Fisheries Research 42 (1999) 6786

A spatially oriented analysis of estuaries and their associated

commercial sheries in New South Wales, Australia
B.C. Pease*
Fisheries Research Institute, NSW Fisheries, PO Box 21, Cronulla, NSW 2230, Australia
Received 30 December 1997; received in revised form 12 March 1999; accepted 2 April 1999

Abstract
Management restrictions, in the form of input controls, on the complex commercial wild-capture sheries within 53 estuaries
in New South Wales (NSW) are currently applied to individual estuaries. The objective of this study was to determine whether
the estuarine commercial sheries of NSW can be grouped into larger, logically related spatial management units than
individual estuaries, based on the similarity of shared environmental and sheries attributes. Although there have been many
attempts to delineate marine biogeographic regions around Australia, there has been no attempt to relate commercial sheries
to bioregions or use them directly in sheries management schemes. In this study, multivariate techniques were used to
analyse spatial relationships among 53 estuaries, based on 8 environmental, 22 commercial shing method and 81 taxonomic
attributes. Principal components analysis of the environmental attributes indicated the presence of three latitudinal estuarine
regions (north, central and south), corresponding closely to the coastal inshore regions previously delineated by studies based
primarily on oceanographic attributes. After stratifying estuaries by water area, multivariate analysis of the sheries attributes
veried the presence of these same three latitudinal regions. Water area and latitude were the primary physical attributes of the
estuaries which were correlated with the delineation of these three regions based on sheries attributes. The management
implications of the results are discussed. Because the regions are delimited by attributes of the commercial sheries, they
provide a useful framework for future research on and management of estuarine sheries in NSW. The method of applying
multivariate analysis simultaneously to attributes of the physical environment and commercial sheries, as described in this
paper, may be useful for identifying regions in other multi-species sheries with complex shing area and effort components.
# 1999 Elsevier Science B.V. All rights reserved.
Keywords: Multivariate analysis; Classication; Regionalisation; Estuarine sheries; Australian estuaries

1. Introduction
The commercial, wild-capture, estuarine sheries
of New South Wales (NSW), Australia are not only
productive, they are also very diverse and complex. In
*Tel.: +61-2-9527-8411; fax: +61-2-9527-8576; e-mail:
peaseb@fisheries.nsw.gov.au

recent years recorded landings of these estuarine sheries have comprised up to 127 taxa, caught with over
20 shing methods. Estuarine sheries contribute
approximately 20% of the total commercial, wildcapture, sheries harvest (30 00034 000 tonnes) from
NSW waters (Pease and Scribner, 1993, 1994; Scribner and Kathuria, 1996). Commercial catches have
been taken from over 90 estuaries in NSW since 1940

0165-7836/99/$ see front matter # 1999 Elsevier Science B.V. All rights reserved.
PII: S 0 1 6 5 - 7 8 3 6 ( 9 9 ) 0 0 0 3 5 - 1

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B.C. Pease / Fisheries Research 42 (1999) 6786

Fig. 1. Map showing the 53 study estuaries on the coast of New South Wales, Australia. Dashed lines show proposed regional boundaries.

B.C. Pease / Fisheries Research 42 (1999) 6786

(Pease and Grinberg, 1995) and over 50% of the total

commercial harvest (including aquaculture) from
NSW waters is comprised of species that are estuarine
dependent (Pollard, 1976).
Management restrictions on these estuarine sheries consist of a complex set of input controls which
are currently applied to individual estuaries. These
restrictions vary considerably among estuaries along
the 1900 km NSW coastline (Fig. 1). The objective of
this study was to determine whether the commercial
estuarine sheries of NSW can be grouped into larger,
logically related, spatial management units than individual estuaries, based on the similarity of attributes of
the physical environment and commercial sheries.
There have been many attempts to delineate marine
biogeographic regions around Australia (Hedley,
1926; Whitley, 1932; Bennett and Pope, 1953; Knox,
1963; Wilson and Allen, 1987). These early bioregionalisations were based on subjective evaluations of
environmental characteristics and the distribution patterns of selected taxonomic groups. They are useful
for taxonomic studies, but the spatial scales are too
large for use with localised sheries. More recent
bioregionalisation studies by Hayden et al. (1984),
the CSIRO Divisions of Fisheries and Oceanography
(1997), the Australian and New Zealand Environment
and Conservation Council (1998), and Pollard et al.
(1998) have been aimed at implementing a system of
representative marine protected areas. Multivariate
techniques have been used in some of these recent
studies to quantitatively summarise the presence/
absence distribution patterns of marine and estuarine
species in a number of taxonomic groups simultaneously. The scale of these recent studies in Australia
is more useful than earlier work but the patterns
may be dominated by marine shelf-dwelling species
that are not generally found in estuaries. None of
the bioregionalisation work in Australia has been
based on attributes of commercial sheries or subsequently used for dening or managing commercial
sheries.
Bioregions have been dened for estuarine shes on
the west coast of the United States (Horn and Allen,
1976; Monaco et al., 1992) using multivariate analysis
of presence/absence species distributions. These studies focused on species distribution patterns and
regression analysis of the relationships between species diversity and environmental parameters. How-

69

ever, the analyses were not related to the commercial

sheries in these estuaries.
Stergiou, 1988, 1989 and Stergiou and Pollard
(1994) recently used multivariate techniques in a
spatial analysis of the commercial sheries catches
from Greek waters and discussed the management
implications of the resulting regionalisation. Because
of the complexity of the estuarine sheries in NSW, I
have expanded on the multivariate techniques of
Stergiou and Pollard (1994) to include simultaneous
multivariate analyses of attributes of the physical
environment, commercial shing methods and commercial catch by taxon.
2. Methods
2.1. Study areas
There are over 900 water bodies along the coast of
NSW which are permanently or periodically open to
the Pacic Ocean (Williams et al., 1998). Most of
these water bodies are small, ephemeral creeks and
drains. In this study an estuary is dened as: ``a
partially enclosed coastal body of water which is
either permanently or periodically open to the sea
and within which there is a measurable variation of
salinity due to the mixture of sea water with fresh
water derived from land drainage'' (Day, 1981). For
the purposes of this study an estuary is considered to
have a single opening to the ocean and includes all
arms, branches, basins and lagoons which are connected to this water body.
During the ve-year period from 1991 through 1995
commercial sh and shellsh catches were consistently reported (i.e. catches greater than zero reported
each year) from the 53 estuaries shown in Fig. 1.
These 53 estuaries were used as the primary spatial
units for this study.
2.2. Physical/environmental attributes
Eight physical/environmental attributes (Table 1)
were used for the initial spatial classication of the
53 estuaries. These attributes were chosen because
they are readily quantiable attributes of the physical
environment that are known to be linked with estuarine species distribution and diversity (Horn and Allen,

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B.C. Pease / Fisheries Research 42 (1999) 6786

Table 1
Physical/environmental attributes used in the principal components analysis
Attributes

Units

Sources

Latitude
Geomorphological type

Degrees and minutes

1. Saline coastal lagoon
2. Barrier estuary
3. Drowned river valley estuary
Square kilometres
Square kilometres
Metres
Metres
Millimetres
Square kilometres

Hydrographic charts
Roy, 1984

Catchment area
Water area
Entrance depth
Entrance width
Average annual rainfall
Seagrass area

1976; Pease et al., 1981; Bell and Pollard, 1989;

Monaco et al., 1992; Pollard, 1994; Gilmore, 1995).
Geomorphology is the only attribute that has not been
quantied in previous studies. I have assigned ordinal
numbers to the three estuary types dened by Roy
(1984). A value of one was assigned to ``saline coastal
lagoons'' which have entrances that are closed under
most conditions. A value of two was assigned to
``barrier estuaries'' which have narrow, restricted
entrance channels in which tides are attenuated. A
value of three was assigned to ``drowned river valley
estuaries'' which have deep, marine dominated
entrance regions with full tidal exchange.
A matrix comprised of the natural log-transformed
physical/environmental value for each attribute for
each estuary was constructed. From this matrix, a
triangular matrix of similarities between all pairs of
estuaries was calculated using normalised Euclidean
distance. This similarity matrix was then subjected to
cluster analysis by group-average linking to construct
a hierarchical agglomerative dendrogram for delineating spatial groups. The matrix of transformed physical/environmental attributes was then ordinated using
principal components analysis (PCA) to view the
spatial relationships. Log transformation was selected
for attributes in each type of multivariate analysis after
plotting the distribution of each attribute and observing the effects of the most common transformation
algorithms (Clarke and Warwick, 1994). All multivariate analyses were carried out on a personal computer using PRIMER version 3.1b and the
implementation of appropriate clustering and ordination techniques was based on the recommendations of
Clarke and Warwick (1994).

Bell and Edwards, 1980

West et al., 1985
Hydrographic charts; Lucas, 1976; West et al., 1985
Hydrographic charts; Lucas, 1976; West et al., 1985
Bell and Edwards, 1980
West et al., 1985

2.3. Commercial fisheries attributes

The commercial sheries catch and effort data for
this study were compiled from the mandatory monthly
catch returns which have been submitted to the NSW
Department of Fisheries by all licensed commercial
shers in NSW since 1940. Since 1990, shers in the
estuarine sheries have been required to submit a
separate monthly return for each estuary shed during
each month. They are also required to list the number
of days shed using each shing method and the
weight of each species or dened species group
caught, along with other information about shing
vessel, crew and catch disposal. For detailed information about the catch statistics collection and storage
system used see Pease and Grinberg (1995).
The annual mean number of days shed per sher
using each of 22 gear types (Table 2) and the annual
mean catch per sher of 81 taxa (Table 3) from each of
the 53 study estuaries during the ve year period
19911995 were used to conduct the spatial analysis
of the commercial shing method and catch data.
Therefore, the sampling unit for these attributes in
each estuary during this period was an approximation
of the average individual commercial sher.
Matrices of log transformed values of days shed by
each shing method for each estuary were compiled
along with matrices of log transformed values of catch
by each of the taxa for each estuary. Triangular
matrices of similarities between all pairs of estuaries
were computed using the BrayCurtis coefcient
(Bray and Curtis, 1957) for effort and catch separately.
The effort and catch similarity matrices were subjected to cluster analysis using group-average linking

B.C. Pease / Fisheries Research 42 (1999) 6786

71

Table 2
Summary of commercial fishing methods used in spatial analysis (mean days fished and standard deviation are the mean and standard
deviation of days fished per year by commercial fishers in all 53 study estuaries during the period 19911995)
Method

Mean days fished

Standard deviation

Nets
Seine, baitfish
Seine, garfish (bullringing)
Seine, beach (fish hauling)
Hoop or lift
Gill net, bottom set
Gill net, flathead
Gill net, splash/retrieve
Gill net, top set
Prawn beach seine (hauling)
Prawn wall net (running)
Prawn danish seine
Prawn set pocket
Prawn trawl

686
754
11 035
114
4604
3337
14 304
25 479
4460
4176
4284
4032
17 616

223
263
836
84
3169
1633
11 144
16 559
576
1608
1266
1127
1767

Traps
Crab
Eel
Fish
Lobster

17 319
6606
5781
1111

1909
2030
1767
410

2644
36
42

488
29
21

760
146

140
98

Lines
Handline
Jigging
Set line
Others
Hand gathering
Skindiving

to construct hierarchical agglomerative dendrograms

for delineating spatial groups. Signicance of spatial
boundaries was tested with one way analysis of similarities (ANOSIM as described by Clarke and Warwick, 1994). Diversity characteristics of methods and
taxa within groups was summarised with the
DIVERSE program (Clarke and Warwick, 1994)
which calculates total abundance, occurrence, Margalef's richness index and Pielou's evenness index at
each site (estuary). Methods and taxa discriminating
spatial groups were identied using the SIMPER
program (Clarke and Warwick, 1994). For each
method or taxon this program calculates the ratio of
average contribution to similarity between groups to
the standard deviation of similarity between groups.
This ratio will be referred to as the ``discrimination

index'' because a high value reects greater usefulness

for discrimination than lower values. Average abundance of each method or taxon within each group is
also summarised by the SIMPER program.
In order to view spatial relationships, the similarity
matrices were ordinated using non-metric multidimensional scaling (MDS as described by Kruskal,
1964). Similarity of estuaries is inferred from their
proximity in these two-dimensional plots. Finally, the
BIO-ENV procedure (Clarke and Ainsworth, 1993)
was used to analyse the correlation between spatial
patterns associated with physical/environmental attributes and the spatial patterns associated with catch by
taxa. This was done by selecting the subset of abiotic
variables which maximise the Spearman rank correlation between abiotic and biotic similarity matrices.

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B.C. Pease / Fisheries Research 42 (1999) 6786

Table 3
Summary of commercial fisheries taxa used in spatial analysis (mean catch and standard deviation are the mean and standard deviation of the
annual commercial catch (kg) from 53 study estuaries during the period 19911995)
Common name

Scientific name

Mean catch (kg)

Standard deviation

Finfish
Anchovy
Biddy, silver
Bream, black and yellowfin
Catfish, estuary
Catfish, forktailed
Catfish, unspecified
Dory, John
Drummer
Eel, pike
Eel, river
Fish, unspecified estuary
Flathead, dusky
Flathead, sand
Flounder, unspecified
Garfish, no bill
Garfish, river
Garfish, sea
Goatfish
Hairtail
Hardyhead
Kingfish, yellowtail
Leatherjacket, unspecified
Longtom
Luderick
Mackerel, blue
Mackerel, unspecified
Morwong, red
Morwong, unspecified
Mullet, flattail
Mullet, pink-eye
Mullet, sand
Mullet, sea
Mulloway
Nanata
Old maid
Pike
Pilchard
Salmon, Australian
Shark, bull
Shark, carpet
Shark, fiddler
Shark, hammerhead
Shark, shovelnose
Shark, unspecified
Snapper
Sole, black
Sole, lemon
Stingray
Sweep
Tailor
Tarwhine

Classes Chondrichthys and Osteichthys

Engraulis australis
Gerres subfasciatus
Acanthopagrus butcheri and A. australis
Cnidoglanis macrocephalus
Arius graffei
Families Plotosidae and Ariidae
Zeus faber
Girella elevata
Murainesox bagio
Anguilla australis and A. reinhardtii
Classes Chondrichthys and Osteichthys
Platycephalus fuscus
Platycephalus caeruleopunctatus and P. bassensis
Family Pleuronectidae
Arrhamphus sclerolepis
Hyorhamphus regularis
Hyporhamphus melanochir
Family Mullidae
Trichiurus coxii
Family Atherinidae
Seriola lalandi
Family Monacanthidae
Family Belonidae
Girella tricuspidata
Scomber australasicus
Family Scombridae
Cheilodactylus fuscus
Family Cheilodactylidae
Liza argentea
Myxus petardi
Myxus elongatus
Mugil cephalus
Argyrosomus hololepidotus
Class Osteichthys
Selenotoca multifasciatus
Dinolestes lewini
Family Clupeidae
Arripis trutta
Carcharhinus leucas
Orectolobus ornatus
Trygonorrhina spp.
Family Sphyrnidae
Aptychotrema rostrata
Class Chondrichthys
Pagrus auratus
Synaptura nigra
Paraplagusia unicolor
Families Urolophidae and Dasyatidae
Scorpis lineolatus
Pomatomus saltatrix
Rhabdosargus sarba

8299
152 855
342 148
1370
1149
16 981
639
649
2018
182 064
70 142
165 550
3040
2064
8955
26 482
11 201
2990
25 707
145
4304
21 098
1163
383 187
5785
63
201
48
110 844
4098
17 182
1 804 851
57 187
627
9066
3211
11 182
2092
7626
555
923
167
45
4339
3783
510
72
1552
385
43 694
29 239

4653
28 427
58 732
1226
1703
5606
600
824
2024
42 386
16 298
10 360
1519
590
3375
11 360
8075
5775
28 770
235
4882
4562
377
45 698
5968
68
176
19
29 210
2438
12 991
155 777
10 131
1144
3755
1701
4131
1225
4718
301
347
56
35
1875
387
156
40
819
292
8565
16 799

B.C. Pease / Fisheries Research 42 (1999) 6786

73

Table 3 (Continued )
Common name

Scientific name

Trevally, black
Trevally, silver
Trumpeter
Trumpeter, unspecified
Whitebait
Whiting, sand
Whiting, school
Whiting, trumpeter
Whiting, unspecified
Yellowtail

Siganus fuscesens
Pseudocaranx dentex
Latridopsis forsteri
Family Latrididae
Class Osteichthys
Sillago ciliata
Sillago bassensis and S. flindersi
Sillago maculata
Family Sillaginidae
Trachurus novaezelandiae

7631
82 449
2704
2664
36 292
137 926
387
37 735
748
32 477

2026
11 313
1348
1725
12 867
19 891
711
11 785
361
8500

Crustaceans
Crab, blue swimmer
Crab, mud
Crab, unspecified
Lobster, eastern rock
Prawn, eastern king
Prawn, greasyback
Prawn, school
Prawn, tiger
Prawn, unspecified
Shrimp, mantis

Phylum Arthropoda
Portunus pelagicus
Scylla serrata
Section Brachyura
Jasus verreauxi
Penaeus plebejus
Metapenaeus bennettae
Metapenaeus macleavi
Penaeus esculentus P. semisulcatus and P. monodon
Family Penaeidae
Order Stomatopoda

152 602
109 770
299
2970
83 714
37 452
587 816
2888
72 778
570

26 196
17 833
195
1737
20 081
17 106
147 738
2465
26 846
215

Molluscs
Calamari, southern
Cockle
Cuttlefish
Octopus
Pipi
Scallop
Squid

Phylum Mollusca
Sepioteuthis australis
Anadara trapezia
Sepia spp.
Octopus spp.
Family Donacidae
Pecten fumatus
Photololigo spp.

738
50 403
1346
16 814
2974
120
49 937

652
16 793
1344
6261
2468
214
6898

Other shellfish
Beachworms
Shellfish, unspecified

Phyla other than Chordata

Phylum Annelida, family Nereidae
Phyla other than Chordata

119
3671

56
1690

3. Results
3.1. Physical/environmental attributes
Cluster analysis of the physical/environmental attributes (Fig. 2(a)) indicated that, at the normalised
Euclidean distance of 4.0, there were three latitudinal
groups of estuaries. A northern group included only
estuaries located between the northern border of NSW
and 328S (Fig. 1). A central group contained most of
the estuaries located between 328 and 358100 S, as well
as the Clarence and Camden Haven Rivers from north

Mean catch (kg)

Standard deviation

of 328S and the Clyde River, Moruya River and Tuross

Lake from south of 358100 S. A southern group contained most of the estuaries between 358100 S and the
southern border of NSW, as well as Wollumboola and
Smiths Lakes from the central region.
The ordination of physical factors by principal
components analysis (Fig. 2(b)) shows the spatial
integrity of the three groups dened by the previous
cluster analysis. Information about the rst ve principal components is summarised in Table 4. The rst
two principal components accounted for most (76%)
of the variance in the data set. Principal component 1

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B.C. Pease / Fisheries Research 42 (1999) 6786

Fig. 2. (a) Dendrogram showing group average clustering of 53 estuaries based on eight physical/environmental attributes. Shading shows
groups delineated at a normalised Euclidean distance of 4.00; (b) Ordination of 53 estuaries by PCA of physical/environmental attributes.
Boundaries and shaded fill delineated by cluster analysis in Part a. Nestuaries north of 328, Cestuaries south of 328 and north of 358 100 and
Sestuaries south of 358 100 .

B.C. Pease / Fisheries Research 42 (1999) 6786

75

Table 4
Coefficients in the linear combinations of physical/environmental attributes making up the principal components (also shown are the
percentage variations explained by the first five principal components)
Variable

PC1

PC2

PC3

PC4

PC5

Latitude
Geomorphological type
Catchment area
Water area
Entrance depth
Entrance width
Mean annual rainfall
Seagrass area
% Variation explained

0.291
0.400
0.401
0.411
0.405
0.404
0.139
0.279
57

0.576
0.117
0.099
0.061
0.228
0.176
0.747
0.015
19

0.141
0.277
0.109
0.400
0.216
0.235
0.054
0.792
12

0.376
0.280
0.650
0.144
0.146
0.381
0.366
0.192
5

0.166
0.728
0.225
0.110
0.566
0.222
0.036
0.094
3

explained 57% of the variance in the data set and was

strongly associated with the physical factors related to
size of the estuary (Table 4). In decreasing order of
signicance these attributes were water area, entrance
depth, entrance width, and geomorphological type
(type 1 saline lagoons included the smallest estuaries
and type 3 drowned river valleys included the largest
estuaries). Principal component 2 explained 19% of
the variance and was strongly associated with physical
factors related to latitude, such as rainfall. The vectors
for estuary size and latitude are shown on the ordination (Fig. 2). Principal components 35 explained
only 20% of the variance and were primarily associated with seagrass area, catchment area and geomorphological type, respectively.
A summary of the distribution of each physical
factor with respect to the regions delineated in
Fig. 2 provides an insight into the way these factors

inuence the multivariate regionalisation process.

Fig. 3 summarises the distribution of geomorphological type among regions. All of the northern estuaries
are riverine barrier estuaries. The central region is
dominated by large lagoon-type barrier estuaries and
drowned river valley estuaries. The southern region
contains most of the small intermittently opening
saline coastal lagoons.
Fig. 4 summarises the distribution of the other
physical factors among estuaries within the three
regions. The lowest values for all of the physical
attributes were generally found in the small estuaries
of the southern region. The highest values for all of the
physical attributes directly related to estuary size
(water area, entrance depth and entrance width) were
generally found in the central region and the values for
these attributes in the northern region were intermediate to the values in the other two regions. Catchment
area is less closely related to estuary size and the
highest values were found in both the northern and
central regions. Average annual rainfall was inversely
related to latitude, with the highest rainfall in the
northern region. Seagrass area followed a similar
regional pattern to the attributes related to estuary size.
3.2. Commercial fishing method attributes

Fig. 3. Distribution of 53 estuaries by geomorphological type

based on Roy (1984) and by region delineated in Fig. 2. Northern
region n12 estuaries, central region n13 estuaries and southern
region n28 estuaries.

Two levels of cluster analysis were applied to the

shing method attributes before a regional pattern was
detected. Analysis of attributes for all estuaries
revealed a grouping pattern based primarily on water
area. Further cluster analysis of the group composed of
``large'' estuaries, resulted in the delineation of two
latitudinal groups with a single boundary at 358100 S.

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B.C. Pease / Fisheries Research 42 (1999) 6786

Fig. 4. Distribution of catchment area, water area, entrance depth, entrance width, average annual rainfall and seagrass area by estuary along
an increasing latitudinal gradient from the northern-most estuary at the origin. Shaded regions are those delineated in Fig. 2.

Cluster analysis of the shing method data for all

estuaries (Fig. 5(a)) indicated that, at a BrayCurtis
similarity of 40% there were three groups of estuaries

which were primarily separated by magnitude of water

area. The group labelled ``large'' contained most of
the estuaries that were generally larger than approxi-

B.C. Pease / Fisheries Research 42 (1999) 6786

77

Fig. 5. (a) Dendrogram showing group average clustering of 53 estuaries based on 22 fishing method attributes. Shading shows groups
delineated at a similarity of 40%; (b) ordination of 53 estuaries by MDS (stress0.10) based on 22 fishing method attributes. Boundaries and
shaded fill delineated by cluster analysis in Part a. Sestuaries smaller than 4 km2 and Lestuaries larger than 4 km2. Dashed line shows the
approximate boundary between estuaries smaller than 4 km2 and those larger than 4 km2.

78

B.C. Pease / Fisheries Research 42 (1999) 6786

mately 4 km2 in water area (only 8 of the 37 estuaries

in this group were smaller) and were located in all
three of the regional groups dened by the previous
analysis of physical attributes. The two estuaries
labelled ``small north'' were less than 4 km2 in water
area and were located in the northern region. The third
group, labelled ``small south'' comprised estuaries
generally less than 4 km2 in water area and located
in the southern region, except for Lake Wollumboola
which is slightly larger than 4 km2 and is located near
the southern boundary of the central region.
The ordination of the shing method data by MDS
(Fig. 5(b)) shows the spatial integrity of the three
groups dened by the previous cluster analysis. Estuary size generally increases from left to right. The
dashed line divides estuaries into those smaller than
4 km2 on the right (except for Swan and Wollumboola
Lakes, which are slightly larger) from those larger
than 4 km2 on the left (except for Brou Lake and Bega
River, which are smaller). A low stress value (0.10) for
the MDS implies that this two-dimensional ordination
provides a relatively good spatial representation of
similarity.
ANOSIM showed that shing effort per sher in
estuaries greater than 4 km2 was signicantly different
(P<0.001) from the effort in smaller estuaries. In
the smaller estuaries approximately half as many
shing methods were used and ``days shed'' was
signicantly lower (Table 5(a)). Parametric correlation (Zar, 1974) analysis (Table 5(a)) showed that

Table 5
Mean diversity attributes of fishing method days per estuarya,b
Estuaries
a. All estuaries
Large
Small
Correlation
b. Estuaries >4 km2
NorthCentral
South
a

Days

Methods

Richness Evenness

335
54
0.87

15
6
0.92

2.42
1.37
0.78

0.84
0.76
0.39

418
108

16
12

2.46
2.31

0.84
0.83

For the 30 large (>4 km2) and 23 small (<4 km2) estuaries.
Correlation is the correlation coefficient of the method attribute
with water area (log transformed).
b
For the regional groups delineated in Fig. 6 within the 30 large
(>4 km2) estuaries. In both a and b: daystotal days fished,
methodsnumber of fishing methods, richnessMargalefs index
and evennessPielou's index.

frequency (days shed) and richness (indicated by

number of methods and richness diversity index) of
methods were highly correlated with estuary size,
while evenness diversity was uncorrelated. SIMPER
analysis revealed that the beach seining method
showed the highest discrimination index (2.35)
between the two size groups and that the number of
days shed with every method was higher in the large
size group.
Further cluster analysis of the shing method data at
the next level of discrimination (only including estuaries generally larger than 4 km2 in water area) indicated that, at a BrayCurtis similarity of 42%, there
were two latitudinal groups of large estuaries
(Fig. 6(a)) which were separated at the southern
boundary dened by the analysis of physical attributes. The ordination of this method data by MDS
(Fig. 6(b)) conrms the spatial integrity of these two
latitudinal groups. In this ordination, large estuaries
from the northern and central regions within the
shaded group are oriented in a vertical latitudinal
gradient with moderate spatial mixing in the centre
of the gradient. The moderately low stress value (0.13)
for this MDS implies that this two-dimensional ordination is an adequate spatial representation of similarity.
ANOSIM was used to test the null hypothesis that
there was no signicant difference in days shed by
each method between large estuaries (>4 km2) within
the three latitudinal regions dened by the previous
analysis of physical attributes. Fishing effort in large
estuaries in the northern region was not signicantly
different (P>0.05) from the effort in large estuaries in
the central region, but shing effort from large estuaries in the southern region was signicantly different
(P<0.001) from that in the estuaries of the northern
and central regions. The average number of days
shed was much lower in the southern region but
richness of methods, as indicated by the number of
methods and richness diversity index, were only
slightly lower in the southern region and evenness
diversity was virtually the same in both regional
groups (Table 5(b)). SIMPER analysis showed
that the splash/retrieve gill netting method provided
the best discrimination (index1.91) between
regional groups and that the average number of
days shed by every method was lowest in the southern region.

B.C. Pease / Fisheries Research 42 (1999) 6786

79

Fig. 6. (a) Dendrogram showing group average clustering by 22 commercial fishing method attributes of 37 estuaries designated as ``large'' in
Fig. 5(a). Shading shows groups delineated at a similarity of 56%; (b) ordination of these 37 ``large'' estuaries by MDS (stress0.13) based on
22 fishing method attributes. Boundaries and shaded fill delineated by cluster analysis in Part a. Nestuaries north of 328, Cestuaries south
of 328 and north of 358 100 and Sestuaries south of 358 100 .

80

B.C. Pease / Fisheries Research 42 (1999) 6786

3.3. Commercial catch by taxa attributes

Analysis of the catch attributes provided results that
were very similar to those previously shown for the
analysis of commercial shing method attributes.
Again, two levels of cluster analysis were applied
before a regional pattern was detected. Analysis of
attributes for all estuaries revealed a grouping pattern
based primarily on water area. Further cluster analysis
of the group composed of ``large'' estuaries resulted in
the delineation of three latitudinal groups with the
same boundaries as those delineated by the PCA of
physical attributes.
Cluster analysis of the catch by taxa data for all
estuaries (Fig. 7(a)) and the resulting ordination
(Fig. 7(b)) revealed grouping patterns similar to those
observed for the shing method attributes; however,
Fig. 7(b) illustrates that the groups which were based
on cluster analysis of catch attributes showed an even
more distinct separation at the 4 km2 boundary than
those based on shing method attributes. The only
exceptions to the 4 km2 boundary were Swan and
Wollumboola Lakes in the ``small'' estuary group,
which were slightly larger than 4 km2, and Pambula
Lake and Bega River in the ``large'' group, which were
slightly smaller than 4 km2. The lower stress value
(0.09) for this MDS implies that this two-dimensional
ordination provides a slightly better spatial representation of similarity than the MDS of shing method
attributes (stress0.10).
ANOSIM showed that catch compositions from
estuaries greater than 4 km2 in area were signicantly
different (P<0.001) from those in smaller estuaries. In
Table 6(a) the catch per taxon and richness of taxa
(indicated by number of taxa and richness diversity
index) were much lower in the small estuaries. Catch
per taxon and richness of taxa (measured by number of
taxa and richness diversity index) were both highly
correlated with estuary size (Table 6(a)) but evenness
diversity was uncorrelated. SIMPER analysis revealed
that silver biddies (Gerres subfasciatus) and at-tail
mullet (Liza argentea) showed the highest discrimination indices (1.81 and 1.71, respectively) between the
two estuary size groups and that the average catch of
every taxon was higher in the larger estuary size
group.
Further cluster analysis of the catch data from
estuaries generally larger than 4 km2 in water area

Fig. 7. (a) Dendrogram showing group average clustering of 53

estuaries based on 81 catch by taxa attributes. Shading shows
groups delineated at a similarity of 42%; (b) ordination of 53
estuaries by MDS (stress0.09) based on 81 catch by taxa
attributes. Boundaries and shaded fill delineated by cluster analysis
in Part a. Sestuaries smaller than 4 km2 and Lestuaries larger
than 4 km2. Dashed line shows the approximate boundary between
estuaries smaller than 4 km2 and those larger than 4 km2.

(Fig. 8(a)) indicated that, at a BrayCurtis similarity

of 39%, there were three latitudinal groups of large
estuaries with boundaries that corresponded exactly
with those dened by the analysis of physical attributes. The ordination of this catch data using MDS

B.C. Pease / Fisheries Research 42 (1999) 6786

81

Table 6
Mean diversity attributes of catch per estuarya,b
Estuaries
a. All estuaries
Large
Small
Correlation
b. Estuaries >4 km2
North
Central
South

Catch

Taxa

Richness Evenness

11 635
1547
0.90

49
18
0.94

5.17
2.40
0.90

0.65
0.61
0.21

11 639
16 555
3638

48
59
34

5.04
5.99
4.00

0.58
0.68
0.70

From 30 large (>4 km2) and 23 small (<4 km2) estuaries.

Correlation is the correlation coefficient of the catch attribute with
water area (log transformed).
b
Mean diversity attributes of catch per estuary within the regional
groups defined in Fig. 8 for the 30 large (>4 km2) estuaries. In both
a and b: catchtotal catch, taxanumber of taxa, richnessMargalef's index and evennessPielou's index.

(Fig. 8(b)) conrms that all three groups are spatially

distinct, except for the inclusion of Clarence River in
the central region. The low stress value (0.11) for this
MDS implies that this two-dimensional ordination is a
good spatial representation of similarity.
ANOSIM was used to test the null hypothesis that
there was no signicant difference in catches from
large estuaries (>4 km2) among the three latitudinal
regions dened by the previous analysis of physical
attributes. Catches from each region were found to be
signicantly different (P<0.001) from catches in the
other regions. The average catch per taxon and richness of taxa (indicated by number of taxa and richness
diversity index) were lowest in the southern region and
highest in the central region (Table 6(b)). Evenness
diversity increased from north to south. Results of the
SIMPER analysis are summarised in Table 7, which
shows the four taxa with the highest discrimination
values and their average regional catches for each of
the three regional comparisons. All of the discriminating taxa except for squid are identied to the species
level and the primary discriminating species were all
nsh. Sea mullet (Mugil cephalus) is a key discriminating species for both the north/south and central/
south comparisons, while silver biddies provided the
best discrimination for the north/central comparison.
The highest catches of most discriminating sh species (sea mullet, silver biddy, tarwhine (Rhabdosargus
sarba), and trumpeter whiting (Sillago maculata)) are
obtained with the beach hauling method.

Fig. 8. (a) Dendrogram showing group average clustering by 81

catch by taxa attributes of 30 estuaries designated as ``large'' in
Fig. 7(a). Shading shows groups delineated at a similarity of 62%;
(b) ordination of these 30 ``large'' estuaries by MDS (stress0.11)
based on 81 catch by taxa attributes. Boundaries and shaded fill
delineated by cluster analysis in Part a. Nestuaries north of 328,
Cestuaries south of 328 and north of 358 100 and Sestuaries
south of 358 100 .

The BIO-ENV technique was used to investigate the

correlation of the MDS ordination of catch by taxa in
the large estuaries (>4 km2) with the PCA ordination
of physical attributes in the large estuaries. The highest Spearman rank correlation coefcient of 0.58
resulted from the combination of latitude and water
area, which indicates that these are the primary

82

B.C. Pease / Fisheries Research 42 (1999) 6786

Table 7
Comparison of regions by top four discriminating taxa based on SIMPER analysis of species data from large (>4 km2) estuariesa
Common name

Scientific name

Discrimination index

Average catch (kg)

Silver biddy
Tarwhine
Squid
Yellowtail

Gerres subfasciatus
Rhabdosargus sarba
Photololigo spp.
Trachurus novaezelandiae

2.17
2.08
1.95
1.88

33
2
1
1

Sea mullet
Mud crab
Bull shark
Old maid

Mugil cephalus
Scylla serrata
Carcharhinus leucas
Selenotoca inultifasciatus

3.19
2.61
2.24
2.01

3830
413
46
39

Sea mullet
Trumpeter whiting
Blue swimmer crab
Squid

Mugil cephalus
Sillago maculata
Portunus pelagicus
Photololigo spp.

2.06
1.97
1.82
1.82

North

Central

South

1155
207
192
435
586
9
0
0
2569
254
447
192

586
8
9
2

Discrimination indexratio of average dissimilarity for this taxon to standard deviation of dissimilarity for the taxon. Average catch is the
average catch per fisher in each of the two regions being compared.

physical factors explaining the shared relationship

between the physical and biological variables.
4. Discussion
Multivariate analysis of the abiotic and biotic variables in this study indicates that the estuaries of NSW
can be grouped into three latitudinal regions, with
boundaries at 328 and 358100 S. The physical factors
that contribute most to this regional structure appear to
be those related to estuary size and latitude. It is
difcult to assign a more specic role to physical
and environmental factors because they tend to be
highly intercorrelated and cause/effect relationships
are poorly understood (Horn and Allen, 1976; Monaco
et al., 1992).
Estuary size is a primary factor in the delineation of
regional distribution patterns of estuaries in NSW, as
well as determining the nature of the commercial
sheries within them. The rst principal component
of the PCA (Table 4) was dominated by ve variables
associated with estuary size: water area, entrance
depth, entrance width, catchment area and geomorphological type. The distribution pattern of these
variables (Figs. 3 and 4) indicates that most of the
largest estuaries, which are typically drowned river
valleys and large barrier lagoon estuaries, are found in

the central region. Most of the smallest estuaries,

many of which are saline coastal lagoons, are located
in the southern region. All of the medium to large
sized estuaries in the northern region are riverine
barrier estuaries.
The initial classication of all estuaries by shing
method and catch by taxa also resulted in a size-based
clustering pattern (Figs. 5 and 7). This pattern in the
spatial distribution of sheries variables is caused
primarily by the fact that shers use signicantly
fewer shing methods in estuaries smaller than
approximately 4 km2 (Table 5). The pattern is
enhanced by the fact that beach seining is not generally used in these small estuaries, whereas the largest
estuarine catches of many sh species are obtained by
this method. Analysis of the sheries variables in
estuaries larger than 4 km2 (Figs. 6 and 8) indicates
that the number of shing methods also plays an
important role in separating the sheries of the relatively small estuaries in the southern region from those
in the other regions. However, the signicant difference of catch by taxa between the large estuaries of the
northern and central regions was not apparently linked
to a signicant difference in shing effort.
The high correlation between richness of taxa and
water area (Table 6), along with the BIO-ENV correlation between water area and catch by taxa, indicate
that estuary size is a primary factor delineating the

B.C. Pease / Fisheries Research 42 (1999) 6786

catch composition of estuarine sheries. The mechanism behind this relationship is complex and undoubtedly involves many abiotic and biotic variables.
Structure and complexity of shing effort is one
important factor confounded with many environmental factors. Horn and Allen (1976) and Monaco et al.
(1992) also found a signicant correlation between
species richness (presence/absence) of shes and
estuary size on the west coast of the United States.
Using multiple regression techniques, Horn and Allen
(1976) identied estuary mouth width as the only
signicant predictor of species number, while Monaco
et al. (1992) found mouth depth to be the best predictor. In the current NSW study, both mouth depth
and width had high coefcients in the rst principal
component of the PCA (Table 4). Mouth depth and
width generally increase with increasing estuary size.
A larger entrance also indicates a greater degree of
marine inuence with greater access to and from the
marine environment, which is generally associated
with higher species richness than euryhaline estuarine
environments. Larger estuaries tend to have greater
heterogeneity of habitats, which also leads to
increased species richness (Gilmore, 1995). Seagrasses provide a particularly complex habitat which
has important nursery functions for many commercially important species (Pease et al., 1981; Bell and
Pollard, 1989). Seagrass area in the estuaries of this
study was regionally distributed similarly to the variables related to estuary size (Fig. 4).
Estuary size is also linked to geomorphology, runoff
and associated entrance opening regimes. Hurrell and
Webb (1993) found a linear relationship between
water area and catchment area of estuaries in NSW.
They showed that the smallest estuaries, closest to the
origin of the relationship, tend to be closed for longer
periods than they are open because runoff is directly
related to catchment area. Estuaries of an intermediate
size are intermittently closed, but remain open most of
the time and the largest estuaries all remain permanently open. Comparisons of the sh communities in
intermittently open and nearby permanently open
estuaries in Australia (Pollard, 1994; Potter and
Hyndes, 1994) and South Africa (Bennett, 1989) have
shown that fewer sh species generally occur in the
intermittently opening estuaries. Therefore, another
factor explaining the signicantly lower species richness in estuaries smaller than 4 km2 is the fact that

83

65% of these are only intermittently open (West et al.,

1985). It is also worth noting that 80% of the intermittently opening estuaries occur in the taxonomically
depauperate southern region. Furthermore, both central coast estuaries which the PCA associated with the
southern region (Smiths and Wollumboola Lakes) are
intermittently opening.
Most of the regional outliers or exceptions in the
analysis of physical/environmental and catch attributes are probably related to variablility in estuary
size within latitudinal regions. The Clarence River
is located in the northern region but the PCA of
physical attributes and MDS of catch attributes both
put this estuary into the central group. Clarence
River is the largest estuary in the northern region
and the fourth largest estuary in NSW. The PCA also
put the two largest estuaries in the southern region
(Clyde River and Tuross Lake) into the central group
and the two smallest estuaries in the central region
(Smiths and Wollumboola Lakes) into the southern
group.
Latitude is the other general physical variable that
appears to play an important role in the classication
of estuaries and the sheries within them. However, as
discussed above, this role is apparently secondary to
that of estuary size. Latitude provides a simple, spatially quantiable variable; however, it is autocorrelated with a large array of environmental variables
such as temperature, rainfall and wind patterns,
including those variables which may actually determine estuary size. Latitude and latitudinally distributed average annual rainfall (Fig. 4) were the primary
coefcients contributing to the second principal component of the PCA (Table 4) and provide the spatial
orientation and consistency of the classication into
northern, central and southern regions. Bucher and
Saenger (1994) also found that average annual rainfall
played a key role in classifying tropical and subtropical Australian estuaries.
The BIO-ENV correlation between latitude and
catch by taxon indicates that latitude is another primary factor delineating the catch composition of
estuarine sheries. However, Table 6 illustrates that
this correlation is not derived from a direct relationship between latitude and species richness. Horn and
Allen (1976) and Monaco et al. (1992) also found that
cluster analysis of sh species (presence/absence) in
estuaries on the west coast of the United States

84

B.C. Pease / Fisheries Research 42 (1999) 6786

resulted in latitudinally oriented groups. However,

multiple regression analysis showed that latitude
was not a signicant predictor of species richness.
These results support the ndings of Rohde et al.
(1993), that Rapoport's rule (species richness generally increases with decreasing latitude) cannot be
generally applied to estuarine or marine teleost shes.
Horn and Allen (1976, 1978) indicated that the
latitudinal structure of estuarine and coastal sh fauna
along the California coast is primarily related to water
temperature and oceanographic boundaries. Recent
coastal bioregionalisation studies in NSW by Ortiz
(1994) and Pollard et al. (1998) relate the inuence of
the East Australian Current (EAC) to multivariate
analysis of coastal marine sh distributions (recorded
presence/absence). They found that the inuence of
this warm, western boundary current (Cresswell,
1987) divides the coast of NSW into three regions
which correspond well with those delineated in the
present study. The EAC ows southward from the
Coral Sea, and typically remains close inshore on the
continental shelf until it reaches an easterly coastal
protrusion, such as Cape Byron (288390 S), Smoky
Cape (308570 S) or Sugarloaf Point (328260 S), where
it diverges from the coast and forms large eddies. Ortiz
(1994) found that the EAC separates from the continental shelf most frequently at Sugarloaf Point and
provides a sub-tropical inuence on the continental
shelf waters north of Sugarloaf Point 90% of the time.
On the continental shelf southwards of Sugarloaf Point
to Beecroft Head (358S), the warm temperate waters
of the northern Tasman Sea are inuenced around 50%
of the time by sub-tropical water in eddies from the
EAC that impinge on the coast. South of Beecroft
Point, coastal trapped waves hold cold temperate
water from the southern Tasman Sea inshore and warm
water from the EAC only impinges around 10% of the
time. Using multivariate techniques, Ortiz (1994) and
Pollard et al. (1998) found that coastal sh species
distributions also t into this model of northern,
central and southern regions with similar regional
boundaries.
Recent multivariate analysis of oceanographic data
for the marine surface (150 m) waters around Australia by the CSIRO Divisions of Fisheries and Oceanography (1997) also resulted in a very similar pattern
of northern, central and southern regions adjacent to
the NSW coast, with boundaries at 328 and 358S.

Temperature, salinity, nitrate, silicate and dissolved

oxygen measurements collected by research vessels,
satellites and surface drifters were used in the classication analysis and the resulting groups were ordinated on two-dimensional maps. The resulting map
provides a visual summary of the EAC processes
discussed by Ortiz (1994).
The multivariate techniques used in the current
analysis result in a bioregional pattern for the estuarine
sheries of NSW that is consistent with the ndings of
other bioregional studies in Australia and the United
States. As early as 1953, Bennett and Pope recognised
that intertidal ora and fauna along the coast of NSW
was associated with three biogeographical provinces
(Solanderian, Peronian and Maugean). More recently,
Pollard et al. (1998) identied three ``major coastal
biophysical regions'' in NSW with similar boundaries
to those identied in the current study. These studies
show general agreement despite the fact that different
techniques were used in each study and sampling bias
occurs in all such bioregionalisation studies. Biological attributes used in the current study are based on the
commercial sher as a sampling unit and incorporate
both quantity and taxonomic composition of the average catch. It is understood that shers introduce bias
by sampling non-randomly with a range of selective
gears (Hilborn and Walters, 1992; Stergiou and Pollard, 1994) and that non-biological, socio-economic
factors such as local traditions, market dynamics and
management strategies inuence a sher's sampling
activity. The other studies mentioned used presence/
absence data collected primarily by researchers. Prendergast et al. (1993) demonstrated that presence/
absence data from large-scale faunal surveys are often
subject to bias caused by variation in recording intensity, which is undoubtedly magnied for highly
mobile marine species. The method of applying multivariate analysis simultaneously to attributes of the
physical environment and commercial sheries, as
described in this paper, provides a useful means of
identifying regions in multi-species sheries with
complex shing area and effort components. These
techniques can be easily applied to other complex
sheries.
The main purpose of this study was to dene and
describe spatial patterns in the estuarine sheries of
NSW. The primary strength of this multivariate
approach is that it incorporates a wide range of

B.C. Pease / Fisheries Research 42 (1999) 6786

physical/environmental, shing effort and catch attributes in the descriptive process. Analysis of the shing
effort data indicates that the sheries attributes are
subject to variability associated with estuary size,
which confounds relationships between sheries attributes and environmental factors. However, along with
the more general regionalisation results, the enhanced
description of shing effort and catch characteristics
of the complex estuarine sheries provides information which is potentially very useful for future management.
With signicantly lower commercial catch and
effort, those estuaries less than 4 km2 in area should
be considered for closure as estuarine harvest refugia.
Because of their small size these estuaries are less
accessible for commercial shing activity than larger
estuaries but more vulnerable to overshing, potential
ecosystem damage and social conict with residents
and recreational users. Pollard (1994) showed that
small intermittently opening lagoons support a lower
diversity of non-commercial sh species than larger,
permanently opening lagoons, but also demonstrated
that they may support signicant quantities of commercially and recreationally important estuarine sh
species. The distribution of these smaller estuaries, as
potential harvest refugia along the coastline, should
provide enhanced recruitment opportunities for many
inshore coastal species (Pollard, 1976). Recent unpublished tagging studies show extensive movement of
species such as yellown bream and blacksh between
estuaries in NSW, indicating that such small estuarine
refugia could potentially enhance stocks of these
species in the surrounding larger estuaries.
The three latitudinal regions which have been
identied by this study should be considered as potential ``management units''. The development of management plans for the State's estuarine sheries should
be structured around a recognition of regional factors.
In fact, the Management Advisory Committee (MAC)
for the estuarine sheries of NSW is currently using
these three ``bioregions'' in a proposed zoning policy
(Zantiotis-Linton, 1998) to reduce social conict in
the shery by restricting the activity of individual
estuarine shers to a single bioregion. Regional factors should also be considered when reviewing and
restructuring input controls. Another recent proposal
by the estuarine sheries MAC to standardise and
simplify the seasonal regulation of gill net soak times

85

in this shery also employs these bioregions, recognising the regional variability in seasonal water temperatures. The three estuarine bioregions described
may also provide a useful framework for future stock
assessment and monitoring of commercially and
recreationally important sh and shellsh species,
recognising that factors such as shing effort and
growth rates vary regionally.
Acknowledgements
The author wishes to thank Dr. John Glaister and
Kevin Rowling for their interest and support, without
which this work would not have reached fruition. I
also wish to thank Trudy Walford for her help with
graphics, Rossana Silveira for her assistance with the
PRIMER software and Christine Allen, Dr. Rick
Fletcher, Dr. Dave Pollard, Dennis Reid, Kevin Rowling and Rob Williams for reviewing the paper and
providing helpful comments.

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