Aust. J . Mar. Freshw. Res.

, 1982, 33, 1039-55

Demersal Fish and Cephalopod Communities

of an Unexploited Coastal Environment
in Northern Australia

S. F. Ruiner and I. S . R. Munro

Division of Fisheries Research, CSIRO Marine Laboratories,
P.O. Box 21, Cronulla, N.S.W. 2230.

Abstract
Fish communities in the south-eastern Gulf of Carpentaria were surveyed before the development of a
demersal trawl fishery for prawns. Distribution data for fish and cephalopods were examined to determine
community composition and species-distribution patterns in an unexploited shallow-water tropical fish
community. Common species were mostly Indo-west Pacific in affinity, and included many species
important in demersal trawl fisheries. Three clusters of site groups were present, comprising site groups from
nearshore ( - 2-5 m), shallow offshore, and deep offshore ( - 15-35 m) areas, respectively. Site groups did
not correspond to clearly defined communities. Species overlaps were frequent. Limits to the distribution of
common species seemed to be related more to physical factors (bottom depth, salinity and sediment type)
than to other biotic factors (distribution of species). The overall picture was of a rich assemblage of fish
species in an environment characterized by gradients rather than by sharp changes.

Introduction
Limited information on the fish and cephalopod communities of soft-bottom
substrates in the coastal waters of northern Australia is available from Taiwanese
trawling studies (Liu 1976; Liu et al. 1978; Lee 1979). No published information exists
for these communities in the Gulf of Carpentaria. Interest during the early 1960s in the
commercial potential of penaeid prawn stocks in the Gulf prompted a biological
survey in the south-eastern Gulf of Carpentaria between 1963 and 1965 (Munro 1972),
to determine the types of communities and the habitats in which prawns were found.
The survey recorded information on all species collected by prawn trawling, together
with data on site depths, water temperatures, salinities and sediment types.
Trawling activity in the Gulf of Carpentaria was minimal before this survey, which
marked the starting point of an intensive trawl fishery for prawns. The fauna thus has
interest as an unexploited shallow-water tropical fish community, both as a base-line
against which fish communities in more heavily exploited areas may be compared and
as the ecological background to the prawn fishery itself.
Species coexistence is a well-known phenomenon in coral reefs (Hiatt and Strasburg
1960; Goldman and Talbot 1976), often explained as a result of habitat complexity or
predictability. Recent work indicates that species coexistence may also be common in
tropical fish communities from more homogeneous environments (Lowe-McConnell
1969; Bradbury 1980), suggesting that habitat complexity may not be a sufficient
explanation in some situations.
In this paper, the fish and cephalopod (squid and cuttlefish) communities of the
south-eastern Gulf of Carpentaria are described, using trawl-catch data from March to
May 1964. Comparison is made with other Indo-west Pacific demersal fish
1040 S. F. Rainer and I. S . R. Munro

communities. Patterns in community composition and species distribution are

examined for evidence of species coexistence, and for relationships with environmental
variables that may indicate habitat partitioning. Squid and cuttlefish are included here
with fish because of their highly mobile habit. Numerical analysis and interpretation of
the data were by S.F.R., taxonomy by I.S.R.M.

Study Area
Munro (1972) gives a general description of the south-eastern Gulf of Carpentaria,
summarized here.
The area surveyed comprised approx. 25 000 km2 south of 16's. and east of 139OE.
(Fig. 1). Extensive shallow areas in the south and south-west shelve gradually to

Fig. 1. Map of the south-eastern Gulf of Carpentaria. Bathymetric contours (5-m intervals) are after
Munro (1967~).

depths of 30-40 m east of Mornington Island. The bottom slopes more steeply from
the eastern shore and from the islands in the west of the survey area. Tidal range is
3 . 3 m at spring tides and 0 . 5 m at neap tides. Water temperatures of inshore waters
( < 10 m) are usually between 17 and 30°C, and offshore waters (10-35 m) usually
between 23 and 29OC. The summer monsoonal rains, which usually begin in December
and may persist until March, reduce the salinity of nearshore waters until about
September. Nearshore waters subsequently become hypersaline, with salinities
exceeding 38%. Sediments are mainly grey sandy muds, often with ridges of coarse
sandy shell. Pockets of fine silt are present close inshore, near river mouths, and coarse
sands are often present in deeper waters.
Sites sampled during March-May 1964 were between 2 and 33 m deep. Bottom
temperatures were between 22 and 31 OC. Bottom salinities were between 26 and 35%,
with nearshore waters usually being several parts per thousand less saline than offshore
waters.
Demersal Fish Communities

Methods
Quantitative data were available for 1301 trawl catches, mostly collected between 29 July 1963 and
1 September 1964. This analysis is restricted to 341 samples collected between March and May 1964, since
sampling coverage of the south-eastern Gulf of Carpentaria was usually patchy at other times.
A brief description of methods for the collection and treatment of physical and biological data is given.
Details are given by Munro (1972).

Physical Data
Water depth was determined by echo-sounding, generally to the nearest 1 . 8 m (1 fathom), without
reduction to a standard tidal datum. Water samples were collected at the surface and near the bottom, using
a Nansen bottle with a reversing thermometer. Salinities were determined subsequently from conductivity
measurements. Sediment samples, collected wlth a pipe dredge, were analysed microscopically for the
abundance of lutite, quartz sands, shell and foraminiferans. The sediments were grouped subjectively on the
basis of the relative abundance of their component sediment fractions. Five sediment types were recognized:
1, fine muds and silts; 2, very muddy sands; 3, muddy sands; 4, dirty sands; 5, clean sands.

Biological Data
Bottom and near-bottom animals were sampled with 'flat' and 'balloon' trawls with headlines
of 16.5 m (9 fathoms) or 18.3 m (10 fathoms), respectively, and 3.8-cm mesh. Trawls were from
15 to 120 min duration (usually 30 min), at a speed of 1 m s-I (2 knots). Penaeid prawns were removed from
the catch. A subsample of about one-tenth to one-quarter of a fish basket (approx. 5-9 kg) was usually then
taken for enumeration, along with selected species from the remainder of the catch. Fish were identified to
species, if possible, cephalopods to genus or higher taxon.

Analytical Procedures
Numerical classification (Clifford and Stephenson 1975) was used to group samples on the basis of their
species composition. The samples of most groups came from geographically restricted areas and were
therefore referred to as site groups. The spatial distributions of site groups and of commonly occurring
species in each site group were then compared with depth, hydrological characteristics and sediment type.
Classification was performed on presence-absence data only, a choice indicated by considerations of cost
and high variability in the abundance data. The TAXON library program MULCLAS (Lance and Williams
1967a, 19676; Dale et al. 1981) was used, with the Canberra metric coefficient of dissimilarity and flexible
sorting with intense clustering. The coefficients I and P were set to 0,625 and - 0.25, respectively (Clifford
and Stephenson 1975;Williams 1976). Double-zero matches were omitted in the calculation of the Canberra
metric. These procedures and values were selected because previous experience with similar data had shown
their ability to produce ecologically interpretable groups.
The data set of species occurrences in 341 samples was reduced by the exclusion of impoverished samples
and rare species. The exclusion of species recorded in fewer than three samples and of samples with fewer
than three species reduced the data set to 315 samples. Distribution maps of species subsequently found to be
of interest were based on this data set. A further reduction of the data set to 303 samples was made for
numerical classification, in order to comply with computer storage limits; here, samples with fewer than 10
species were excluded. The 10-group level of classification produced geographically discrete entities
amenable to simple ecological interpretation.
Species occurring commonly in these site groups were selected for further analysis, retaining only species
with a constancy of 50% or more (i.e, species that occurred in at least half the samples) in at least one site
group. These species, here termed 'important', were then classified into species groups, using the same
procedures as for the classification of samples into site groups. A high proportion of the species groups
contained non-conformist members. Subsequent consideration of species distributions was therefore made
on a subjective basis.

Results
The samples retained for mapping and classification contained 173 taxa of fish (166
identified to single-species level) and seven taxa of cephalopods (none identified to
species). The 10 site groups differentiated (designated A-J) each contained 7-77
samples. Three major clusters were apparent among the site groups (Fig. 2). Site
 S. F. Rainer and I. S. R. Munro

groups A, B and C were located in nearshore waters; site groups D , E and F were
somewhat further from land, but still in relatively shallow waters; site groups G-J were
located in the deeper offshore waters (Fig. 3). Boundaries between the three major
clusters were diffuse, due to spatial overlaps between them. Site groups within the
major clusters usually had overlapping distributions.

Environment of Site Groups

Differences in the geographic locations of the three major clusters were reflected by
different mean bottom depths for their component site groups (Table 1). Mean bottom
depths were between 2.9 and 4 . 2 m for the nearshore site groups, between 5 . 0 and
10.6 m for the shallow offshore site groups, and between 15.6 and 21 . 9 m for the deep
offshore site groups.
4.0-

-
a,
2 3.0 -
9
Fig. 2. Dendrogram of relationships among 10
.-+-r - site groups from numerical classification. Site
L
-
.-m groups are ordered from A to J in increasing
.-E 2.0 - mean depth of the component sites. Number of
V1
.- sites and total number of species included are
P given for each site group.

A C B D E F G I H J
Site group
7 4 0 39 15 37 38 77 19 17 1 4
No. of sites
48 94 73 121 63 109 72 88 1 5 5 8 7
No, of species

Water temperature, salinity and sediment type also differed between the major
clusters. Mean temperatures near the bottom had a range of 7 . O°C (22.8-29.8 OC) in
the nearshore site groups, compared with a range of 3 . 1O C (25.7-28.8 OC) in the other
site groups. Mean salinities were lowest and most variable in the nearshore site groups
(means 27.8-30.7%; 6 0.6-1.4%), and highest and least variable in the deep offshore
site groups (means 32.5-33.52; 6 0.3-1 .1%,). Surface temperatures and salinities in
most site groups were little different from bottom values. The greatest differences
(0.7OC) between surface and bottom temperatures were in site groups A and G.
Salinities differed most (0.5%) between the surface and the bottom in site groups A
and J. These differences suggest that the water column was probably stratified in the
area of site group A, sampled in March, but was probably well mixed at sites sampled
offshore or during April and May.
Sediment samples showed a predominance of fine sediments close inshore (Fig. 4).
Coarser sediments were mainly restricted to offshore areas, although some inshore
areas contained a wide range of sediment types. The mean values for the sediment types
of samples within each site group conformed to the general pattern of increasing
coarseness with distance from shore. High values of 6 indicated high small-scale
variability in sediment type among the nearshore site groups and in the deep offshore
site group J.
Demersal Fish Communities 1043

Samples from site groups with similar spatial distributions tended to have similar
environmental characteristics. Site groups whose samples were collected early in the 3-
month period (site groups A, H and J) had higher mean water temperatures and lower
salinities than those collected later in the period (site groups B and I). For near-bottom
values of temperature and salinity, the differences within the 3-month period were up
to 7 .O°C and 2 . 9Ym for nearshore site groups, and up to 3 . 1OC and 0.3960 for deeper-
water groups.

Fig. 3. Location in the south-eastern Gulf of Carpentaria of site groups A-J defined by classification (a-J,
respectively), samples omitted from classification (k), and all 315 samples (I).

Community Characteristics of Site Groups

The richness of fish species (number of species per sample) varied markedly among
the site groups (Fig. 2). Site groups comprising a greater number of samples tended to
have a greater number of species. Exceptions were site groups D and H, the richest site
groups, which had 121 and 155 species recorded from only 15 and 17 samples,
 Table 1. Physical characteristics of site groups in the south-eastern Gulf of Carpentaria in March-May 1964
Values are given as the mean 1 s.d. for values within the 10 site groups derived by classification (A-J). Group K comprises samples omitted from the classification. The
+
number of samples used for each value is given in parentheses where it differs from the number of samples in a group (see Fig. 3). Sediment grade values are for sediment types
as described in Methods

Site Depth Temperature ( O C ) Salinity (%,) Sediment

group (4 Surface Bottom Surface Bottom grade
Demersal Fish Communities 1045

respectively. Site group G, on the other hand, had only 72 species from 77 samples. Site
groups with similar depth also differed considerably in species richness. In shallow
waters, for example, site group D (mean depth 5 . 0 m) was considerably richer than site
groups B and C ( 4 . 1 , 4 . 2 m). Similarly, in deep offshore waters, site group H (19.7 m)
was considerably richer than either of site groups I or J (21.5, 21.9 m). Overall, no
differences in species richness were apparent between the three major clusters, nor any
relationship between species richness and water temperature, salinity or sediment type.

Fig. 4. Distribution of major sediment classes in the south-eastern Gulf of Carpentaria. (a) Fine muds and
silts; (b) very muddy sands; (c) muddy sands, (d) dirty sands; (e) clean sands. Squares are 18-km grids in
which at least one sample of a particular class was collected; data are for all samples collected during
1963-1965, with n samples in each class.

Species Composition of Site Groups

Of the 180 species present in the 303 sites analysed, 61 were present as important
species in at least one site group (Table 2). Although most of these species were
important in only one or two site groups (24 and 11 species, respectively), no species
were unique to a particular site group. Fifty-four of the species were present in at least
half of the site groups (i.e, five or more), and eight species were important in five or
more site groups.
The various site groups differed considerably in their heterogeneity. The ratio of
important species to total species in a site group was highest in site groups A, E and G
(0.38, 0.32, 0 . 3 1, respectively). A low ratio of important to total species occurred in
site groups D and H (0.07 for both). These ratios, when considered with the total
species richness, indicate that site groups A, E and G were relatively homogeneous. Site
groups D and H, both with a high total species richness, seemed to be relatively
heterogeneous groupings with no strong identity of their own. Each major cluster
contained only one relatively homogeneous site group, suggesting that subdivision of
the clusters on the basis of species composition was not justified.
The important species included some whose distributions clearly matched the three
major clusters. The shallow inshore site groups consistently contained specimens of the
 S. F. Rainer and I. S. R. Munro

Table 2. Important species of site-group clusters, percentage constancy and the number of samples in each
cluster in which the species were recorded (N)
Species are arranged within each cluster in decreasing value of N; constancy values of 50% or higher are given
in bold type

Species Percentage constancy in site group N

A B C D E F G H I J
Species important mainly in inshore site groups
Johnius carutta 100
Polydactylus specularis 71
Secutor ruconius 14
Thryssa kammalensis 86
Otolithes ruber 57
Eleutheronema tetradactylum 100
Ilisha indica 86
Cynoglossus bilineatus 29
Thryssa hamiltonii 71
Leptobrama spp. 100
Stolephorus tri -
Johnieops vogleri 71
Harpodon translucens 86
Trichiurus haumela 71
Pellona ditchela 29
Absalom radiata 71
Nematalosa come 57
Herklotsichthys sp. 57
Leiognathus splendens 71
Escualosa thoracata 57 - 5 - - - - - - - 6
Scomberomorus queenslandicus 5 7 - 5 2 7 3 - - - - - 6

Species important mainly in shallow offshore site groups

Saurida undosquamis - 44
Loligo spp. + squid 71 31
Lovamia fasciata - 21
Sillago sihama 14 5
Leiognathus sp. nr brevirostris 29 97
Caranx bucculentus 14 13
Pseudorhombus arsius - -
Leiognathus sp, nr blochii 14 54
Pomadasys maculatus - 49
Paraplagusia australis 14 28
Apistops caloundra 14 -
Apogonichthys poecilopterus - 3
Euristhmus lepturus - 5
Acentrogobius caninus - -
Alepes kalla 43 31

Species important mainly in deep offshore site groups

Paramonacanthus spp. 14 10
Nemipterus tolu - -
Tripodichthys blochii - -
Equulites leuciscus - 18
Elates ransonnetii - -
Eutherapon theraps
Platycephalus harrisii
Gastrophysus scleratus
Takifugu whitleyi
Demersal Fish Communities

Table 2 (contd)

Species Percentage constancy in site group N

A B C D E F G H I J

Scolopsis regina
Upeneus sundaicus
Equulites bindus
Sillago maculata
Pegasus volitans
Engyprosopon grandisquama
Apistus carinatus
Pseudorhombus spinosus
Rhynchostracion nasus
Callionymus belcheri
Amphot istius kuhlii
Priacanthus tayenus
Pelates quadrilineatus
Selaroides leptolepis
Platycephalus tuberculatus

No. of species important

in site group
Total No, of species

threadfin Eleutheronema tetradactylum, the anchovy Thryssa kammaleasis and the

jewfish Otolithes ruber. These species occurred infrequently in samples from deeper
waters. Other species that were important in individual nearshore site groups, such as
the beach salmon Leptobrama spp. and the Bombay duck Harpodon translucens (site
group A), the anchovy Stolephorus tri (site group B) and the herring Ilisha indica (site
groups A and C), also occurred infrequently in other site groups. Differences in species
composition between the shallow and deep offshore site clusters were less clear. Species
such as the soldierfish Lovamia fasciata and the flounder Pseudorhombus arsius
occurred frequently in site groups E and F, but rarely elsewhere, and the flathead
Elates ransonnetii, the grunter Eutherapon theraps, the blowfish Gastrophysus scleratus
and the butterfly bream Nemipterus tolu were frequent in samples from site groups
G-J. On the other hand, species such as the saury Saurida undosquamis, the
leatherjackets Paramonacanthus spp, and the tripodfish Tripodichthys blochii were
frequent in samples from most of the site groups D-J, and many other species were
distributed variously among the site groups of the two clusters of offshore sites.

Species Distribution Patterns

The distribution of important species in the period from March to May 1964 thus
appeared to follow a variety of spatial patterns, only some of which coincided with the
three major clusters of sites in the south-eastern Gulf of Carpentaria. Most species
occurred frequently only within a limited range of depths (Table 3). Species such as
Eleutheronema tetradactylum (Fig. 5a) and the jewfish Johnius carutta were found
most frequently in shallow water ( < 10 m). Species widely distributed in offshore
waters included species such as the flathead Platycephalus harrisii (Fig. 5d) in waters
deeper than 5 m, and Nemipterus tolu (Fig. 5e) in waters deeper than 10 m.
Other species were distributed in patterns suggesting relationships with physical
variables in addition to depth. Eleven species, including the trevally Alepes kalla in
 S. F. Rainer and I. S. R. Munro

shallow water (Fig. 5i), and Paramonacanthus spp., otherwise widely distributed, were
absent or infrequent in nearshore sites to the south, an area to which Stolephorus tri
Table 3. Patterns of distribution and mean depths for 61 important species in the south-eastern Gulf of
Carpentaria, March-May 1964
Depths are given as mean +1 s.d. Species whose distribution is illustrated in Fig. 5 are indicated with ah
asterisk

Species Depth Species Depth

(m) (m)

Species whose geographic distribution is related to depth (39 species)

Escualosa thoracata 3+ 1 *Saurida undosquamis
Herklotsichthys sp. 3+1 Pseudorhombus arsius
Leptobrama spp. 4+2 Equulites bindus
*Eleutheronema tetradactylum 4+2 Takifugu whitleyi
Harpodon translucens 4+2 Pegasus volitans
Otolithes ruber 4+2 *Platycephalus harrisii
Thryssa hamiltonii 5+2 Amphotistius kuhlii
T . kammalensis 5+3 Engyprosopon grandisquama
Johnius carutta 5+4 Selaroides leptolepis
Trichiurus haumela 6 +4 Apistus carinatus
Cynoglossus bilineatus 6+4 Elates ransonnetii
Absalom radiata 6 5 + Eutherapon theraps
Pellona ditchela 7 6 + Pseudorhombus spinosus
Leiognathus sp. nr brevirostris 7 5 + Rhynchostracion nasus
Secutor ruconius 7+5 Scolopsis regina
Polydactylus specularis 10 6 + *Nemipterus tolu
Johnieops vogleri 10+7 Gastrophysus scleratus
Leiognathus sp. nr blochii 11 + 6 Sillago m&ulata
*Acentrogobius caninus 1113 Platycephalus tuberculatus
Lovamia fasciata 11 + 5

Species of limited geographical distribution related to factors additional to depth (22 species)
Abundant in the south (1 species)
*Stolephorus tri 5+3
Least abundant in the east (3 species)
Nematalosa come 3+2 * Leiognathus splendens 11+5
Ilisha indica 5+4
Least abundant in the west (7 species)
Paraplagusia australis 9+4 CaNionymus belcheri
Apogonlchthys poecilopterus 12+4 Priacanthus tayenus
Apistops caloundra 12+5 Pelates quadrilineatus
Euristhmus lepturus 12+5
Least abundant in the south (1 l species)
Scomberomorus queenslandicus 4+ 1 Sillago sihama
*Alepes kalla 5 3+ Equulites leuciscus
10f 7 Paramonacanthus spp
{2gOAsp. 10+6 Upeneus sundaicus
Pomadasys maculatus 11+7 Tripodichthys blochii
Caram bucculentus Ilk3

Asquid also comprise specimens of Loligo sp.

(Fig. 5c) was largely restricted. These species were presumably controlled by salinity,
sediment type, or a related factor.
Some nearshore species occurred less frequently west of Bentinck Island, as did a
few offshore species. These species included the bullseye Priacanthus tayenus, the
Demersal Fish Communities 1049

cardinalfish Apogonichthyspoecilopterus and the otherwise widely distributed tongue-

sole Paraplagusia australis (Fig. 5h). No important species were restricted to the west
of Bentinck Island, but there were several species that were frequent in samples from
western or south-western waters and infrequent or absent in samples from eastern or
south-eastern waters. These were two nearshore species, the herrings Herklotsichthys
sp. and Nematalosa come, and one offshore species, the ponyfish Leiognathus
splendens.

........

Fig. 5. Location in the south-eastern Gulf of Carpentaria of samples containing important species
illustrating distribution patterns given in Table 3. (a) Eleutheronema tetradactylurn; (b) Acentrogobius
caninus; ( c ) Saurida undosquamis; ( d ) Platycephalus harrisii; (e) Nemipterus tolu; (f)Stolephorus tri;
( g ) Leiognathus splendens; (h) Paraplagusia australis; ( i ) Alepes kalla.

Species from genera within the same family often had overlapping distributions. The
five species of Clupeidae, for example, were allcharacteristic of nearshore waters, and
four of the five were important in site group A. The three species of Engraulidae were
similarly characteristic of nearshore waters. The six species of Leiognathidae, mostly
wide-ranging species important in the site groups of both inshore and offshore waters,
included four species important in site group E and three species in each of site groups
B, F and G.
Five families included genera with more than one important species. The flounders
Pseudorhombus arsius and P. spinosus were characteristic of shallow and deep offshore
samples respectively, with little overlap in site groups. The flathead Platycephalus
tuberculatus was largely restricted to a single deep offshore site group (J), and the
congeneric P,harrisii was present in samples from all offshore site groups, including
 S. F. Rainer and I. S. R. Munro

site group J. Differences in the distribution of the other congeneric species were less
marked. Considerable overlap existed in the distributions of the two anchovies Thryssa
hamiltonii and T. kammalensis, both of which were important in nearshore site groups.

Discussion
Comparison with Indo-west Pacijic Demersal Fish Communities
The fish species from the south-eastern Gulf of Carpentaria considered here are
generally typical of those recorded from similar locations in other tropical areas, and
are strongly Indo-west Pacific in affinity. Most of the species have been recorded in
other coastal areas in northern Australia (Marshall 1964; Grant 1975), but relatively
few have a distribution extending into warm temperate waters. In Moreton Bay, for
example, the species collected in trawling studies by Young and Wadley (1979) and
Stephenson and Burgess (1980) included only 16 (27%) of the important fish species
from this study. This compares with a study of fish from Papua-New Guinea waters
(Munro 1967b), which included 38 (64%) of the important fish species, and another
study of the fish from Sri Lanka (Munro 1955), which included 29 (49%) of these
species.
Little information is available on the community structure of similar unexploited
demersal stocks. Pauly (1979) determined the probable composition of demersal fish
stocks in the Gulf of Thailand before intensive trawl fishing. He concluded that small
specialized prey species such as Leiognathidae, Gerridae and Mullidae comprised
about half the virgin stock, and generalist species such as flatfishes occurred only in
very small quantities. Direct comparisons with the present study are difficult, since
biomass data are not available for the Gulf of Carpentaria, but the same relationship is
evident for those groups at least. Large zoobenthos feeders such as rays (e.g.
Amphotistius kuhlii) were abundant in both studies, whereas large predators were
relatively uncommon. However, an intermediate predator (sensu Pauly 1979), Saurida
undosquamis, occurred in more samples than any other single taxon in the Gulf of
Carpentaria, whereas Saurida spp. were relatively unimportant in the Gulf of
Thailand. Polydactylus specularis, Paramonacanthus spp. and Tripodichthys blochii
were almost as common in the south-eastern Gulf of Carpentaria as S. undosquamis;
neither these nor related species appear to have been significant in the virgin stocks of
the Gulf of Thailand. Insufficient data are available on the ecology of the Australian
fish stocks to assess the significance of these differences.

Species Distribution Patterns

The distributions of important species and patterns of depth, hydrology and
sediment types were examined for similarities or differences that would indicate which
features of the environment were significant in determining the distribution of
individual species.
Most nearshore species seemed to be constrained less by depth per se than by
sediment type and hydrology. The nearshore species restricted to shallow waters (mean
depth < 5 m) followed the distribution of fine muds and silts. Those nearshore species
that also occurred in deeper waters were less restricted around the margins of the
survey area and followed mainly the distribution of very muddy sands. Variant
distributions for Alepes kalla and Scomberomorus queenslandicus suggested that they
were avoiding the reduced salinity water in the south, an area apparently preferred by
Stolephorus tri.
Demersal Fish Communities

Most of the species occurring commonly in the offshore waters were widely
distributed. In the case of widespread species such as Saurida undosquamis or
Paramonacanthus spp., the only limitations to their distributions seemed to be the low
salinity of the shallow waters in the extreme south. The northern limitation to the
distributions of four species, including Leiognathus sp. nr blochii, suggests the
avoidance of areas with relatively coarse sediment. Eleven species, including
Platycephalus harrisii, appeared to avoid areas containing very muddy sands or fine
silts. One species, Platycephalus tuberculatus, was apparently limited to relatively deep
waters with clean sands.
A few species showed distribution patterns that were not easily explained on the
basis of the physical data collected. These include eight species that were absent or rare
in the east or west of the survey area. Thus, one i&portant species, Leiognathus
splendens, was common in the area bounded by Mornington and Bentinck Islands, and
seven were absent or rare in the west of the survey area. These included Apogonichthys
poecilopterus and Paraplagusia australis. The distribution patterns of two nearshore
species, the herrings Ilisha indica and Nematalosa come, showed a variant pattern
whose limitation was apparently not related either to sediment type or to hydrology.
The distributions of most species during the 3-month period examined were similar
to what is known of their distributions for a full year. Species avoiding the low-salinity
water in the south of the survey area, including Suurida undosquamis and
Paramonacanthus spp., were usually more widely distributed. In addition, the flounder
Pseudorhombus arsius and the mackerel Scomberomorus queenslandicus had different
distributions at other times. Pseudorhombus arsius was more widely distributed in
offshore waters, rather than being limited to shallow offshore waters, Scomberomorus
queenslandicus, known to be a seasonally migratory species, was more frequent
offshore rather than inshore, and the records for March-May were probably for
juvenile individuals.
The distribution of most species accorded with ecological preferences known for
other areas. Differences were usually in the absence of a species from a particular
habitat. For example, Thryssa kammalensis and Eleutheronema tetradactylum have
been recorded as occurring in coastal or shallow coastal waters (Munro 1955, 1967b).
In this study, they were restricted to a nearshore habitat. Similarly, the restricted
distribution of Stolephorus tri was not in accord with occurrences ranging from coastal
waters to estuaries and river mouths (Munro 1967b). Several species found only in
offshore waters in this study are known to occur elsewhere in estuarine areas or in river
mouths (Munro 1955, 1967b). These included the species Euristhmus lepturus,
Eutherapon theraps, Pelates quadrilineatus, Apogonichthys poecilopterus, Sillago
maculata and Pseudorhombus arsius. These differences may arise for a variety of
reasons, including the limited sampling period used in the analysis, the absence of
samples from very shallow waters ( < 2 m), and limitations due to the type ofcollecting
gear used.

Determinants of Community Composition

The three clusters of site groups differentiated by numerical classification each
contained species that were restricted to a particular cluster. Since the distribution
patterns of most species were not limited by these site groups, the clusters of site groups
could not be considered as discrete communities. Site groups within each cluster,
differing as much in species richness as in species composition, showed a high degree of
spatial overlap. The environmental characteristics of the site groups also overlapped,
 S. F. Rainer and I. S. R. Munro

indicating that environmental factors were relatively unimportant in influencing the

composition of each site group. The overall picture is thus one of a rich assemblage of
fish species in an environment characterized more by gradients than by sharp changes.
In this situation, only limited conclusions about the determinants of community
composition are possible.
The occurrence of many coexisting species is usually taken to imply fine partitioning
of resources, particularly space (Sale 1978; Robertson and Lassig 1980). A
considerable degree of overlap was apparent in the distribution of fish species that
probably had similar patterns of resource utilization. This was exemplified by the
Clupeidae and Engraulidae, families whose members are likely to feed predominantly
on zooplankton and which together contained eight species characteristic of nearshore
waters. Similar overlaps were seen in the distribution of genera within other families,
and of other congeneric species. The existence of large areas with apparently similar
physical characteristics indicates that habitat complexity is low over much of the
south-eastern Gulf of Carpentaria. Resource partitioning between overlapping species
is, therefore, probably made on the basis of resources other than space. The scale of the
study reported here is too large to resolve this question. Smaller-scale sampling and
studies of resource partitioning would seem to be essential requirements in
understanding the ecology of this fauna.

Acknowledgments
Many people have assisted with the collection and analysis of the data used in this
paper. We particularly thank the master of the survey vessel Rama and the field staff at
the Karumba field laboratory. Programming assistance by N. Ridgeway is gratefully
acknowledged.

References
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Munro, I. S. R. (1955). 'The Marine and Fresh Water Fishes of Ceylon.' (Aust. Dept External Affairs:
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Munro, I. S. R. (19676). 'The Fishes of New Guinea.' (Dept Agriculture Stock and Fish: Port Moresby.)
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Pauly, D. (1979). Theory and management of tropical multispecies stocks; a review, with emphasis on the
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territorial damselfishes from the Great Barrier Reef. Bull. Mar. Sci. 30, 187-203.
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Manuscript received 11 January 1982, accepted 10 May 1982

Appendix
Systematic list of 61 species of cephalopodsand fishes sampled from the south-eastern Gulf of Carpentaria, and
the total number of occurences (C N ) in at least 50% of sites of at least one site group
Ph., Phylum; CI., Class; O., Order; F., Family; sp., species. Species linked by brackets were not
differentiated at the time of sampling.-C N not separately determined

Species ZN

Ph. Mollusca
C1. Cephalopoda
Squid (includes Loligo sp.)
F. Loliginidae
Loligo sp.
Ph. Chordata
C1. Elasmobranchii
0 . Rajiformes
F. Dasyatidae
Amphotistius kuhlii (Miiller & Henle, 1841)
C1. Teleostomi
0 . Clupeiformes
F. Clupeidae
Escualosa thoracata (Valenciennes, 1847)
Herklotsichthys sp.
Zlisha indica (Swainson, 1839)
Nematalosa come Richardson, 1846
Pellona ditchela (Valenciennes, 1847)
F. Engraulidae
Stolephorus tri (Bleeker, 1852)
Thryssa hamiltonii (Gray, 1832)
T. kammalensis (Bleeker, 1849)
 S. F. Rainer and I. S. R. Munro

Appendix (Contd)

Species T N

0. Myctophiformes
F. Synodontidae
Saurida undosquamis (Richardson, 1848)
F. Harpodontidae
Harpodon translucens Saville-Kent, 1889
0. Siluroidiformes
F. Plotosidae
Euristhmus lepturus (Giinther, 1864)
0. Polynemiformes
F. Polynemidae
Eleutheronema tetradactylurn (Shaw, 1804)
Polydactylus specularis (de Vis, 1883)
0. Perciformes
F. Theraponidae
Eutherapon theraps (Cuvier, 1829)
Pelates quadrilineatus (Bloch, 1760)
F. Priacanthidae
Priacanthus tayenus Richardson, 1846
F. Apogonidae
Apogonichthys poecilopterus (Cuvier, 1828)
Lovamia fasciata (Shaw, 1790)
F. Sillaginidae
Sillago maculata (Quoy & Gaimard, 1824)
S . sihama (Forskdl, 1775)
F. Carangidae
Absalom radiata (Macleay, 1881)
Alepes kalla (Cuvier, 1833)
Caranx bucculentus Alleyne & Macleay, 1877
Selaroides leptolepis (Cuvier, 1833)
F. Nemipteridae
Nemipterus tolu (Valenciennes, 1830)
F. Leiognathidae
Equulites bindus (Valenciennes, 1830)
E. leuciscus (Giinther, 1860)
Leiognathus sp. nr blochii (Valenciennes, 1835)
Leiognathus $p. nr brevirostris (Valenciennes, 1835)
L . splendens (Cuvier, 1820)
Secutor ruconius (Hamilton-Buchanan, 1822)
F. Pomadasyidae
Pomadasys maculatus (Bloch, 1797)
F. Scolopsidae
Scolopsis regina Whitley, 1937
F. Sciaenidae
Johnieops vogleri (Bleeker, 1853)
Johnius carutta Bloch, 1793
Otolithes ruber (Bloch & Schneider, 1801)
F. Mullidae
Upeneus sundaicus (Bleeker, 1849)
F. Pempheridae
Leptobrama muelleri Steindachner, 1878
i L . pectoralis (Ramsay & Ogilby, 1887)
F. Callionymidae
CaNionymus belcheri Richardson, 1884
Demersal Fish Communities

Appendix (Contd)

Species I: N

F. Trichiuridae
Trichiurus haumela (Forskil, 1775) 45
F. Scombridae
Scomberomorus queenslandicus Munro, 1943 11
F. Gobiidae
Acentrogobius caninus (Valenciennes, 1837) 34
F. Scorpaenidae
Apistops caloundra (de Vis, 1886) 62
Apistus carinatus (Bloch & Schneider, 1801) 61
F. Platycephalidae
Elates ransonnetii (Steindachner, 1876) 114
Platycephalus harrisii (McCulloch, 1914) 124
P. tuberculatus Cuvier, 1829 13
0 . Pleuronectiformes
F. Bothidae
Engyprosopon grandisquama (Temminck & Schlegel, 1846) 76
Pseudorhombus arsius (Hamilton-Buchanan, 1822) 96
P . spinosus McCulloch, 1914 58
F. Cynoglossidae
Cynoglossus bilineatus (LacBpkde, 1802) 83
Paraplagusia australis (Rendahl, 1922) 86
0 . Tetradontiformes
F. Triacanthidae
Tripodichthys blochii (Bleeker, 1852) 165
F. Monacanthidae
Paramonacanthus oblongus (Temminck & Schlegel, 1850) 204
i Paramonacanthus sulcatus (Hollard, 1854)
F. Ostraciidae
-

Rhynchostracion nasus (Bloch, 1758) 58

F. Tetraodontidae
Gastrophysus scleratus (Gmelin, 1788) 82
Takijiugu whitleyi (Paradice, 1927) 110
0 . Pegasiformes
F. Pegasidae
Pegasus volitans Linnaeus, 1758 89