide water table. On the In

sheltered beach interstitial fauna are mostly concentrated high on the shore where some drai~Qge occurs. Nevertheless zonation in this case is still mostly across the shore and not vertically in the sediment.

seasonality

are

becoming

clearer

as

feeders, diatom filter

non-selective

deposit

feeders,

result of ecophysiological studies on responses, preferences and tolerances of species (Jansson, important meioraunal 1967, Wieser, 1968a; 1975; Wieser Hartwig et al. , et al. , 1974; 1977;

feeders, carnivores and even feeders. studies have Various aggregation shown of meiofauna in response to specific bacterial strains (Gray, Johnson, 1970) or where there are high levels of organics generally or
localized

~lieser, Scheimer, 1977). A wide range of food items recorded for meiofauna
.

has

been

(Lee et

small scale concentrations al., 1977; McLachlan et al.,

including detritus,

diatoms,

bacteria,

protozoa,

other

meiofauna

(Jansson, Fenchel,

Feeding

and dissolved organics 1969; 1968b; MCIntyre, 1982). 1978; Giere, 1975, categories bacterial include

Hogue, Miller, 1977b; Moore, 1979ai prey 1981") concentrations of or (Fenchel, Feeding 1978). rates have been estimated directly for a copepod (Rieper, 1978). have also lJeen 'l'he re growth, studies on ]?roduction and

330 population dynamics of important species (pawlak, 1969; Lasker et a1., 1970; Hall, Hessle r, 1971; Feller, 1980). Most round reproduction McLachlan 1977c; species have year times
i

supply particulate generally microfauna fuelled organics

adequate
considered

dissolved

and

organic food. which

Meiofauna are to subsist on the in turn are largely

with generation Giere,

ranging

1-3

months

(McIntyre, 1969

by dissolved and particulate flushed into the sand (McIntyre

Lasker et al., 1970; Bartsch, Schmidt, 1979).

1975;

et a1., 1970). ~1easurements of benthic metabolism have attempted to partition oxygen uptake between these different of the interstitial fauna (Munro et 0.1., lS78; Dye, lS80a; 1981). Rates of mineralisation of organics lJy the interstitial system have also been measured in experimental sand columns in an attempt to understand these processes better and estimate nutrient cycling et 0.1., Boucher, 1970;
( r-1c In t y r e

Over the past decade ecological studies on sandy beach meiofauna have advanced from the more general surveys of the sixties. More attention has been paid to the complex of factors interstitial system and been bet ter (Gray, Rieger, composing the results have

components

analy sed stat istically 1971; Hulings, 1971; McLachlan, 1978; The basic pattern

Hulings, Gray, 1976; Hogue, Miller, 1981).

Chamroux,

1976;

McLachlan

et 0.1., 1981c).

of meiofaunal response to different sediment types is well documented and to chemical features of the responses system are clearer. t-10re interstitial
ItlOr k

Vernberg, ratio

Coull

(1974)

estimated

the of as

metabolic activities of ciliates to Hteiofauna to macrofauna

1:0.5:2.1

is

needed

on

biotic

interactions

inc1udirg predation.

competition,

feeding

and

sandy for 0. sublittoral sediment and 1:0.54:0.04 tor an exposed Dye beach. (lS81) partitioned lJenthic

metabolism to the energetics

interstitial

lJetween microfauna,

meiofauna

and macrofauna
rJith

on two exposed beaches in

regard largely

metabolism still

and benthic system has as a black

the

1:0.23:0.04 and 1:0.27:1.85, the latter beach having an exceptionally rich

on

ratios

been

regarded

macrofauna.
partitioned Cape East

t-'JcLachlan

et

0.1.,

box and few authors beyond the estimation biomass. On exposed

have progressed of abundance or beaches, and in

(19810.)

benthic lJeaches

metabolism amongst the

interstitial

fauna

and

macrofauna

as

most other sediments, meiofaunal numbers fluctuate within an order of magnitude of 106m-2 and dry biomass values range 20-4400 mg.m-2. Some figures are compared in Table 2.
.

1:0.4. On a west Cape lJeach receiving a input, high kelp lJiomass ratios of macrofauna:
3.5:1:1.5 meiofauna:

bacteria

were

these 1:1:4.7.

ratios

(Griffiths et al., 1983) while productivity for \-Jere

Clearly biomass tends to be higher in

intertidal the great on exposed vertical beaches extent of It may be conc 1uded that ou r knowledge of the interstitial system of open sandy beaches has advanced considerably over the
past

the

because

these systems prov ides greatest habi tat availability and high percolation rates

two

uecades.

~Je have

yooc1

331

Table

2. Abundance and dry biomass values for meiofauna recorded from marine sands.
Source
Local i ty

Abundance sublittoral 10G.m-2

sublittoral lO6.m-2

Biomass
100 - 600 mg.m-2

'ilieser

(1960)

Buzzards Bay

Coull

(1970)

Bermuda

33 - 259 mg.m-2

McIntyre, Murison

(1973)

South Africa

exposed intertidal lO6.m-2

279 - 1 092 mg.m-2

McLachlan

(1977a, b)

South Africa

exposed intertidal lOG.m-2

20 - 3 360 mg.m-~

')

McLachlan et al., (1977b)

South Africa

sublittoral 10o.m-2 sand 105-l06.m-2 exposed intertidal lO6.rn-2

350 - 1 045 mg.m-2

Fenchel

(1978)

sublittoral

50 - 1 700 mg.m-2

Mc~achla~ et al., (1981b)

South Africa

~oo - 4 000 mg.m-2

Sten~on-00zey, (1983)

Griffiths

South Africa

exposed intertidal lO6-l07.m-2

1 000 - 44 000 mg.m-2

:;eneral

idea

of

the

distribution

patterns

of

patterns relation

stitia1 respect the excellent work describing in co-workers attention it deserves.

fauna in interstitial of features of interto many this However, in climate. of Riedl and wate r flow of

interstitial

meiofauna processes

and in

most

other

intertidal

sands (e.g. Figs. 3 and 4a).

This viater

through beach sand has not recei ved the The patterns water input, percolation and pulsing disturbance described by Riedl, Machan a considerable (Fig. 5 ) add (1972) degree of comprehension to distribution

flow is the superparameter controlling interstitial climate and the distrifauna in ueach of interstitial !.Jut ion uacteria and meiofauna sands. Both tend to concentrate in .the therefore r.1TL and below moist sand at and above the sand surface where there is bigh water minimum input, percolation and aeration, currents stagnation and pulsing

332

Under sheltered are not too severe. strata become the aerated conditions much flatter, the interstitial fauna concent rate s close to the sand su rface and the overall dimensions of the system take on a more two dimensional nature.
MAXIMUM
REACH

all organic it receives, materials whether these dissolved be or particulate. In this process bacteria most are mostly at food with
important,

the

top

of

with rneiofauna the interstitial

chain. higher

There is little exchange trophic levels, especially beaches. of By the removal and materials
nutrients

OF STRONG

-MHWS -MTL -MLWS

.-MHWS

on exposed the release

INPUT CURRENTS

mineralisation

of organic
inorganic

and the

micro- and meiofauna are responsible for the 'purification' of large volumes of sea water flushed through the interstitial 3. system.

MAXIMUM AERATION AND INPUT CURRENTS

-MTL

-MLWS

Hhile

THE MACROFAUNA not usually

as

quantitatively

MAXIMUM PULSING

INPUT AND MINIMUM DISTURBANCE

-MHWS -MTL
-MLWS

important in energy flow as the interstitial fauna, the macrofauna of

sandy beaches are often abundant some cases, attain
densities.

and, in
l1igh

exceptionally

-MHWS
MAXIMUM PERCOLATION MINIMUM PULSING DISTURBANCE AND

The ir main feature on open beaches is the Ligh degree of mobility displayed by all species. These animals may vary from a few mm in length to lGcm and consequently a variety of methods have been used to sample them. Generally sand is passed through a O.5mm or lmm screen, screens though of 2mm in areas of coarse or even 4mm have sand, been

-MTL -MLWS

I MAXIMUM INPUT
MAXIMUM
MINIMUM

AERATION A ND
DISTURBANCE'
. .

~.
.. ...
. ... '.

-MHWS
..'

..~/. .
.

'

:,:<,:..

. '.

.' .

-MTL

used. Ci'hus not all .wor ke r s 11a ve sampled the same components of the community. The macrofauna community consists of

...::..<

-MLWS

FIGURE

5. Diagrammatic representation of

those organisms too large to move between the sand grains and generally in the size range lmg-2g dry tissue mass. Sucl1 species make up shifting populations in the intertidal and surf zones Hhen evaluating them of open beaches. it is best to consider the population across an intertidal strip of beach as a contracting and whole, this unit expanding as veach profile changes and

some important interstitial parameters related to water flow in an intertidal h~gh energy beach. (After Riedl, Machan, 1972). The interstitial system appears to be very efficient at mineralising virtually

333

tides

pass

Consequently

from springs to neaps. all further discussion of

SHELTER
...

EXPOSURE .
;.:.: <;?.~.U

abundance and biomass of macrofauna populations consider metre wide will strips of beach and not square metres, stated otherwise. All authors unless should present their results in this way, at the same time stating the width of the beach. ~he macrofauna of sandy beaches includes most major invertebrate taxa although it has long been recognised that molluscs, crustaceans and polychaetes are the most important (Rees, 1939; ~iatkin, 1942;
South'\..;ard,

STAC EANS

~:tf:ti~t:i::!t:tt:;:::i:tti::;::::::::::~:::;~:;:::::;::::::;:;::::;:::;::;;::::;;;:::;;:;:;:;::;:;:::;;.:.;.:::;:;.;.:.;.;.

~
~
1952,

MOLLU

POLYCHAETES

Responses major FIGURE of the 6. invertebrate sroups to exposure the gradient on sandy beaches.
(Stephen,

1953;

SChuster-Diedricks,

1956; Dexter, tendency

Sourie, 1969,

1957;

Pichon,

1967;

is 1972) . a 'I'h ere / for crustaceans to be more abundant on tropical beaches or more exposed beaches and molluscs to De more exposed less and/or abundant on (McIntyre, temperate beaches 1968,

1928, Rapson, 1931, 1932; Holme, 1954; Figueras, 1954; Edgren, 1956; Ansell, 1961; 1959: Hade, MCIntyre, 1967; Smith, 1970; Brown, 1971b; 1971 ; Ansell et al., Irwin, 1973; de Villiers, 197 2b ; 1974; ~lcLachlan , Hanekom, 1979;
McLacblan, t1cLachlan
198 2b) .

Van et

cler

Horst, 1979b;

a1.,

1979: Lebe r,

1970; Seed, Lowry, 1973; although there are many

this

Dexter, 1981) exceptions to

and polychaetes are sometimes more these either of taxa abunoant than (Brown, 1981; 1983). t'icDermot t, Inoeed, Dexter;s (1983) work suggests the dominate most crustaceans that exposed beaches and polychaetes the most beaches with molluscs reaching abundance intermediate in ). (Fig. situatior.s In te nns oi 6 biomass, howeve r , molluscs are usually et important most ('I'revallion al. ,
ITiaX 1 ITiU;iL

Bro'vm (1982) re vi e'\-;ed i--lOrk on whelks or the genus Bullia and p~nsell (1983) has covered the genus Donax. There have been some general accounts of crustaceans em sandy beaches (McIntyre, Lopez, EpelcJe-Aguire, 1963; 1975:
Kamihira,

1979)

and

many

studies

sheltered

individual

species or groups.

on Amphipods on the (Croker,

have been extensively studied east coast of the United states 1971;

1967a, 1967b, 1968, 1970; Dexter, 1967, Sameoto, 1969a, 1969b; 1970; Craig, 1973; Croker

1970;

Dexter, et

McIntyre,
l'.nsell

1974, 1976; Eleftheriou, 1976; McLachlan, 1977a; al., 1972a; Bally, 1981; 1981).

Bousefield,

Shelton, Robertson,

et al., 1975; Scott, Croker, 1976; Holland, Polgar, 1976; Donn, 1980) and elsewhere (Williamson, 1951; Barnard, 1963; Fincham, 1981). 1971, IsOpods, have also 1974, been 1977; widely Kamihira, studied both air- and 1971, 1974,

~here have been many studies on the biology and ecology of dominarlt bivalves sandy beaches and gastropods of

water-breathers,

(Fish, 1970; Jones,

334 1979; Dexter, 1977a; K1apow, Fish, Fish, 1972; Kens1ey, 1972, Ferrara, Ho1dich, Brown, 1973; Johnson, Glynn et a1., 1975; Che1azzi, Ferrara, 1976b; Eleftheriou et a1. , 1980; 1972; 1974; ranges trom one (Gau1d, Buchanan, 1956; McLachlan et a1. , 1981b) to 82 (Vohra, this number decreasing exposure. divided beaches into exposure and, based 1971), with Bally (1981) three degrees of
increasing

1974; 1976a, 1978; Ho1anov,

Hendrickson, 1980). Mysids have been investigated by Fishe1son, Loya (1968), Brown, Talbot (1972) and Woo1drigde (1981) and cumaceans by pike, Le Sueu r (1958) and Corey (1970).

survey of 105 average numbers of species, abundances and c1ry biomass values for these (Table 3) . TABLE 3. Summary dry of mean macrofauna1 biomass values based
Bally, 1981).

on a literature beach studies, listed

Hippid studied.

crabs

have

been (1976)

extensively reveiwed while (1944), Emerita

abundance

and

Efford

on 105 beach

surveys. (After

distribution

in the genus

other studies include A1ikunhi (1955), (1969), Thomassin Matthews Di11ery, (1965, 1966, 1972), Efford Knapp (1970), Wenner (1977), Henner, (1979). (1979) and Subramoniam Fusaro Ocypodids have also been widely studied (Crane,
:' e 11 0 \.;' s,

High energy No. spp. Abundance m-2 Abundance m-1 Biomass g.m-2 Biomass 9.m -1 Sand particle diameter urn 11 400 20045 2.26 871 310

Medium energy 17 752 34571 1.97 170 257

Low energy 30 1710 2797867 6.23 63 238

1941; 1975; 1973;

George, Jones,

Knott, 1972;

1965; Hill, Wolcott,

Hunter, 1978;

vannini, 1976; IS80c; Robertsor" Robertson, pheiffer, Pfeiffer, 1981; Hails, 1981; Hill, 1982; Wada, 1982;
~.'lcLa

chl an,

While with et

Ya z i z ,

1982

and

Ca in e

(1974 )

and

du
on

Preez There

(1981) have

have also

studied been

portunids.

studies

and abundance decrease (Angus, 1979; l'lcLachlan al. , 1981b) individual size increases, yielding high biomass values exposure

diversity

cnidarians echinoderms (Buchanan, 1966; Scheibling, Ferber, 1971;

(Kastendiek,

1982) ,
Laurence,

1982;

even at 10w"er abundances in some cases. Though most classifications of beach exposure are very subjective, the above trend s clear. highest remain The biomass values come from very exposed dissipative beaches, e.g. 6621 9 ashfree dry mass.m-1 in the Bast Cape (McLachlan, 1977a) and 25735g dry mass m.-l in Peru (penchaszadeh, 1971). These high biomass values are, without exception, due to filter feeders (Donax,

Dexter, 1975; 1977b) , a1., et (Clark 1962; Hock, 1981 ; Hilson, 1970; (New, 1981; Brown, 1982) and insects 1968; Craig, 1970; Orth et al., 1977; Griffiths, 1980; Chelazzi Stenton-Dozey,

Dexter,

Ebert,

polychaetes Longbottom,

et a1.,

1983).

Bally all

(1981) summarised beach macrofauna

the

results to

of

surveys

1981.

Emer ita) . Exposed Leaches are thus not necessarily sparsely inhabited.

The numbe r of spec ies found

on a beach

335 Species diversity normally increases o-1cLachlan,

y r e

l.1 c

from high to low tide marks

1971 ;
l-1cLachlan

1977a) and may decrease again around the break point (McGwyyne, 1980) and then Day et increase offshore (Field, 1971; However, aL, 1971; Christie, 1976). on a beach dominated by kelp input, maximum macrofauna diversity occurred around the drift line at the top of the beach (Stenton-Dozey, Griffiths, 1983). ~he distribution of macrofauna along the beach is generally patchy, the comb i ned result of movement and sorting by the ash, or localized food concentrations S .,-1 species biological aggregations of (Loesch, 1976; 1978; Bally detail it to 1957;
AChutankutty,

Lawrence, Ferber, 1970; Bally, Albeit, 1981, 1978; et ale , 1981b; Lopez-Cotelo

et a1., 1982) beach slope (McLachlan et a1., 1981b) sand moisture (Hayes, 1977; Salvat Hithers, (Brown, 1964, 1966, 1967; Bally, 1981; 1977) 1964; food in the surf water Rapson, 1954; \jade,

1968; 1'1cLuskyet a1., 1975; Ansell et a1., 1972a; Nair, 1978; McLachlan et a1., 1981a; Hutchings et a1., 1983) and dynamic changes such as due to storms (Scott, 1960; Brown, 1971a). Effects of
d recJg i ng &nd beach nou r i shmen t have al so

been investigated tidal Marsh,

Saloman

in the inter- and sUD1978; Turbeville, MahacJevan, 1982~

(Reilly, Bellis, 1982;

et

Thurn, Allen, 1975; Moueza, Chessel, 1976; Saloman, Naughton,

Culter, ~redation

ale f

Hayes,

1977;

disturbance

1982) , while effects of have been assessed

Fusaro, 1980; Brown, 1952). (1981) investigated patchiness in on west Cape be a common beaches and showed occurrence ir! all

experimentally in a shel tered sand flat by Hoodin (1981). r--lany authors have monitored organic or chlorophyll levels in the surf without
correlations UncJ oubt 'v.ith high ec11:l
;

LJeing
with

0_ble to
faunal
hO\-lever,

species at all tide levels, often havins 101-102m. proportions of This sampling suggests transect that line
strategies

demonstrate
abundance.
1Jeaches

Liomass that on

must most

receive and oeaches

may if not

be

unreliable,

high it

inputs

of particulate

organics

particularly

replicated.

Dauer,

is

noticeable

Simon (1975), however, showed that by pooling the results of a whole line transect, error due to patchiness can De Hartnoll (1983) has red uced by 10-15%. quantified on a little size the species area relationship sandy beach. He found sheltered increase inc reased
tha t for

feeding very wbere rich filter populations occur, phytoplankton Glooms have !.Jeen recorded in the surf (New Zealand
Le'.;in et

Rapson,
al. , al.,

1954; 1981a;

Hashington
South
Sou

1979a;

.;:.frica
t h Am e r i c a

l'1cLachlan et

in diversity ubove
accu ra te

as

sample and

Gianuca, ~opulations

1983).
uevelop

Alternately,

rich on after al.,

0.2Sm 2
sampl ing

in

upwelling

sugge sted

an

agg ,"2gate area

of O. 5m2

is adequate.

resions, probably feeding phytoplankton acJvected shorewards upwelling cycles (HutChings et 1983) . Of the physical particle considered size the

Di

st

r i-bu t ion

on

the

beach

factors. macrofauna or

and abundance of organisms has been related to many Factors said to influence the include content sand grain size and/ 1970; (Longbottom,

factors, wave action and <jenerally been have most important although

organic

336 Eleftheriou, Nicholson (1975) show that grain size alone can not characterize a beach. As particle size, beach slope and wave action are closely related, defining the former two fixes the latter 19800.) . (Davies, McLachlan, 1972; et McLachlan significant macrofauna
( 1981 b) ale , correlations

20 (a)

4000

115 en LU C3 LU g, 10 u. 0
II) 5

3000

I I I I" E

2000

obtained between

LU

a:

e"
~eC1'

~~~/ /

LU II) :J 1000 Z
:E

en a:

l>'~ flJ/

species numbers and abundance

"e~

and both grain size and beach slope, but not wi th wave acti on est imates for beaches on the south and east coast of South Africa (Fig. 7). This suggests that it is not wave action but rather steep slope and coa rse sands that 1imi t the fauna. very exposed beaches (in terms of wave heiCjht) often have richer faunas than less exposed beaches where the latter have coarse grains and steep slopes. Finer sands result in flatter slopes as does heavier wave action. 'i'he flatter the slope of a beach the more
eve r: y 1 wa v e e n erg y

:J z 0

C11 ~(;>o/

c:,Q
-""

(\J" ~.Y~ 0

4'0

~o
BEACH

~o
SLOPE

\0
4000

20
(b)

115 en ~LU C3 LU g, 10 u. 0 LU co 5
~:J a:

\' 3000 I

I
E
2000 en a: :E

LU co

:J 1000 Z

is dissipated

in the

Z 0 100 500 Md (;Jm) 1000

surf zone and intertidal and this is the Thus a very exposed crucial factor. beach can support a rich mac rof auna if opposed as to dissipative it is relective McLachlan, (e.g. 1977a; Maitland
Copali s beach,

FIGURE

7.

TIle species

relationships diversity

be t.,leen

macrofaunal

anu

abun-

beach,
Lewi n

dance

Indeed, high energy et al., 1979a). dissipative conditions seem optimal for the development of a high lJiomass of

and the slope (a) and particle size (b) of ueaches in South Africa. (After ~icLachlan et al., 1981b).

filter feeders. Much less is known of macrofauna ecology intertidal. Fauna is below the generally absent around the break point but increases in abundance
onshor e and

1971, Dexter, 1978; 1970, 1972; Hill, Shin, Hunter, 1979; 1981, 1982; Kastendiek, 1982). The zone inside the break point bas been termed the inner turbulent zone and the area outside this (out to where wave effects on the bottom

and diversity
wi th

are
zone. Masse

negligible)

the

outer

turbulent

offshore

in

accordance

less

turbulent

substrates and (Clark, Milne,

conditions, less organic higher Morgans, 1955; 1971; 1976;

coarse levels 1962; Day et Hasse,

Biomass values are seldom high. (1970, 1971) recorded "number of

Field, Barnard, 1963; al., Christie, 1971;

10031 m-2 and biomass 2.5-11.7g 1048 dry mass m-2 at various localities in at depths 1,5 Sm. the ['ledi te r ranean
McIntyre, Eleftheriou

(1968) recorded an

337
aver-age

dry

intertidal

subtidal

biomass of 1. 3g .m-2 in the the and 3.7g.m-2 in with 62 species on the beach The

the midshore in New Zealand. In South scheme, Africa with Dahl's rnodifications, has been found acceptable
(~lC Lach lan,

and 116 off the Scottish shore. poorest zone was just below LHS.
t-1uch has

1980b;

McLachlan

et

0.1.,

1981b)

been

written

on The was

intertidal earliest a zonation that

AIR BREATHERS
WATER BREATHERS

distribution scheme for

patterns. sandy beaches

serious attempt

at providing

I MIOLITTORAL
II
RETENTION

SUBLITTORAL
FRINGE

of (1952) (Fig.8). very Dahl This was suggestion by to an earlier similar Davenport (1903) . Based on the distr icrustaceans bution in northern of temperate and Sou t h Am e r i can be a c h e s he proposed essentially three zones equivalent to the three zones of rocky shores. These zones were: fringe characterized subterrestrial ocypodid warm areas crabs in amphipods in cold areas; tal it rid mid1ittoral zone characterized the by cmd the by

RESURGENCE SATURATION 1,1 \,1

I I I

. .
.

B~<6~~~S

~~~~~

SAL VAT'S ZONES

ELWS

FIGURE

8. Diagrammatic

representation zona t i on
on

of

cirolanid isopods (although these may be absent in cold temperate areas); and the fringe with a mixed fauna Sl.:blitt.oral often characterized Ly l1ippid craus in
the tropics and haustorid

schemes of macrofauna beaches. Salvat (1964, 1966,

sandy

1967) Dased

and

other

alternate

scheme

on

described an physical

amphipods

areas. in temperate Dahl's zones were thus derined biologically. many

factors. He clelineated four zones on the ueach, Lased on water saturation B) . (Fiy. Pollock, Humon (1971 ) used were studying, exce~t that they suudivided the zone of dryinSj and they extended L,ones strata their into covering the 'whole interstitial system 3). Salvat's Table 1 and Fig. system bas Deen found reliable in describing zonation of macrofauna, (see particularly isopods (Hithers, Bally, 1981). Both these 1977; authors this they system for a Massachusetts beach

authors have tried varying results. scheme with tbis Gauld, 3uchanan (1956) round an overlap on west of talitrids and ocypodids Subsequently African showed (1972) Singapore found
et 0.1.,

Excir-olana

recorded shores and also Lower zones in this zone. correspondence. Philip less India, Vohra (1972) in and Jaramillo (1978) in Chile

in Dahl's

scheme useful and Escofet. found it fairly suitable Uruguay and beaches in Brazil, for J.~rgentiria . Hood (1968), however, found circolanids replaced by a sphaeromid on
(1978)

admi tted, lloweve r, tho. Doundar ies t are not sharp and that oones Sjrade into each (1976) otber. Eleftheriou, Jones pointed out that differences in inter-

338 tidal !..Jetween distrib'.ltion different areas and species make eurydicid isopods unreliable indicators of zonation as proposed by Dahl. They did not, however, attempt to use Salvat's scheme. Views on zonation of macrofauna on sandy the larger one in the eulittoral (e.g. Amphidesma spp. in New Zealand, Rapson, 1952; Donax spp. in India, Ansell et a1., 1972b; Donax spp, in the Last Cape, MCLachlan, 198Gb). Besides level, patterns at intraspecific zonation
i:.onation

community

beaches are thus highly conflicting, a situation not surprising in a dynamic environment wi th a highly mobile fauna. It may safely be said that zonation, in sense, has never the classical been proved on a sandy beach, i.e. sharp boundaries have not Further, individual clearer zonation than whole. Indeed, Brown of the opinion distinguishable breathers breathers been demonstrated. species show much the fauna as a is are air (pers. comm.)

recorded generally size most taxa

has Leen in several species. his '1' takes the form of L-.onation of and bas ueen recorded in typical of open beaches, crustaceans al. ,
\J 001

classes

including (Alikunhi, ~1cLachl an

molluscs and 1944; Glynn et et a1., 1979b; Lagardere,

1975;
g e ,

d rid

1981; 1982;

Ansell,

1980;

Brown,

Haley, lS82).

This may !..Je to due

that only two zones on sandy shores

in the supralittoral and water below them. However, two boundaries are clearly and indisputably open !..Jeaches, namely on the visi!:Jle drift line and the edge of the saturated sand where the water table reaches the to Dahl's surface. These correspond although boundaries. Howeve r , the boundary between Salvat's zones of retention and resurgence are not obvious considerable evidence that is there meaningful midlittoral These beaches. on zonation schemes are Within these summarized in Fig. 8. zones most species sbow subzones and subdivision made can be of the many

differential sorting of the sizes in the swash or to active miS:]ration to areas differentially suitable to the orsanism at different life stages. Zonation as the researcher records is, however, only the uistribution it

of the t!sammolittoral fauna c.L an instant in time and most species undergo tided migrations of some sort. Ty 1?i call y t his involves a simple movement up and dOl-ln allowins the animal to stay in the swash zone Y/here conditions for feedins are opitmal and predation minimal. In molluscs this movement involves rio endogenous rhythms but simply a series of responses to changing lJhysical conditions, most
the tides

the

th !..Jeach ,Ii

there is often blurring of boundaries. Isopod species usually exhibit a clear zonation near the top of the shore and midlittoral (Withers, 1977; Eleftheriou, Jones, 1976; Bally, 1981; McLachlan et al., 198+b; Dexter, 1983). Bivalves, often with two also exhibit zonation, species occupy ing a beach, the sma ller species in the sublittoral fringe and

notably
thixotropy Turner,

the

degree

of the Belding, 1969;

and of saturation (~1ori, 1938; sand Ansell, 1957;

'1'rueman,

In some molluscs' it may be TIIOre complex, e.g. Donax serra in the Last Cape, where
Trevallion,

1971).

there is not normal tidal migration but rather semilunar movement up and down the shore to occupy a position near mean

339

tide
tide 1979c)

level during

during neaps have Movements

springs related also

and to been

near et periods recorded

low 0.1, of

compress
the may with be

so

that

at

high into a

tide narrow

most

of

U1cLachlan

invertebrate com~ressed

population

on the beach
strip between to into

calm storm and ( Le be r, 1 98 20. ) crustaceans

considerable Further, migration, in

overlap

.
these and to night as behaviour 1963, 1965, shown at springs rhythms more include (e.g. are complex. entry mysids, increased opposed (Papi, 1972; to

species.
tidal

audition

normal
the

movement

In generally ~hey into activity neaps, pardi,

endogenous have been plankton and during and 1963; orientation Enright,

subtidal bas been recorded in winter Donax parvula (Leder, 1982b). Temporal changes in Deach

in

the

macrofauna

anphipods

isopods),

have been communities

in most i nd i v i dual

recorded both whole in (Sanchez et 0.1. 1982 ) and ,

spec of ies these popul at ions. 'J..'he

uramatic

are those

assoc-

0.1., 1968, 1969; Cubit, 1969; Fincham, 1970, 1973; Alheit, Naylor, 1977; McLachlan 1976: r'1acQuart-110ulin, et 0.1. , 1979c; Harsh, Branch, 1979: Hager, Croker,
et 1981) activity it as prevents the tides

Hammer

et

iated with the monsoon in India \lhe r e most of the fauna Gisappears during this period (Ansell et 0.1., 19720.; D\JiveCli et 0.1., Longer years 1954;
1978)

1973; have coe,

McLusky also 1955; been

et

0.1.,

1975). several

1980;
0.1.

Fusaro, 1980;
advantage

Jaramillo Ugolini,
increased that s~randed

1980; Scapini, of
is being

term fluctuation

spanning

'l'he during the retreat.

springs

.
may

Dramatic

recorded (Rapson, Davis, Von Blaricom, changes in the long the

animals

Ansell,

Trueman costs

(1973)

calculated in

the

form of occasional mass Ttlortalities, e.g. Donax spp. (Orton, 1929; Fitch, 1950; Loesch, 1957: JOhnston, 1966, Nel, 1968; Grindley, 1968; de poi to most
sorJous

term

also

take

energy

it ,,;as more Emerita and ?~ofitable for the animals to migrate than to attempt to maintain position. behaviour has several Tidal migratory

of migration concluded that

Dor:ax and

Villiers, 1974) such as related uinoflagellate blooms. been and reported in for several

Seasonal chanses have species studied

communities.

advantages: it allows animals to stay in the zone optimal for feeding, i.e. the edge of the swash zone; it keeps
thern

. ~J.

an

unstructured

and

physically

too shallow for many fish predators but just immersed enough not to be fully open to birds: it helps to prevent animals being supralittoral enables them stranded
for m s ,

controlled habitat most species tend

like a sandy beach, to be unspecialized (Brown, generalists with board niches Talbot, 1982; McGwynne, 1972: Brown, 1980;
the Bally, 1981). Because of this, physical

on the shore; in
e .g . Ty 1 0 s , it

nature
mobility amongst unlikely, of LJe

of
the

the

environment the
sandy

to

move

d9wn

the

beach

during the low tide to feed on debris.

This

and the competition

fauna is not possibility can

of

fauna,
beach the

though competition

migration
on

naturally
and

has all

marked

exploitation

ef fect s

zonation

zones

ruled

out.

Hilson

(1981)

340 demonstrated negative interactions in a dense assemblage of deposi t feeding polychaetes, but this was in a very sheltered beach. Trophic include feeders groups among the macrofauna predators, filter and, on less and
.

predators.

scavenger/ (suspension) deposit usually of

As the bulk of the fauna are usually filter feeders, the size of beach populations is probably closely related to the richness of inshore waters in particulate organic material (Ansell et al., 1972a; Hutchings et al., 1983). In some areas large amounts of mac rodebr is may cast ashore, particularly ue adjacent to kelp beds. In such situations vast populations of supralittoral scavengers develop (Hayes, 1974;
Griffiths, Stenton-Dozey,

exposed

shores, feeders

feeders. dominate

Filter

consist Donax

mostly

bivalves

The

tellinaceans 0f
of the

high

energy beaches, most other suspension beach filter

spp., are, unlike family,

(Ansell,

members feeders is not feeders,

1983).
in sandy

Deposi t feeding

common

1981).

except in very situations. re sheltered In cases \vhe the fauna is impoverished, however, such steep very coarse, beaches, as on supralittoral scavengers may dominate as a result of the virtual absence of other species (Gauld, Buchanan, 1956; Dye et al., 1981; Hoo1dridge et al., 1981). On
shel te red Arenicola, beaches deposi t and feede rs, e. g. Callianassa Scolelepis,

In other areas carrion input may be high, e.g. in the form of cnidarians, and in such cases midlittoral such as Bullia reach high (McGwynne, 1980; Brown, 1982). food webs scavengers abundance

have been given for beaches in South America (Koepcke, Koepcke, 1952; Gianuca, 1983), the Cape (Brown, 1064; Griffiths Peninsula et
t'Iacrofaunal

may be very important or even dominate (e.g. Saloman, Naughton, 1978: McDermott, 1983) . The tropic structure of the beach macrofauna is therefore normally dominated by mobile filter feeders. be more forms may feeders usually Even where other abundant, filter dominate biomass McIntyre,

The main predators on the macrofauna as listed by several authors are birds, fishes and crabs (Coe, 1955; Loesch, 1957: Masse, 1963:
.

a1. , 1983) and et a1. , 1981a)

the

E;ast

Cape

(McLaCfll

an

Hade, 1975).

1967;

Penshaszadeh,

Olivier,

Eleftheriou, (Dexter, 1979: 1976). P.s there is little or no primary production on the beach, the macrofauna is dependent on food imports ad jacent systems,

The former two v/ill ve discussed being later. Crabs, the tJart of benthos, represent a form of predation resident in the veach/surf system. Du Preez (1981) has made a detailed study of the swimming crab Ovalipes on East Cape beaches and shown it to lJe an important preda tor on Donax and Bull ia. Seve ral authors Iilentionthe importance of both ghost crabs and swimming crabs as predators on ueaches (Loesch, 1957; Koepcke,
Virnstein 1953; (1977)

from

i.e. the land and the

sea (Brown, 1964: Ansell et al., 1972a; McLachlan et al., 1981a: Griffiths et the most is sea The al. , 1983). important filter and and and supplies for particulates and scavengers for and deposit feeders carrion

other

Ansell
showed

et

al.,

1972a).

the

importance

macrodebris

of

crabs

in

controlling

macrofauna

341 communities sand in gastropods burrowing by predation in subtidal Chesapeake Bay. Naticid are important predators on but are not generally 4. PhYTOPLANKTON Blooms of diatoms persistent a typical or sporadic, feature

in
now

surf sandy

zones, ueach

appea r to ue

bivalves

uf many

found on open ocean beaches. mortality on sandy This includes not only mass poisoning (de due mortalities to being cast Villiers, 1974) but also
"natural"

Besides

predation, times

surf zones. Rich surf phytoplankton blooms have been reported from the Hashington coast, U.S.A., (Thayer, 1935; Lewin, Norris,
,

can at beaches.

be

important

1970),

the Gulf coast of

above the shore by storms (Brown, 1971a; Penshaszadeh, Olivier, 1975; t:1cLachlan Young (1982) et a1., 1981a). McLachlan, suggested the importance of upwelling temperature drops sudden in mobility as a factor contriretarding to..; a r d s this. Ansell et a1., bt.:.':ing and (1978) mortality that have shown where rate 1S fairly constant,
coefficient,

the (Gunter, U. S. A. 1979; Gunter, Lyles, 1979), New Zealand (Rapson, 1954; Cassie, Cassie, Lewin, 19GO; Norris, 1970), South Africa (McLachlan, Lew in, 1981) America and the east coast of South
(Gianuca, 1983). These blooms are mostly cor.lposed uf masses of cells LJelonging to the of one or tvJO species

genera

Chaetoceros,

Asterionella,

Aulacodiscus The
anc1

and Anaulus. described by Gunter (1979) Lyles (1979) are Chaetoceros

the blooms
Gunter,

mortality production lists

caused

are of

related.
cornrnercial

the longivity and Ansell (1983) exploitation

dominatecJ

strips

hugging

the

ueaches.

cases
by

Eeavy

rains

leach

nutrients

from

the

or Donax

..7h ere

heavy

mortality

may be

Ilian.

land and these, lJeing retained near the beach by calm seas, apparently cause the blooms. These blooms take the form of strips S-6m \JicJ for miles along the e shore. All the other reported surf zone blooms occur as patches uf phytoplankton
~wi thin

distribu~ion and diversity of beach macrofauna by physical largely determined is prime among which factors, are wave
l~

rnay~

be

concluded

that

the

~well

c:levelopec1

surf

zones.

action,

particle

size

and

beach

slope.

these, have

those

uff

the

~Jas hi ng ton

Of coast

Food input probably determines abundance or both scavengers and filter feeders. most pronounced feature The clegree macrofauna animals is a bigh mobility and tidal migrations beaches. macrofauna The on exposed of main role the is to process food imports,

been

most

intensively

studies

coming mainly from the surf, and in turn

predators.

to be consumed by terrestrial and marine They are thus in the middle

of the food chain.

342 retained
often clearly retention

in

the

stranded appears in described composi

surf on
to the long tion

by wave act ion and the beach. This

be surf term of the a mechanism zone. changes blooms for Lewin in wi th before

Subsequently paid the b)

more

attention and Lewin to

has

been

to the ecology
!Jlooms. described zone in in Jijina, physical relation

distribution
(in prep. of at

of
a, the 13

features
blooms

(1974)
species Aulacodiscus

coastal beaches

Washington !Jest developed

and
on

Oregon.
long flat

littorii

dominant

Blooms were
beaches beaches

armatum dominant 1950 and Chaetoceros after 1950. Other studies looked at light utilization (Lewin, Mackas, 1972) , nitrate reductase activity (Collos, Lewin, 1974), daily periodicity (Lewin, Rao, 1975), C:N ratio (ColIos, Lewin, 1976), respiration (Robertson, Lewin, 1976),
a1
.,

while steeper beaches and near rocks has poorer blooms. were Low air and Hater temperatures associated increased diatom with densities were
densities.

while
inversely They

nutrient concentrations proportional to

concluded that the type

chemical
and

composition
the cIa y co at

(Lewin
( Le win

et
et

of

beach

and

its physical very important

characterisin bloom

1 9 7 9 a)

tics

were

al., 1980) of the surf Chaetoceros and Asterionella.

diatoms

development.

f.1cLachlan

(1980d) where surf

suggested
trap lJeach out that

that blooms cells

generated

Lewin et al., (1975) analysed envi ronmental conditions associated with the finding two years, the blooms over populations relatively constant diatom (1978) Lewin over this period. evaluated surf. seasonal Nitrate concentrations fluctuated widely and in relation to season, upv/elling phytoplankton growth and ammonium seemed to be an important nitrogen source, during summer. particularly Lew in et controlling factors cycle of nitrate in the the

form

circulation

develop
nutrients,

and on

pointing

this

is

most

likely (i.e.

long

beaches

vf

flat

slope
1.1cLachlan!

I-.ewin

dissipative beaches). (1951) described blooms

of Anaulus birostratus from the Bast Cape, South Africa, and shov/ed tha t the blooms were positioned over rip currents by <.lay. All surf bloom torming species
show a daytime buoyancy and this results

a1., (1979b) measured

by Siliqua and concluded
clam could

ammonia
that

excretion
the razor

in tbe collection of diatom cells as a foam or scum over rip currents. Presuffiably the opposing forces of incoming waves and outgoing rip current keep them in position. ~Jhile c.:iiatom cells may travel all around the surf cells, the rip current area bottle-neck
f.1cLachlan, blooms in

populations
regenerate

of

Washington

beaches
of Le win

significant

amounts

zone. into the surf ammonium (1977) summar ised knowledge of blooms, pointing of the Diatoms

circulation
may act as a

these

out the interdependence the food

for
Lewin

buoyant
(1981) in

material.
also not nutrient showed that

diatoms and
are the main

razor clams. of the razor

clams while the clams in turn regenerate inorganicnitrogen which is utilized by the diatoms.

developed is low,

water

very high
concen-

nutrients.

Though

tration
inorganic

supply

is are

constnat

as

nutrients

continually

343 draining out of the beach. Nutrients species commonly occur in well developed

are generated on the beach both by the filtration of water and mineralisation fauna of organics by the interstitial (McLachlan, 1980a, 1982; McLachlan et al., 1981a) and by excretion of the macrofauna
(Proscb,
~1cLachlan ,

surf zones on gently shelving beaches. Diurnal buoyancy of the diatoms results in their concentration over rip currents by day. Nutrients are derived from many sources, l..>u the beach is probably most t important. Inorganic nitrogen generated as ammonium
l..>y

1983).
and Cape beef!

Both of these

sources

are important

macrofauna

and

have been shown to generate East amounts of nitrogen on beaches. Subsequently wind has

significant

shown to be of overriding importance in controlling sbort term changes in blooms in the surf (Romer, Sloff, pers. comm.). 2stimates of primary production in surf

as nitrate Ly the interstitial fauna is constantly draining the into surf and passing through rip currents where the lJlooms are situated. In this way the lJlooms are positioned in a site of steady nutrient influence controlling circulation
more of the

zooplankton

excretion

and

supply. on
.wave

Hind Lloom

has a pronounced development by

~le need

zones off beaches have been made by (1960) and Edwards CaSSle, Cassie (1973a). The latter obtained figures of 10495 and phytoplankton off shallow beaches. productivity blooms
Schaefer,

action

patterns. interaction

and to

surf know
wind,

between

5597 kcal.m-2.y-l for primary production in tbe two in

now

waves,

Estimates

sheltered Venezuelan primary of phytoplanktoD surf under way

(Leviin,

and blooms. \Je also need more reports on the sources of primary production in surf zones that do
f!utrients

not support these diatom blooms. 5.

L;OOPLANK'I'ON

are

figures

1983) and we shou Id soon ha ve fo r biomass and production of

these rich algal communities.

a1., (1982), Hilce, and Quinlan et ale (1983) Quinlan (1983) free living reported masses of on algae, Pilayella, sand on oall-liKe
ReceDtly h1ilce et bottoms and in the surf off beaches in This species Nahan t bay, Hassachusetts. peak and biomass in summer reached by fragmentation and reproliferated generation. uncertain
.the

from the literature. papers Hhile several taxonomic list species taken off sandy beaches, few were sampled in the surf zone or include ecological notes. Brattegard (1969 and subsequent \,orks) described mysic1s from shallow vlater off the Bahamas including some from sandy habitats and similar accounts are given vy Higley, Burns (1971) for the Atlantic coast of the U.S.A. and Bacescu (1975) for Tanzania. Bowman (1971) described coast a by and of copepods the uff the and one lay

Studies on surf virtually absent

zone

zooplankton

are

but

It's nutrient source is nutrient recycling within

beach/surf

system

is

considered

southeast recognised dominated

U.S .A. and this

important.
~... .J.L.

'coastal
one'coastal'

association'

may

be

concluded

diatom

blooms

that rich surf consisting of one 0r two

'estuarine'

species.

Outside

'shelf'

'oceanic'

associations

344 by dominated respectively. studied four and Sander, and seven Moore species (1978) copepods outer limit of distribution near the outer limit of rips. He postulatecJ that detrital foocJ in suspension c1ecreases outsicJe tl1is zone and that zonation was mainly related to nearshore, tl1e wave-induced Several circulation ~atterns. have recently been clone

inshore

offshore

around the West Indies as shallow as 10m and recorded average numbers of 100500.m-3 with other groups making up 11-22% of Youngbluth total zooplankton numbers. (1979) studied zooplankton

studies

within lkm of the coast of puerto Rico at about 10m depth. He recorded numbers of 41-7568.m-3 of which 65-84% were copepods. Settled volume biomass values averaged 0.086ml.m-3 and highest numbers were recorded at night. Moran (1972) found the benthic mysid, Gast rosaccus sanc tus, to occu r in shoals along the shore in the top lcm of sand, but moving into the top I-2m of water at night
exhibited

on surf ~one ~ooplankton in the Last Cape. In IIIOst of tl1iswork the use of bas attempted to avoid the such as normal loss of large forms mysids, a problem often not appreciated by workers usins small nets. Cockcroft nets (1979) sampled inside the breakers at 1m depth, just outside at 3m and further out at beach. outside
efficiency Abundance

large

10m off Greatest the

exposed a moderately biomass occurred just

(although

breakers

netting .surf).

in

the

Mediterranean.

It

\-las very

Fincham Britain, studying isopod

'Vlooldridge

zonation offshore. a clear (1970), studying amphipods in

Macqua rt -t-10u 1 in

(1977), and an
ancJ

after dark when dry biomass values ranged 6-980mg.m-3. Mysids dominated biomass
increased

low in the considerably

two in

benthic
the

mysids
ran.ean,

i.jedi ter

mean

(1981), the East Night

mysid

in

studying a Cape, found time

benthic

(21% at 1m, 83% at 3m and 8496 at 10m; 63% ) hith copepods averagins only At site prawn, 13% . the the 1m
africana,
\Jas

patterns.

similar planktonic

Macropetasma

very The

impor tan t althoug h poo rly dominant species overall Mesopodopsis slabberi.
distinctly l1igher tban

sampled.

crustaceans, activity of benthic mysids, amphipods ancJ particularly isopods, is .well known (Alheit, Naylor, 197G) . The most detailed account of su rf zone zooplankton yet published is the work of Clutter (1967) at La Jolla. He studied zonation of four benthic the nearshore and five pelagic mysids out to 17m depth. All species clear zones, bottom. A
species,. where rip

was the mysic1, Biomass was

in l\.lgoa

offshore

Bay

and

about

the

same

as

local

estuaries,

indicating

recorded in the general

richness of zooplankton in the surf ~one. Subsequently detailed occurs zones. this Cockcroft (1983) has macJe a which

stucJy of

Macropetasma,

formed

schools

and occupied

in vast shoals in Bast Cape surf Growing up to 7cm in length,

occurring on or near the large species dominated

elongata, and peakecJ had its

inside the surf while the most abundant

Metamysidopsis ~urrents dispersed

prawn is really a nektonic form. exhibits clear zonation with It a juveniles inshore ancJ adul ts out as far as 20m ciepth. by day They and shoal bottom disperse near the at night,

345 being most abundant inside the breakers. Numbers recorded averaged 21-37.m-3 giving a dry biomass of 0.9-1.8g.m-3. This species uses the surf zone as a nursery area and, being an opportunistic omnivore, it feeds on detritus, phytoplankton and small crustaceans. Shoals tend to move inshore at night and to concentrate around phytoplankton
blooms by day.

1000.m-3 just behind the breakers 10.m-3 further offshore. It

and even

reached densities exceeding 15000.m-3 (mainly juveniles, dry biomass I.02g.m-3) behind the lJreakers at mysid species seldom Other 10.m-3. Dry more than zooplankton biomass averaged about 0.25g.m-3 behind the breakers and 0.015g.m-3 offshore, an order of magnitude lower. times. totalled Mesopodopsis virtually moving abundant population juveniles even into formed vast shoals which

(pers. Eomer comm. ) is food chains around surf blooms, including small
zooplankton.

investigating phytoplankton and large disappeared at and by day (presumably

He finds that zooplankton is concentrated around blooms by day but more dispersed at night, this resulting in a slight decrease in biomass after (Macropetasma, Crustaceans dark. mysids, of
copepods)

to deeper

water)

night. consisted breeding large

and then became This inshore largely of

though

account and and

for about

85%

estuaries.

occurred at sea numbers often moved This study indicates

biomass

siphonophores,

that the inshore

area and surf ~one bas

others for the total numbers :emainder. in a bloom during the day are about 2000 zooplankters.m-3 with a dry lJiomass of "') higher values, up However, O. 8g .m-.).
cbaetognaths,
~1 e a n

a mysid dominated zooplankton cOIllTIlunity of very high biomass. Abundance and biomass in this region are an order of magnitude higher than offshore and even higher than recorded in most estuaries. Hany have probably underestimated the contribution of the larger and more mobile zooplankton through using small nets. Clearly our knowledge zooplankton is abysmal. beyond species lists
abundance. extrapolate
\Jb i 1 e

_':/

to

750g.m"", swarms

have

been

recorded

dense

of plankters,

in particularly

authors

where prawns were abundant. Hooldridge plankton of (1983) Algoa studied Bay from the zoojust behind

the breakers to about 4km offshore (20m depth) using a net of 1.5m diameter. y;h salps, medusae, ctenophores and i I e others biomass. accounted crustacean were often making Fifteen common, up species crustaceans 80% of of mysids dominated, about

of surf zone Few papers go counts or of is difficult amount to of small

it the

from

hard uata currently available, three points seem clear: i) the zooplankton of
exposed su r f zones may be ve ry r i cll wi th

for more than 90% of biomass, with copepods second

high dominate
are

lJiomass

values,

ii)

large"

in importance. A single species of mysid, Mesopodopsis slabberi, was by far the dominant with species, average especially numbers close around inshore,

crustaceans

mysids, penaeids) the biomass (as they can cope of these areas) but are to sample, iii) the re

(e.g.

with the turbulence

<.1ifficult

346 clear offshore zonation patterns and diu rnal changes in abundance. Huch wor k remains to be done in this field, in particular relating distribution and abundance to wave induced circulation patterns and available food in the nearshore zone. 6. FISHES studied better than Though far zooplankton, we still know comparatively little of the ecology of fishes from surf zones off open beaches, largely because of the difficul ties of sampling pearse et al., in these environments. (1942) list fish caught by seine net off beaches more Carolina and North extensive Gunter
( 1 963 ),

recruitment, juveniles in sandy Macer, beaches

growth and the shallow (Riley,

feeding subtidal

of off 1966;

Corlett,

Edwards, steele, 1967; 1968; poxton et al., 1982) . Lockwood, 1974; Among these fish, plaice (pleuronectes platessa), dabs tLimandia limandia) and

turbot (Scophthalmus maximus) are the most important. summer, after In large numbers move into metamorphosis,
fin e san d bay s
'vI

her e

the

wa t e r

is

1e s s

than 3-5m (Jeep. There they feed on demersal plankton an<.1 benthos, including mysids and the siphons of Tellina and tentacles of }?olychaetes. Horta1i ty is high and the populations of these juveniles through deeper winter. United (1958) In the States Gunter sampled surf fish off the 'l'exas coast F recording fewer species off open beaches than inside tbe D3.rrier. He mostly caught small and juvenile fish which spring were most and aLundant in summer. McFarland (1963) zooplankton 'v,asthe primary for the Texas. found that food source generally autumn water until again at decline they the rapidly move onset into of

reports et

of al.
( 1967

fishes

in

surf

zones of the Un ited States (1958)

S c h a e f fer ,

are given by
1970 ), Cup k a

(1960), McFarland (1983). Around the Europe much and

(1972) and McDermott coasts of Britain attention

to juvenile has been paid as nursery using beaches flatfish grounds (r1ace r, 1967; Edwards, Steele, et a1., 1970; Steele, 1968; Edwards Edwards, 1970; Steele et al., 1970b; Jones, 1973; Gibson, 1973; Thijssen et al., 1974; Lockwood, 1974; Pox ton et al., surf the 1982). In South Africa Rossouw (1983) has studied elasmobranchs from zones with particular emphasis on

surf fish at Mustang Island, Planktivores were the most

abundan t trophi c 9roup and even mullet t (Mugil cephalus) and benthic feeders took large amounts of zooplankton at times. He recorded 47 species with an average (;f four in winter (biomass 25.81b/acre) an(J 16 in summer (biomass

annulatus, sandshark Rhinobatos Lasiak (1982, 1983) has made a deta ile(J

investigation

of surf

zone ichthyofauna

communities and Romer, (pers. comm.) has investigated food chains around surf phytoplankton blooms. In the cold temperate and Europe shallow
impo r tant as nu r se ry

103.21b/acre).

High

primary }?roduction

in the surf, he concluded, was more than suffici"ent to support the fish community. MOddle, Ross (1981) also recorded midwater planktivorous fish dominating the surf zone in the northern Gulf of Mexico.
Armitage,

waters of Britain sandy bays are

areas for f la tf i sl1

and

several

authors

have

studied

Alevizon

(1980)

347 recorded the diet of the Florida pompano (.Trachinotus carolinus) and found Donax to be an important item. et ale , (1960) recorded 71 Carlisle species off Californian beaches with an average of 284 specimens per beach seine surfperch, Amphistichtus The haul.
argenteus,

Edwards (1973b) sampled with a beach seine net off two Venezuelan beaches and recorded 49 species (31 at one beach and 32 at the other) which he di vided into plankton feeders, fish carnivores, benthos feeders and berbivores estimated unpolluted a at (Mugil). 5g.m-2 and beach. beach Fish biomass dry mass at 19.m-2 at was the at

and

the orbina,

Henticirrhus

polluted

undulatus, were dominant but flatfish, skates, rays and sandsharks (Rhinobatos) were also common. Best catches were They made a taken on the low tide. detailed study of the biology of the surfperch including reproduction, growth and feeding and Emerita analoga,
the diet.

Plankton feeders and benthos feeders most were important. Penchaszadeh (1983) found mysids and other motile to be the main prey of crustaceans fishes off Venezuelan beaches. In Australia described the Lenanton utilization et of al. surf (1982) zone

found the mole crab, to be the main item in

accumulations lif macrophytes and their associated amphipods as nursery feeding the the areas for species. In the 0+ year classes of three fish

and mysicls were the dominant foods though fishes, particularly anchovies, were During sampling important. also 71 species of fish were recorded.
(1983)

Schaefer (1970 ) studied feeding of striped bass, Morone saxatilis, in Amphipods Island. surf at Long

Africa, Rossou\-J has studied elasmobranchs with particular reference to the biology
(1983)

Eastern

Cape,

South

of

the

sandshark,

Rhinobatos

annulatus.

f-';cDermott

sampled

the fauna of a of

b irth

This species comes into the surf to give ins u mm e r. Juveniles stay in the surf but adults
on

barrier beach

in New Jersey by means

r,love
Iilysius,

offshore
small

in
Donax

cores and seine netting. The dominant benthic organism and main food for the polychaet, Scolelepis the fish was squamata, which reached densi ties up to in a 40000.m-2 (=50g dry mass.m-2) band approximately 20m wide. Of 26 fish species only seven were common and one This species, Menidia menidia, the silverside, made up 85% of fish of numbers and 40% fish mass and of its diet. Scolelepis formed 70"-80% (1982a) recorded four fish species feeding on beach macrofauna, including
Leber

winter.
and

'.I.'heyfeed

various rays, other elasmobranchs, duckbill Hyliobatis spp., come into very shallow water to prey on Donax spp. The se ,
lither benthic prey.

Amongst

together

wi th lither large

rays are very

benthos.

import ant preda tors on the

As

resident.

in the case of flatfishes in I..;uropean waters, the <jreat success of skates and rays in these environments is probably due them to to their forage flattened in very shape shallow allowing water,

the silverside.

Donax and Emerita were

thus being able to take even the macrofauna that undergo tidal migrations.

the main prey items.

348

Romer (pers. cor,1m.) is food chains around surf

investigating phytoplankton

controlled by winds. six feeding categories, narL1e 11' : benthic feeders, planktivores, uetritivores, }?iscivores, herbivores and omnivores.
Herbivores

blooms. He has found that mullet t, Li za richardsoni, grazed directly on the phytoplankton and are taken in turn by large r predatory fishes. The mu llet t as well as other juvenile fish in the surf also prey extensively on zooplankton. Lasiak (1982, 1983) has made the most study to date of surf fish detailed seining on two Algoa Bay communities, beaches At over a period of three she years. recorded the less exposed beach

Lasiak

(1982) distinguished

owed their occurrence

to the presence of

some rocky reefs r,ear the sampling site at the less exposed beach, and were virtually absent at the more exposed beach. The mullet, Liza richardsoni, was the only detritivore, but in Lasiak's study fed mainly on zooplankton and may thus be considered a planktivore.
}ler simplified

66 species and 29 at the more exposed numbers site with teleosts dominating dominating biomass. elasmobranchs and Seven common species were resident while species intermediate were or 0ther sporadic in occurrence. The community showed no overall seasonal trends but a degree very high of short term variability which seemed to be la.rgely coupled to the effects of wind on the biological zone as several surf to parameters were closely correlated wind and intensity sampling. Studies over 24h clear trends though maximum
direction

groups feeding to include,

planktivores,

thus 0e benthic feeders, piscivores and omnivores.

may

At both 0eaches Iilotile organisms (mysids, prawns, fish, cephalopods and zooplankton) dominated the food base and these foods were taken 01' all feeding groups includirJg 0entbic feeders. Almost tunistic
summa rises

dll

species

employed

oppor-

feeding strategies. Table 4 the biomass, composition ared of this surf zone fish

food consumption
assemLJlage.

prior showed tended

to no to

abundance,

biomass occur

and around

diversity

often

consecutive variable.

twilight. days proved concluded

Sampling on to be very that the surf

~~estill know very little of surf zone icbtbyofauna and the danger of sampling in heavy surf will proLably keep the

She

zone fish assemblage was a highly variable community whose structure and were probably dominated by dynamics factors while biological abiotic interactions were of secondary importance. As most food organisms were highly motile, their availability would be strongly influenced by surf conditions which in turn are determined by 'wind effects on the wave regime. Hence short term variability is largely

To really resource partitioning and niche structure of surf zone fishes some
situation understand

that

\Jay.

more accurate means of sampling is necessary whereby tbe actual location of fish within can lJe made. the
could spatial three

uifferent perhaps
In of

parts
SCUBA

of the surf

counts are
the distriin in the terms zone surf of in

solution.
and lJe better

this way
fish understood of

bution

feeding

partitioning dimensions.
fish

this

However,
\"ill lJe more

zooplankton
important

as food for

349 Table 4. Biomass, composition and food consumption of fishes in less exposed surf (After Lasiak, 1982). zones in the East Cape. FISH Number of Species Feeding group Biomass g.m-l (dry mass) Dominants 78 60 10 10 158 3 4 2 2 Other 6 5 4 0 Species or group Macrobenthos
I-1y sids/pr awns

TOTAL PREY TAKEN Amount consumed g.m-l.y-l 269 657 200

Benthic feeders Planktivores piscivores Omnivores Total

L.ooplankton Fish Total

-B1
1798

at night and studies not employing after dark sampling will miss this. Surf zone ~ish communities with are few highly

there are few published reports on the ecology of birds on sandy beaches. Most papers simply record counts of waders and other birds along sandy shorelines. Fitch (1950) recorded gulls dropping

dynamic species.

assemblages

resident clams to break gull recorded

c 1 alllS
.

They are variable

in space and

time and short term variations in wind, conditions, are controlling surf probably major factor controlling a Most species show a these communities.
11igh

them and Brunton (1978) predation toheroa of (1980) dead veitch recorded beaches records (Kuyken, 1978). in New Zealand and have Leen made in and 1978) Britain Prater, Davies (1978)

birds similar Belgium

(CadLJury,

on

and

degree of opportunism in feeding, consequently all feeding types may heavily sChooling on zooplankton when and they crustaceans

predate larger

are locally abundant. Planktivores, and benthic feeders with opportunistic are therefore p1anktivore tendencies, a major component of surf zone
important are as Fish energy transformers in surf zones and, because they are highly motile, are also important in the export of energy from ichthyofauna.

sanderling overwintering on recorded beaches, British while Schneider, Harrington (1981) demons t ra tecJ depl et ion of benthic .i?reyat a migratory stopover Burger et al. (1977) for shorebirds. compared the utilization of an open beach, a sheltered ueach and a salt marsh several
ad j acent

by

migrating
to beaches,

birds.

~here

are
te rns

reports 1977;

of Lirds Davies, and

breeding 1981;

on or

the beach/surf zone system.

pa r ti cula r ly

(Fuchs,

Randall, (Summers,

7. BIRDS
brief mention of birds as predators on sandy Koepcke, 1952; beach benthos (Koepcke, Vlhile Brown, 1964; Leber, 1982; Vader, 1982) some authors have made

McLachlan, 1982) Hockey, 1980). In South Africa records

plovers

there are several count Pringle, for the western Cape.

350

cooper

(1977) conducted counts over 18

amounts
beach and

of

insects
cast

!Jlowing ashore.

onto
On

months which revealed eight migrant and seven resident species. Summers et al. (1977) reco rded higher bi rd numbe rs on the west coast than the south coast of the western cape, presumably due to greater food abundance in the former area. 99,6km They recorded 1267 birds over of sandy beacbes and 8319 bi rds shores. on the Twelve beaches

car r ion

the kelp

dominated

on the \ve st coast birds are the main predators and were to take estimated 40% of herbivore standing stock (Griffiths et al.,
1983 ). Hoc key eta 1. (l 9 8 3 ) s u mm a r i sed

beaches

the rdle of shorebirds

on sandy l.Jeaches

over 120,4km of mixed wader species occurred

with sanderling (Calidris alba), whitefronted sandplovers (Charadrius marginatus) and black oystercatchers (Haematopus Resident moquini) most important. 7,3km-l and species averaged

in Soutb Africa. Tbey found diversity and al.Jundanceof shore!Jirds to increase with latitude. Birds consumed 10-49% of production macro!Jenthic different in areas and return faeces, feathers
beach system.

significant and

amounts

of

carcases

to

the

migrants 15,2km-l in summer. Summers, cooper (1977) recorded fewer oystercatchers on beaches than rocky shores and Summers, Haltner (1979) recorded mass changes in waders, showing most species to be heavier in winter. On the Natal coast Joubert (1981) made counts

Moran, Fischelson (1971) recorded two plovers, Charadrius hiaticula and C. feeding alexandrius, mysid on the
Ga s t r 0 sac

plovers

sanctus. c u s 'J.'hey found tbese to bunt at random, regardless of areas

the size of the mysids, preferring

over 10 months, recording 14 species and densities of 39km-l, with waders making up only O,52km-l. Terns
roosted on the beach numbers. In the Eastern Cape (1980) counted birds over
wit h

in relatively

high

where mysids were densest. However, (1 S81) reanalysed Schneider their data to show that c. hiaticula selectively takes larger sid s . Schneider (1982) studied the foraging behaviour of
lilY

turnstones McLachlan on three 17

0f

(Arenaria

interpres)

feeding

et al. beaches species

0f

on Donax variabilis on a florida beach and Pienkowski (1982) described diet and feeding of two vlovers on a sand flat in
Northumberland. night and
hey '.c' fed d u r i n 9 cla Y and

12

months,
9e

recording

a v era

n u m l.J r s e

1 8 , 9 k 1<1-1

which

waders

estimated

bird

accounted biomass

consumption rates for

from four

They for 47%. and also food standard metabolic species which

volychaet worms and taceans dominated in the diet. et al. of in (1980) detailed ana lysed on

crus-

Myers

the l.Jeach

dominant
.

predation crustaceans

sanderlings

made up 95% of numbe rs From thi s they estimated that birds were taking 32% of available macrobenthic production from these beaches each year. This was mainly- in the form of sand mussels (Donax spp.), mussel siphons, and mysids. Birds also took significant

la!Joratory

bey showed how several ex per i men t s . 'i.' variables affect a sanderling's capture by prey Capture is determined rate. density, the sampling rate and sampl i ng

efficiency. penetrability

Sampling of tbe

rate

depends

on and

substrate

3S1 sampling size, inversely 10mm efficiency depth with prey depends and risk depth, on prey substrate varied very flow part
!Jetween

of

them. larger

The macrofauna are food web including their own

penetrability. or shallower

Prey being

zoopl ank ton, fishes and !Ji rds, while the interstitisal fauna constitute food web in the sand (McIntyre, Murison, 1973; McLachlan, 1977 a; a1.,1981a). 'l'he only McLachlan et links !Jetween

60% of prey

taken and

much less below 10mm. Larger prey were more vulnerable than smaller prey but small prey were never rejected. Prey risk was higher where prey organisms occurred close together. Capture rate could be predicted on the basis of prey size, density and sand penetrability. Fo!:" prey it was best to be small, deep in the substratum and not near other individuals. Prey species had no effect (Excirolana vs. Emerita). However, this was all done with dead prey. Most feeding although ac wrack was Jdone wi thout visual cues sanderling may use visual cues e.g. when feeding on beach Although probe locat ion by

these two food \4ebs is the common input of particulate (and probably dissolved) organics to both and the possible washing
(l-1U

of micro-organisms
tal., 1 9 78 )

from the sand

n roe

to provide food for

more sheltered In situations, however, deposi t feeders and Iuay take large sand swallowers in amounts et al. , amphipods of interstitial fauna. Oliver (1982) recorded phoxocephalid feeding

filter

feeders.

times,

insects.

'liasrandom it could also be affected the feeding patterns of nearby birds.

Hading sa.ndy birds

on a var iety of foods including nematodes and Cro ker (1967b) recorded a baustorid umphipod feeding on meiofauna, both cases being in sheltered lagoon sands. On beach Alheit, a (lS82) juvenile Scheibel recorded demersal fish feeding copepods. as BOl-/eve r , are mostly negligiule on look separately at the on
open
Iliac rof

harpacticoid exchanges

are

impo

rtant

preda to r s

on

these

beaches,

being

efficient during

croppers of

beaches
auna

of

~ne

macrobenthos

periods

food

lov: :.ide. On beaches where high inputs concentrate macrofauna of macrodebris along the drift line above the reach of ~arine predators, bi rds may be the most impo rta:-, t predators. Even bi ras that may be or.ly beaches :oos:. the on important 'rdhe the faecal deposits of re large flocks may add significant of nutrients locally. amounts

web and interstitial energetics.

8.2 t-iacrofauna

Several

authors

have

Itlen t ioned

aspects

of 1Jredation or food chains sandy L>eaches (Pearse et Bonnet, 1946; Koepcke, HedSJ?eth, 1957; Brown, a1., 1970b; Philips, 1973a, 1973b; Reilly,

relating to al., 1942; 1952;

Koepcke,

1964; steele et 1970; Edwards, Bellis, 1978;

8.

FOOD HEBS AND ENERGY FLOW

8.1 Introduction The ~acrofauna and interstitial open sandy beaches form two communities with little fauna of separate no energy

Ansell et al., 1978; Reilly, 1979; McLachlan et al., 1981a; Leber, 1982; Gianuca, 1983). Macrofauna food chains start and end in the sea, wi th the land playing only a negligible rele, mainly
i n t 11e t 0 rTi1 0 fin s e c t s b low i n g 0 n tot 11e

or

beach and land crabs, bi rds and mammal s

,,~') .:J:) -

moving onto the beach to feed. Open beaches are characterized by the absence (Brown, 1964) and of attached plants primary production by the benthic microflora is generally negligible. On temperate beach, however, a sheltered (1968) found viable Steele, Baird

herbivorous fishes. (1978) uistinguished

macrofauna, detritivores
f.1cLachlan

Ansell two
(2)

et
':jroups

a1 ., of and
only

(1) and et

herbivores
separated

carnivores.

0.1., (19810.)

filter

feeders

and

scavengers/predators

diatoms mixed to 20cm depth in the sand primary production but low and They measurable. considered vertical mixing due to wave action to be the main the shallow limiting factor. In beaches, subtidal off two Venezuelan Edwards benthic (1973a) primary recorded production
significant

on L;ast Cape Leaches, where l1eposit feeders were of IJesJligible importance.

On 0.1.

kelp

uomi no. ed t and

Leaches

Griff iths et
herbivores,

(lS83)

uistinguished

carnivores
possible of

filter
and sandy

feeders.
trophic

'l'he

oou

sources

':Jroups

on listed in ~able 5.
macrofauna

Geaches

are

due

to

both ~re extremely important in surf water, altbough largely ingored to uate, '.,ould fall in cate(4) . On TIIOst sandy (1) and sories beaches toocJ source (1) is 01' far the most important, with food sources (3) (4) Leing and next. On sheltered shores, and
0 f

micro- and kcal.m-2y-l

macro-algae (2943 and 6500 beaches) two but at the se areas are hardly typical of open beaches and the latter area VIas subject primary production to heavy pollution.

zooplankton,

yvhi

ch

may very column be water the in important, particularly where surf phytoplankton blooms occur. Unfortuna::.ely primary

however,
impo

particulate and fauna

once

cJetritus

production bloom

figures have yet

a surf
be

phytoplanktor.

for to

living
Til s t 0

filicroflora

may

oe

primary

Edwards in the water beaches off Venezuelan 2934mgc.m-2.d-l.

publ i shed. (19730.)

estimated

prodllction

column
as

in the sediments, and unuer such conditions deposit feeders sucl1 as Arenicola,
incorporatec1
Callianassa

rtance

and

Scolelepis

may

be

abundant.

Different sources
C:: v'i"a:ls.

authors have partitioned food fo r sandy beaches in a variety

Bro.vln (1964)

Hhile
Le<1ches

inputs
Lave are

or particulate
yet to
lJe

Til at

to some

JdeaSUreu,

distinguished

an

estiTi-lates requirements

available

Lased
TLiacrofauna.

on

the In

supply and a erratic of macrodebris supply of particulates while regular Hayes (1974) recognised dissolved and organics, stranded microparticulate plankton and large organic masses. Edwards (1973a) distinguished three main pathways by. which energy travels from primary foods to fishes; (1) via zooplankton to plankton feeding fish, (2) via benthos to demersal fishes and (3) algae to benthic directly from

of

the

India particulate 56141 kJ.m-2y-l 4291 kJ.m. -2y-l and 38765 m-ly-l at tively.
accounted consumption

feeCiers accounted for for and carnivores beach on Shertallai 235kJ. respecthus

kJ.m-l.y-l and Cochin beach particulate for feeders

1978).

the food Sl% of ., on average (Ansell et 0.1 In tbe L;ast Cape, filter feeding were estimated to assimilate

macrofauna

353 Table 5. Food sources and trophic groups of macrofauna on open sandy beaches. FOOD SOURCE 1. 2. 3. particulate organics including phytoplankton Deposited detritus, benthic diatoms, interstitial fauna FEEDING GROUP Filter feeders Deposit feeders
Scavengers

Stranded debris - plants animals Other fauna

herbivores carnivores

4.

Predators

109421

kJ.m-.l.y-.L,

93%

of

the

total with the

the beach for scavenger/predators

macrofauna,

and fly larvae (Muir, 1977; Koop, Field, 1980, Stenton-Dozey, 1981; Griffiths, 1980; Griffiths, Stenton-Dozey, 1981 ; Koop et al., lS82a; Griffiths et al. , 1983). Biodegradion of macrophytes by the macrofauna In sheltered amounts of
deposited: high

making

up

remai ning 7% (McLachlan et al., 19 81a) . / This does not include zooplankton organisms feeders and which very are
abundant.

mainly filter Scavengers

is thus very important.

are only important where there are large inputs of debris or where filter feeders physical conditions or the absence of food (e.g. East Coast of South Africa, Dye et al.,
a reabsen t becau se of

unstable

localities or where large particulate organics are deposit feeders rllay reach

1981; Wooldridge et al., 1981). Huge inputs of macrophytes characterize

(Eleftheriou, values McDermott, 1983) On 1976; most beaches organic content of the sand
lJiomass

MCIntyre,

is beaches adjacent to kelp beds or similar areas. In southern California Hayes (1974) estimated these inputs to amount to 473kg wet (ca.78kg dry) algae -1 -1 m.y (Macrocystis), while on \Jest Cape beaches adjacent to kelp beds

very

low,

e.g.

0,2-0,4mg.g-l

sand

in Scotland and India (Steele, 1976) and undetectable amounts in the \Jest cape and Bast cape (BrOvlD , 1971a; McLachl2.n, 1977a)
.

Predators on the other benthos include crabs, starfish and gastropods. These normally abundance, occur making in up relatively 5-10% of low biomass.

values range around 2000kg

m-l.y-l a1., 1983; Hayes consume input, this (Koop, Kield, 1982a; Stenton-Dozey,

wet Ecklonia
KOOp et Griffiths,

1980;

Griffiths et aI, 1983). While (1974) found Tylos punctatus to only 4-5% of the for estimates taken by edible kelp macrofauna amphipods

As many are supralittoral forms, e.g. Ocypode, they may be the only macrofauna where condi tions are too dynamic macrofauna to survive for intertidal (Gauld, Buchanan, 1956; Dye et al., 1981). swash

consumption being

in the West Cape range 9-74%, isopods,

'l'heygenerally take both living benthic prey and stranded carrion.

354 Energy flows through different species These food chains end in fishes, birds, crabs and other invertebrate predators very man. iew studies and have estimated different juvenile ant the flatfish importance are the of these predators. cropping Scotland In most importsiphons of

and trophic efficiencies vary widely. Turnover (P/B) ranges from 0.3 for tenuis on a temperate beac h Tellina (Trevallion, spiculum on 1971) a to 10.3 for Donax beach tropical Indian

(Ansell et a1., 1972b). Ansell et a1., (1978) found turnover on these tropical beaches generally ten times higher than on cold temperate Scottish beaches. Edwards (1973a) recorded average P/B's macrobenthos, of 2.8-3.5 for the t-1cLachlanet al., 0.5-5.0 molluscs for and (1981a) used P/B's of production of Koop, crustaceans, and the P/B for Ligia somatic

predators,

bivalves, palps of polychaetes and whole prey. Bival ves may then put more ene rgy into regeneration of siphons and less into reproduction. Trevallion (1971) up to 1.3% of total estimatec.1 that energy annual expenditure in 'J.'ellina may go into regenerating tenuis siphons. the most In India crabs were considered
important predators, with fishes and man less impor tan t and birds virtually absent (Steele, 1976; Ansell

Field (1981) estimated

dilatata at 3.75 times per year. Assimilation 100%) vary efficiencies widely, commonly (A/C being X very

et al., 1978). Edwards (1973a) estimated fish to account for -/1-8996 of

production accounting at for his 13% t'vVD beaches, on one and starfish crabs for

low for scavengers/grazers feeding on abundant macroalgae, e.g. 64% for 'rylos (Hayes, 1974), 5.5% for Ligia (Koop, Field, 1981) and 30-50% for Talorchestia and fly larvae, (Stenton-Dozey, 1980). Koop et a1., (1982a) average overall estimated an efficiency of for assimilation 22% invertebrate kelp grazers while Ansell et a1., (1978) and Edwards (1973a) assumed assimilation efficiencies of 80% Griffiths,

on the other beach. McLachlan et predation al. , (lS81a) partitioned on east cape Leaches. Birds took 32% of intertidal macrofauna production al., 1980), fishes were (t-1cLachlan et estimated predators to take 55% and Genthic (mainly crabs, Ovalipes) 10%

13%

with Elan accounting for less than 3%. DU Preez (1981) has subsequently made a detailed of
La

for

carnivores

et Ansell efficiencies through the as 15-16% detritivores

and 60% for herbivores. (1978) estimated a1. , of transfer of energy

chain (P/ A X 100%) for herbivores and and 25-30% for carnivores food

investigation

of the predation

Ovalipes
s i a k

fish
beaches

on Donax and Bullia and and Rossouw (1983) have studied predation. kelp On dominated
of the Hest Cape, 110weve r, bi rds (Griffiths

in India while Edwards (1973a) estimated p/ 17% C X 100% as 15% for benthos, for demersal fish and 11-15% for other fish at his unpolluted site. Ecological efficiencies calculated from et a1. (1981a) were 9-19% macro-fauna with a mean of 17%. McLachlan for the

are the major a1.,1983).

predators

et

Evaluating our knowledge of sandy l>each food chains as a whole is made difficult by only not the small number of quantitative studies, but also the absence of data on zooplankton. The

355 macrofauna cannot be considered in isodescribing these food lation when chains. production, Surf zone also untimately zooplankton falling prey To conclude this section on macrofauna energetics it appears that five main beach types can be recognised in terms of macrofauna energy flow patterns (Table 7). Information on surf zone zooplankton in different zoogeographic regions is unfortunately not available to allow incorporation into this scheme. It may Le expected, however, that as zoopolankton and macrobenthic filter feeders utilize the same food resources, their
similar

to fishes and being exported from the system, is probably more important in terms of energy for example, surf flow. In the East Cape where we have some idea of

zooplankton biomass, zone a pattern emerges if larger different mobile crustaceans, such as mysids and prawns, are included with zooplankton

total

biomass

vv'Ould Le

roughly

out to 500m from the beach (Table 6). Table 6. Summary of approximate faunal dry biomass for an average beach and Cape. surf zone East (From in the McLachlan, 1983) BIOMASS SJ .m -1

in filanycases.

8.3 Interstitial Fauna Less attention bas

been

paid

to

interstitial energetics, early as 1942, Pearse et that the interstitial beaches

functions,

although 2.S al. , proposed systems of sandy

CA7E;GORY

had c1igestive and incubating the nlicrofauna mineralising

!'Jacrobenthos

1500 1100 5 300

Zooplankton

Birds Fishes

materials in the sand. Little was made subsequently to evaluate or quantify this until quite recently. system This consists typically of bacteria, protozoans and organic attempt meiofauna constituting their own tood web in the sand.

The food chain is centred around filter feeding feeding benthos and zooplankton particulate detritus and phytoon plankton and being taken in turn by fishes, birds and crabs. As zooplankton turnover is 6-7 times tbat of the macrofauna, the centre of gravity of the food chain is clearly in the surf and not on the beach. HcFar land (1963) came to a similar conclusion, showing that plankton was the predominant fish food the The off Texas beaches, greatly exceeding benthos as an available food source.

Dissolved and fifle }?articulate organics form the Lase chain, of this food although in sheltered situations the benthic microflora might also play a role. McIntyre et al., (1970) were the first to demonstrate that interstitial

fauna '2.)uld subsist on uissolved organics. Using two experimental sand columns, they maintained populations viable microand meiofaunal in the second

column on the effluent leaving the first column. They suggested that bacteria were the prime utilizers of uissolved
organics, utilizing

surf zones he studied were characterized by high primary productivity in summer.

these

org ani c

mo le-

cules wben they adsorb

onto sand grains

356 Table 7. Major beach types in terms of macrofauna energetics. Predominant Sources Food Dominant Trophic Group Filter Feeders

I-1 c r 0 f au a

n a 1

Type 1. High turnover, particulatebased food chain

2. Low turnover, particulate-

Latitude Tropical

Biomass Low to moderate

Predators

Particulates

Invertebrates Fishes

Temperate

Particulates

Low to high

Filter Feeders

Fishes,

Birds

Invertebrates

based food chain. Temperate Stranded macrophytes

r:Joderate

3 .

r.1 c rod a

e !J r i s

Scavengers/ Birds Grazers

based

food

chain. 4. Sediment-based food chains in

sheltered
1'empe rat e

Deposited detritus microorganisms and benthic microflora

or Tropical

Low to high

Deposit Feeders

Birds,

Fishes

Invertebrates

beaches. 5. Carrion-based food chains in

extremely posed exsteep

Temperate or Tropical

S t ran

d 2

(I

c 2. r r

ion

:",0'(/

SCQvenge~s/ Birds, Crabs

Predators

beaches. where they can be digested with extracellular enzymes. The meiofauna in turn fed mainly on these bacteria, accountinCj for approximately 5% or bacterial consumption.

some
directly dissolved

1.1 i 0 f au e

n a 1

species

may
!Jroad

feed

meiofauna

or in a sense <10 form the top of the interdetritus, microflora

on

organics,

Fenchel (1972) considered bacteria the only primary decomposers of particulate plant detritus with their decomposition rate limited by Consequently he trophic levels oxygen and nutrients. suggested that higher food

stitial food we!J as most species feed on bacteria, protozoa or other meiofauna (McIntyre, Murison, 1973; McLachlan et 1981a). ale , The position of protozoans in this system has !Jeen little studied in open beaches; \/h i 1 e some forms may have a similar trophic to status are even predatory on meiofauna. However, in a broad sense they probably occupy a .tJosition interbacteria,

in the interstitial

others

web are dependant on initial processinCj Although of plant detritus by bacteria.

357 mediate between, and overlapping with, bacteria and meiofauna in interstitial food chains. Three within herbivores trophic the and types may be
recognised predators, with only

(1977) and Dye (1983).

Another approach sand

has been the stuuy of experimental columns in the laboratory. Several authors

meiofauna, bacterivores,

have maintained laboratory sand microcosms for extended periods (McIntyre et al., 1970; pugh, 1976; Boucher, et al., 1977; McLachlan Chamroux, Hormald,
19S1c)

the Hydrozoa and Turbellaria exclusively predatory (Tietjen, 1980). Swedmark (1964) however, listed four categories, predators, diatom and epig rowth feeder s, detritus Besides the side red eaters direct that and suspension of feeders. on congrazing meiofauna (1980)

1976;
~tirling,
.

Chamrou x

microfauna,

Tietjen meiofauna

stimulate

al. , circuit system has been maintained for 16 months on soluble amino acids (Boucher, Chamroux, 1976) confirming the utilization of dissolved by microorganisms organics and their subsequent However, structure indicated soluble
the In only this

et

1979; closed A successfully

bacterial activity in the interstitial system by (1) mechanical breakdown of det:itus, bacterial nutrients secretion bacterial conveying making for c010nisation, to it more susceptible (2) excretion of (3) use, microbial

consumption by the meiofauna. in meiofaunal the via

}?ath to and

changes that organics

trophic case

community diversity) of not

were

(decreased
!Jacteria

utilization was
the

of mucous thereby attracting (4) growth and movement nutrients and oxygen. efficiencies (A/ X C have been estimated

llIeiofauna.

!Jacteria

meiofauna

estimated the carbon proportions (1970). There

Assimilation

100%) for meiofauna

to account for 95% and 5% of input respectively, the same y r e as f 0 u n d by 1.1 In t c eta 1 .

in the range 6-26% while transfer efficiencies (P/ X 100%) were A for three nematode species 79-97% (Tietjen, 1980). For turnover (p/) B a ratio of 10 has been widely used for meiofauna
Because 'working of

have !Jeen few detailed

studies of in situ un -cases that

interstitial energy budgets open ueaches. The three

(Gerlach, 1971).
the d iff icul ty minute of scale when of

with

the

inhabitants

w'arrant attention concern a comparative study of Scottish and Indian beaches, studies on high energy beaches in the Cape Eastern and studies on beaches
...

fauna, investigations the interstitial energetics interstitial have of generally taken the form of 'black box I studies has of tbe systertlas a \Jhole. This been or done by oxygen measuring consumpbenthic commonly

receiving

high

kelp

inputs

in

the

Western Cape. Munro

et

al.

"(1978)

Illade

a
of

detailed

comparative stitial" beaches beaches sand The

investigation on

interstitial tion in partition

dynamics

tropical

interIndian

situ and then attempting to this. Benthic oxygen consump-

(Md 150-190u!l1) and Scottish (Md 250um) using small and large

tion measurements have been discussed and reviewed by Lassere (1976), Pamatmat

columns and in situ measurements. -tropical epi}?sammic beach lacked

35'S

diatoms which were common on the temperate beach. Dissolved organics were estimated to make up 80% of the input on the Scottish beach and 39% on the Indian beaches. Interstitial community respir(164gC.
vJinter

organic levels were 500-1000 ug C.l-l in Scotland and 900-3700 ug .1-1 in

India. I-loweve r , surf flushing of the Indian ueaches way remove significant
quantities of bacteria, thus making them available to macrofauna filter feeders and decreasing the quanti ty availal:Jle to
the meiofauna.

ation on the tropical beach m-2 .y-l) was nine times the values and twice the summer the temperate beach (mean y-l) with microbial estimated to be an 72 and former 15g.c.m-2.y-l

values for 42gc.m-2. production in the

gc.m-2.y-l

average annual of (using the latter in respiration figures based on an assumed conversion efficiency of 45% and the assumption that the contribution of Meiofauna meiofauna was negligible). biomass was, however, much higher on the Scottish the 8) . beach (273-523mg. m-2) than Indian beach They postulated (24-60mg.m-2)(Table that more vigorous

In the Last Cape the interstitial fauna consists essentially of meiofauna, protozoans and bacteria to considerable depth in the sand (Dye, 1979a; NcLachlan et al., 1979a).
f 1 uc t ua t ions in

Dye (1980) showed tidal

uenth ic oxyg en consuTIlj,.!-

tion as a result vf tidally induced changes in pore moisture and interstitial water flow, lowest values being recorded when the sand dried out. Dye (1979a) investigated biological oxygen demand in sand cores in as these gave higher the laboratory values than in situ techniques. He showed that caused disturbance a significant increase in oxygen uptake. study Dye (1981) more cJetailed partitioned this oxygen uj?take l:Jetween bacteria, protozoans and meiofauna on two beaches (TaLle 9). In a later beach sand, using intact

flushing of the sand on the tropical Indian beach both boosted respiration bacteria from the sand and striped grains. tropical beach received more The organics and had a higher interstitial consumption rate than the Scottish beach as a result of increased water flow by wave flushing. Surt oxygen

~able 8. Biomass, production and respiration of interstitial organisms on Scottish and Indian beaches. (After Munro et al., 1978). All values in gc.m-2.y-l. INDIA Production 0 72 2 SCOTLAND Biomass
0 31 0.02 4

Respiration Algae Microfauna

Meiofauna

Respiration

Production
7 15 2

Biomass

0 160 4

0.2

359 Partitioning of interstitial oxygen uptake on two East Cape beaches. (After Dye, 1981).
MODERA'l'ELY EXPOSED BEACH

Table 9.

Biomass %

02 uptake %

VERY EXPOSED BEACH Biomass % 02 uptake %

Meiofauna Protozoa Bacteria

75 19 6

18 18 64

79 17 4

21 24 55

In the West Cape very rich interstitiul populations develop on beaches wi th high Koop, Griffiths (1982) kelp inputs. recorded dry biomass of 241, 200 and 663 g.m-l of macro- meio- and microfauna respectively. Bacteria were concentrated at lower tide levels, probably subsisting on small particles of broken down
fa.eces Koop

rap i d 1 Y .

1:' beg

rea t

s i g n i f i can ceo

these (1974)

microbes recorded in

indicates why such little

\-lhere

Hayes kelp

breakdown

looked at only one grazer, Tylos Dunctatus.

California

he

kelp of

resulting from feeding and macrofauna and Ti1eiofauna.

8.4 Conclusions It may be concluded that intertidal sandy beaches are important in the processing ot organics whether these be as DON, from POH or larger animal and plant toods genere.l1y remains. These COllie

flow much

carbon a1. , (1982a) quantified Bacteria on through this system.

et

the stranded fronds were responsible initial degradion. Breakdown

for of

the kelp resulted in high concentrations of leachates (up to 5640 mgC.l-l) in the sediment below the kelp. Over 90% of this was utilized by bacteria after seeping through 1m of sand and 23-27% of carbon in the kelp was converted to bacterial carbon. The ratio of increase

the sea and in turn most production returns there. The interstitial fauna alv;ay s more important are than the macrofauna of in this mineralisation organic materials, even wbere a rich macrofauna occurs (Table 10). The
interstitial

system

is

Criven

by

in bacterial biomass to kelp carbon used giving conversion a was 58. 7% , efficiency of 29.4% if bacterial biomass is
~1u h c

50% carbon
of the

(Koop et al. , 1982a) faeces of invertebrate

soluble and !?articulate being via the main }?ath bacteria attached to sand grains to meiofauna. The intermediate rale of protozoans is uncertain. 0n exposed
organics,

input s

of

grazers were also converted to bacterial grazers these have low carbon as assimilation
remaining mineralised

beaches there is little excbange between this sytem and the mac rofauna although the importance of microorganisms flushed from the sand as food for the lilacrofauna needs investigation. further The concentrations of meiofauna ilushed from

efficiencies. The kelp was of the 73-77% by quite microbes the

360

Table 10. Approximate partitioning of total benthic assimilation between filacrofauna and interstitial fauna in four geographic areas based on data in Munro et al. (1978) (1982). Ansell et al. (1978), McLachlan et al. (1981a) and KOOlY, Griffiths,

ASSIMILATION LOCALITY AND BEACH TYPE Macrofauna

[kJ.m-l.y-l (%)] Interstitial fauna BEACH WIDTH (m)

Scotland, temperate

30 000

(23 )

S9 000

(77)

60

India,

tropical

427 000

(38)

703 000

(62)

60

East Cape, warm temperate

116 000

(37)

236 000

(63)

GO

West Cape, temperate with high kelp input

42 000

(3)

1599 000

(97)

63

exposed beaches here sand of the measu~ed by McLachlan (1977a) and found On sheltered beaches to be negligible. a greater size spectrum of organisms may occur, filling the gap between interstitial fauna and larger macrofauna and thereby facilitating energy flow by predation. In such environments deposit feeders may also ingest significant quantities of interstitial fauna. Inshore ments zone food are chains in beach environhowever centred in the surf than the intertidal, even

9. may

NU'I'RIE;N'l'

CYCLING
been specula ted tha t beaches in recycling importance

It has

long
(jf

Le

organic mineralising materials coming from the sea (Pearse et al., 1942) . This is effected Ly the nutrients

Ly

consuming fauna including nitrogen then e.g. excreting this

orSjanic anu in

nutrients

phosphorous and inorganic form,

some areas PO4. In NH4' groundwater seepage can, bo\vever, also supply considerable amounts of inorganic nutrient via the Leach to the surf zone. This can take the form of Goth artesian aquifers ariel 'jeneral seepage around the
area

rather

where rich intertidal macrofaunas occur. Our area of least understanding at the moment is these dynamic food chains of the surf zone and in particular the role zooplankton and larger motile of
crustaceans.

meets this

the permanent water taLle the sea. Johannes (lS80) fauna to Le very important in Hestern where
\,her e

Australia

submarine

grounell,Jater

361 discharge ranged 1-5m.y-l and connitrogen (Chamroux, 1965; McIntyre et al., 1970; pugh, 1976; Boucher, Chamroux, Stirling, McLachlan 1976; 1979; et is Cahet, Munro the 1976; Wormald, et al., 1978; Gener ally activity

tained up to 380 ug at 1-1 N03-N. Nutrient regeneration by the macrofauna, zooplankton, birds and fishes has been virtually (1979b) and respectively excretion in Siliqua and Washington and ignored. Lewin et al., (1983) ammonia Prosch, McLachlan investigated

al. ,

19 SIc) . dominant

nitrification

in Leach sand and organic nitrogen is efficiently mineralised to nitrate. Increasing organic loading tends to raise equilibrium levels and it is usually only at very high levels that a beach can not cope and anoxic conditions and ammonia appear (Oliff et al., 1970; pugh, 1976). Boucher, Chamroux (1976) recorded total Itlineralisation of amino Munro acids adcJed to tbeir sand columns while et recorded al., (1978) 70%

bivalves of the genera Donax on beaches in the East Cape, Sou tb

Africa. In both cases very large populations of these bivalves develop and nut rients regene rated by them could supplying significance in be of phytoplankton requirements in the surf Hayes Tylos to (1974) found zone. only on 4-5% of kelp input beaches while Griffiths, Californian Stenton-Dozey (1981) and Koop et al. consume (1982a) estimated kelp was that 9-74% of stranded by the macrofauna. consumed

organics, mineralisation of natural mostly in the top 5cm of thei r columns.

~1cLachlan et mineralisation
65% when amino these

al. (1981e) found 35% of natural organics in ancJ this

were

50em sand columns

increased

to

Not all of this is assimilated, however! may efficiencies and low assimilation comparatively a small result in being kelp nitrogen prop6rtion of regenerated directly by t.he macrofauna. The contribution of other faunal components must also be important in this regard but have yet to be investigated.
Generally .3andy
important

acids

acJded.

Most been has

of

sanc.1 column artificial to as in

.studies

have f 1 O\J flow

somewhat been

water situ

unrelatecJ

rates, gence, rather

there has been continuous submerand inputs have been constant than tidal or pulsing. Except-

the
in

interstitial
has been

system

of
more

beaches

thought

nutrient

regeneration

the

macrofauna.

Water

filters

than through

ions are pugh (1976) and McLachlan et al. (lS81c), although even these experiments Jid not incorporate pulsing flows. Filtration rates have ranSjed 25-1000 1.m-2.cJ-l, covering a variety of flows both intertidal and subtidal.
t.lcLachlan

the sand, as a result of tides and waves both in the intertidal and the subtidal, and DOM and P 0l'1 are removed Ly tbe interstitial fauna and mineralised. Several studies of experimental systems

earlier sand col umn \vor k with a model of water filtration and nutrient regeneration by sandy ueaches. This was based on a regression model a predicting of ancJ filtration particle range. tide as function slope sand

(1982)

followed

up

of sand columns or model beaches have looked at mineralisation and/or nutrient mostly concentrating on regeneration,

volume size,

beach

362 Coupled degree

to th i s \Jas an est ima te

of the in

of mineralization

of organics

to rocks and subject to larye kelp inputs are interesting for comparison. It may therefore be concluded that sandy beaches can mineralize most oryanics tl1ey receive, the l>ulk of this l>e ins
done very efticiently fauna. (80-100%) This l>y the interstitial

relation to sand particle size and the average distance. The filtration filtration distance is about 35% of the (Riedl, distance intertidal horizontal Machan, a 1972) and average mineralisation At rate of lO41.m-l.d-l was estimated to be at least 60-80%. filtration

is water

accomplished

uy

the l?rocess of

and an organic nitrogen level of 0.15mg. 1-1 this predicted regeneration of 0.9g N03-N m-l.d-l. An attempt to by by estimate total beaches in the nutrient production East Cape was made

filtration through the ueach and will be greatest where steep beaches and short period waves result in high filtration volumes (e.g. in the tropics) . Hhere vast macrofauna populations occur tiley
also
contrilJute

McLachlan et a1. (1981a).

They estimated

may

significantly

to

nitrogen excretion by the macrofauna and interstitial fauna of the beach and surf 350gN.m-l.y-l 591gN. at and zone -1 .y-1 respectively. on Thus, even m as these with very high beaches such biomass, the interstitial macrofauna fauna are more important in nutrient regeneration, accounting for 63%. Hhile figures approximate, the above only suggest that the mineralising activities the of the benthos alone can replenish nutrient pool of the surf zone in a matter of days or weeks. Koop bial et al. (1982b) quantified micro-

nutrient birds and

important.

regeneration.

Zooplankton,

iishes

must

also

ue

very

1'-10st organic

nitrogen
0.

and will

phosphorous rapidly

supplieCl

to

IJeach

ue

to returned the surf as nitrates, ammonia and phosphates. Thereafter the fate of these nutrients will depend on physical conch t ions in the surf 20 ne . Hhere cellular circulation patterns occur there may Ge a high degree of retention, coasts or reflective whi Ie on irregular
beaches,

exchange

VIi th the open sea may

regeneration of nitrogen from stranded kelp over an eight day cycle. In this time the microbes incorporated

Ge much more rapid. As I.lany surf zones are characterized l>y high primary productivity, utilization of Geach generated nutrients in the surf 2.one way be very important and is in need vf more Hayes (1974) and Koop et al. attention. (1982b) consider such Ilutrients of impurtance little in the surf Vlbile McLachlan et al. (1981a) suggest that these nutrients may l>e sufficient to surf phytoplankton Glooms on maintain long open beaches. 10. This
TmJARDS A l-10RE;

94% of the kelp nitrogen, which normally strands at an average input of l2gN.m-l.d-l. However, in the long term the beach can not act as a nitrogen sink and this must return to the sea. In this case this could supply roughly one quarter of the nitrogen of phytoplankton in the requirements kelp adjacent

beds. Hhile hardly typical of open beaches, even on the west coast (Bally, 1981), these figures for beaches adjacent

revieVl

has

HOLISTIC APPROACH attempted a functional

363

look at our present knowledge of the ecology of beach/ surf zone env ironment s as a whole, fauna and incorporating flora, all aspects of flow and energy

Energy flovl and nutrient cycling in a long open lJeach typical of the Last Cape is illustrated in rig. s. It This is a of rich system energy and producers (benthos, decomposers Energy flows to be boundaries.
\v'i tb

considerable

flows

nutrient cycling. published studies

Unfortunately, most of beach environments

have not attempted such a holistic approach and have tended to investigate one or more components in isolation. In the East Cape research on beach

includes (phytoplankton), consumers zooplankton, fishes) and

I!utrients.

(interstitial fauna). within this system appear than across its However, although exchange

greater

envi ronments attempting a the nutrient ba sed

1 9 8 Oc ) :

has been unique in holistic approach towards quantification of energy flow and cycling zone under the concept This body of a is
ecosystem.

across the dune/beach boundary is negligible, exchanges across the outer boundary
quantified.

of tbe surf cells remain to be

'.L'hemairl trophic patbway is

beach/surf

on two contentions (1) that the sand with

(HcLachlan,

from tine organic materials (POM phytoplankton) via L.ooplankton filter-feedin';j Ti\acrofauna to The interstitial
important

and and

beach

together

the water

of the envelope

fishes. and Dm1.

nitrogen

fauna is also extremely POM

and of inorganic beach

of the surf zone form an ecosystem where internal energy flows are greater than exchanges across boundaries and (2) that boundaries, this system has definable these being
mar

in

mineralisin<;J

The

total

c.mount
lJy the

regenerated

surf

60 ne

fauna is not yet known, Lut must De SUDar,d interas the Iliacr of auna stitial iauna alone regenerate can enough to replenish the zone surf nutrient pool within a few weeks to days. Altbouyh not indicated in Fig. 9,

1 and via r d

line on the the drift outer limit of and the gin

stantial

surf circulation cells on the seaward margin. The central concept is that the circulation cells that typify the surf zones 1973) of open beaches (Inman, Brush, imply a certain measure of

wave action probably represents tilemost important factor controlling this system, as it is responsible for transporting water, cells sand. not only biological ancl needed to information is understand beach/surf ecosysteIlls. Ijater chemical movement, processes the. key
occurring

integrity retention

or

surf

zone

water within

and this

zone. that

of This is in opposition to the idea high energy beaches are purely

materials

materials,

sediment biological and driving surf circulation and pumping water through Leach

between sea interfaces and land. Clearly, however, this can not apply to poc ket beaches, lJeaches assoc iated with

Clearly,

rocky shores or beaches dropping directly into deep water where there is
no real .surf zone. The kelp input beaches of the South African west coast, example, as interfaces function for between adjacent kelp beds and the land.

to here,
surf

all dynamic is the realm

'.i.'he circulation

of the pbysical discreteness of boundaries and

oceanographer.

exchanges

across

them

364

SEA
I

SURF
EXPORT

BEACH
OTHER PREDA TERS 1837
3684

DUNES

8861

I
I 2468

45:70
IMPORT 8337 600
1

EXPORT 2155

I I I I
.
=

I I
:

I I
I

.
I

I i

t I

!
I !

8-t I
I I
236325

:612~
. 89269

I I I \ I \1 ~ I I ... i I

I I I I i I I I
J ~
LINE

DRIFT

FIGURE 9. Energy flow (solid lines) and nutrient cycles (broken lines) for a typical East Cape beach, largely after McLachlan et al. (1981a). All values in kJ.m-l.y-l except nutrient values in g inorganic N. need to be quantified. essential that physical biologists Beaches need
study

It is therefore oceanographers, work this together. areas only must fashion. geographic

ecosystems,

evaluated
knowledge environment.

and this only can be in the presence of a detailed of the physical and chemical

and be

chemists in

in

different studied

to
the

Biologists

in particular must not intertidal fauna, but

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venture out into the surf zone. In the dynamics organic of fine materials, phytoplankton and zooplankton lies the key to a more realistic understanding of the function of sandy 0each/surf zone

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Amspoker, A.M. of inte r tidal Sc r ipps Beach, U.S.A. Bot. Mar.

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Ansell

366 part 1. Sarsia 39, Southern Florida. 17-105. Brown AC (1964) Food relationships on the intertidal sandy beaches of the Cape peninsula" S. Afr. J. sci. 60, 35-41. Brown AC (1971a) The ecology of the sandy beaches of the Cape peninsula, South Africa Part I: Introduction, Trans. Roy. Soc. S. Afr. 39, 247-279. Brown AC (1971b) The ecology of the sandy beaches of the Cape peninsula, South Africa part 2: The mode of life of Bullia (Gastropoda; prosobranchiata), Trans. Roy. Soc. S. Afr. 39, 281-319. Brown AC (1973) The ecology of the sandy beaches of the Cape peninsula, South Africa Part 4: Observation on two inter-tidal isopoda, Eurydice longicornis (StUder) and Exosphaeroma t runcati telson (Barnard) , Trans. Roy. Soc. S. Afr. 40, 381-404. Brown AC (1982) The biology of sandy beach whelks of the genus Bullia (Nassariidae), Oceanogr. Mar. BioI. Ann. Rev. 20, 309-361. Brown B (1982) Spatial and temporal distribution of a deposit-feeding polychaete on a heterogeneous tidal flat. J.exp. mar. BioI. Ecol. 65, 213-227. Brown R (1981) Saccocirridae (Annelida: Archiannelida) from the central coast of New Sou th Hales, Aust. J. t1ar. Freshwat. Res. 32, 439-456. Brown AC and Talbot MS (1972) The ecology of the sandy beaches of the Cape Peninsula, South Africa Part 3: A study of Gastrosaccus psammodytes Tattersall (Crustacea: Mysidacea), Trans. Roy.
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