Download Ecology of tropical soft-botlom benthos

Ecology of tropical soft-botlom benthos: a review with
emphasis on emerging concepts *
Daniel M. A10ngi
Australian Institute of Marine Science, P.M.B. No. 3, Townsville M.C., Queensland 4810, Australia.
(Ree. 9-1-1989. Acep. 9-111-1989)
Abstraet: A rcview of the tropical soft-bottom literature reveals that several general concepts in benthic
ecology. formulated mainly from temperate work, are either in need oí modification oc are no! readily
applicable to tropica l benthic ecosystcms. Severa! coocepts emerge from the pIesent tropical líterature
suggesting that in comparison with temperate cornmunities: (1) species diversity and faunal densities are no!
necessarily grcater in the teopies, (2) environmental stress (excluding anthropogenic input) is genera1ly
more sevcre, (3) ¡nfaunal communities are composed of proportionately more small opportunistic species
(4) predation by demersal fishes and crustaceans is more intense, (5) microbes may be a carbon sink in
sorne shallow-water habitats. notably mangroves. (6) production is general1y high, but breeding and reproduc­
tíon are frequently nor continuous and,(7) the distributlon and abundances al' tropiCal benthos, like most
other communities, rcflcct temporal and spatial mosaies of major regulatory factors (competition, predation,
food supp!y, environmcntal disturbances).
Severa! tropical marine ecosystems such as rnangroves and coral reefs are unique, and other environments
such as continental she!ves possess several common features which distinguish them to sorne degree from
their tempcrate counterparts. To confirm, reject or modify these emerging coneepts, severa! aspects oC
tropical benthic ecosystems require further study. including effects of wet season activity, physiological
tolcrances, nutrient recycling, secondary production, bentluc-pelagíc coupling and pollution. Such infor­
mation and emerging conccptualizations are neeessary to permit proper and ¡nformed conservation and
management of these uniquc ccosystems.
MDst research on benthic cornmunities has
been conducted in temperate, and to a lesser
extent, subtropical and boreal latitudes (Gray
bottom
benthic assemblages in
(Longhurst
lhe tropics
& Pauly 1987).
Kecent reviews of tropical estuarine and
198 1). Lack of benthic research in tropical
marine ecosystems (De La Cruz 1986; A10ngi
regiDns is due to rnany reasons, including dis­
1989a; Hatcher, Johannes
tance of major oceanographic centers from the
tropics, lack of funding and inadequate facili­
ties.
Research into tropical marine benthic eco­
these systems for the future of mankind, both
as significant contributors to the global carbon
cycle and as nursery grounds for commercially
systems has, over the past four decades, been
viable
sporadic and, until recently, conducted mainly
Hatcher, Johannes
during
short-term
developed
visits
countries.
by
Most
workers
& Robertson 1989)
have c1ear1y underscored the importance of
species. Indeed,
as
pointed
out
by
& Robertson (1989), conser­
from
vation and management of tropical soft-bottom
early (pre-1970)
systems is critical, but the information neces­
work was necessarily taxonomic or consisted of
sary
compilations of faunal lists. Hence, little com­
conservation strategies is insufficient.
to pennit
the development of
proper
prehensive work exists detai1.ing factors affect­
The purpose of lhis review is to briefly
ing the distribution and abundance of soft-
examine the existing tropical infaunalliterature
*
reefs, see review of A10ngi 1989a) and to make
(but gene rally excJuding mangroves and coral
Contribution No. 451 from thc Australian lnsti­
tute of Marine Sciencc.
evident how unique these cornrnunities are, to
86
REVISTA DE BIOLOGIA TROPICAL
formulate a preliminary synlhesis of eeological
concepts presently emerging from the study of
infaunal cornmunities in the trapies.
Table I provides a comprehensive, but not
faunal and macroinfaunal assemblages in tropi­
cal intertidal habitats. Abundances of hetero­
trophic microbial communities are not included
because nearly all microbial slUdies to date in
the tropics have utilized antiquated techniques
such as plate counts. Only recent studies have
used modern, more eeliable, techniques 5uch as
diceet counts by epifluorescence microscopy
and uptake of radionucHdes to estimate micro­
bial activity (Aiongi I 988a.b). Data on tropical
microalgal abundances (as chlor9phyll a) does
exist, bUl most of the informatian is incompa­
rable because of lack of standardized units (e.g.,
,..g Ch! a per g_1 sediment dry weight, per g_1
sediment wet weight, per wet ce, per dry ce,
etc). However, lhe comparable intertidal data,
mostly from mangrove and coral reef sediments,
1 O ,..g ChIa g_l
(see
review
conditions
are
more
Early tropical studies of intertidaJ macroin­
exhaustive, compilation' of densities of roeio­
densities
environmental
quiescent.
Abundances. community composition and
intertida! zonation.
suggests low «
algae and plant detritus (Grelet el al. 1987), and
where
Tropical Intertidal Assemblages
algal
abundances are greater in coral reer lagoons
where sed,iments are rich in micro· and macro·
DW) miero­
Alongi
1989a).
fauna were concerned mainly with zonation of
'species associations', c1assifying low, mid and
high intertidal zones on the basis of the domi­
nant speeies (Dahl
1953; Gauld & Buchanan
1956; Berry 1964; Mclntyre 1968; Vohra
1971). This reOects the application of Thorson's
(1957) parallel-bottom community concept
-dominant during this time period- to tropical
intertidal habitats. Later studies (e.g. Dexter
1972,1974,1979; Shehon & Robertson 1981)
focussed
habitats
on
in
comparing
terms
of
different
species
intertidal
composition,
abundance, biomass and diversity.
It is elear that tropical intertidal habitats
differ
greatly
in
macrofaunal
community
composition from temperate intertidaJ assem·
blages. Tropical sandy beaches and sand f1ats
are dominated by crustaceans. mainly crabs,
and bivalve molluscs and tropical mudOats have
a high proportion of polychaetes and micro­
crustaceans, whereas temperate intertidal habi·
tats consist mainly of polychaetes and gastropod
molluscs (see Swennen, Duiven
& Spaans 1982;
Vargas 1987). Why? Sevoral behavioural studies
The data summarized in Table I must be
have suggested that tropical crustaceans and
considered with caution owing to the different
sampling techniques and sieve sizes used, and
rapid escape mechanisms than their temperate
sediment
depths.
pre-1975
papees examined did nol gjve sufficient infor­
In
fact,
many
mation to assess densities per unit area or vol·
ume. Nevertheless, it is reasonably elear that
the highest faunal densities occur in sheltered
habitats, whereas the lowest densities are found
in exposed, coarse sandy beaches. As reflected
in lhe tables, most tropical intertidal studies
have been conducted on sandy beaches.
Densities of meiofauna and small «
macroinfauna
in
tropical
5 mm)
unvegetated
tida!
flats and beaches are generally greater than in
mangroves
where
densities
of
both
faunal
groups average
from
< 500 individuals 10 cm-' and
< 10 to several hundred animals m -',
respectively (Grelet el al 1987; Alongi 1989a).
bivalves are more motile and possess more
counterparts
to
avoid
high
temperatures,
salinity and dessication in the tropics (Ansell
&
Trevallion 1969; McLusky el al 1975; Jones
1979). Ocypodid crabs and teUinid bivalves,
espeeially of the genus Donax appear to be
particularly well adapted to life in lhe tropical
intertidal and severaJ studies have documented
their unique life histories (Wade 1967; Ansell &
Trevallion 1969; Ansell
& Trevallion 1969;
Ansell
el
al 1972;
McLusky el al 1972).
However, differeoces do exist amoog tropical
intertidal regioos, as showo in the comparative
studies of Dexter (1972,
Shelton
into
1974, 1979) and
& Robertson (1981) and must be taken
account
before generalizations can be
made.
The lower densities in mangroves are probably
due to a variety of factors, including possible
negative
Seasonality and effeets of monsoons.
effects of maUgI'ove-derived tannins
and lower concentrations of interstitial oxygen
Maoy of the above·cited studies were con­
and surface microalgae. In cuntrast, infaunal
ducted during only one season, further limiting
TABLE 1
Densities of meiofauna (No. lndilliduals 10 cm-1) and macroinfauna (No. Individuals m-2) in sorne tropical intertidal
habitats x = mean: range
Macroinfauna
Reference
x = 4245;0-40,064
Gauld & Buchanan 1956
¡¡ = 2626; 420-3815
¡¡ = 1405; 395-1010
x = 1305; 176-3360
¡¡ = 2688; 2656-2720
McTntyre 1968
¡¡ = 1307; 968-1960
x=208; 127-368
Meiofauna
Locality
Habitat
Gold coast. Africa
sandy beach
Porto Novo,lndia
estuarine sand bar
marine sandy beach
moderately exposed
sandy beach
Lebanon
Tunisia
Moroeeo
sandy beaches
x=27;7-68
x= 196; 101-409
x= 148; 29-570
Shimmary Beach,
Panama, Caribbean
Sea
carbonate beach
x
Naos Is. Panama
Pacific ccast
quartz sandy
beach
x = 2439;1507-4067
Singapore
sheltered sandy
beach
x=5614; 1678-9236
Andaman & Nicobar
Istand. Indian Dccan
sandflat
600-800
Rao 1975
Shertallai. India
sandy beach
X =467; 215-1337
Muruo, WeUs &
Mclntyre 1978
Cebu. Phillipines
sandy beach
x= 739;417-1142
Natividad 1979
Texas. USA
sandy beach
Hulings 1971
�
193; 25-463
x= 2700; 335-6616
Dextcr 1972; 1974;
1979
Vohra 1974
Shelton & Robertson
1981
x = 219;118-381
Rama Sarma&
Chandra Mohan 1981
Bay of BcngaL India
coarse sandy beach
Malaysia
mudflats
Sungei Bulok
Kuala Selangor
x = 129
x =479
Broom 1982
Surinam, S. America
exposed mudflat
x=245
Swennen, Duiven &
Spaamo 1982
Mombasa. Kenya
sandy beaches
x = 1790; 1013-2·560
i =993; 592-2736
Ansari, Ingole &
Parulekar 1984
Punta Morales, Gulf
of Nicoya, Costa Rica
mudflat
¡¡ =898; 653-1078
Gulf of Aquaba.
Arabia Sea
seagrass beds;
fringing coral sands
x=2369;922-4239
Gulf of Nicoya,
Costa Rica
mudflat
Queensland.
Australia
sandflat
x =841; 324-4000
de la Cruz & Vargas
1987
x = 1547; 815-4106
Grelet et 011987
x= 13,827;
3787-41086
Vargas 1987
Alongi 1988b
comparisons
between
and
arnong
intertidal
habitats. Extensive studies conducted in the
Indo-Paeifie (Ganapati
& Rao 1962; Trevallion
&
& Solís-Weiss
extant (J ackson 1973; Heek 1979; Young
Young 1982; Ibañez-Aguirre
1986).
el al 1970; Ansell el al 1972; Natividad 1979;
Despite variations in sieve sizes and sampling
& Chandra Mohan 1981; Broom
gea!. it is clear that the eastem Indian coastline
1982; Rarkantra & I'arulekar 1985) have,
however, examined seasonality and the effects
(e.g. Bay of Bengal) harboes a numerieally rieh
of
stressed western eoast (Datta
Rama Sanna
monsoons
season
on
intertidal
infauna.
Whidl
(or seasons) organisms exhibít theír
infauna compared to the more highly polluted,
& Sarangi 1986).
As discussed later, species diversity is gene rally
highest abundances appeaes to be dependent on
low refleeting
whether the habitat in question is in the wet or
species. Outside India, the data indica te poor to
dry tropies. Meiofauna and maeroinfauna suffer
moderately
increased mortality during monsoons in the wet
1972a,b) with numerieal riehness apparently
related to various stresses such as low salinity
tropies due to lhe fluetuations in salinity and
sediment erosiono In the dry tropics, organisms
usually attain peak densities in autumn and
and
lhe dominanee of opportunistie
rieh
oxygen
infaunal
assemblages (Wade
and low plankton produetion
(Holm 1978).
The
winter; lowest abundances occur in spring and
Indian
studies
reveal
that
tropical
eover
subtidal infauna respond negatively to freshwater
Infaunal eommunities also respond differ­
inputs during the monsoons, but recolonize
quiekly and are dominated by small, surfaee
summer
when
the absenee
of cloud
results in sediments wanning to > 30° C.
ently depending upon lhe intensity and fre­
quency of the monsoons. For example, in India
where cyclones are very intense and prolonged,
meiofaunal
densities
northeastern
Australia
decrease
rapidly.
In
where monsoons are
deposit- and suspension-feeding polyehaetes
and bivalves. As with temperate benthos,
eoastal lagoons in the tropies have also been
poorly studied with most studies eomprised
of
faunal
lists
rather
lhan
information
on
generally not as prolonged or as intense as lhose
density. biomass and species diversity (Moore
in Soulheast Asia or on lhe Indian subeon­
el al 1968: Holm 1978). The study by Holm
tinent, meiofaunal densities in tropical man�
(1978) indicates a potentially rich epi- and
grove estuaries were most abundant during the
infauna in moderately flushed lagoons Iined
summer
monsoons occur in different seasons in these
vegetation. Densitites of
with
submerged
maerofauna ranged from 1059 to 4700 per m'
in a seagrass flat (Moore el al 1968). In other
different
tropical lagoons not well flushed, high saline
wet
season (Alongi
1989a).
These
eontrasting results may refleet lhe faet lhat
climatologieal
zones.
Interestingly,
both meiofaunal and macroinfaunal cornmuni­
waters up to 300
ties appear to be able to repopulate quickly
Priee
following
climatologieal disturbanees in lhe
tropies. The magnitude of seasonal fluetuations
is also dependent upon distanee from lhe
equator. Habitats bordering lhe equator, for
instance, in Malaysia (Broom 1982) exhibit less
seasonal variability lhan tropical assemblages
closer to lhe poles, refleeting lhe faet lhat
elimate and its effeet on intertidal and shallow
subtidal bentho, varie, greatly wilhin lhe tropies.
o
/00 are common. Jones,
& Hughs (i 978) found a moderately
abundant maerofauna (> 500 m-') in a saline
lagoon
nn
lhe Saudi Arabian
coast of lhe
Arabian Gulf.
Diveesity Cas number of species)
was low ( < 15). These moderately high densi­
ties, despite the high salinity, were attributed to
eomparatively high primary production.
No! surprisingly, lhere is very Httle informa­
tion on funetional relationships among tropical
subtidal assemblages. In one of the few ,ueh
studies, Blaek
& Peteeson (1988) examined
possible adult-Iarval interaetions and competi­
Tropical coasta! assemblages (estuaries, lagoons,
seagrass beds and neushore subtidal).
tion between suspension.feeding bivaJves and
smaller
found
Most
studies
of benthie eommunities
tropical estuaries and shallow «
in
infauna in Western Australia,
that
the
suspension-feeding
They
bivalves
exhibited no evident competitive interactions
50 m) near­
with, or provided any positive enhancement of,
shore regions have been conducted in India
the establishment of the smaller infauna. These
(Table 2). Comparatively few eomprehensive
results, if found in other tropical subtidal
habitats, would help to explain the coexistence
benthie studies of tropical seagrass beds are
89
ALONGl: Tropical soft-bottom benthos.
TABLE 2
Densities 01 meiolaulUl (No. Individuals 10 cm'1) and macrolaurw. [No. m'l) in some tropical shallow·water, coastal habitats
(estuarine and marine) x�mean, range
Meiofauna
Macroinfauna
Reference
x = 240; 148-433
Wade 1972a
Locality
Habitat
Depth
1m)
Kingston Harbor,
Jamaica
marine
muds
6-16
Shatt-al-atab,
Arabian Gu1f
carbonate
muds
7·15
x =227; 24-599
Saad & Arlt, 1977
Karwar. central
west coast of
India
estuarine
muddy sands
and mud
6·'
i= 1098; 1022-1250
Ansari 1978
Madovi,Cumbarjua
Zuari estuanes.
Goa, India
estuarine
sand to mud
4·7
Nannada Estuary,
India
estuaríne
sand to mud
3-19
x=53;0-3164
X =592; 0-4049
Krishna,Godavari,
Mahanadi, Hooghly
rivers,E. coast
India
marine muds
15-50
x =813; 502·2149
x=1529;90-4785
Aman et al 1982
Carrie Bow Cay,
Belize
Thalassio
meadows
1·2
i=12,167
Young & Young 1982
Carrie Bow Cay,
Belize
bore sand
1·2
;=16.750
Young & Young 1982
Gautami-Godavari
estuaries,India
estuaríne
muddy sands
<10
x = 1386; 245-2194
South Gujarat
estuaries, India
estuaríne
sands to mud
< 10
x = 138; 0-5334
x = 122; 0-880
Vellar Estuary.
Porto Novo,India
estuarine
sand
2·5
x=90; 33-213
x= 5 1 (premonsoon)
x =87 (monsoon)
x = 46 (postmonsoon)
x=94; 78- 1 1 2
¡¡ =80 (premonsoon)
x=99 (momoon)
x =75 (postmonsoon)
Fernanoo. Khan &
Kasínathan 1983
Gulf of Nicoya,
Costa Rica
marine
mUddy sands
+mud
6-65
x"" 1269; 0-8744
Maurer & Vargas
1984
Goa,Coast,
India
estuarine
muddy sands
10
i = 1703; 475-310 (premonsoon)
Harkantra &
Parulekar 1985
\Vest Bonga!,
India
estuarme
.and,
i =620; 100-2420
Dalla & Sarangi
1986
<
x = 1653; 492-4382 (total)
x =528; 93-2120 (monsoonal)
of �lIrface suspension-and deposit-feeders in
tropical seafloors.
Varshney. Govinúan
& Desai 1981
Kondalarao 1983
r=810; 71·1495 (postmonsoon)
10
Parulekar,
Dhargelkar &
Singba1 1980
Govindan, Varshney
& Desai. 1983
limiting inter-shelf comparisons. Macroinfaunal
densities naturally vary widely depending upon
sediment type and water depth. There is very
little information on meiofaunal densities and
biomass on tropical shelves (Table 3).
Tropical continental shelves
Benthic
nental
inforrnation
shelves
on
tropicaJ
conti­
have likewise be en obtained
mainly by Soviet and Indian scientists work­
Despite the lack of data, tbere appear to be
several characteristics common to most if not
all tropical shelves which distinguish tbem, to
a limited degree, from temperate continental
ing in the lndian ocean (Table 3). Consequently,
shelves:
sampling procedures (different sediment depths;
1. Faunal composition appeaes to be domi­
the same problems regarding sieve sizes and
dry weight
vs.
wet weight biom.ss) prevail.
nated by large epibentbos, lancelets (e.g.
TABLE 3
Densiries (No. JOcm'l) and biomass (mg DW 10 cm-') ofmeio/auna and macroinfauna (No. m-l and
g DW or g WW m'l ) on sorne tropical continental she/lles. x =mean: range
Locality
Depth (m)
Guinea & Senegal
shelves, West Afriea
Mozambique Channel,
S.E. ACríea
N.W.lndian shelf
Density
Macroinfauna
Hiornass
Density
Meiofauna
Biomass
2().350
<
31
Reference
15-\386
1.5-74.2
Longhurst 1958,1959
J2.3ll
5.8-25
Makarov & Averín 1968
x =20;
Neyman 1969
20-350
I().IOO (WW)
S.W. Indian shelf
2().350
West Pakistan shelf
20-200
Persian Gulf
;;; =5;
().IO (WW)
196-224
2().50
Bay of Bengal
-2().170
-x= 163;
13-963
Cap Blanc, Spanish
Sahara
Coa Coast, India
Savich 1970
x = 25
30-35 (WW)
Neyman &
Kondritskiy 1974
Ansari et 01 1977
;<=2.5;
x= 517:
x=92;
0.2-11
58-1644
O,Of-36.5
x=8124;
x= 26.3;
2.4-94.4 (WW)
3().1830
2().840
10-15
1635-35200
:i = 1217
Nichols & Rowe .1977
Ansari, Parulekar &
,.250-2925
West Indian shelf
10-70
Andaman Sea.
Indian Ocean
1().200
Malacca Strait.
Indian Ocean
8().320
Arabian Sea
Andaman Sea
Bay of Bengal
10-250
10-250
Hong Kong Shelf
x=708;
543-985
¡ :::::248 ;
68-438
¡=149;
7-200
2().l50
"--
-
¡= 14:
3.6-32.8 (WW)
;: = 9;
1-27 (WW)
x = 258;
118-528
= 1.4 (DW)
=0.9 (DW)
Jagtap 1980
x= 10;
1-37 (WW)
Parulekar.l Ansari
x = 15;
0.2-71 (WW)
Ansari &. Paro lekar
x =0.9 (DW)
Rodrigues,
Harkantra.
Parulekar 1982
;¡ =0.8 (DW)
=0.2 (DW)
13-23
>;:= 1 1 . 5;
5.5-24.5 (WW)
;¡ = 1.8 (DW)
x= 101:
55-183
1981
1981
x= 35;
12-155 (WW)
Shin tl Thomspon
1982
Ea!t African
Shelf(l2-170N)
lO-lOO
x =3.5;
0.5-17.6 (DW)
Dornain 1982
Pero shelf
35-350
x=45.3;
1-247 (WW)
Rosenberg el al
Java. Indian
Ocean
5-30
1983
Warwick &.
Ruswahyuni 1987
x=463;
llO-739
BranchioSloma) and 1aege Foearninifeea such
al watee-column tumovee (Bukatin
as Marginopora and Alveoline/la (Buchanan
1982;
1958:
Longhuest
1958: Gosselck 1975);
2. Across shelf variations are due to severa!
Longhuest
1983;
Alongi
el al
1989b);
3. Infaunal communities aee composed mainly
of small (
< 5_0 mm) oppoetunists that aee
ate production and sedi.mentation, infringe.
sueface deposit-and suspension-feedees_ This
appears to be an adaptive strategy lo re­
ment of water movement and lack of season-
spond quickly to erratic inputs of nutrients,
factoes, including gradients in depth, caebon­
91
ALONGI: Tropical 8Oft-bottom bcnthos.
low water content (i.e. hlgh compaction) of
mostly quartz and carbonate sand and gener­
'
ally warm ( > 20 C) sea temperatures
(Longhurst 1959:
Harkantra el al 1980;
Alongi 1989b);
4. One common
feature
of subtropical and
tropical shelves is an
abundant demersal
fisheries,
shrimps
including
(Ghana,
penaeid
Buchanan
Leone, Longhurst
prawns
1958;
and
Sierra
1983; India, Harkantra
however, densities of bacteria, meiofauna and
macroinfauna are comparatively high due to a
more
stable
seafloor
phytoplankton
and sedimentation of
blooms
caused
by
nutrient
runoff from lhe Amazon (Table 4). AlIer
&
AlIer (1986) conc1uded lhat lhe hlgh degree
of physical disturbance coupled wilh low food
inputs appear to be the major factors regulat­
ing
benthic infauna OR lhe
Amazon shelf.
In contrast, densities of lhe major groups
el al 1980; Georgia Bight, Hanson el al 1981;
are higher on lhe central Great Barrier Reef
Gulf ofMéxico,
shelf .probably due 10 lhe more stable seabed
and gre.ter food input, particularly from export
Yingst and Rhoads 1985;
central Great Barrier Reef, Alongi 1989b),
Demersal populations are probably abundant
on these shelves due to the predominantly
shallow
sandy
bottoms
and
abundant
epibenthic and surface infaunal prey;
5. Tropical
continental shelves are generally
of detritus from individual reefs of the GBR on
the outer shelf. Alongi ( I989b) conc1uded that
intermediate levels of food input and physical
disturbances from trawling and storms regulate
benthic standing crops and processes on lhe
shallow, driven by ¡ntermittent ¡ntrusions of
central shelf.
upwelled, nutrient-rich water and by estua­
rine outwelling of detritus (probably of low
Although sedimentary organic carbon and
nitrogen concentrations on the Amazon shelf
nitrogen content) and input of reef detritus
are on average much higher than on the central
of
quality.
GBR
tropical
compared to levels on temperate shelves and
generally
Bentltic
hlgher
communities
nutritional
thrive
in
shelf (Table 4), !he arnounts are low
upwelling regions off Panama (Lee 1978),
other tropical shelves influenced by upwelling
Peru (Tarazona e l al. 1988) and northeast
Africa (Nichols & Rowe 1977), and in areas
(Longhurst & Pauly, 1987).
Iimitation and moderate to
Detrital food
hlgh levels of
of massive estuarine outwelling such as off
physical
India (Neyrnan 1969; Ansari el al. 1977).
features of the benthic regimes of both of these
tropical continental shelves.
Interestingly, lhese shelves are cornmonly
disturbance
appear to be common
subjected to anoxia (with deleterious effects
on lhe benthos) if estuarine inputs and
upwelling events occur on a massive scale,
too frequently, or occur simultaneousiy wilh
periods of stagnant water mass, for instance,
off lhe Amazon (Aller
& Aller 1986) and off
Peru (Rosenberg el al 1983).
Emerging concepts
It is hoped lhat the reader has developed, at
thls stage, sorne appreciation of tropical sea­
floor habitats, the assembIages lhat tive in
them, and how general concepts in benlhic
ecology (diversily, stress, disturbances, see
Gray 198 1) may have evolved wilh fewer
Comparison of two tropical benlhic regimes:
lhe Amazon and central Great Barríer
misconceptions and to a more advanced state,
if tropical information was obtained and/or
Reef continental shelves.
considered earHer.
In trus section, I discuss
The comprehensive studies of Aller and AlIer
how sorne major preconceptions in ternpe·
rate benthlc ecology, when applied lo Ihe
( I 986) on the Amazon shelf and by Alongi
tropics, are either in need of modification or
(1989b) on
are not valid.
the central Great
Barrier Reef
shelf provide a rare opportunily to compare
and contrast tropical benthlc biocoenoses from
l . Species diversity and faunal densities are
different biogeographical provinces. The inner
IlOI greater in the tropic� The tropics is not a
Amazon shelf is dominated by massive runoff
homogenous part of !he biosphere. Thus it is
not surposing that lhe greatest range of faunal
and sedimentation from the Amazon river and
neae the over mouth, resulting in an unstable
seabed causing either very low or totalIy absent
densities and species diversity occurs here.
Older ecological studies conducted primarily in
infaunal communities. Away from the mouth,
rainforests and on coral reefs indicated hlghest
REVISTA DE BIOLOGIA TROPICAL
92
TABLE 4
Comparison ofmajor ¡n[aunal groups and sedimentary choracteristics on rhe Amazon (Aller & Aller
and central Great Barrier Ree! (Alongi J 989b) continental shelves
Depth range
Amazon
GBR
O-52
15-46
(m)
1986)
x= 7.5 ; 1 .3-2 1 x 10'
Bacteria (g'1 DW)
Meiofauna (No. 1 0 cm'l)
x =282; 1l-2045
x= 1520; 810-3255
Macroinfauna (No. m-l)
x= 1038; 59-3953
x = 2530; 2061l-5406
organic e
(% DW)
x =0.6; 0.07-0.84
¡¡=0.25; 0.05-0.71
organic N
(% DW)
x = 0.10; 0.01-0.14
;¡ =0.06; 0.02-0 . 1 3
6.0
4 .2
-----
e/N ratio
Major SOUIces of detritus
¡ntermittant phytoplankton blooms;
[iver runaff of pLant detritus
river runaff of mangrove + terrestrial
debris; low phytoplankton input; reef
detritus + mucus
Seafloor
unstable. fluid muds to mid shelf; a
high physical disturbances from IaIger
river discharges: inner shelf muds to
outer shelf quartz sand
stable. compacted sand with low Hl O
content; across-shelf gradient of
terrigenous to carbonate facies;
physical disturbances from trawling
and cyclones; river input limited
to inner shelf
species richness per habitat in the tropics, espe­
disturbances than temperate habitats, aceount­
cially in the lndo-Pacific region (see MacArthur
ing for the wider variation in species diversity
1 972). Sandees (1968, 1969) hypothesized Ihat
of tropical benthos. For instanee} in rny experi­
physically stable, berugo envirooments such as
ence, intertidal regions in the tropies are fairly
the tropics favor development of more complex,
inhospitable
plaees,
where
organisms
are
with
eommonly subjeeted to very high ( > 300 C )
stressed and/or younger environments where
temperatures, desiccation, low dissolved oxygen
levels, low food supp1y, ehemic.al plant defenses
species-rich
communities
compared
physieal faetoes dominat. in regulating com­
munity evolution. Even eaclier, Thoeson (1955,
and monsoons. These conditions are gene rally
195 7) suggested that species richness increases
not borne
markedly toward the equator for epifauna of
Alongi (1987) found 10w to moderate species
hard·bottoms, allhough
infaunal
diveesity and low number of species per habitat
assemblages.
.r or
These hypolheses have had Iheir detraetoes
ments in northem Australia, supporting Hilary
& Waltees 1979), but modifieation
Moore's (1972) earlier conlention Ihat tropical
and refinement of Ihe stability-time idea by
intertidal organismsare, on average, subjected to
(e.g. Abele
not
foc
by temperate organisms. lndeed,
free-living nematodes
in mangrove sedi­
Huston (1979) has attained a general level of
greater physieal stress than their temperate
acceptanee. Huston's hypothesis is not eentered
eounterparts. Olher benthie studies have found
around comparisons
10w or moderate species diversity in the tropies
of
temperate-tropical
speeies diversity, but in predieting variations in
again depending on degree of physieal stress
within-habitat diveesity as eaused by varying
(Johnson 1 970; Wade 1972a,b; Spight 1977;
levels of disturbance. The model is latitudinally
Maurer
independent and has provided sorne relief from
1987).
& Vargas 19 84; Warwiek & Ruswahyuni
Ihe old ideas of tropical vs. temperate diveesity
gradients.
[ propose that benthie habltats in the tropies
are subjected to a wider range of environmental
2. Envíronmental stress is generally greater in
the tropies. [n addition to the evidence eited
aboye} other examples can be provided
to
.A, T,ONGI:
Tropical soft·bottom benthos.
support this supposition. Shallow inshore eom­
munities in regions of ¡ntense wet season activ·
ity and upwelling are frequently smothered by
massive riverine sedimentation (AUcr & Aller
1986) or by erosion of estuarine mudbanks
(Seshapp. 1953). They are frequently stressed
by low sa!inity (Goodbody 1961) and by anoxia
eaused by Ihe impingement of coasta! water
masses during he.vy continental runoff and
subsequent diatom blooms (Harper et al 1981;
Nichols 1976a,b: Rosenberg et al 1983). It is
of interest to note that most physiologieal
studies have indicated tbat tropical organisms
are generally not more tolerant to environmen·
ta! stress than temperate organisms (Mayer
1914; Seholander et al 1953).
3.
Tropical infaunal assemblages are composed
of proportionately more opportunistic, small­
sized species. Nearly all of the benthic studies
cited earHer suggest that conditions on tropical
seafloors eommonly lead to the development of
pioneering seres and retardation of the estab·
Jishment of large, equilibrium assemblages.
Low food supply (oligotrophy), freshwater
input, heavy sedimentation or erosion, anoxia,
poUution, compaction of carbonate sediments
and ctimatic disturbanees (cyelones and
monsoons) are generally more typiea! of sub­
tropical and tropical shelves and coastal areas
than in temperate latitudes. Pioneering assem·
blages are well adapted to respond quickly to
patehy food inputs as well as to repopulate
after disturbanee, Ihat is, they exhibit a high
degree of resilieney. The faet Ihat many such
organisms are surface or water·column feeders
,uggests advantages in eompeting for Ihe lirnited
rood supply depositing to Ihe seafioor. The
relative abundance of surfaee infauna! feeders
can lead to greater rates of predation by epi­
fauna and demersal organisms. Indeed, as men·
tioned eartier, demersa! fishing industries are
reasonably well developed on most shallow,
warm water shelve, (Longhurst & Pauly 1987).
4. Predation is more intense in the trapies,
This early eeologieal paradigm (MaeArthur
1972) appears to hold true for tropiea! benthos.
Nearly all of Ihe evidenee i, eireum,tantia! and
based 00 studies of predation pressure on
intertida! gastropods (Bertness et al. 1981 and
referenees within). Yermeij (1978) argued Ihat
narrow or oeeluded apertures, thiekened shells,
93
low spires and more pointed seulpture on
molluse shells increases from temperate to tro­
pica! seas. Experiments conducted in Panarna
and New England e1early indicate that shell­
erushing predation of intertida! gastropds is
greater in Ihe tropies. Interestingly, greater
predation appears to modify behaviour of
tropiea! intertida! gastropod" sueh as eurtailing
Iheir foraging time (Bertness, Garrity &
Levings 1981).
OnIy one similar experiment has been con­
dueted with tropiea! mudflat infauna (Vargas
1988), Ihe results of which suggest that Ihe role
of maeropredators (birds, erabs, fish) in con­
trolling eommunity strueture is relatively minoro
However, a large body of correlative evidence
indicates a positive relationship between in­
fauna! biomass and total fish eateh on many
subtropieal and tropical shelves (Longhurst
1959; Savieh 1971; Harkantra et al 1980;
Rosenberg et al. 1983; Tarazona et al. 1988).
Additiona! eireumstantia! evidenee is provided
by Ihe greater proportion of predatory
nematodes in coral reefs and mangroves in
Austra!ia (Alongi 1987, 1989b) in eomparison
with temperate nematode cornmunities.
Other trophic relationships sueh as eompeu­
tion, cornmensalism and amensalism remain too
poorly understood for tropiea! infauna to
elueidate pattems and to compare wilh trophie
interactions among temperate infauna.
5.
Microbes may be a sink for carbon in some
Reeent
earbon flux models of mangrove and coral reef
benthos suggest an enormous rate of reeyeling
between microbes and sedimentary carbon
pools, implying that mierobes are a sink or long­
term ,hunt for earbon in Ihese systems (Stanley
et al 1987; Alongi 1988a,b,e; 1989b; Boto,
Alongi & Nott 1989). This sink hypolhesis
(Alongi 1988e) evolved from a large body of
circumstantial evidence obtained mainly from
mangroves in tropica! Australia: (1) bacteria!
standing erop is very large and produetivity
is high, often exeeeding Ihe rate of tata!
primary produetion, (2) mieroa!ga! biomass is
10w 10 moderate, (3) Ihe biomas, of protozoan,.
meiotauna and small ( < 5 mm) inlauna are
low eompared to olher benthic systems, (4)
the above- and below-ground partieulate
earbon pools are large, but the below-ground
DOC pool is low to moderate in size, (5)
correlative evidence reveals relationships of
tropical shallow-water benthic systems.
REVISTA DE BIOLOGIA TROPICAL
94
bacterial
numbers
and
growth
rates
with
wide variation of PIB ratios for meiofauna and
organic carbon, but not with supposed grazers,
the faet that this information was derived from
and, (6) amino acid and DOC fluxes across
temperate organisms. Bacterial production rates
the
are high in the tropies, ranging from 0.01 10
5 g e m-' d-' (Alongi I989b,e). Produetion
sediment-water
interface
are
negligible
unless the benthos is poisooed. After poisooing,
fluxes of amino acid. and DOC from tropical
estimates
mangrove sediments are large, ranging from
0.08 to 0,2 and from 0.3 to 2.4 g C. m -'
have been considered in another ceview (Alongi
1989b) and will not be detailed further. Need­
d-t, respectively. This latter evidence indicates
less to say lower ¡nfaunal biomass and general1y
no net export but large intemal recycling of
warmer temperatures are probably at least two
nutrient pools by microbenthos.
for bacteria
in tropical sediments
I
reasons for the higher baeterial standing erop
and productivity in the tropics.
In addition, �-glutamic acid, a non-protejo
amina acid found in sorne anaerobic bacteria
but not derived from mangrove root hairs,
leachates, detritus or bacteria damaged during
porewater extraction, is a major component of
the interstitial dissolved free amino aeid pool,
sugge.ting
marker
that this amino aeid may be
lar
natural
baeterial mortality
a
and
reeyeling of bacterial intraeellular pools. In faet,
the composition of major amiDa acids found
free in pare waters in mangroves and coral
reeh and those found in intraeellular pools of
sorne sulfate-reducing bacteria are very similar
(Alongi
1988c).
bacteria, replenishment of the
DOC pool would have to depend on eontribu­
tiaD from bacterial consumers. However, based
on the abov� ¡nfacmation, this mlght suggest
that the eotice microbial-meiofaunal foad chain
is, in tum, a sink for carbon particularly in
subsurface sediments where consumer densities
decline
rapidly in
(1976) for the lBP results, AnseU el al 1978;
Parulekar, Dhargalkar & Singbal 1980). The
study by Ansell et al (I978) indieates macro­
faunal turnover rates nearly ten times greater
in SW India than in Seotland with annual PIB
ratios ranging from 6 to 136. They suggest that
these higher turnover rates reflect a lack of
temperature
eompensation (as suggested
by
early physiologieal studies) and higher rates of
predation on tropical organisms.
A larger volume of data indieates that in
comparison
with
tempera te
counterparts,
tropicar species exhibit greater metabolism and
If protozoa and meiofauna consume large
quantities of
Several studies have estimated production
of tropical maeroinfauna (see review of Steele
the absenee
of biogenie
structures Ctubes, burrows, sorne oxidized root
hairs). This raises two questions: to what extent
are protists and meiofauna consumed by higher
organisms? Aow productive are they and
their potential consumers?
growth rates on a per weight basis, and shorter
lifespans and greater marta lity (Nayer 1954;
&
& Ditte1 1982; Parulekar
Edwards et al 1970; Lewis 1971; MeLusky
Stirling 1975; Epifanio
1984; Riehardson 198 7). Breeding and repro­
ductive activity are not conttnuous for most
tropical benthie speeies with the life eyde of
most
organisms
activity
oc
interrupted
excessive
heat
by wet season
(Le.,
mortality
I
hibernation or eneystment) (Penzias 1969;
Ansell et al. 1972a,b; Govidan et al. 1972;
Vohra 1971) and lunar eycles (DeVries et al.
1983a).
Tropical
erustaeeans,
Callinectes spp., frequently
for
example,
undergo one or
more reproductive peaks superimposed on a
year-Iong background of eontinuous, low -Ievel
6. Bacterial and' infaunal production in the
tropics is high, but breeding and reproduction
are frequently not continuous. Valid estimates
of benthie seeondary produetivity exist only
for the two most size-divergent groups, bacteria
and macroinfauna. Many workers have indirect­
Iy estimated meiofaunal produetivity using
breeding (DeVries et al. 1983b).
7. The distributian and abundances of infaunal
benthos re/lect temporal and spatial mosaics af
the major regulatory factors.
lt is interesting
to observe arguments in the literature concern­
ing whether competition, predation or distur­
annual tumover rates and mean biomass values.
bance is the major factor regulating cornmunities.
These
unrealistic to any ecologist who works in the
estimates must be considered highly
questionable
if not invalid beeause of the
Such
views
natural
are
groundless
environment.
and
Recent
intuitively
papers
(c.g.
95
ALONGI : Tropical soft-bottom benthos.
Dayton 1 984, Menge
& Sutherland 1987) have
interplay
regulating
adopted
varying
more
reaHstic
between
portrayals
envuonmental
force s
of
the
under
eonditions.
Not
Another danger is that summarizing
tropical
literature
mainly
by
the
considering
abundance data ignores Ihe fact Ihat species
may function disproportionately as predicted
from field experiments condueted in tropical
from their abundances or biomass. In any
event, one cannot expect tropical and temperate
and
temperate deciduous forests are similar to tro·
surprisingly, many of these ideas have developed
roeky intertidal areas where the relative ease
success
of
simple
manipulations
yielded concrete results (Menge
& Lubchen·
cho 1 98 1 ; Garrity, Levings & Caffey
Menge el al.
1986: An,ell
have
1986;
& Morton 1987).
One of the more recent attempts to realisti·
cally
model
community
model of Menge
regulation
is
the
& SutherJand ( l 9 87). AppHca·
ble to all habitats, the model predicts that, { l )
benthic ecosystems to be similar anymore than
pical rainforests.
Usted beIow are tapies in urgent need of more
study:
1 . effects of freshwater discharge, storros and
other climatic disturbances;
2. benthic·pelagic coupling
predation and competition are absent o r weak
in
stressful
environments,
in moderate
(2)
habitats, predation pressure is stiU weak, but
on
continental
shelves, including rate of detritus sedimen·
tation;
nutrient recycling (C,N'p) and geochemistry;
4. microbial biomass and productivity;
3.
sessile animals are less affected by stress leading
5. physiological tolerances, especially to temo
competition,
ameliorates
6. effects of pollution on cornmunity structure
'predator bottlenecks') in benign environments.
7. secondary production of meiobenthos aod
to population densities great enough to initiate
exeessive
and
(3)
competition
predation
for
space
(excluding
Testing in soft·bottom environments is needed
to aecept, reject or modify the model, but it is
an important first step in realistically grappl·
ing with food web interactions as affected by
changes in the environment. Many of these
changes can be as ephemeral as a tidal cycle,
perature and salinity;
and function;
macroinfauna;
8. trophic studies particularly demersal
feeding
and
groups; and
interactions
among
fish
benthlc
9. multidisciplinary, system·level studies
further accentuating the concept of communi­
combining analyses of water mass movement
with sediment chernistry, geology and ecoIo·
many factors constantly changing in time and
rare.
ties regulated and structured by a patchwork of
space.
gy. Studies of Ihe entire benthic regime are
Wilh
increasing
population
growth
and
anthropogenic input to coastlines in tropical
CONCLUS10NS AND RECOMMENDATIONS
This review was written to provide counter­
generalizations
and
examples
to
concepts
countries,
information - eoncerning the
structure
and
on. benthic
and proper management and conservation of
ingrained in the general soft-bottom literature­
season replication) aod mostly from temperate
infonnation
these unique ecosystems.
ACKNOWLEDGEMENTS
concepts formulated from often unreliable data
(poor methodology, lack of site', sample and
accurate
communities is urgently needed for inforrned
The author is supported by Marine Sciences
dynamics of infaunal assemblages. It is probable,
and Technologies Grant No. 86/0708 and
the Australian Institute of Marine Science. 1
coocepts will be modified or proveo wrong to
patiently
that these counter-generalizations aod emergiog
sorne degree in the future oc, in the opposite
thank our Iibrarians, particularly B. Betts for
finding
many
references,
Truscott for typing Ihe manuscript.
and
l
K.
also
veio, be uncritica11y accepted as fact. They are
oversimplificatioos, but the poteotial benefits
thank l.A. Vargas and two anonymous review­
in benthic ecology by considering Ihe tropical
component, outweigh Ihe risks.
and l. Tietjen over the years were instrumental
of attempting to improve upon general concepts
ers for their suggestioIls on the manuscript.
Discussions with K.G. Boto, A. Robertson
in fonnulating maoy of my ideas presented in
this review.
96
REV ISTA DE BIOLOGIA TROPICAL
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