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Estuarine, Coastal and Shelf Science (1985) 2lr 5ll-537

Annual Variation in the Nocturnal Nekton


Assemblage of a Tropical Estuary

NormanJ. Quinno and Barbara L. KoiisD


I Fisheries Departmento and Department of Chemical Technologgb, Papua New
1
Guinea Unioersity of Technology, Lae, Papua New Guinea
I
i Receiaed 23 October 1983 and in revisedform 4 March 1985

Keywords: Oceanography; Fourier analysis; time-series; salinity; fish; water


temperature; regression analysis; Papua New Guinea

A nocturnal demersal nekton assemblage was sampled fortnightly for two years
at five sites in the Labu estuary using a 3 m beam trawl with a3.2 cm mesh net.
Forty-eight species were caught, totalling 31 458 individuals with the five most
abundant species comprising over 950,1 of the catch. Using multiple regression
techniques with Fourier transformations, the mean number of species, S, the
mean abundance, N, and mean weight, W, wete found to conform to a regular
annual cycle with maxima in April and May. Seven of the 11 most abundant
spicies demonstrated regular annual cycles of abundance. S, N and W were
greatest in the wider, middle sites and lowest in a shallow, stagnant side branch
of the estuary. Catch weights and abundances were significantly correlated with
physical data.
Salinity and temperature values in the estuary exhibited an annual cycle with
maxima occurring in February/March. The annual thermal variation of surface
water outside the estuary followed a similar cycle. The salinity at the mouth of
the Markham River is lowest during January/February, which corresponds
with the rainy season in the Markham River catchment. Significant annual vari-
ation existed between years in estuarine bottom salinity and salinity values in
Labu Bay.
Several species exhibited a greater variation in abundance and mass between
years than within years. This supports the hypothesis that in the tropics
between-year variation in coastal marine biotic communities is greater than
within-vear variation.

Introduction
Knowledge of the fish inhabiting Papua New Guinea estuaries is largely confined to lists
of species (Munro, 1967; Liem & Haines, 1977; Haines, 1979; Berta et al., 1975;
Collette, 1983), taxonomic descriptions (Collette, 1982) and studies of the biology of
barramundi, Lates calcarifer (Moore, 1982; Moore & Reynolds, 1982; Reynolds &
Moore, 1982). Studies of fish of the Huon Gulf and seasonality of Papua New Guinea
estuarine fish are non-existent. There are only a few published accounts of the fish
faunas of the tropical estuaries of this region, e.g. Singapore (Thia-Eng,1973) and North
Queensland (Blaber, 1980).
The importance of estuaries as feeding and nursery grounds for fish and the need to
assess the effects of existing or potential alterations by man have resulted in increased

5ll
027 2-77 14185 I 1005r l + 27 $03.00/0 @ 1985 Academic Press Inc. (London) Limited
512 N. J. Quinn {s B. L. Kojis

attention to these areas throughout the world. Recently interest in mangrove communi-
ties in Papua New Guinea has been stimulated by the prospects of three major develop-
ment schemes the Purari hydroelectric power scheme, proposals by Japanese
-
companies to develop the large unexploited mangrove resources of the Papuan Gulf and
the Lae port expansion.
Mangrove and coastal ecosystems in Papua New Guinea are relatively undisturbed.
Local villagers use mangroves as a source of building material and firewood, but
the population density is low and the effect is small. Industrial use of mangroves is
also limited; only 19400 tons of mangrove bark were exported from I89l to 1970
(unpublished report of the Office of Forestry, undated, entitled Mangroae Stands:
Papua New Guinea). There has been no recent increase in mangrove exploitation.
The only full-time commercial fishery is in the Gulf of Papua where increases in
catches are recent (1981:. 32.8 tonnes fis}e, 4-7 tonnes crabs; 1982: 68'7 tonnes fish,
3'0 tonnes crab) (Anon., 1983). As utilization of these virtually untouched coastal
ecosystems is inevitable, a baseline of information is necessary from which comparisons
can be made.
This study describes the trawlable demersal fish and crustacean communities of the
Labu estuary adjacent to the proposed expansion of the Lae wharf and discusses spatial
and temporal changes in abundance and seasonality in relation to variations in abiotic
parameters.
Additionally, the fauna from the estuary is compared with that of a subtropical estuary
(Serpentine Creek, Brisbane, Queensland, Australia) (Figure l).

t40 "E

Popuo
New
Guineo Loe
lVorkhom

Porl
expo ns ion
t
Vrl qqe

Huon
Gui f
A
I

North
Queenslond
lt
lkm Auslro lio

Bri s bo ne

140"8

Figure l. (a) Regional map showing Lae, Papua New Guinea and Brisbane, Australia.
(b) Labu estuary trawling sites. Trawl sites are numbered. Sites where hydrological
data were collected are indicated with X.
N octurnal nekton assemblage aariation 513

Materials andMethods
Physical data sampling
Sampling was carried out fortnightly from June 1980 to May 1982 from 17.00 to 23.00h
at seven sites. Dissolved oxygen, temperature and salinity were measured in situ using a
'Kahlisco' conductivity meter with thermistor and a dissolved oxygen meter at each site.
Measurements were taken on the surface and near the bottom at the completion of each
trawl. S7ater tr€ursparency was measured using a 20 cm diameter Secchi disc. Rainfall
data for the years 1973 to 1982 were obtained from the Civil Engineering Department at
the Papua New Guinea University of Technology.

Biological sampling
On the basis of a pilot survey from March to June 1980 five sites were sampled on a
fortnightly basis from June 1980 to May 1982. A 3 m beam trawl with a stretched mesh
of 3'2cm, identical to that used by Quinn (1980), was trawled heading upstream for
about 300m at each site for 15 min at approximately one knot. The beam trawl was
chosen because it is an active fishing gear that yields quantitative results which can be
compared with work done at lower latitudes in Queensland, Australia.
All sampling methods are biased because any one type of equipment is more suitable
for catching some species than others. For example, in this study the beam trawl caught
mostly bottom fish while surface feeding (e.g. gar) and schooling (e.C. mullet) fish were
not caught at all, although they were present in the estuary.
Sampling was carried out on 4l out of a possible 48 dates. Inclement weather and
other difficulties account for the missing sampling sets. $Tithin each sampling date not all
sites were trawled owing to snags, the presence of too many jellyfish (Catostylus sp.) and
requests by villagers to forego trawling a site when numerous gill nets had been set.
Approximately 97o/, of sites were trawled within the sampling dates.
To reduce collecting variability, trawling began at Site I about l-2 h after sunset.
Replicate trawls were not taken owing to time limitations imposed by a suspected diel
variation in the fauna. This is discussed in greater detail in Quinn & Koiis (1983). Lunar
phase was considered a controlled variable since sites were trawled each fortnight during
new and full moons (Quinn & Kojis, 1984).
Captured fish were counted, standard length (S.L.) measured to the nearest centi-
metre and returned to the water. Samples of 12 species comprising the most abundant
taxa were collected and brought back to the laboratory for length/weight measurements.
From these data a power curve equation for the length/weight relationship was derived
using a Texas Instruments power curve programme. Veight estimates were made by
taking the median value for a species' size class and using the length/weight equation for
that species.
Unfamiliar fish were preserved for subsequent identification. The genus Upeneas had
two morphologically similar 'species' that were difficult to separate in the field; one is
probably a new species (Johnson, personal communication). Authorities used in the
identification of fish were Munro (L967) and Carcasson (1977). Specimens were lodged
with the Queensland Museum, Brisbane, Australia.

Analysis

D escriptia e statistics
The mean total number of individuals per trawl is designated by N, the mean total
514 N. J. Quinn {x B. L. Kojis

weight by lV and total number of species by S and are termed sample parameters. The
abundance and weights of individual species are identified by n and z{, respectively. In
temporal analyses the means were determined across sites, while for spatial analyses
means were calculated through time. The amplitude of the coefficient of variation (C.V.)
was used as an index of the degree of heterogeneity of the estuarine water and biota.

H armonic r egr e s sion p erio di ci ty


In ecological studies two types of non-random time-series patterns can occur. The first
may involve a series of observations which show repetition although no apparent regu-
larity. These are difficult to characterize and often reflect randomly fluctuating or
irregular hydrological conditions. The second, and the one considered here, involves a
series of observations with regular repetition through time. Biotic cycles of this form are
believed to be caused by regular variation in abiotic and/or biotic factors and usually
persist for extended periods.
\Ve fitted our data to a Fourier transform model using sine and cosine terms, because
they provide a powerful tool for describing periodic phenomena. Bliss (1958) has pro-
vided a comprehensive description of harmonic regression analysis with examples from
agriculture. A similar type of analysis was used by Maddock & Swann (1977) to investi.-
gate trends in sea temperature and climate, by Hamon & Kerr (1968) to investigate
the time variation in the EaSt Australian current and by Quinn (1980) for the hydrology
of a sub-tropical estuary. Only regressions over the 95o/o confidence level were accepted
and plotted.
Studies of periodicity using mathematical models are uncommon and relatively recent
in marine ecology. Platt & Denman (1975) in their review of spectral analysis in ecology
cited only I I studies, of which seven dealt with marine organisms. Bulmer (197 4) used a
mixed model, which included periodic terms, to analyse a l0-year cycle in Atlantic
salmon angling records. Recently these techniques have been applied to the Australian
marine fauna (Stephenson, 1978, 1980a, 1981; Stephenson & Burgess, 1980; Stephenson
et al.,l982a,b; Gilmour & Stephenson, 1983).
Multiple regressions
Studies analysing the effect of several abiotic parameters on fish (Risk, 1972; Oviatt &
Nixon, 1973; Quinn, 1980) and prawn (Stephenson, 1980b; Stephenson & \Tilliams,
1981) populations have only recently been analysed using statistical methods.
As the composition of the nekton may be the result of physical conditions during the
previous several months, regressions were also performed on lagged physical data. In
this analysis only temperature and salinity are used to estimatey values, the abundance
or weight of nekton populations. The lagging interval used ranged from one to five
fortnights. As the lag increases, the number of valid observations declines. Given the
limitations of the existing data set, five fortnights (2.5 months) was the longest lag
examined. If more biological knowledge about the populations had been available,
further analyses with increased lags might have indicated which stages of the life cycle
are most affected by certain abiotic factors (Stephenson & \Tilliams, 1981).

V ariations betw een y ear s


Using a paired r-test, variations in physical and biological data between years were
investigated. Similar lunar phases from June 1980 to May 1981 were compared with
June 1981 to May 1982. Owing to missed sampling dates in either of the years, a subset
of 15 lunar pairs were available for comparison.
Nocturnal nekton assemblage aariation 515

Results
Phgsical data

Estuarine sites
The mean salinity and coefficient of variation for the five trawl sites per sample time are
plotted in Figure 2. The highest recorded bottom salinity was 30'5%o in February 1981,
while the lowest was 3'9%o in October 1981 (range 28'6%o; annual mean over all times :
18.2%o; C.V.:28'60/.; N:38). Surface and bottom salinities followed annual cycles
which peaked in February (Table l). Mean surface salinity varied from 0'6%o to 26'6%o
(range 26'0%'; r : 8'0%; C.V. : 75'7o/"; N : 39). The salinity range between sites was
greatest in August/September.
Bottom temperatures averaged over sites for each date were lowest in August 1980
(26'5 "C) and highest in November 1980 (32'3 "C), a range of 5'8 "C (Figure 3)' Both the
surface and bottom temperatures followed an annual cycle that peaked in February. The
surface temperatures ranged from a maximum of 32'l 'C in November 1980 to a
minimum of 24'4 'C in August 1980 (range 7'7 "C; x : 29'2 'C; C.V. : 7'0'/.; N : 40).
Bottom water temperature was slightly warmer (x : 29'6 'C; C.V. : 4'9'A;N : 39).
Dissolved oxygen values followed no annual cycle (Figure 4). Mean surface values
ranged from 2'6mgl-1 to saturated at about 6'2mgl-1 at 30'C (range 3'6mgl-1;
i:3'7 mgl-1; C.V.: 23'5'/,; N:24). Bottom readings varied from 0'2mgl I to
3'8mgl-1 (range 3'6mgl-1; x :2'lmgl-1; C.V. :45'7'/Lt N:24.). r$fater trans-
parency varied from 0'9 to l'7m G: l'2m; C.V.: l3'l%) with no annual cycle
(P>0.05) (Figure 5). Hydrological differences between sites were calculated from the

c
.9
,o
o
o o
o\
c
9
.9
=
.:
o o
U) O

ASON J JA JFMA
t980 I 98r 1982

Figure 2. Times series of surface and bottom salinity mean values for five sites in the
Labu estuary. -, predicted salinity values; observed values; -----, coeffrcient
of variation.
-,
516 N. J . Quinn Cx B. L. Kojis

TABLE 1. Regression coeffcients for hydrological observations from Labu estuary - hve
site mean

Month of
maximum
A B M iAo R2 P N value

Surface temperature,
oC
- 1.13 -1.67 0.01 29.06 0.53 <0.001 39 Early Feb.
Bottom temperature,'C -0 6l 1.26 -0 0l 29.97 0.41 < 0'001 38 Feb.
Surface salinity, %o -2.38 - 6.98 -0.12 11.67 0.61 < 0.001 38 Feb.
Bottom salinity, %o -0.47 -5.76 -0'22 t:u 0.56 < 0.001 38 Late Feb.
Surface dissolvedO, mg I t 0.17 N.S. 23
Bottom dissolved O, mg l-l 0.r2 N.S. 23
Transparency, m 0.13 N.s. 37

Where l, B, Mo and constant are variables in the equation:


Y -- \A^ + Asin( zn-J-\ * r.o, ( r"=! \ *r,,t
\ 25.64) \ 25.64 )
Trigonometric functions are in radians.

c
O ;.9o
o
+
o
o
d
E 80 .9
F
;
60 oo
O

Figure 3. Time series of surface and bottom water temperature mean values for five
sites in the Labu estuary. -, predicted temperature valuesl observed values;
----, coefficient of variation. -,

mean over time (Table 2). As expected the site nearest the mouth of the estuary had the
highest mean bottom salinity, 20'0%o, while sites further upstream had lower salinities
with site 5 having the lowest at L7.2%o. Mean surface salinities followed a similar pattern
ranging from9'60/oo at the mouth to 6'6%o at site 5. Coefficients of variation were similar
for all sites with the surface salinities coefficients being over twice as high as bottom
salinities.
N octurnal nekton assemblage aariation 517

i c
.o
o
E a
'a
c o
o
0 o
0
E .o
o
.9
o
o
i5 U

JJJ A SOND J FMAMJ J ASONDJ FMA


1980 r98t t982
Figure 4. Time series of surface and bottom dissolved oxygen mean values for five sites
in the Labu estuary. observed valuesl - - - -, coefficient of variation.
-,

c
E
,9
2 40 _e
o
c
o
ol
o 200
c
c .9
o
Fo J J A S O N D J F M A M J J A S O N D J F MA
ni;
o
o
t980 l98r t9 82 O

Figure 5. Time series of water transparency mean values for five sites in the Labu
estuary. observed valuesl - - - -, coefficient of variation.
-,

Mean surface temperatures were approximately uniform (29.5'C) for all sites (Table
2). Mean bottom temperature for site 4 was about I 'C higher than the other sites. This
site was furthest upstream and in an isolated section of the estuary with no tributaries.
The coefficient of variation was less for bottom than surface temperatures.
Mean surface dissolved oxygen readings were uniform around 34mgl-r (Table 2).
Sites I and 2 had similar mean dissolved oxygen readings of about 3 mg I - 1, while sites 3
and 5 were slightly lower. Site 4 had the lowest dissolved oxygen readings and occasion-
ally hydrogen sulphide odours were present. Bottom reading were more variable than
surface readings.
Mean transparency ranged from l'1 to l'3 m (Table 2). Turbidity was slightly greater
at site I than at the other sites, probably owing to the increased effect of tidal mixing.
There was greater variability in the hydrological observations through time than
through space (Table 2). The bottom temperature C.V. was l'3o1" between sites and
518 N. J. Quinn €s B. L. Kojis

T tstn 2. Summary of hydrological data for five sites in Labu estuary

Physical data Mean C.V. N

Site 1
Surface temperature,'C
oC
29.3 7.9 4l
Bottom temperature, 29.5 5.6 39
Surface salinity, %o 9.5 64.3 40
Bottom salinity, %o 19.8 25.7 37
Surface dissolved oxygen, mg I 1
4.1 27.6 20
Bottom dissolved oxygen, mg I 1
3.4 49.8 20
Transparency, m 1.1 26.0 37

Site 2
Surface temperature,'C 29.4 66 41
Bottom temperature, oC 29.5 5.5 39
Surface salinity, %o 8.3 7t.5 40
Bottom salinity, %o 182 32.4 37
Surface dissolved oxygen, mg I - 1
37 222 23
Bottom dissolved oxygen, mg I - I 2.9 57.1 23
Transparency, m 1.2 75.5 37

Site 3
Surface temperature,'C 29.4 6.6 4t
Bottom temperature, oC 295 5.5 39
Surface salinity, %o 8.3 71.5 40
Bottom salinity, %o 18.2 32.4 37
Surface dissolved oxygen, mg I 1
3.7 22.3 23
Bottom dissolved oxygen, mg l- 1
2.9 571 23
Transparency, m r.2 15.6 37

Site 4
Surface temperature, oC 30.0 6.4 40
Bottom temperature,'C 30.4 5.3 38
Surface salinity, %o 6.6 9t.7 38
Bottom salinity, %o 187 24.9 35
Surface dissolved oxygen, mg I I
3.8 229 23
Bottom dissolved oxygen, mg l- 1
0.9 738 23
Transparency, m 7.3 t7.4 37

Site 5
Surface temperature, "C 287 7.8 4l
Bottom temperature,'C 29.4 5.5 39
Surface salinity, %o 6.7 879 39
Bottom salinity, %o 17.2 33.7 36
Surface dissolved oxygen, mg I - 1
)) 26.5 )7
Bottom dissolved oxygen, mg I 1
t.4 61.8 23
Transparency, m 7.3 15.5 38

Mean ztalues through sites


Surface temperature,'C 29.2 1.4 5
Bottom temperature, 'C 29.6 1.3 5
Surface salinity, %o 7.9 l3'9 5
Bottom salinity, %o 18.4 4.6 5
Surface dissolved oxygen, mg 1- 1 3.7 4.6 5
Bottom dissolved oxygen, mg I - 1 2.3 42.1 5
Transparency,m 1.2 62 5
N octurnal nekton assemblage aariation 519

TABLE 3. Comparison between years of hydrological observations in Labu estuary


using a paired l-test

Year One Year Two

Standard Standard Degrees of


Physical data Mean ertor Mean error freedom

Surface temperature,'C 29.2 0.40 29.0 0.67 t4 N.S. 0.50


Bottom temPerature, oC 29.6 0.23 29.9 0.51 t3 N.S. -0.74
Surface salinity, %o 6.2 0.98 8.8 2.08 t3 <0.09 - 1.87
Bottom salinity, %o 15.8 0'84 20.9 r.33 t2 <0.01 -3.49
Dissolved O, surface, mg l-t 4.2 0.59 3.6 0.30 5 N.S. O.75
Dissolved O, bottom, mg 1- 1
24 0.32 2'0 0.47 5 N.S. 0.56
Transparency, m 1.2 0.05 t-3 0.04 t3 N.S. -O.49

E
E

c
'o
t

ro t5 ?o ?5 30 35 40 45 50
JF M A M J J A SO N D

Figure 6. Weekly rainfall for 1973-81with 95%, confidence limits.

about 5.51o through time. Salinity was similar with over six times the variability over
time as space. Bottom dissolved oxygen was the only parameter with nearly equal varia-
bility inspace and time. Both surface dissolved oxygen and transparency had a greater
temporal than spatial variability.
Comparisons between years using a paired t-test demonstrated significant differences
for surface and bottom salinity values (Table 3). The mean surface salinity during year
one was 6.2%o and rose to 8'8%o during year two. Similarly, for bottom salinity the second
year mean was higher (21'0%,) than for year one (15'8%0).

Rainfall
The mean weekly rainfall for the 10 years 1973-1982 is plotted in Figure 6 (I:74'5 mm;
C.V. : 20'2o/o; N : 519). The mean weekly rainfall for 1981 and 1982 was 7l'6 mm and
58.9 mm, respectively. The annual rainfall varied between 260 and 520 cm during the
lO-year period. The recorded rainfall from June 1980 to May 1981 was significantly
greater than from June 1981 to May 1982 (N : 12, t:3'21, P<0'01). Neighbouring
520

Tesrr 4. Analysis of variance of mean weekly rainfall

Source of of
Degrees Mean sum
variance freedom ofsquares

By year
Year 8 12855.585 1.2r4
Residual 458 4000.1 I I
Total 466 4152 136

By week
rVeek 5l 4927.297 r.773
Residual 335 4t99.870
Total 386 4279.189

localities such as the Siassi Islands (120 km north-east of Lae) and stations up the
Markham Valley reported drought conditions during these latter months.
There was a significant difference in mean weekly rainfall between years (P<0.02),
but there was no significant difference in mean weekly rainfall (P>0.05) between weeks
(Table 4). Rainfall was variable and could be considerable during any week of the year!
(Figure 6).

Biological ob serzt ations

D es uiptizt e statistics
In 4l trawling sets from June 1980 to May 1982 (24 months) 3L 458 individuals,
weighing over L77 kg, from 38 fish species, nine crustacean species and a scyphozoan
were caught. Over 95o/o of the individuals belonged to five species (Table 5) while 25
bony fish species were represented by more than l0 individuals. Most of the fish caught
were juveniles. These were typically brackish water forms capable of tolerating a wide
range of salinity by secreting a thick mucous layer over the skin (Equula), by possessing
large scales (Gerres, Lactarius) or by having other modifications to minimize osmotic
changes (Thia-Eng, L973). Laryer fish were relatively few.
No truly freshwater fish were caught. Scatophagus argus, Toxotes jaculator and
Anodontostoma chacunda, which could tolerate fresh and estuarine conditions, were col-
lected occasionally. The common marine fish that entered the estuary were the anchovy
(Stolephorus), the snapper (Lutjanus), the grunter (Pomadasys) and the carangid
(Caranx). Only large individuals of some of the less common species were caught, such
as Arothron reticularis.
The length/weight equations for 12 fish species were very highly significant (Table 6)
and were used to estimate catch weight. Eleven species of fish and prawns, each with a
weight greater than lo/o of the total, account for about 97t/o of the total mass caught
(Table 7).
The overall mean number of individuals caught per night (1.25h total trawling time)
was 796 individuals weighing 4'3kg with a mean of 14.0 species and a catch rate of
1
3'4 kg h- trawling.
From interviews with the natives it was found that 31 of the 38 fish species and five
crustacean species caught in the beam trawl were eaten. The Appendix contains the
scientific names of the catch and an indication of those locallv eaten.
Nocturnal nekton assemblage aariation 521

TABLE 5. Rank order and per cent abundance for species trawled in the Labu estuary.
Authors of species are given in Appendix along with an indication of which species
were eaten

Rank Number of %of Cumulative


order Species individuals total

1 Equulaequula 22106 70.3 70.3


2 Metapenaeus demani 3259 10.3 80'6
3 Secutor ruconius 2860 9.1 89.7
4 Gazza achlamys I 199 3.8 93.5
5 Ambassis interruptus 584 1.9 95.4
6 Polydactylusmicrostomus 190 0.6 96.0
7 Pseudosciaena weberi 186 06 96.6
8 Apogon amboinensis 169 0.5 97.1
9 Lactarius lactarius t2t o.4 97.5
l0 Caranx sexfasciatus t02 0.3 97.8
II Gerres filamentosus l0l 0.3 98.1
12 Apogon hyalosoma 83 0-3 98.4
13 Palaemonidae 75 0.2 98.5
L4 Lutjanusjohni 64 0.2 98'7
l5 Pomadasys argyreus 61 0-2 98.9
16 Upeneus sp. 37 0.1 99.0
17 Anodontostomachacunda 37 0.1 99.1
l8 Stolephorusbataoiensis 3t 0.1 99.2
l9 Upeneus oittatus 3t 0.1 99.3
20 Setipinna papuensis 28 0.1 99'4
2l Lutjanus ehrenbergi )') <0.1 >99'4
20 L. maxweberi 2l <0.1
23 Eleotris c.f. macrolepis l8 <0'1
24 Archamiaburoensis l7 <0.1
25 Arothronreticularis l5 <0.1
26 Lutjanusargentimaculatus ll <0.1
for each of the remaining species, fewer than ten
individuals were caught

Total 31458

H armoni c p erio di city analy si s


As with the abiotic data, multiple regression incorporating the Fourier transformation
was performed on each of the 1l most abundant and massive species to ascertain their
conformity to an annual cycle and to determine the period of peak abundance or mass.
Seven species had annual cycles of abundance (Table 8). Five of these species were most
common in the five months ]une to October. Pseudosciaena ueberi had no cycle, but sig-
nificantly increased in abundance through time. The most abundantfish, Equula equula)
is most abundant in July (Figure 7). Secutor ruconius, rank order 2,was most abundant in
February/March while Metapenaeus demani peaked in November/December.
Five species had changes in the weight of the catch that corresponded to annual
changes (Table 8). Three of these species also had maximum peaks from June to August.
The most massive population, Equula equula, has its greatest catch of individuals and
mass in July/August. Similarly, the largest catches of prawns are in November/
December. Catches of Arothron reticularis, weight rank order 2,were too infrequent and
irregular to fit the model.
522 N. J. Quinn U B. L. Kojis

Tenrr 6. Length/weight relationships for Labu estuarine fish

Species P N

Ambassis interruptus l'50 x 10-s 3.19 o.992 <0.001 t7


Ano dont o s toma chacunda 1.03 x l0-s 3.22 0.995 < 0.001 25
Apogon hyalosoma 6.82 x l0 6
3.38 o.954 < 0.001 30
Equula equula 5'75 x 10 s 2-89 0.977 < 0.001 248
Gazza achlamys 6'13 x 10 s 2.79 0.965 < 0'001 112
Gerres filamentosus 3.48 x l0-s 2.95 0.994 <0.001 12
Lactarius lactarius 3 01 x lO-s 295 0.992 <0.001 42
P o ly d a c ty lus mi cr o s t omus 3.48 x 10 s 2'92 0.974 <0.001 10
Pomadasys argyreus x l0-s
1.74 313 a 996 < 0.001 22
Pseudosciaena weberi 1 96 x l0-s 301 0.970 < 0.001 34
Secutor ruconius 4.53 x 70-s 2.30 0'780 < 0.001 64
S tolephorus bataztiensis 5.57 x l}-s ).61 0.986 < 0.001 18

Tesrt 7. summary statistics for estuary nekton populations with the highest weights

Rank Number Mean


order of catch o/
number sampling per Catch of
by times sampled weight total
Species weight trawled date (g) (g) weight

Equula equula I 41 469.7 96280 59 1


Arothron reticularis 2 1l 115.5 2320 14.5
Metapenaeus demani 3 41 35.6 73t0 45
Gazza achlamys 4 32 41.5 6850 4.2
Secutor ruconius 5 37 28.5 5830 3.6
Polydacty lus microstomus 6 33 27.4 4385 2.7
Ambassis interuptus 7 38 r8.8 3865 2.4
Lurjanus johni 8 22 77.4 3570
Lutjanus maxweberi I 5 3l .l 2690 1.7
Lactarius Jactatius i0 29 90 1855 1.1
Pomadasys argyreus t1 23 8.1 I 660 1.0
Apogon hyalosoma T2 7.2 r495 0.9
Himantura granulata 13 I 5.8 r200 o.7
Apogon amboinensis I4 )6 5.2 1075 0.7
Gerres filamentosus 75 24 5.2 1070 o.7

The total weight of the catch, total number of fish and number of species had an
annual cycle with the grearest carch occurring in June/July (Table g).
In parametric statistical testing, the most basic assumptions are: (l) that data are
normally distributed, and (2) thar variances are homogeneous (Sokal & Rohlf, 1969). For
significance testing in cyclical regressions it is necessary that the raw data be normally
distributed since significance testing involves paramerric distributions (Bliss, 1970).
Recently, Gilmour & Stephenson (1983) specifically noted that: (l) residuals, or rhe
unexplained proportion of the variation in data after cyclical regression, should be
normally distributed, and (2) that the magnitude of residuals should be independent of
predicted (explained) components. The reason for transforming the data was that for the
N o cturnal nekton as s embl ag e a ari ati on 523

Tenrr 8. Regression coefficients for annual cycles in Labu estuary nekton

Peak
Species month A B M ll2Ai P

Catch weight
Equula equula Jul/Aug 275'0 2u4 y 269.7 < 0.001 0.68
Arothron reticularis N.S. 0.20
Metapenaeus demani Nov/Dec 17'1 t5.7 <0.01 o.42

ry:' -r,
Gazza achlamys N,S. 0.13
Secutor ruconius
P o ly da c ty lu s mi cr
Ambassis interruptus
ost omu s
f"t7tr{",
r 30.4 < 0.01
21.8 <0.05
N.S.
0.40
0.38
0.24
Lutjanus johni N.S. 0.10
N.S.
L, maxwebei
Lactarius lactarius
Pomadasys argyreus Y- j' r 86 N.S.
N.S.
0.16
0.40
O.23
Number of individuals
Equula equula Jul/Aug 56.6 43.4 r.3 74.0 <0.001 0.75
Metapenaeus demani Nov/Dec -7 1 15.7 < 0.01 O'42
Secutor ruconius Feb/Mar 11.9 14.8 < 0.05 0.32
Gazza achlamys N.S. o.t4
Ambassis interruptus N.S. 0.19
Polydacty lus microstomus Iun 0'8 0.1 < 0'001 0.55
Pseudosciaena weberi -0.8 <0.001 0.64
Apogon amboinensis Jr'rr/errg 0.5 -0.1 t-7 < 0.001 0.62
Lactarius lactarius JuVAug 0'4 04 0.6 <0.001 0 61
Carnax sexJasciatus N.S. 0.18
Gerres fi,lamentosus Aug/Sep 0.3 0.5 <0.05 0.31
Summary parameters
ll/ Jul 123 5 t33 0 427.8 < 0.01 O.54
s Jul 15 t.7 12.8 <0 001 0.58
N Iun 25'8 80 I <0.01 0.50

A, B are the cyclic coefficients in the Fourier equation, M is the linear coefficient in the
Fourier equation, and ll2Aoisthe constant value in the equation.
Peak month is the period of predicted maximum values.
There were 25 64 sampling periods in a year.
Where 1,8, M and constant are variables in the equation:
y : I 12 Ao + A cos(2t 125'64) + B sin(2t125'64) + M (t).
Trigonometric functions are in radians.

o
o
j
!
.=
!
.= 200
o
too _erayv.\ s
o o --a
E
f
z
JJ A JF M DJ F
r980 I 981 r982

Figure 7. Time series of the mean number of individuals of Equula equula for five sites
intheLabuestuary.-,meannumberofindividualspersite;--------,predicted
nunber of individuals.
524 N. J. Quinn €c B. L. Kojis

Tesrp 9. Regression coefficient for catch weight of ,Equula equula and transformed data

Month of
maximum
Transformation occurrence M iAo

Square root Jul 6.479 5'229 0.189 t5.43 0 738 <0.001


4th root Jul/Aug 0.733 0.604 0.022 3.87 0.762 < 0.001
log (N+ 1) Jul 0.295 0.245 0.009 3.32 0-776 < 0.001
None Ju1/Aug 275-0 275-0 7.7 269.7 0.683 <0.001

most effective use of this technique the data should be normally distributed. A further
desirable feature of the transformation used was that the explained variation in data
should be as great as possible, i.e. r should approach 1.0 (Gilmour & Stephenson, 1983).
The fourth root and log (n -t 1) transformations reduced the skewness and kurtosis to
that approximating a normal distribution and, hence, were used here. It is appreciated
that the choice of transformations is somewhat arbitrary and that other transformations
might have been used, such as those described by Box and cox (1964). However, rhe
chosen transformations were considered reasonable in meeting the assumptions of the
mathematical models.
Recently, Gilmour & Stephenson (1983) investigated two approaches for selecting
Box-Cox transformations to apply to data used in cyclical regressions, but neglected to
determine experimentally the effect the transformation had on the period of greatest
occurrence (commonly termed phase angle or I-u"). Since the primary purpose in the
use of cyclical regressions in ecological studies is to elucidate periods of greatest occur-
rence) extended investigations of various transformations to increase r without know-
ingly increasing the certainty of the phase angle are likely to be of marginal use and were
not pursued.
Equula equula catch weights were used to assess the effect of transformed data on the
determination of annual cycles. The use of any of the transformations, square root,
fourth root and log(n -t 1), resulted in increasing the correlation coefficient without
appreciable changes in the month of maximum occurrence (Table 9). Transformed data
enhanced the fit to the cycle (increasing r) without altering the phase angle. By increasing
the correlation coefficients, several smaller populations exhibited annual cycles. As these
species are uncommon, the use of the transformations does little to increase our
knowledge of the estuary fauna.

Multiple regressions with catch data


Stepwise, forward inclusion regressions between N, S and IZ resulted in several signifi-
cant relationships. S was correlated with bottom salinity, N with surface temperature
and W with surface salinity (Table l0).
Four species abundances had a significant relationship with unlagged hydrological
values (Table ll), with a mean r of 0'61. Equula equula, the dominant species, and
Lactarius lactarius were highly significantly correlated with surface water temperature.
Apogon amboinensis correlated well with bottom water temperature and Pseudosciaena
weberi with bottom salinity. All of the regressions included only one hydrological
variable as a predictor.
Nocturnal nekton assemblage oariation

TABLE 10. Catch summary values regressed against physical data

Surface Bottom Surface Bottom


temperature temperature salinity salinity Linear
coefficient coefficient coefficient coefficient coefficient Constant r

Number of species -0.19 r6.3 0.35 <0.05


Number of individuals -81 89 3137.5 0.45 <0.01
Total weight -169.5 5580.5 0.44 <0.01

The weight of each species was related to hydrological values (Table l2). Equula
equula weight, rank order l, was negatively correlated with surface salinity while
Lutjanus johni, rank order 8, was positively correlated. Surface temperature was a weak
indicator of Metapenaeus demani (rank order 3) and Lactarius lactarius (rank order l0)
weight, with only 11076 and 20o/o of the variance explained respectively. The mean r
value was 0'47, explaining 22"1; of the variance, about 15 f,u less than that explained with
regressions on species abundances. Both the catch weight and number of individuals of
L. lactarius were correlated with the same hydrological observation. The only other
species with correlations for both weight and abundance was E. equula, but it was
correlated with different variables.
There are three types of complications to be expected in effecting regressions between
a given time set of hydrological data and a given time set of catch data:

(1) The relationship may not be linear, and if not, exponential, logarithmic or power
curve fitting may be required.
(2) The data may not be normally distributed and may require transformations which
may in turn convert a non-linear to a linear relationship.
(3) The catches may be influenced by several climatic conditions at an earlier time, and
this is referred to as the lag effect.

Hydrological observations lagged at fortnightly intervals were regressed against the


number of individuals of commonly caught species (Table I l). Bottom and surface water
temperatures were the most common significantly correlated hydrological variables - l0
and nine times respectively. Then followed surface (seven times) and bottom (twice)
salinity. It is curious to note that the variable with the greatest temporal homogeneity
(C.V., 4'9o/o), bottom temperature, was the most common significantly correlated
hydrological variable. In other words, the 'best' hydrological parameter from which
abundances can be predicted is the parameter that varies least throughout time and
which probably reflects the small range in abundance. The model lag was three
fortnights with eight significant correlations. A single fortnight lag had six correlations,
two fortnights had three, four and five fortnights, five correlations. There is no apparent
consistency in hydrological variables ofprevious fortnights affecting abundances.
The transformed catch weights of Equula equula and Metapenaeus demani were
regressed on lagged hydrological variables (Table 13). Correlations with bottom salinity
(lagged five fortnights) and surface salinity (lagged three) and bottom temperature
recurred in each regression for E, equula. Two regressions were significant with trans-
formed catch weights of M. demani and both included surface temperature. As suspected
TenrE I I . Regression coefficients between numbers of individuals and hydrological variables u
N)

Physical Surface Bottom Surface Bottom Linear


Species data lag' temperature temperature salinity salinity coefficient Constant P

Eguula equula 0 - 17.02 601 3 0.55 < 0.01


Secutor ruconius
1
-rt:o 465.3 o.35 <0.05
3
3 8.58
-237.7 o'45 <0.02
:.?
4 r0.54
-295.9 0.50 <0.0t
ro
Metapenaeus demani
5 10.26
-287-5 0.51 <0.01
2
-2.61 96.6 0.35 <0.05
3
-412 138.5 o.43 < 0.05 a5
4
-4.80 159.9 0.55 < 0.01
Gazza achlamys
5
) -3.83 t28.5 0.62 <0.01
0.91
-l.t 0.48 <0.01 l-
3 0.85
-,3.2
0.1 0.45 < 0.05 x
,o
4 r.20 0.65 < 0.01
Ambassis interruptus
5 0.38 1.8 o.52 <0.01
3
- 0.98 16.7 0.40 <0.05
4
-t.42 0.19 43.r 0.71 < 0.001
5 _ ru 40.6 0.58 < 0.001
P o ly d a c tjt lu s mi cr o s tomu s
N.S.
Apogon ambionensis 0
-o'28 -0'04 10.I 0 56 <0.001
I 027 86 0.52 <0.01
2
-0 39 12.l 0.64 <0.001
3
-5', 10.5 0.41 <0.05
4
-0.31 9.9 o.54 <0.01
Gerres filamentosus
5
-o.26 8.4 0'45 < 0.01
1
-o.21 7.0 0.41 < 0.05
Lactarius Iactarius
3
- 0.38
*0 yu 0.9 0.59 <0.05
0 23 7.2 0.63 < 0.001
I -0 05 l.l 0.41 < 0.05
Caranx sexfasciatus
N.S.
Pseudosciaena ueberi 0
I
- 0.10 -0.06 1.0 0.65 < 0.001
0. l5 -0.24 4.O 0.60 < 0.001
3 0.29 -0.18 -4.5 0.68 < 0.001

'Each lag represents one fortnight.


TABLB 12. Regression coefficients between catch weight and hydrological variables

Physical Surface Bottom Surface Bottom Linear


Species data lag temperature temperature salinity salinity coefrcient Constant P 2
:r
I
Equula equula 0 -3t.4 716.6 0.65 <0.001 s
Arothron recticalnris 2 -57.4 t799'2 o.?7 <0.05 :
3 -rs.n t275.8 0.41 <0.05 hr
5 - 91.9 17.9 2521.8 o.54 <0.01 N
Metapenaeus demani 0 3.6
-7t.5 o-34 <0.05 6
4 -t.t 299'l 0.41 <0.05
5 - 10.3 344'5 0.05 <0.01 sI
Secutor ruconius ,
3
4
16.8
20-9
l' 6'6
466.5
o'37 /<0.05
0.46 f .o.os
0.54 / <O.01
6
sI
- 585.5
5 18.7
-4-2 -520.r 0.50 <0.01
Gazza achlamys 4 5.5 0'67 <0.001 o
P o ly d acty lus mi cr o s t or/tus 0 0.6 36.9 0.34 <0.05 is
Lutjanusjohni 0 1.6 - l.lJ 35.1 0.57 <0.001
I 2.2 t.o
-22.6 0.4r <0.05
2 2.L -23.7 0.46 <0.01
3 --9 5 2-4 270.2 0.65 <0.001
Pomadasys argyreus 3
-2.2 70.8 0.51 <0.01
5
-1.4 45.6 0.17 <0.05
Ambassis intenuptus 4 -7.4 238.3 0.64 <0.001
5
-9.5 303.r 0.59 <0.001
Lactarius lactarius 0 -2.5 82.O 0.35 <0.01
I -2.6 85.4 0.51 <0.001
Lutjanus maxuteberi : N.S.

ut
{
t\)
iJl
i\)
00

Tesrr 13. Regression coefficents between carch weights and lagged hydrological variables
3
:-?
Species LaE Transformation a' Variable Constant P o
Equula equula 0 37.16 bottom salinity 13r.83 rn
5 square root t.2t bottom salinity
- o.57 < 0.05
2 square root bottom temperature
F
-3.71 !-
3 square root -0.62 surface salinity 116.50 0.84 < 0.001
5 4th root 0.14 bottom salinity X
,o
2 4th root -0 41 bottom temperature
3 4th root -0.07 surface salinity 15.13 0.89 <0.001
5 log(n + 1) 0.06 bottom salinity
3 log(n + 1)
-0.03 surface salinity
4 log(rz+ 1) bottom temperature 6'66 091
-0.16 <0.0001
Metapenaeus demani 0 0.36 N.S.
I 4th root 0. 15 surface temperature
3 4th root 0.03 surface salinity 0.70 <0.05
I log(zf -183
1) 0.09 surface temperature
- l.l8 0.54 < 0.05

"Q is the coefficient for the hydrological variables


Nocturnal nekton assemblage z;ariation 529

the relationship is clearly complex. It is postulated that hydrological parameters along


with interactions of other species, such as the presence of predators, need to be con-
sidered in the models. This analysis was not attempted owing to small catches of known
predators of the dominants.

V ariation betw een y ear s


The data from similar lunar phases for two years (June 1980 to May 1981 and June 1981
to May 1982) were compared using a \Tilcoxon matched pairs non-parametric test to
determine if there was a significant difference between years in the total catch
abundance, mass) or number of species caught (Table l4). No significant difference was
present for N and IZ although both means were higher during the second year. The

TABLE 14. Comparisons between years of catch summary statistics from Labu estuary
using a paired r-test

Year one Year two

Degrees
Standard Standard of
Mean error Mean error freedom P

Mean total number of individuals


per site per time 77.5 7.4 92.0 t0.2 15 N.S. t-23
Mean total weight (g) per site
per time 442.1 58.6 479.9 60.9 15 N.S. 0.s4
Nur,'rber of fish species per time t2.4 0.5 r4.3 o.7 15 <O.O5 2 27

TABLE 15. Comparisons between years of number of individuals of abundant species


from the Labu estuary using a paired r-test

Year one Year two

Degrees
Standard Standard of
Mean error Mean ertor freedom P

Equula equula 936 9.8 t36.2 19.1 15 <0.01 2 93


secutor ruconius 18.7 9.6 72.2 4.4 15 N.S. 0.58
Metapenaeus demani 16.8 3.9 t2'8 2.7 t5 N.S. 0.01
Gazza achlamys 5.7 24 48 1.0 8 N.S. 0.32
Ambassis interruPtus 3.8 1.0 2.8 0.6 l5 N.S. 0.31
P oly dactg lus mi cr o st omus 0.8 0.2 l.l o.3 15 N.S. 0.32
Apogon amboinensis 15 0.4 0.5 0.2 15 < 0.05 2.27
Gerres f.lamentosus 0.6 0.2 05 0.2 15 N.S. 0.40
Lactarius Iactarius 0.7 0.2 0.6 02 15 N.S. 1.0r
Caranx sexfasciatus 07 0.2 0.3 0.1 15 N.S. 1.38
Pseudosciaena weberi 0.0 0.0 1.9 0.5 15 < 0.001 4.38
Pomadasys argyreus 0'4 0.1 0.3 0.1 t5 N.S. 0 77
Lutjanus johni 0.5 0.1 02 0.1 l5 <0.01 1.83
530 N. J. Quinn €s B. L. Kojis

Tesrn 16. Comparisons between years of catch weight of selected species from the
Labu estuary using a paired l-test

Year one Year two

Degrees
Standard Standard of
Mean error etror freedom

Equula equula 404.3 79'0 624.1 95.6 t5 <0.05 2.2t


Arothron recticularis t68.4 73.8 52.5 28.6 l5 N.S. t-45
Metapenaeus demani 29.4 6'7 44.3 8.7 15 N.S. r.53
Secutor ruconius 32.4 18.0 27.3 8.4 l5 N.S. 0'25
Gazza achlamys 278 8.9 46'8 13.9 8 N.S. 0.95
P o ly da c ty lu s mi cr o s t omu s 23'6 7.7 20.0 4.6 15 N.S. 0.43
Lurianus.iohni 28.9 8.4 3.5 1.8 t5 <0'01 2.82
Pomadasgs argyreus 15'9 10.0 3.9 2.1 l5 N.S. 1.15
Ambassis interruptus 226 b.9 20.5 5.2 15 N.S. 0'32
Lactarius lactarius 79 2.4 11.1 33 l5 N.S. 0.86
Apogon amboinensis 29.7 2.1 1.6 l5 <0.01 3.06
Gerres filamentosus 7.1 2.4 5.3 t.7 t5 N.S. 0's3

number of species per time) S, was significantly different with 12.4 species during the
first year and 14.3 during the second.
A comparison of the abundances of the 13 most abundant species between the two
years found three significant differences. The dominant, Equula equula, was significantly
more abundant during the second year than during the first (Table l5). Pseudosciaena
weberi was caught only during the second year and Apogon amboinensis was less abundant
during the second year.
Catch weight for most species was similar between years; only three species had sig-
nificant differences (Table 16). Catch weight for Equula equula was greater during the
second year, while the catch weight for Apogon amboinensis was greater during the first
year. These results were consistent with those for abundances. The weight of Lutjanus
johni was greater in the first year, suggesting that larger individuals were caught that year
as there was no significant difference in numbers of individuals between years.

Discussion
The hydrological study of the Labu estuary shows the existence of minor physico-
chemical gradients associated with the penetration of tides and runoff. As the estuary is
relatively short and shallow, a saline environment exists in all sections throughout the
year. The tidal currents are only strong in the lower reaches resulting in little turbulence
and vertical mixing in other regions. The upper reach is a zone of reduced dissolved
oxygen and salinity and increased temperature. Spatial variation in hydrological param-
eters is less than temporal variation. It is likely that the estuary's homogeneity and
limited extent reduce the number of niches available for utilization by fish.
Variations in temperature and salinity in the Labu estuary follow annual cycles.
Although there was no distinct dry season at the rain gauge in Lae, it is hypothesized that
the rainfall in the higher altitudes of the Labu watershed was more seasonal which
N o cturnal nekton as s emb lag e a ari ati on 53r

resulted in the annual variation of hydrological parameters in the Labu estuary. $7hile
cycles in salinity have been recognized in tropical estuaries, it has been commonly
thought that the temperature remains nearly constant all year (Rodriguez, 1975).
Although tropical estuarine temperature variation is less than that observed in subtropi-
cal waters (Quinn, 1980), there is still a regular annual cycle to which fish could respond.
There is no apparent cycle in dissolved oxygen or turbidity.
Temperature/salinity cycles are probably related as lower temperatures occur during
periods of lowest salinity in the estuary. The months of July/August in Lae are charac-
terized by cloudy days with prolonged periods of rain. This agrees with the observation
by Egborge (1972) in a study of the hydrology of the Oshun River, Nigeria, that lower
temperatures in flood periods occur because of increased cloud cover.
The Labu estuary is stratified with significant differences between surface and sub-
surface hydrological values. The differences between near substrate and mid-river sur-
face temperatures are similar to results from other studies; for example, 3'0'C in a
Ghanaian river (Thomas, 1966) and 1.0'C in a Malayan stream (Bishop, 1973). This is
expected in sheltered coastal waters where there is little wind-generated water move-
ment to mix the less dense freshwater on the surface with higher salinity water below.
Additionally, the small tidal range (maximum range l'1 m) does not encourage mixing.
Despite its limited extent, the estuary was occupied by at least 38 species of bottom
fish from 26 different families. Thirty one of these species were eaten by local villagers.
Some small demersal fish species were probably missed owing to the mesh size and other
sampling considerations. Schooling fish (e.g. mullet) and surface feeders (e.g. gar) were
present in the estuary, but not caught.
The fish recorded are generally typical of those recorded from similar locations in the
Indo-west Pacific tropics, except for the absence of members of several families such as
Platycephalidae, Cynoglossidae, Sillaginidae and Soleidae. The assemblage is appar-
ently much less diverse that that of Trinity Inlet, Queensland (16'55'S), with 54 species
(Blaber, 1980), and Ponggal estuary, Singapore (1'N), with 80 species (Thia-Eng, 1973).
The use of fine mesh seines in these studies undoubtedly increased the number of species
caught. Only six species (11%) were common with the Trinity Inlet and five species
(6?i,) with Ponggol estuary.
The assemblage was dominated by Equula equula, representing 70'3oio of all individ-
uals caught and 59.19i, of the mass. It was the most ubiquitous teleost occurring in all 41
trawl dates. No published studies of E. equula are available, but in a detailed study a
related species, Leigonathus breairostris, was found to spawn throughout the year in the
Gulf of Mannar (8"N, 79'E) with individual fish possibly spawning more than once a
year (James & Badrudeen, 1975). The presence of juvenile E. equula (S.L.<55mm)
throughout the year in Labu estuary suggests that it could have a similar reproductive
biology.
The biology of Pomadasys argyreus is reasonably well known. In eastern India' it
grows to a maximum of 45 cm (S.L.) weighing 1'5 kg, becomes sexually mature at l3 cm
and spawns in February (Konchina, 1977). This suggests that spawning would occur
around August in the southern hemisphere. Estuary catches of P. argyrezs declined
throughout the study and had no annual cycle. Individuals caught were mostly juveniles,
3-l I cm, whose presence was inversely related to water temperature of the previous two
monrhs suggesting that in a more favourable habitat the greatest abundance would occur
in the cooler season of June to August. The estuary is probably used only as a nursery
with breeding probably occurring elsewhere.
532 N . J . Quinn {s B. L. Kojis

The main feature of the life cycles of some coastal fish is the division into a juvenile
phase which is largely estuarine and an adult phase which is primarily marine. It is com-
monly thought that most species breed on the continental shelf, because only a few small
specialized forms have adapted their entire life cycles to the variable conditions of
temperature, salinity and turbidity characteristic of estuaries (Vallace & van der Elst,
1975). Generally, there is a seasonal movement of adult populations into the inshore
spawning grounds. Fish in pre-spawning and partially spawned condition tend to move
in and out of the lower reaches of estuaries with the tides, while post-spawners generally
leave the system.
Lowe-McConnell (1977) outlined four zones from inshore muddy water to clear, deep
water with characteristic fish populations and suggested that broad shelf communities
are highly diverse, while shelf communities near large rivers with seasonal outflows
generally were less diverse and contained dominants. As the continental shelf of the
Huon Gulf was extremely narrow (<2km) and limited in extent (<20km long), it is
tikely that this assemblage has small populations of many species if not lower diversity.
To test whether the fish assemblage of the continental shelf of the Huon Gulf is less
diverse than the 350 + species recorded from the much larger shelf of the Gulf of Papua
(9'00'S, 143"27'E to 9'02'S, 146"38'E) (Kailola & Wilson, 1978) an extensive trawling
study of the Huon Gulf is suggested.
If the biology of tropical estuarine fish is governed by the seasonality of flooding
(Lowe-McConnell, 1977), then the irregularity of the local rainfall patterns is likely to
result in local anomalies in the reproductive biology. In India with its regular monsoons,
Sreekumari (1977) noted that the beginning of rains brings changes in the environment
that act as a definite stimulus to spawning even in species llke Stolephorus zollingeri,
which are continuous breeders. In New Ireland, Papua New Guinea, Dalzell (1980) also
observed that spawning in S. deztisi and S. heterolobu.s occurs at a low level all year round,
but is greatest at the time of change in monsoon season. Catches of S. bataaiensfu in the
Labu estuary were too small for any generalized observations. Flowever, it is likely that
using a small mesh net would result in a greater catch. Spawning would most likely occur
around April to May and October/November to December, at the time of the slight
change in seassns.
Although generalized temperature and salinity preferences have been shown for
various estuarine species (Copeland & Bechtel, I974), as a whole these organisms show a
wide tolerance for short-term changes in these parameters. This could help to explain
the general lack of importance of temperature and salinity as critical variables in the
multiple regression analysis. The annual salinity and temperature range was small owing
to, respectively, heavy rainfall throughout the year and the equatorial position of the
estuary. The multiple regression technique was limited in its application as the abiotic
parameter variation was perhaps too subtle to accommodate sampling and annual vari-
arions. Other factors such as biological interactions could complicate such an approach
to determination of predictive factors (Oviatt & Nixon, L973).lt is possible that trophic
relationships and reproductive cycles are of critical importance in the spatial and
temporal distribution of estuarine populations. As in other low latitude estuaries, the
Labu estuary is dominated by juvenile stages of a small number of species.
Fluctuations in the abundance of higher latitude marine species throughout a year is
well known. Considerable within-estuarine species variability in annual abundances has
been noted (Livingston et al., 1976) and occasionally related to climatic cycles over
several decades (Dow, 1977). ln the tropics, less is known about within- and
Nocturnal nekton assemblage zsariation 533

between-year variarion in estuarine biota. In this study hydrological variation was


greater between years than within years and a few species experienced significant
changes between years.
The hydrological and biological consequences of exceptionally wet or dry years are
largely unknown. It is obvious that periods of high rainfall will have a tremendous effect
on the hydrodynamics and sedimentation of the estuary. The biological effects of low
salinity will also be considerable. On the other hand, drought conditions are also likely to
have an ellect as evaporation would increase the salinity. A severely reduced runoff
might also allow sediment to block the mouth as occurs in South Africa (Blaber, 1973)
and effect the assemblage composition.
During the second year of this study, drought conditions were experienced in the
Markham River valley and higher salinities were experienced in the estuary. The signifi-
cant population variation between years could be related to the hydrological variation.
llowever, it is possible that some species have a 'supra'-annual cycle and these cycles
could have accounted for the community changes. Stephenson (1980c) noted many
macrobenthic species had cycles of three to six years. The testing of this hypothesis using
two-year data should only be considered as preliminary.
Pseudosciaena weberi (S.L. 3-10 cm) was caught only during the second year of
sampling. This catch variation differs with pseudosciaenid populations in India which
are prolonged breeders and resident in the estuary throughout the year (Nair, 1977).
llowever, it has been suggested that within the tropics fish populations vary greatly from
year to year owing to changes in abiotic and biotic factors (Lowe-McConnell, 1979). The
salinity in the estuary was higher during the second year and the population
demonstrated a correlation with salinity values of the previous several fortnights. The
difference between years may be related to other unmeasured abiotic factors such as the
injection of nutrients into the system) or by biotic pressures such as the alteration of a
preferred habitat for predators and subsequent decrease in their populations. Such
influences may impose a between-year variation in the relatively aseasonal estuarine
environment that restricts efforts to determine stable annual patterns in such
environments.
It is evident from the foregoing that the distribution of juvenile fish in Labu estuary
cannot be satisfactorily explained solely with regard to physical factors. It is likely that
the community structure depends on small environmental changes interacting with pre-
dation and other biotic pressures. In this study, large piscivorous fish such as sharks,
carangids and sciaenids were not caught in any of the sites. The absence of such pred-
ators from the shallow sampling sites may increase the attraction of these areas as sanctu-
aries for juvenile fish, as was suggested for a north Queensland estuary (Blaber' 1980).
Predation by birds on juvenile fish in African estuaries has been shown to be high
(Blaber, 1973). However, few birds were observed in the Labu estuary, possibly because
their plumage is commonly sought by local villagers for decoration. Intense human
predation probably results in small bird populations and consequently bird predation of
fish was probably not important.
This work uses the same sampling device, a 3 m beam trawl, as previous work (Quinn,
1980; Quinn & Kojis, 1981) carried out in a similar estuary at approximately the same
longitude (150'E), but with a 20' latitudinal difference. It is commonly thought that
species diversity increases as latitude decreases (Pianka, 1966). However, along the East
Australian/Papua New Guinea coast, this does not appear to be true. Vhile 45 species
of fish were caught in the subtropical Serpentine Creek, only 39 were caught in this
534 N. J. Quinn Cx B. L. Kojis

study. This was in spite of a longer sampling programme for the latter study. The
subtropical estuary was characterized by three species, Spheroides pleurostictus, Gerres
or.)atus, and Sillago maculata, each representing about 20, 17 and l7o/o of the total
abundances respectively (Quinn, 1980). The tropical estuary was dominated by a single
species, Equula equula, representing 70'3% of the catch. The sub-dominant fish Secutor
ruconius occurred in 9'loft of the catch with Metapenaeus demani representing l03%.
Thus, the low latitude, tropical Labu estuary is less diverse than the subtropical
Serpentine Creek.
MacArthur (1965) suggested that the number of species within a habitat can be expec-
ted to increase with productivity, with structural complexity and lack of seasonality of
resources. As there are no studies of the productivity of the Labu estuary, no com-
parisons can be made. Less seasonal variation of resources in the Labu estuary has not
resulted in increased diversity. It is possible that the Serpentine Creek estuary is a more
complex habitat than the Labu estuary and thus alters the latitudinal effects.
It is further postulated that the diversity of the estuarine assemblage is also related to
diversity of the shallow-water fauna of the Huon Gulf. Moreton Bay is a large shallow
bay with a variety of habitats and physico-chemical environments which support a large
nekton community which has been described by Bradbury (1980) and Stephenson and
Burgess (1980). As there is no study of the demersal fish of the Huon Gulf, it is imposs-
ible to compare the open water stocks available to both studies and relate it to estuarine
nekton diversity. Ifowever, the area of shallow water is known to be small in the Huon
Gulf and a narrower range of habitats and physicochemical environments is likely. It is
suggested that future studies investigate the community structure of the demersal
nekton in the waters < 180 m of the Huon Gulf. Populations studies should focus on the
suspected dominant family, Leiognathidae.
Alternatively, according to present theory, high diversity can be maintained only in a
non-equilibrium or sub-climax state (Patrick,1967; Odum, 1969; Slobodkin & Sanders,
1969; Louchs, 1970). According to this 'intermediate disturbance hypothesis' (Connell,
1978), diversity declines during long stable interludes due to competitive elimination,
resulting in resource monopolized climax assemblages composed of few species. Distur-
bances such as cyclones and wide ranges of temperature and salinity interrupt and set
back this process of competitive exclusion. Organisms are killed, populations decimated
at various scales of frequency and intensity. If the intensity and frequency of disturbance
is sufficient to affect all, or most species, then the assemblage will return to pioneering
stages and diversity will be low. If, however, intensity and frequency of disturbance are
of a magnitude which affects only certain species, thus acting in a selective manner as a
'pruning device' to prevent resource monopolization, diversity will attain a maximum
relative to the extremes of no disturbance or severe disturbance.
The Labu estuary is a relatively disturbance-free habitat for euryhaline species and
diversity is low. As the estuary is not directly connected to the Markham River, annual
variation in salinity is not affected by heavy rains far up the valley and hence the annual
variation in salinity is small. Only locally intense rains are likely to greatly reduce the
salinity and serve as a disruptive influence. As less than average rain fell during most of
the sampling period, it is impossible to assess how much the bottom salinity would drop
and to what extent the freshwater intrusion would affect euryhaline populations. Further
sampling during and after a year of heavy rain is necessary to determine the extent the
community is altered after an unusually large lowering of salinities.
Nocturnal nekton assemblage uariation 535

Acknowledgernents
\We are indebted to the people of Labu Butu village for
their assistance and cooperation.
$7e would like to gratefully acknowledge the Papua New Guinea University of Tech-
nology for financial assistance for this study. Additional financial support was provided
by the Papua New Guinea Harbours Board and the South Pacific Commission.
Messers R. Adams, K. Bakoma, M. Blowers, R. Hancock, K. Kapi, M. Matmillo, M.
Sappu, H. Silver, D. Stewart, C. $flright and T. Yamelo assisted with the fieldwork. Mr
J. Johnson of the Queensland Museum kindly assisted with fish identification. Mr S.
Frusher, Department of Primary Industry, Dr A.J. Bruce, Northern Territory Museum
of Arts and Sciences, Mr P. Davie, Queensland Museum and Dr P. Rothlisberg, CSIRO
helped identify the Crustacea. Prof. W. Stephenson kindly assisted with computer
analysis performed at the lJniversity of Queensland. Both he and Prof. J.M. Thomson
constructively criticized portions of the manuscript. Dr G.R. Huntsman and an
anonymous reviewer offered useful suggestions.

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Appendix: List of organisms caught in the beam trawl

Locally
Species Family eaten

Ambassis interraprzs Bleeker 1956 Chandidae Yes


Anodonto stoma chacunda (H.-B.) 1822 Dorosomidae Yes
Antennarius hispfdzrs (Bloch & Schneider) l80l Antennariidae No
Apogon amboinens,s Bleeker 1858 Apogonidae Yes
A. hyalosoma Bleeker 1952 Apogonidae Yes
Ar chamia buroensi s (Bleeker) I 956 Apogonidae Yes
Arothron reticularis (Bloch & Schneider) 180 I Tetrodontidae No
Autaous grammepomus (Blecker, I 849) Gobiidae No
Callianassa c.f . karumba Callianassa No
Charybdis helleri Portunidae No
Caranx sexfascicrzs Quoy & Gaimard 1824 Carangidae Yes
Eleotrisfuscus (Bloch & Schneider) 1801 Eleotridae Yes
E. c.f. macroleprs (Bleeker) 1875 Eleotridae Yes
Epinephelus tauztina (Forskhl) L7 7 5 Epinephelidae Yes
Equula equula (Forskil) I 775 Leiognathidae Yes
Gazza achlamgs Jordan & Starks 1917 Leiognathidae Yes
Gerres filamentoszs Cuvier 1829 Gerridae Yes
Glossogobius circumspectus (Macleay, 1884) Gobiidae No
Harpodon translucens Saville-Kent 1889 Harpodontidae Yes
Himantura granulata (Macleay) 1883 Dasyatidae Yes
Lactarius lactarius (Bloch & Schneider) l80l Lactariidae Yes
Lutjanus argentimaculatus (Forsk6l) 1775 Lutjanidae Yes
L. ehrenbergi (Peters) 1869 Lutianidae Yes
L. johni (Bloch) 7792 Lutjanidae Yes
L. maxweberi? Popta, 1921 Lutjanidae Yes
Metapeneaus demani Penaeidae Yes
Monodactylus argenteus (L.) 1758 Monodactylidae Yes
Mur aene so x ciner eus (ForskAl) I 775 Muraenesocidae Yes
Oratosquilla nepa Squillidae Yes
Oxyurichthys tentacularis? (C. & V., 1837) Gobiidae No
Palaemonid prawns Palaemonidae Yes
Penaeus semisulcatus Penaeidae Yes
Platax orbicularui (Forskil) 1775 Platacidae Yes
(Bleeker) I 85 I
P oly dacty lus micro stomus Polynemidae Yes
Pomadasys argyrens (Valenciennes) 1833 Pomadasyidae Yes
Portunus pelagicus Portunidae Yes
P seudo sciaena we beri (Bleeker) 187 7 Sciaenidae Yes
Scatophagus argus (L.) 7766 Scatophagidae Yes
Secutor runconius (H.-8.) 1822 Leiognathidae Yes
Setipinna papuezsrs Munro 1964 Engraulidae Yes
S tol ephorus bataoi ensis Hardenberg I 933 Engraulidae Yes
Tetraroge barbara (Cuvier) 1829 Tetrarogidae No
Thalamita sp. Portunidae No
Toxotes jaculator (Pallas) 1766 Toxotidae Yes
Triancanthus indicus Regan 1903 Triacanthidae No
U peneus ait t atus (Forskil) 1 775 Mullidae Yes
Upeneus sp. Mullidae Yes
V aruna litt er at a (Fabricius) I 798 Grapsidae No