Australian Joumai of Ecology (1995) 20, 81-107

Studies on fish movement dynamics in a tropical floodplain

river: Prerequisites for a procedure to monitor the impacts
of mining
K. A. BISHOP,' R. W. J. PIDGE0N2 AND D. J. WALDEN^
^Freshwater Biology Consultant, 4 Sugar Creek Road, Bungwahl, NSW 2423, and
Mlligator Rivers Region Research Institute, Office of the Supervising Scientist,
Jabiru, NT 0886, Australia

Abstract Towards the end of the Wet season in the tropical coastlands of northem Australia,
there are dramatic upstream movements of many fish species in some seasonally flowing
streams. These movements are considered to be a part of refuge-seeking migrations. Aspects of
the dynamics of the movements in Magela Creek (in the 'Top End' of the Northem Territory)
downstream from the Ranger Uranium Mine have been examined with a range of techniques
(mainly direct observation) to facilitate the development of a possible procedure for monitoring
impacts of the mine on the fish community of the creek system. Data on diel pattems of
movements validated that monitoring, for 1 h at midday at a single point adjacent to the mine,
reflects day-to-day changes in total diel movements. To help identify the location of any
impacts arising in the future, information on upstream progress rates, longitudinal changes in
movements, and movements between the creek and lowland billabongs, were used to (i)
demonstrate the creek-long continuity of movements and (ii) indicate the possible sources and
destinations offish approaching the mine. Marked differences in sources were apparent for two
groups of species: terapontids originating from the lowland creek channels, and chequered
rainbowfish and ambassids originating from the floodplain and lowland billabongs. Identification
of the relative contributions from these habitats will require additional monitoring effort.

Key words: fish movement dynamics, migration, mining impacts, monitoring, rivers, tropical
floodplain.

INTRODUCTION latter community-based studies yet, generally, ese con-

General background
The tropical freshwater fish fauna of northem Australia
contains few species exploited commercially or artisanally,
and as a result, the fauna has been studied little in
comparison to those from other tropical regions. There
have also been few studies specifically on movements of
the Australian tropical fish fauna and these are limited to
either movements near estuaries (Kowarsky & Ross 1981;
Russell & Garrett 1988) or the impact of released waste-
waters (Bishop & Walden 1990). In non-specific studies,
information on fish movements has been derived from
investigations focusing on life-cycle components (Beumer
1979; Davis 1985; Griffin 1987; Hogan 1994) or changes
in community characteristics (Beumer 1980; Bishop 1987;
Bishop et al, 1990). Knowledge of and concepts about
movement dynamics have increased the most from the
Accepted for publication October 1994.
cepts are still very much in an early stage of development.
In contrast, much work has been reported on fish
movements in other tropical regions, with the result that
the understanding of the movement dynatnics of many
species is relatively advanced in, for example. South
America (e.g. Lowe-McCormell 1964; Junk 1984; Quiros
1989) and Africa (e.g. Williams 1971; Welcomme 1974,
1975, 1979; Benech & Quensiere 1983a,b). However, as
noted by Cambray (1990) for Africa, many of these
studies have centred on conspicuous movements of adtilts
of relatively large fish species.
Where studies on movement dynamics identify regular,
extensive travel to and from spawning, feeding or refuge
areas, then, following definitions and discussions by
Heape (1931, in Myers 1949) and Harden Jones (1968),
the movements can be more accurately termed as
'migrations'. Knowledge of migration dynamics is fun-
damental to the understanding of how animals survive in
areas with marked fluctuations in conditions. Accordingly,
such knowledge provides the basis for their sustainable
82 K. A. B I S H O P ET AL.
management through, for example, the identification of
important migration paths and/or destinations containing
critical resources.
As will be demonstrated in this paper, this knowledge
can also be integral in the design and interpretation of
biological monitoring studies based on fish. Advocates of
the use of fish in monitoring (Hendricks et al. 1980;
Hocutt 1981; Karr 1981; Karr & Dudley 1981; Fausch et
al. 1984) contend that the advantages of using fish
outweigh the disadvantages, but caution that an awareness
of the disadvantages is essential from the outset. High
mobility is commonly seen as a major disadvantage for
the use of fish in monitoring, but in this paper we present
studies necessary to develop a monitoring procedure
based on this feature. Lake (1986) considered that the
apparently successful use of fish in monitoring river
'health' in the United States is not transferable to
Australia because species richness in Australia is generally
Billabong
Catchment boundary
Fig. 1. Location of the Alligator Rivers Region and Magela Creek in the Northern Territory. Major habitat zones in the Magela Creek
catchment are shown.
low and the communities are modified by introduced
species. However, these charaaeristics are only typical of
communities found in southern Australia, because in
tropical northern Austraha, very few introduced species
are present and species richness is high by world standards
(Bishop & Forbes 1991).
Specific background
Towards the end of the Wet season in the tropical
coastlands of northern Australia, there are conspicuous
upstream movements of many fish species in some
seasonally flowing streams. Upstream movements of a
range of fish species in Magela Creek were first detected
in 1979 during the early ecological studies on fish in the
AUigator Rivers Region of the Northern Territory (Bishop
el al. 1986, 1990, in press). Subsequently, these move-
ments were found to occur regularly in the second half of
East Alligator River
estuary
Magela
floodplain
Lowlands
Escarpment/
Arnhemland
Plateau
Magela Creek drainage system
 F I S H M O V E M E N T D Y N A M I C S AND M O N I T O R I N G 83

the Wet season and, in some species, often on a dramatic and reflected conditions in a broad geographic area
scale. Information on seasonal changes in the composition downstream of the Ranger mine. In comparison with
of fish communities in a range of habitats (Bishop et al. other mine-related monitoring underway (Humphrey et
1986, 1990; Bishop 1987) led Bishop and Forbes (1991) al, 1990), the monitoring of fish movements was con-
to hypothesize that these movements were a part of sidered to provide, potentially, the ultimate assurance
refuge-seeking migrations, important survival strategies that environmental protection measures were performing
enabling fish to reach permanent headwaters in and near satisfactorily.
the Arnhem Land escarpment and so survive the Dry Bishop and Walden (1988) envisaged that procedures
season when conditions deteriorate downstream, in the for the detection of mining impacts on movements would
lowlands and floodplain. Bishop and Eorbes (1991) be developed to relate to three time frames: rapid
presented a provisional model of fish migration in Magela detection based on observations of avoidance behaviour
Creek and stated that the migrations were a predictable (1 day); short-term detection of effects within the move-
response to the temporal and spatial patchiness of ment season based on predictable relationships between
resources in this seasonally flowing stream. daily movement rates and hydrology (1 week); and long-
A large uranium mining operation. Ranger Uranium term detection based on relationships between measures
Mine, was established adjacent to the lowland section of of yearly movement rate and hydrology (1 wet season).
Magela Creek in 1980 (Fig. 1). A major environmental The rapid detection procedure has been described by
concem associated with mining is the management of Bishop and Walden (1990) and some analytical procedures
excess water that accumulates each year as a consequence for the short-term and long-term detection have been
of the seasonal, monsoonal rainfall. These waste waters examined by Bishop and Walden (1991).
are complex and contain elevated concentrations of Field investigations to facilitate the development of
naturally occurring substances (radionuclides, heavy the detection procedures involved three approaches: (i)
metals and suspended solids) and exotic chemicals routine monitoring by direct observation of the numbers
(hydrocarbons and process chemicals) that could pose of fish moving past the mine; (ii) a number of studies on
environmental threats if they were allowed to drain
uncontrolled from the mine site. To date a number of
controlled and semi-controlled releases have been made
Floodplain
to Magela Creek. Humphrey et al, (1993) noted that the
concentrations of mine waste substances in the receiving Fish trapping site
waters are sufficiently low at present that ecological data
gathered from these waters are still regarded as 'baseline',
unaffected by contemporary mining. They also noted
that mining in the Magela Creek catchment was likely to
continue for another 50-100 years and thus the present
phase of mining is an early one.
Gauging station
One of the recommendations of the Australian Govem- GS8210009
ment enquiry into the establishment of the mine (Fox et ong
al, 1977) was that the release or seepage of mine waters
from Ranger should not be allowed to discourage or
impair fish migration. Movements might be affected
directly as a result of any avoidance responses of the fish
towards mine waters introduced into the creek channel.
Of great concem would be any indirect effects that arose
from impacts on recruitment processes (reproduction
and growth), which are most likely to occur in depositional
environments, such as lowland billabongs and the flood-
plain, where harmful substances may accumulate.
Consequently, in 1983, investigations began on the
possibility of developing a procedure for detecting the
future impact of the Ranger Uranium Mine on fish
movements. This development was viewed as important
because, if a reliable monitoring procedure became
available and movements are indicative of migratory Fig. 2. Location of movement sampling sites (MX, MI, GD, CA,
activity, it would provide a means to focus on a critical PO), Ranger Uranium Mine and billabongs in the Magela Creek
survival strategy which was directly ecologically significant study area.
 84 K. A. B I S H O P ET AL.
aspects ofthe movement dynamics of major species; and
(iii) a study of observer bias.
It was considered strategically important to maximize
the pre-impact data-set in order to increase the power
of the routine monitoring procedures in future. Con-
sequently, routine daily monitoring commenced in 1985,
before the procedure was fully validated. There was
some risk in this process but early examination of available
data indicated that it was unlikely that the procedure
would not be validated. Monitoring was standardized
temporally to 1 h at midday and spatially to one point,
site PO (Fig. 2), on the westem channel ofthe creek near
Ranger.
In this paper we describe the studies on movement
djrnamics. Apart from their use in validation, an important
additional function was to improve the understanding of
the movement process thus enabling an evaluation of the
ecological significance of any changes detected in the
future. The studies included an examination of: (i) diel
changes in movements, in order to verify that 1 h obser-
vations at midday provide a representative sample of
total diel movement (i.e validation of the temporal
standardization of the routine monitoring); (ii) longi-
tudinal changes in movements, in conjunction with
tracking studies to determine the continuity of move-
ments and progress rates of moving fish; and (iii) sources
and destinations of moving fish, including movements
between the creek and lowland billabongs. The data
required to validate the spatial standardization (i.e.
westem channel) are presently hmited and will not be
examined here.
Where sufficient information was available, the data
and analyses reported in this paper are for the four most
abundant species, Melanotaenia splendida inornata,
Ambassis spp., Leiopotherapon unicolor and Amniataba
percoides, recorded in the routine monitoring (Table 1).
Data and analyses are also presented on Hephaestus
fuliginosus, a species valued by recreational anglers and
important for the interpretation of data from other
studies.
METHODS
Study area
Magela Creek is a seasonally flowing stream located in
the Top End of the Northern Territory approximately
250 km east of Darwin (Fig. 1). The region is in the
Wet-Dry tropics of northem Australia and many of the
streams flow only seasonally. The headwater section of
Magela Creek hes in the Arnhem Land Aboriginal
Reserve and drains the sandstone Amhem Land plateau.
The stream then flows north across a lowland section of
low rehef for 29 km (average channel gradient of 1.5 m
km"') and then enters a large floodplain area, eventually
draining into the estuary of the East Alligator River a
further 40 km from the lowlands. The lowland and
floodplain areas lie within Kakadu National Park, a
world heritage area. In the incised escarpment area ofthe
headwater section, the stream is permanently flowing
with frequent deep waterholes along a 9 km stretch.
Downstream from here, in the lowland section, the
stream has a braided, shallow, sandy channel system
which dries out almost completely during the Dry season.
Adjacent to the channels there are a number of shallow
lagoons (backflow billabongs), mostly on small tributaries,
which are also seasonal. The floodplain contains a number
of large, deep, permanent waterholes (billabongs) and a
vast area of seasonally inundated swamp. The floodplain
billabongs are not connected by distinct creek channels.
The Ranger Uranium Mine is situated near the middle
of the lowlands section of the creek system, 12 km
upstream from Mudginberri Billabong which lies at the
start ofthe floodplain zone.
Techniques for characterizing movements
Most species of fish moving in Magela Creek are highly
visible in the shallow, generally very clear waters within
the sandy, lowland creek channels, and this enabled their
movements to be observed directly. As recorded in some
African rivers (Bell-Cross 1976, in Cambray 1990), most
small species and juveniles of larger species travel within
the stream margin, where velocities are lowest and large
predators are seldom found, and this also facilitated
observation. Accordingly, direct observation was the
technique most widely used in the studies reported in
this paper, although the exact nature ofthe this technique
varied with the study objectives, and the character ofthe
sites being studied (Table 2). The more traditional
mark-recapture approach for studying movements was
considered impractical because the primary moving fish
were small species, and very large numbers would have
had to be marked due to the immense numbers observed
moving.
In a review. Harden Jones (1968) included direa
observation as one of seven groups of techniques used to
study fish movements. However, this technique has been
used infrequently worldwide mainly because conditions
in rivers and the nature of movements are seldom con-
ducive to direct observation. This is apparently not the
situation in some rivers in the Americas, where direct
observation has been used commonly in the tropics
(Petrere 1985) and occasionally in temperate regions (e.g.
Parker & Franzin 1991).
The primary direct observation technique used in the
studies reported here, as well as in ongoing routine
monitoring, was that of a stationary observer, adjacent to
the creek/charmel, making coimts of fish moving up-
 F I S H M O V E M E N T D Y N A M I C S AND M O N I T O R I N G 85

Table 1. Fish species recorded moving in Magela Creek past the Ranger Uranium Mine (site PO) from 1985 to 1993

Relative abundance
Family/taxa Generally abundant Common Rare

Qupeidae
Nematalosa erebi
Osteoglossidae
Scleropages jardini
Plotosidae
Neosilurus ater -I-
Neosilurus hyrtlii -f-
Porochilus rendahli
Belonidae
Strongylura kreffti
Melanotaeniidae
Melanotaenia nigrans
Melanotaenia spleruUda inomata
Atherinidae
Craterocephalus mananae
Craterocephalus stercusmuscarum
Pseudomugilidae
Pseudomugil tenellus
Synbranchidae
Ophistemon gutturale -I-
Ambassidae
Ambassis spp,
Denariusa bandata
Centropomidae
Lates calcarifer
Terapontidae
A mmataba percoides
Hephaestus fuliginosus +
Leiopotherapon umcolor
Syncomistes butleri +
Pingalla midgleyi +
Apogonidae
Glcssamia aprion
Toxotidae
Toxotes chatareus +
Gobiidae
Glossogobius giurus -I-
Eleotridae
Hyseleotris compressa
Mogumda mogumda
Oxyeleotris lineolatus
Oxyeleotris nullipora

All species are likely to be potamodromous except Lates calcarifer which is catadromous. Species shown in bold are the those examined in
this paper.

stream or downstream across a fine perpendicular to the
water flow over a prescribed time (hereafter termed
movement counts). Such counts were standardized by
the observer being in an elevated position (2-3 m) and
using polarizing glasses (for more details see Bishop
1987; Bishop & Walden 1990),
Depending on the objectives of the study component,
movement rates were calculated from movement counts
for any or all ofthe following four parameters: upstream
movement, downstream movement, net movement (up-
stream movement minus downstream movement) or
movement activity (sum of the upstream and the down-
stream movement). These movement parameters were
also derived from observations made at night using
torchlight and at dusk/dawn using an infra-red light
source and television equipment.
Indirect observations were used on the floodplain as
fish movements were obscured by aquatic vegetation.
This necessitated the use of fishtrapping, a technique
commonly used to study fish movements in tropical
86 K. A. B I S H O P ET AL,

Table 2. Nature of techniques used to characterize fish movements and their use in the components of the study

Components of the study

Diel changes in movements Longitudinal changes in movements Sources and destinations of moving fish

Longitudinal Diversions,
Longitudinal changes in outflow,
Extent of Representativeness Progress rates continuity of mean movement backilow and
Nature of technique diurnalism of midday counts based on tracking movements rates crossflow channels

IndireCT observation
Fishtrapping
(entire diel cycle) +'
Direct observation
Day
Observer still near
creek/channel: +'' + ''
Observer mobile
along creek/channel
Multiple observations
in billabong margins
Night
Observer mobile
along creek using
torch +'
Dusk/dawn
Observer still
near creek + *"
Observer still
near creek using
infra-red light and
television equip. +'

a to c identify study sub-components: a, fishtrapping on the flood plain; b, movement counts associated with the fishtrapping; c, movement
counts through dusk and dawn.
d, data originated from the study on the longitudinal continuity of movements.
e to g identify study sub-components; e, diversion of moving fish from the creek (data utilised from three occasions); f, outflow and backflow
channels; g, crossflow channels.

freshwaters (e.g. Lowe-McConnell 1964; Bayley 1973;
Benech & Quensiere 1983a,b). As the fish trap used was
directional, the movement parameters described above
could be derived from the resultant data.
Diel changes in movements
Diel changes in fish movement are a potential source of
error for monitoring, especially when observations are
made at only one time of day, as is the case with
temporally standardized routine monitoring. A reliable
method for determining whether the 1 h movement counts
at midday provide a representative sample of total diel
movement is the examination of the association between
counts made during this hour and counts made over 24 h
on the same day. This method is very labour-intensive
due to the high level of effort required to make 24 h
observations. To reduce this effort, emphasis was first
placed on establishing the extent of diurnalism, and from
this, the need for observations at night could be deter-
mined.
Extent of diurnalism
Three studies were undertaken to obtain information on
the extent of diurnalism.
Fishtrapping on the floodplain Before the commencement
of routine monitoring, an attempt was made in 1984 to
determine diel movement pattems using a large fish trap
located at the head of the floodplain (Fig. 2). The trap
could catch fish moving upstream and downstream with
the use of 25 m long, trap-wing nets (mesh size 14 mm,
height 1.8 m) which served to funnel fish into separate
upstream- and downstream-facing cages.
The trap was located in the western lobe of the
floodplain head. This lobe is approximately 300 m wide
and the trap was located 200 m from the westem bank.
During the Wet season this area was generally covered
 F I S H M O V E M E N T D Y N A M I C S AND M O N I T O R I N G
by less than 1 m of water and contained a medium
density of aquatic vegetation. Water velocities at the trap
were generally low (approx, <0,1 m s ') although higher
velocities (approx, 0,2-0,3ms ') occurred along the
western bank.
The trap was set over seven 24 h periods (1500h on
3.4,84 to 1500h on 6,4,84, water depth range 0,90-0,76 m
at the trap; 1200 h on 16,4.84 to 1200 h on 18,4,84,
0,46-0.43 m; and 1200 h on 2.5,84 to 1200 h on 4.5.84,
0,16-0,14 m), The trap was cleared every 3h during
these periods and fish were identified and counted,
Diel changes were summarized over the seven 24 h
periods by determining the median numbers captured
during each 3h session along with their 95% sign-
confidence intervals. Movement activity in sessions with
dark throughout (darkness, sessions commencing 2100,
2400 and 0300 h) was compared with activity in sessions
with light throughout (daylight, sessions commencing
0900, 1200 and 1500h) using the H test statistic of the
Kruskal-Wallis /j-sample test.
Movement counts associated with fishtrapping During
the fishtrapping, large numbers of many species of fish
were observed moving upstream along the westem mar-
gins ofthe floodplain at site MX (Fig, 2), approximately
1 km upstream of the trapping site. In this area strong
flows were present as a result of Magela Creek discharging
into the head ofthe floodplain, A preliminary examination
of the fishtrapping showed that the fish trap was not
effectively targeting the primary movement route. To
obtain some comparative diel data on this route, limited
movement counts were made at site MX, Counts at
night were made by torchlight while repeatedly traversing
in a downstream direction along a 10 m length of the
bank. Fish suddenly appearing in this traversed section
were recorded as moving fish and their orientation up-
stream or downstream was taken as indicative of the
direction of their movement. This traversing method was
used so as to minimize the possibility that night lighting
would alter movement pattems.
Counts were made on 5.4,84 (1330-1430 h and
1835-1906h), 6.4.84 (0645-0745 h) and 7.4.84 (0020-
0053 h). Results were summarized so that they were
comparable with the fishtrapping data which were
gathered every 3 h. Some data gathered around dusk and
dawn were summarized into 10 min intervals so that the
time noticeable changes in movement rates occurred
could be accurately determined. Approximately 1 year
later on 10.4.85 (0645-0745 h and 1755-1855 h) addi-
tional data were gathered for the latter purpose.
Movement counts at dawn and dusk An efficient method
for determining if movements are primarily diumal is by
examining changes in movement rates during dawn and
dusk, the periods when light intensity is changing most
87
dramatically, Diumalism would be indicated if rates
noticeably increased and decreased during these respec-
tive periods. In 1988, consistently high movement rates
of fish at site PO provided an opportunity to examine
changes in rates during these periods.
To observe fish moving in low-light conditions, we
used a low-light, infra-red sensitive television camera
(WV1900 National Panasonic Moonlight Camera),
monitor and a 750 nm infra-red light. This form of
illumination has been shown not to disturb the behaviour
of some species of fish in laboratory situations (Beach
1978), The camera was encased in a water-proof housing
and placed partially in the water at right angles to the
creek bank. The camera housing, protruding 0,5 m into
the creek, acted as a barrier to fish moving along the
creek edge, and as a result, all fish moving along this edge
had to pass in front of the camera. The effective viewing
zone (i.e. bottom to surface waters) with the camera was
0 to 1,0 m from the front ofthe camera housing and this
was the zone through which the majority of fish passed
and were counted by use of the monitor.
Dawn observations were made on 8,4.88 (0520-0800 h)
and 19.4,88 (0510-0800 h) and the first sign of light
appeared at around 0610 h on these days. Dusk obser-
vations were made on 4,4,88 (1710-2029h) and 7,4,88
(1750-2049 h) and the last sign of light occurred at
around 1930 h on these days. These data were summarized
into 10 min intervals to accurately determine the time
that noticeable changes in movement rates occurred.
Representativeness of midday counts
The level of representativeness of midday counts was
investigated by an analysis of movement counts made on
15 days (30.5,84,6,6.84,20.2.85,11-15.3,85,19-23.3,85,
4.4.90 and 21,3.91) through all daylight hours (0700-
1800 h; 0700-1900 h on 12,3,85), hereafter referred to as
'ADH counts'. All of these counts were made in the
vicinity of site PO and analyses only focused on upstream
moving fish. The majority of these counts required some,
but generally minimal, data interpolation and this was
done assuming linearity of changes in rates between
times.
One-hourly summaries of the data were made giving
11 one-hour counts for each ADH count. The one-hour
counts were summed giving the total count of fish
observed moving upstream for all daylight hours, here-
after referred to as the 'total ADH count'. Spearman-
rank correlations {r^) were used to examine the association
between total ADH counts and the one-hour midday
counts. Midday counts were considered to be represen-
tative of total ADH counts if the correlations were
significant at the 5% level.
In order to define diumal pattems of movements, the
proportion of the total ADH count occurring in each
88 K. A. B I S H O P ET AL,
hour was determined. To avoid large variation in propor-
tions due to small sample size, only ADH counts greater
than 10 fish moving per day were examined, and
accordingly, the number of ADH counts used was less
than 15 for some species. The data were summarized
over days by determining the median proportions for
each hour along with their 95% sign-confidence intervals.
Overall differences between medians for each hour were
examined using the Kruskal-Wallis ^-sample test. Non-
parametric techniques were used because sample sizes
were small and some of the data were not normally
distributed.
Longitudinal changes in movements
Observations made in preliminary tracking studies, and
other studies on compositional changes in fish com-
munities (e.g. Bishop 1987; Bishop et al, 1990), led to the
hypothesis that fish movements at site PO were a part of
creek-long migrations rather than independent, local
movement activity. To test this hypothesis, the extent of
longitudinal continuity of the movements was determined
by examining the level of association between changes in
movement rates across a series of adjacent sites. To
enable a more detailed interpretation of detected associ-
ations, particularly the nature of time lags between sites,
progress rates of upstream moving fish were determined
through short-distance tracking studies.
Progress rates based on tracking
Progress rates of fish moving upstream were determined
in 1985 and 1988 by timing the passage of fish along
marked intervals of the creek (in 1985: 10 X 10 m intervals;
in 1988: 5 x 5 m intervals). The mean observed progress
rate and its coefficient of variation were then calculated
using the number of intervals traversed as the sample
size.
In 1985 observations were made on three days
(21-23.4.85) along a 100 m section of the creek com-
mencing 400 m downstream of site PO. In 1988 obser-
vations were made on two days (20-21.4.88) along 25 m
sections of the creek commencing 50 and 100 m upstream
of site PO.
Longitudinal continuity of movements
The section of creek between the floodplain and site PO
was studied so that information was available on the
relationship between movements in the floodplain and
those in the creek downstream of the Ranger Uranium
Mine. Movement counts of upstream moving fish were
made daily between mid-morning and midday at five
sites along the creek (Fig. 2) over the following periods:
site MX, 14.2.89 to 2.6.89 (2 days of missing data); site
MI, 7.2.89 to 2.6.89; site GD, 7.2.89 to 2.6.89; site CA,
14.2.89 to 2.6.89 (12 days missing); and site PO, 17.1.89
to 16.6.89 (1 day missing). Counts at site PO arose from
routine monitoring, and accordingly were made over 1 h.
Counts at the other sites were made over 10 min in order
to reduce labour costs. All movement rates were expressed
as numbers moving upstream per hour.
The level of association of changes in movement rates
between adjacent sites was examined by calculating
time-lagged cross correlations between data gathered at
the adjacent sites (using MINITAB software package
procedure CCF [cross correlation function]). The primary
time lag between sites was taken as the lag at which the
maximum significant (P<0,05) correlation occurred.
Spurious correlations were minimized by ignoring sig-
nificant correlations that did not have any neighbouring
significant correlations (significant correlations at lag = 0
were exempt from this condition). Secondary lags were
identified in a similar manner; however, an additional
condition was that they had to be separated from the
primary lag only by insignificant correlations.
The determined time lags between adjacent sites were
taken to be estimates of the travelling times between the
sites. From these times the upstream progress rates could
be calculated from the distances between the sites (MX
to MI = 1500 m; MI to GD = 4500 m; GD to CA =
4000 m; CA to PO = 3500 m). To compare these cal-
culated progress rates (CPR) with the observed progress
rates determined in the tracking study (OPR), a relative
progress rate (RPR) was calculated as: RPR = CPR/OPR.
As cross-correlations could not be calculated with missing
values included, the following data restrictions and inter-
polations were introduced. Site MX: data from 14.2.89
to 2.6.89 could only be used. Additionally, data on two
successive days (19-20.3.89) were linearly interpolated.
Site CA: due to a large gap in the data due to high water
levels, the data from this site had to be analysed in two
sections. CAl: the data used were from 14.2.89 to 18.3.89
inclusive (i.e. mid-Wet movement), and data on 15.2.89,
12.3.89 and 17.3.89 were linearly interpolated. CA2: the
data used were from 1.4.92 to 2.6.89 inclusive (i.e. late-
Wet movement). Site PO: data for 20.3.89 were linearly
interpolated.
Sources and destinations of moving flsh
Key information in helping evaluate any impacts detected
in the future is the location of the origin of the impact,
and the locality of resultant impacts. Accordingly, in
monitoring fish movements it would be valuable to have
knowledge of the sources and destinations of the moving
fish. This was primarily investigated by examining
longitudinal changes in mean daily upstream movement
rates in the 1989 wet season as derived from the
longitudinal continuity study described previously.
 FISH MOVEMENT DYNAMICS AND MONITORING 89

Decline in movement rates between sites was taken to

indicate loss of moving fish, while increases in rates were
taken to indicate gains of moving fish. These gains and
losses were, in turn, taken to indicate the probable
destinations and sources of moving fish,
A second study involved the collection of data on fish
movements between the creek and the lowland billabongs.
From this study it could be determined whether the
billabongs were involved in the movement process.
Movement data were collected on a number of occasions
in billabong outflow charmels and the creek immediately
downstream in order to estimate diversion rates of moving
fish from the creek to the billabongs. Observations at
other times were restricted to outflow, backflow and
crossflow channels of billabongs. The location of sites
referred to below is given in Fig. 3.

Diversion of moving fish from the creek

Occasion 1 Two successive one-hour movement counts

were made during the late afternoon of 2.5.83, one in the
outflow channel ofthe Gulungul Billabong 50 m upstream
of its confluence with Magela Creek, and the other in
Magela Creek 50 m downstream ofthe confluence on the
westem side of the westem channel.

Occasion 2 Movements between Magela Creek and

(iii) Coonjimba Billabong were examined during a 10 min
period on 22,2,89, This count was made in the outlet
channel of the billabong 10 m upstream from its con-
fluence with Magela Creek, The diversion rate of fish to
the billabong from the creek was estimated by comparing
a 10 min count made a brief time before and a short
distance downstream from the confluence.

Occasion 3 Movements between Magela Creek and

Georgetown Billabong were examined on four consecutive

Fig. 3. Schematic details of areas where data were collected on fish

movements between Magela Creek (MC) and the lowland billa-
bongs, (i) GL, Gulungul Billabong; a and b, observation sites in
Magela Creek and the outflow charmel in 1983 (diversion; occasion
1); c, sites of multiple observations in the outflow channel in 1988;
d, observation area in backflow channel in 1988, (ii) CJ, Coonjimba
Billabong; a and b, observation sites in Magela Creek and the out-
flow channel (diversion; occasion 2), (iii) GN, Georgetown Billa-
bong; Oj site of simultaneous observations in MC and the outflow
channel (diversion; occasion 3), (iv) DA and IM, Djalkmara and
Indium Billabongs; a and b, observation sites in the margins of
Djalkmara Billabong; c, observation area in the crossflow channel
leading to Djalkmara Billabong, Scale for each figure as per (iv);
flow direction is indicated within Magela Creek; crossflow and
backflow channels, which transport water otily at high flows, are
indicated by the heavy dashed lines. The biUabongs are indicated
by the light dashed lines.
90 K. A. B I S H O P ET AL,

days during the 1989 Wet season (24-27.2.89). Ten
minute movement counts were made on the westem
bank of the western peripheral channel of Magela Creek
5 m downstream of the confluence with the billabong
outlet channel. From this position movement of fish both
into the Georgetown outflow channel and straight up-
stream could be observed. The proportion of fish that
entered the outlet channel was recorded.
Outflow channels
One-hour counts of fish moving to and from Gulungul
Billabong were made at mid-morning approximately
weekly over a 10 week period during the 1988 Wet season
(4.3.88, 11.3.88,18.3.88,24.3.88,31.3.88,8.4.88, 15.4.88,
22.4.88, 28.4.88 and 6.5.88). Counts were made from the
eastem side of the outflow channel approximately 200 m
100
80
60
Dark
40
I 20
upstream from the Magela Creek confluence with the
channel.
Backflow channels
On 5-6.4.88 Magela Creek backflowed into Gulungul
Billabong. On these days observations were made to
determine whether fish were moving out of the billabong
upstream towards the creek, and at what flow rate of
Magela Creek the backflowing commenced/ceased.
Crossflow channels
With a high discharge on 10.3.89, Magela Creek was
over-topping its batiks and flowing through all five
crossflow channels to Djalkmara and Indium Billabongs.
This presented an opportunity to determine whether fish
were attracted from the billabong to the plume of Magela
Creek water. Crossflow channels A, B and C (sensu
Anon. 1985) combined into one channel before entering
Djalkmara Billabong. At the entry point to the billabong,
the channel was 25 m wide, had an average depth of
0.1m and contained a high density of sedges. The number
of fish observed over a 10 s period at a series of sites along
the banks of the billabong was recorded. Counts were
made at 11 sites, spaced 5 m apart, along the bank across
which the crossflow channel entered the billabong. Five
of these sites were direaly exposed to flow coming from
the channel. Counts were also made at 12 sites, spaced
5 m apart, along the bank of the billabong directly
opposite the bank at which the crossflow charmel entered

o
cd
1200 1800 2400
rhi
0600
the billabong. A schematic diagram showing the location
of counting sites in the billabong is given in the Results.
To confirm whether fish were moving out of the billabong
towards the creek, observations on fish movements were
made while traversing the crossflow channels.
140 r(b)

120
RESULTS
100
TO

80 Diel changes in movements

Dark
60 - Extent of diurnalism

40 [ T Eishtrapping on the floodplain Only sufficient numbers

ofAl. splendida (n = 1310) and y4w^ax5j'5 spp. ( = 1127)
20

were captured during fish trapping to allow an attempt to

0 c
^ T , _ n ^ "1 identify diel movement pattems. Very few specimens of
1200 1800 2400 0600 the terapontid species were captured, but direct obser-
Time of day (h) vations indicated that small numbers of L, unicolor were
present on the downstream side of the trap, and some
Fig. 4. Diel pattems of movement activity of (a) M, splendida and
temporarily entered the cage trap.
(b) Ambassis spp. as determined by fishtrapping in Magela Creek at
the head of the floodplain zone. Medians are based on trapping data Diel changes in movement activity of Af. splendida and
collected over 7 days. The vertical bars are 95% sign confidence Ambassis spp. are summarized over the seven 24 h periods
limits. Shaded area shows period of apparent complete darkness. in Fig. 4. The dominant direction of movement was
 F I S H M O V E M E N T D Y N A M I C S AND M O N I T O R I N G 91

downstream for both species (96,4 and 79,0%, respec- M. splendida

60000
tively).
Activity of M. splendida in darkness was significantly
40000
less(// = 22,96, d.f. = l , P < 0 . 0 1 ) than in daylight. On
average, activity in daylight was approximately 30 times Dark
greater than that in darkness. Activity decreased in all 20000
periods from the 1500-1800 h trapping session through
dusk to the 2100-2400 h session. Correspondingly, ac-
0
tivity consistently increased from the 03000600 h session 1200 1800 2400 0600 1200
through dawn to the 0600-0900 hr session.
12000
Activity of Ambassis spp. in darkness was also sig- Ambassis spp.
nificantly less {H = 7.94, d.f = 1, P<0,01) than in 10000
daylight. On average, activity in daylight was approxi- 8000
mately 6.4 times greater than that in darkness. During
the majority of periods the activity decreased from the 6000
1500-1800 h trapping session through dusk to the 4000 Dark
2100-2400 h session, and correspondingly, increased from
the 0300-0600 h session through dawn to the 0600-0900 h 2000
sessions. Exceptions to this pattem consistently occurred 0
during periods of lower water level. Differences in 1200 1800 2400 0600 1200
patterns with changes in water level are shown in Fig. 5.
CO
200
L u nico ior
150

14 r (a) 100
12 Dark
c
10 50
E
8
^ 6 Dark 1200 1800 2400 0600 1200
CO 4 80
c A. percoides
^^ 2
60
1200 1800 2400 0600 1200
40 Dark "

140 (b)
20
o 120

100
1200 1800 2400 0600 1200
I 80 10
Dark
60 H. futiginosus

40

20
Dark
?200 1800 2400 0600 1200
T i m e of d a y (h)
0
Fig. 5. Changes in diel pattems of movement activity oi Ambassis 1200 1800 2400 0600 1200
spp, determined by fish trapping at different water depths, (a)
Time of day (h)
Water level 0,14-0,16 m, means based on trapping data collected
over 2 days are shown, (b) Water level 0,43-0.90 m, medians based Fig. 6. Diel patterns in movement activity of five fish species
on trapping data collected over 5 days are shown. Shaded area determined by movement counts at site MX, 1 km upstream from
shows period of apparent complete darkness. The vertical bars are the fish trapping site. Shaded area shows period of apparent
95% sign confidence limits. complete darkness.
92 K. A. BISHOP ET AL,

Movement counts associated with fishtrapping In contrast vations on splashing sounds produced by predacious
with fishtrapping, vastly more movement activity was fishes (e.g. L, unicolor, Megalops cyprinoides, Lates cal-
apparent and the upstream direction of movement was carifer and Arius leptaspis) ambushing schools of small
more dominant for aff species at site MX: M, splendida, moving fish provided supporting evidence of lack of
20758 vs 23.4 per 3h sampled and 99.3% upstream; movement during night hours. Such sounds reduced
Ambassis spp., 4497 vs 20.1 and 100%; L, unicolor, 57.8 vs greatly between 1900 and 1930h and no sounds were
0.0 and 79.2%; A, percoides, 24 vs 0.0 and 100%; H, apparent during the session of complete darkness.
fuliginosus, 3.8 vs 0.02 and 60%. On three occasions the movement activity of either Af.
Diel changes in movement activity for this suite of five splendida, Ambassis spp. or L, unicolor was consistently
species are shown in Fig. 6. No movement was detected high so that changes in activity could be examined
for any of the species during the only session of complete during dusk or dawn. Changes in activity on these
darkness sampled (session commencing 2400 h). No occasions are shown on Fig. 7. During the dawn of
movement was detected for any of the terapontid species 6.4.84, the movement activity of M, splendida was at a
for the dusk session (session commencing 1800 h). Obser- low level when counts were first possible at 0645 h.
Activity increased from this time onwards, particularly
after 0705 h. Similar pattems of increasing activity were
apparent for L, unicolor during the dawn of 10.4.85.
4000 r- (a) During the dusk of 10.4.85 the activity of Ambassis spp.
M, splendida
showed a consistent decrease between 1755 and 1845 h.
3000
Movement counts at dawn and dusk The only species of
2000 the standard suite observed moving in this study were
M, splendida, Ambassis spp. andL. unicolor. These species
1000 were virtually moving only in an upstream direction and
changes in their movement rates during dawn and dusk
0 are shown in Fig. 8.
0645 0655 0705 0715 0725 0735 No movements were recorded before first light (around
0610 h) for M, splendida and 50 min after first light for L.
100 L. unicolor unicolor. Within 20 min after first movements, the
movement rate of M. splendida increased to approximately
80
30 times the initial movement rate and then decreased at
60

40 Dawn
M splendida
20

0
0
0645 0655 0705 0715 0725 0735
(C) - 0500 0600 0700 0800 1700 1800 1900 2000 2100
12000
Atnbassis spp.
^ 12 0 0 0 - Ambassis spf>. Ambassis spp.

8000 500

0500 0600 0700 0800 17O0 1800 1900 2000 2100

4000
L. unicoior L. unicoior

0
1755 1805 1815 1825 1835 1845 CO
0)
T i m e of d a y ( h ) ^ 0500 0600 0700 0800 1700 1800 1900 2000 2100

Fig. 7. Changes in movement activity of three fish species deter-
mined by migration counts at site MX through dawn and dusk on
selected days, (a) M, splendida, 6 April 1984, dawn; (b) L. unicolor,
10 April 1985, dawn; (c) Ambassis spp., 10 April 1985, dusk.
T i m e ot d a y (h)
Fig. 8. Changes in upstream movement rates through dawn and
dusk for three fish species at site PO. Averages for 2 days are given.
Shaded area shows period of apparent complete darkness.
 F I S H M O V E M E N T D Y N A M I C S AND M O N I T O R I N G 93

times to half of this level. Movement was recorded in all Representativeness of midday counts
but one 10 min period after first movements for L.
Midday counts were significantly correlated with total
unicolor. One to five Ambassis spp. were observed moving
ADH counts for M. splendida {r, = 0.93, P<0,01),
in each 10 min period up to 10 min before first light. The
Ambassisspp. {r, = 0,80,P<0,01),L, unicolor {r, = 0,63,
movement rate then increased rapidly to reach a maxi-
P<0.05)a.nd A. percoides{r, = 0,88,P<0,01), indicating
mum level 30 min later of 1350 times greater than that
that midday counts were representative of total ADH
occurring in complete darkness. The minimum movement
counts. Insufficient data were available on the movement
rate recorded after this time was approximately 60 times
of H. fuliginosus.
greater than that recorded before first light.
Diurnal patterns of movement of each species are
Last movements occurred 30 min before last light
summarized in Fig, 9, The overall differences between
(~1930h) for M. splendida and the greatest reduction in
medians of the one-hour counts were significantly dif-
rates occurred 10 min before this, Ambassis spp, were
ferent for M. splendida{H = 34,93, d,f. = 10,P<0,01),
recorded moving in every 10 min period, however, 30
Ambassis spp, (H = 18,47, d,f, = 10, P<0,05) and
min after last light only one to three individuals were L. unicolor (H = 18,53, d,f, = 10, P<0,05), but not for
recorded moving per period. Movement rates oi Ambassis A. percoides (H = 15,33, d,f, = 10,P>0,10),
spp, decreased most dramatically 30 min before last
light. L. unicolor was not recorded moving during dusk
observations.
Longitudinal changes in movements

Progress rates based on tracking

0.3 M. splendida

0.2 Observed upstream progress rates of tracked moving fish

are summarized in Table 3, Three species/size groups
0.1 were recognized in relation to differences in mean pro-
0.0 gress rates. Slow moving (0,116-0.130ms"' or 4800-
0800 1000 1200 1400 1600 1800 5380 m day"'): adults of Af. spiendida and Ambassis spp.
0.9 -. Ambassis spp. Moderate moving (0.156-0,175 m s ' or 6460-7250m
o day"'): juveniles (i,e, <50 mm SL) of//, fuliginosus and
o
0.6 - ' . ,

A. percoides. Fast moving (0,206-0.228ms"' or 8530-

f =
E 0.3
9440 m day'): adult L. unicolor and A. percoides.
o For Af, splendida and Ambassis spp,, schools rather
o 0.0
E
^ .
1
1
1 1 1 than individuals were followed, and on average they
X 0800 1000 1200 1400 1600 1800 contained 90 and 250 individuals (adults only), respec-
Q 0.2 L. unicolor tively. Approximately 80% of these schools contained
2 both species and on average ambassids out-numbered
o 0.1
Af, splendida by 2,5:1.
0.0
Only adult L. unicolor were followed: one individual
o 0800 1000 1200 1400 1600 1800 and two groups (4 and 10 individuals), which additionally
A. percoides contained/I, percoides. Data on .4. percoides were collected
o
O in 1985 and 1988, In 1988 primarily juveniles were
moving (schools ranging in size from 10-15 individuals
c
(0
with L. unicolor present), and in 1985, adult fish only
were followed (schools ranging in size from 3-19 in-
0800 1000 1200 1400 1600 1800 dividuals). The progress rate estimated for the adults was
0,2 r
H. tuliginosus greater than that determined for the juveniles. Data on
H. fuliginosus were based only on two juveniles.
0.1

0.0 Longitudinal continuity of movements

0800 1000 1200 1400 1600 1800

Time of day (h)
Fig. 9. Diumal changes in upstream movement rates offivespecies
in Magela Creek near site PO, Values shown are the median
proportions ofthe total count made over all dayhght hours (ADH),
Dotted lines indicate 95% sign confidence hmits. Sample size (n)
and total number of fish (f) are shown.
Melanotaenia splendida and Ambassis spp, were the only
species of the standard suite which were consistently
moving in sufficient numbers to allow the examination of
relationships in movement rates between days and sites.
Day-to-day changes in upstream movement rates of
these species at the five sites along the creek are shown in
94 K. A. B I S H O P ET AL,

Figs 10 and 11, respectively. Cross-variable correlograms Sources and destinations of moving fish
used to examine associations between changes in move-
ment rates between adjacent sites are given in Figs 12 Longitudinal changes in mean daily movement rates
and 13, respectively. Significant cross-correlations were
Daily upstream movement rates at each of the five
apparent between movement rates of M, splendida at
counting sites for the suite of five species are shown in
every pair of adjacent sites. The result was similar for
Fig. 14. Changes in mean movement rates between sites
Ambassis spp., except for the site pair GD-CA, which
are summarized in Table 5.
showed no significant cross-correlation in the late-Wet
season. Travelling time estimates and calculated upstream For M, splendida and Ambassis spp. the movement
progress rates for both species are given in Table 4. rates consistently decreased between sites MX and CA
and then increased between sites CA and PO. Accor-
Travelling times for M, splendida could be estimated
dingly, the floodplain downstream of site MX would
for each creek section between adjacent sites. All calcu-
appear to be the primary source of fish passing site PO,
lated progress rates were less than the observed progress
while a secondary source existed between sites CA and
rates determined in the tracking study. This difference
PO. The greatest and second-greatest decrease in rates
was greatest, and hence progress rates lowest, in the most
occurred between creek sections M X - M I and M I - G D ,
downstream creek sections, M X - M I and M I - G D . For
respectively. For M, splendida, these were the creek
the creek section GD-CA in the later part of the Wet
sections in which the lowest and second-lowest calculated
season, a secondary peak (= lag) in movement was
progress rates occurred, respectively. Progress rates were
apparent 3 days after the primary peak.
also low in both of these creek sections {or Ambassis spp.
Travelling times ior Ambassis spp. could be estimated
For the three terapontid species, the general pattem
for each creek section except the GD-CA section in the
was of consistent increases in movement rates from sites
late-Wet season. As for Af. splendida, all calculated pro-
MX to PO (i.e. opposite to that recorded for M, splendida
gress rates were less than the observed progress rates.
and Ambassis spp.). This result indicates that the creek
This difference was greatest for the most downstream
between sites MI and PO appeared to be the source of
creek sections, M X - M I and M I - G D in the middle part
the majority of fish passing site PO {L, unicolor 78%; /I.
of the Wet season. Slower progress rates occurred in the
percoides ~66%; H, fuliginosus 91%). The only exception
most upstream creek section (CA-PO) in the late-Wet
to the abovementioned pattem was a small decrease in
season.

Table 3. Mean observed upstream progress rates of moving fish determined by tracking

Species codes*

Data/parameter MS AM LU AP HF

1985 observations
Number of fish 80
Number of groups 7
Number of intervals traversed by all groups
(10,20,30 m) 28,6,3
Mean progress rate
(ms-') 0.228
(m/day"')-* 9440
Coefficient of variation of progress rate 22.8

1988 observations
Number offish 2450 6190 15 40 2
Number of groups 27 25 3 4 1
Number of intervals traversed by all groups
(5 m) 112 98 15 20 5
Mean progress rate
(ms-i) 0.116 0.126 0.206 0.175 0.156
(m day-')' 4800 5220 8530 7250 6460
Coefficient of variation of progress rate 27.6 22.3 27.7 24.0 35.3

Species codes: MS, M. splendida; AM, Ambassis spp.; LU, L. unicolor; AP, A, percoides; HF, H, fuliginosus, 'Progress rates determined fo
11.5 daylight hours of mo\ ir](; each day.
 F I S H M O V E M E N T D Y N A M I C S AND M O N I T O R I N G 95

Table 4. Estimated travelling times and calculated upstream rates between sites M X and M I for L. unicolor. Such a
progress rates of Af. splendida and Ambassis spp. as determined in decrease indicates that many fish moving upstream from
the longitudinal continuity study site M X appeared to remain downstream of site M I
within Mudginberri Billabong.
Fish species
Ambassis spp. For L. unicolor and A. percoides, the greatest absolute
Parameter/section of creek M. splendida
increase in movement rates occurred between sites M I
Travelling times (i.e. lag in days) and G D , while for H. fuliginosus, the greatest increase
Sites MX-MI 2 1 occurred between sites G D and CA. When adjusted in
Sites MI-GD 2 4 relation to the length of creek between sites, the greatest
Sites GD-CAl 1 0 increase in rates occurred between sites M X and M I for
Sites GD-CA2 0,3
A. percoides.
Sites CAl-PO 0 0
Sites CA2-P0 0 6
Diversion of moving fish from the creek
Calculated progress rate (m day ')
750 1500
Only M. splendida and L. unicolor were moving in
Sites MX-MI
Sites MI-GD 2250 1125 sufficient numbers to warrant an examination of the
Sites GD-CAl 4000 >4000 available data. The data on these species are summarized
Sites GD-CA2 >4000,1333 in Table 6.
Sites CAl-PO >3500 >3500 The highest diversion rates for M. splendida were
Sites CA2-PO >3500 583 recorded during Occasion 3 (Georgetown Billabong).
The observations on this occasion were made immediately
Relative progress rate
0.16 0.29
following a moderate flood event early in the 1989 Wet
Sites MX-MI
Sites MI-GD 0.47 0.22 season and flows consistently decreased over the four
Sites GD-CAl 0.83 0.77 observation days (Fig. 15). Large numbers of fish were
Sites GD-CA2 >0.83,0.28 moving within this shallow, peripheral branch of the
Sites CAl-PO >0.83 >0.67 creek on each observation day and the dominant direction
Sites CA2-PO >0.83 >0.11 was upstream. On each day the majority, if not all, of the
fish were diverted into the outlet channel of the billabong.
= travelling times could not be determined as no significant
cross correlations existed or significant correlations did not meet
The diversion rate increased as the creek discharge
conditions for the identification of primary lags. Where two values decreased, and it was apparent that movement up the
are given, the first is derived from the identified primary lag and the creek was prohibited by emerging sandbanks, a result of
second is derived from the secondary lag. Refer to text for site falling water levels.
codes.

Table 5. Changes in mean upstream movement rates betweenfivesites along Magela Creek during the 1989 Wet season

Gain or loss rate per species

Section of creek MS AM LU AP HF

Absolute change (n h ')

Sites MX-MI -7450 -34675 -2.78 + 1.07 + 0.26
Sites MI-GD -3825 -9629 + 9.58 + 1.10 + 1.35
Sites GD-CA -1857 -597 + 5.76 + 0.93 + 2.03
Sites CA-PO + 2514 + 3737 + 0.93 + 0.93 + 0.18

Change relative to movement rates at site

MX (%) -37% -66% -38% + 243% + 236%
Sites MX-MI -19% -18% + 132% + 250% +1227%
Sites MI-GD -9% -11% + 80% + 211% +1845%
Sites GD-CA + 13% + 7% + 13% + 211% + 164%
Sites CA-PO
Change relative to the distance between sites:
(nh^' km 1)
Sites MX-MI -4960 -23120 -1.85 + 0.71 + 0.17
Sites MI-GD -850 -2140 + 2.13 + 0.24 + 0.30
Sites GD-CA -460 -150 + 1.44 + 0.23 + 0.50
Sites CA-PO + 720 + 1070 + 0.27 + 0.27 + 0.05

table 3. Refer to text for site codes. A gain is indicated by + and a loss is indicated by .
96 K. A. BISHOP ET AL,

Table 6. Movement rates within Magela Creek and diversion rates to billabong outlet channels for M, splendida and L, unicolor

Within Magela Creek

Diversion rate of
Discharge at Upstream Downstream upstream moving fish to
GS8210009 movement rate movement rate billabong outlet channel
Date (m's') (nh') (nh') (%)

Af. splendida. Occasion 1 (Gulungul Billabong)

2.5.83 8.3 1104 16.0

M, splendida; Occasion 2 (Coonjimba Billabong)

22.2.89 27.0 12 864 24 5.5

M, splendida; Occasion 3 (Georgetown Billabong)

24.2.89 8.0 1640 0 80
25.2.89 6.5 2933 165 95
26.2.89 5.0 1935 90 100
27.2.89 4.5 1495 48 100
L. unicolor; Occasion 1 (Gulungul Billabong)
2.5.83 8.3 62 32.0

200000
600000
150000
400000 -
100000

50000 200000 -

120000 200000 r

80000 -
100000
2 40000

L
E 120000 E 120000 r (c)
>
o 80000
E 60000
4000O
E
CO
CD
05
Q. Q. 120000 r (d)
80000 r ( d )
3
80000
40000
40000

100000 r
120000 (e)
50000 80000

40000
Feb Mar Apr May 0
Feb Mar Apr May
Fig. 10. Day to day changes in the upstream movement rates of
M. splendida at five sites along Magela Creek during the 1989 wet Fig. 11. Day to day changes in the upstream movement rates of
season, (a) MX (flood plain), 0 km; (b) MI, 1.5 km; (c) GD, 6.0 km; Ambassis spp. at five sites along Magela Creek during the 1989 wet
(d) CA, 10.0 km; (c) PO, 13 5 km. Distances given are distance season, (a) MX; (b) MI; (c) GD; (d) CA; (e) PO. Distances up-
upstream from the floodplaui. stream from the floodplain (site MX) as for Fig. 10;
 F I S H M O V E M E N T D Y N A M I C S AND M O N I T O R I N G 97

The lowest diversion rate for M. splendida was recorded
during Occasion 2 (Coonjimba Billabong). The obser-
vations on this occasion were made during a moderate
flood event early in the 1989 Wet season (Fig. 15). It is
notable that this lowest diversion rate occurred within
the occasion when flows were greatest and upstream
movement rates within the creek were greatest.
Only one diversion rate estimate was available for L.
unicolor and this arose from Occasion 1 (Gulungul
Billabong). The observations were made within a week
after a major flood event, the last for the 1983 Wet season
(Fig. 15). The diversion rate estimate was twice that
recorded for M. splendida during this occasion.
Outflow channels
Flood events in the 1988 Wet season were all minor, and
four of these events occurred within the period of weekly
observations in the Gulungul Billabong outflow channel
(Fig. 15). No other flood events occurred during any
observation, or within the period after the week 6
observation. Base-flow water levels fell drastically from
this time until the week 8 observation. Only M. splendida,
Ambassis spp. and L. unicolor were moving in sufficient
numbers to warrant an examination of the available data.
The data on these species are summarized in Table 7.
For the first four observations small to moderate
numbers of M. splendida were moving and the dominant
direction was upstream. The dominant direction of
movement was downstream from week 5 to the end of
the study, and moderate to very large numbers were
recorded moving. Very large numbers were recorded
moving downstream on week 8, and this corresponded to
the end of the period when base-flow water levels had
dropped rapidly. Schools of these fish were followed
downstream along the outflow channel, and it was noted

1.0
Section MI-GD
0.5

0.0 _tllllII. lii_l

!--
-0.5 .-

1 1
1 1
20 0 ' 5 10 15 20

1.0

Section GD-CA2
0.5 -

o
u 0.0
c
o
yi -0.5 -

o 1 1 1 1 1
-1.0
O 0 5 10 15 20

1 .U
1, Section CAl - P O Section CA2-P0
-II
0.5

0.0 111.,
-
MMMMM II Ilai
0.5 -
1 1 1 1 1
10
10 15 20 15 20

Lag(d) Lag (d)

Fig. 12. Cross-variable correlograms used to determine travelling times between sites along Magela Creek for upstream moving M. splendida.
Dashed lines indicate the critical value for significance at P = 0.05; the correlation identifying the primary lag is indicated by *; CAl:
observations at site CA in mid wet season; CA2: observations at site CA in late wet season.
98 K. A. B I S H O P ET AL,

that when they came to the Magela Creek confluence,
they turned into the creek and proceeded upstream.
Movement rates ior Ambassis spp. were low during the
first six observations, and most movement was in an
upstream direction. On week 7 large numbers were
recorded moving upstream; however, after this time the
dominant direction was downstream. Extremely large
numbers were recorded moving downstream on weeks 8
and 9, the times which corresponded to the end of the
period when base-flow water levels had dropped rapidly.
On week 8 schools of these fish were followed downstream
along the outflow channel and, as for M, splendida, when
they came to the Magela Creek confluence they turned
into the creek and proceeded upstream.
The dominant direction of movement for L, unicolor
was downstream for all observations except weeks 2 and
3. The greatest rate of downstream movement occurred
within the period when base flow water levels were
falling at the greatest rate (weeks 6 to 8).
Back/low channels
Large numbers of Af. splendida and Ambassis spp. were
observed moving upstream from Gulungul Billabong to
Magela Creek through the backflow channel when back-
flowing occurred across a number of days (Fig. 15). The
extent of these movements was not quantified. The

Section MX-MI
0.5

00 Illiilll
*
.ill
05

1 n 1 1 1 1 1
15 20 20

1.0
Section G D - GA1 Section GD-GA2
0.5
o
o
0.0 llll.
* "" " "
Il.ll..
41
_

i? -0.5

o 1 1 1 1 1 1 1 1 1 1
O 0 5 10 15 20 0 5 10 15 20

1.0
Section CA1 - P O Section CA2-P0
0.5

0.0 It. "~

IlllllIilla-..
-0.5

-1.0 1 1 1 1 1 1 1 1 1 1
10 15 20 10 15 20

Lag(d) Lag (d)

Fig. 13. Cross-variable correlograms used to determine travelling times between sites along Magela Creek for upstream moving Ambassis spp.
Dashed lines indicate the critical value for significance at P = 0.05; the correlation identifying the primary lag is indicated by *; CAl:
observations at site CA in mid wet season; CA2: observations at site CA in late wet season.
 F I S H M O V E M E N T D Y N A M I C S AND M O N I T O R I N G 99

backflowing was estimated to start and cease when flow DISCUSSION

in Magela Creek reached 25 m^ s"'.
Diel changes in movements
Crossflow channels

The observations about the crossflow channel entering Extent of diumalism

Dialkmara Billabong were made at the commencement
of a maior flood event in the middle of the 1989 Wet
season (Fig. 15). Patterns in the distribution and abun-
dance of M. splendida and Ambassis spp. along the
margins of the billabong and in relation to the crossflow
channel carrying water from Magela Creek are shown in
Fig. 16.
The markedly greater abundance of M. splendida in
the vicinity of the channel entry area provides good
evidence that this species was attracted to the flows
entering the billabong. The evidence for ambassids is
less clear. Observations along the length of the crossflow
channel(s) revealed that large numbers of both species
were moving upstream towards Magela Creek proper.
These movements were not quantified.
60000
The three studies undertaken (fishtrapping, FT; move-
ment counts associated with fish trapping, MCAFT; and
movement counts through dawn and dusk, MCTDD)
yielded a range of evidence that indicates movements of
the examined species were essentially diurnal. The
greatest amount of evidence for this was collected for the
most abundant species, M. splendida and Ambassis spp.:
greatly reduced or no activity in darkness (all studies),
increase in activity as light intensity increased (studies
FT andMCTDD for both species and study MCAFT
for M. splendida only), and a decrease in activity as light
intensity decreases (studies FT and MCTDD for both
and study MCAFT for Ambassis spp. only). None of the
terapontid species displayed activity at night in study
MCAFT. Additional evidence of diumality was obtained
for L. unicolor in studies MCAFT and MCTDD where
increases in activity coincided with increases in light
intensity.

50000 V
300 r (a)

May

(b) ,B
o 100 - 1
(0
U i \ 1 i
0
Feb
-1 Apr
J
May
03
15000 D
C
(C)
300 h

200

100

Feb Mar Apr May

Fig. 15. Daily changes in discharge at gauging station GS8210009

0 5000 10000 15000
Distance upstream from site MX (m)
Fig. 14. Mean daily upstream movement of five species along
Magela Creek between site MX and site PO (13 500 m) for the period
7 February 1989 to 2 June 1989. {m) Ambassis spp.; () M.spendida;
(A) L. unicolor; (T) A. percoides; () H. fuliginosus.
on Magela Creek during periods when movement counts v?ere
made in the vicinity of backflow billabongs. (a) 1983, (b) 1988 and
(c) 1989. Arrows indicate occasions when counts were made. In
1983 and 1988 counts were made only near Gulungul Billabong;
observations in backflow conditions are indicated by B. In 1989
counts were made near Coonjimba Billabong (CA) and Georgetown
Billabong (GN), and within and near Djalkmara Billabong (DA).
 100 K. A. B I S H O P ET AL.

Data arising from fishtrapping contrasted greatly with
those from movement counts in that rates were much
lower and the downstream direction was dominant in
study F T These differences probably reflect inefficiencies
of fish trapping in comparison to movement counts, and
the possibility that the fish trap was not located on a
primary movement route. Bishop (1993) suggested that
the fish moving downstream to the fish trap were
previously upstream-moving fish from the primary route
which were unable to proceed further upstream. If this
was the case, then the diurnality observed in the fish-
trapping was a reflection of movement activity on this
route.
It is a common observation that different tropical fish
species migrate along rivers at differing times of the day
Table 7. Movement rates of three fish species in the outflow
channel of Gulungul Billabong during the 1988 Wet season
Movement rates (nh'')
(e.g. Lowe-McConnell 1964; Benech & Quensiere 1983a;
Silva & Davies 1986). Arnold (1974) stated that most
upstream movements are confined to daylight hours.
Smith (1985) indicated that this is particularly the case
for smaller species and Jonsson (1991) commented that
diurnality of upstream movement is usually associated
with large numbers moving, turbid waters or very high
flows.
Only a limited amount of information is available on
diel movement patterns of Australian freshwater fishes,
and this only arises from temperate southeastern
Australia. Mallen-Cooper (1994) indicated that in the
Murray River the terapontid, Bidyanus bidyanus, and the
clupeid, Nematalosa erebi, primarily migrate upstream
during daylight periods. In contrast, Mallen-Cooper
(1992) noted that the percichthyid, Macquaria ambigua,
moved throughout the night and day, but more so at
dusk and dawn (i.e. it appeared to be primarily crepus-
cular). Data presented by Harris (1983) from the Nepean
River indicated that the juveniles of another percichthyid.

Species/date and week Upstream Downstream Net'

N
M. splendida
4.3.88 167 19 148
11.3.88 2 34 1 Inflow from
33
18.3.88 3 545 0 Magela Creek North bank
545
24.3.88 4 437 380 (originating fram the
57
31.3.88 5 862 2760 cross-flow channel)
-1898
8.4.88 6 642 970 -328
15.4.88 7 3108 3160 -52
24.4.88 8 570 11990 - 11 420
100
28.4.88 9
M. splendida
590 834 -244
50
6.5.88

Ambassis spp.
4.3.88
10

1
60 82 -22
0 ill I I . . .
0 0 0 20 Ambassis spp.
11.3.88 2 427 5 422
18.3.88 3 5 0 5
24.3.88 4 0 0 0
31.3.88 5 0 0 0
8.4.88 6 0 0 0 Djalkmara Billabong
15.4.88 7 4492 80 4412
22.4.88 8 1665 78805 - 77 140
28.4.88 9 1273 13 695 - 12428 100
6.5.88 10 231 239 M. splendida
50
L. unicolor 0
4.3.88 1 103 119 -16
11.3.88 2 66 12 54
18.3.88 3 15 8 7 20 Ambassis spp.
24.3.88 4 47 54 -7 I I . . .
0^-
31.3.88 5 63 65 -2
8.4.88 6 115 208 -98
15.4.88 7 31 386 -355
22.4.88 8 224 325 -101 South bank
28.4.88 9 3 10 m
26 -23
6.5.88 10 4 5 -1 Fig. 16. Distribution and abundance of Af. splendida and Ambassis
spp. along the margins of Djalkmara Billabong in relation to a
'Net downstream movement is indicated as negative. crossflow channel carrying water from Magela Creek.
 F I S H M O V E M E N T D Y N A M I C S AND M O N I T O R I N G
Macquaria novemaculeata, may also primarily be crepus-
cular.
It can not be inferred that all species moving upstream
along Magela Creek are likely to display diumalism. The
species examined may not necessarily consistently display
diumalism, as a number of studies have shown the same
or different populations of one species, or even the same
individuals, displaying either diumalism or noctumalism
(e.g. Eriksson 1978). This dualism may be a result of
differences in environmental conditions such as light
intensity or the availability of cover.
Notable in the above context was the result in study
FT that the movement activity of Ambassis spp. tended
to be higher during night hours on days when water
levels were lower. Of the species examined, it is apparent
that Ambassis spp. is more inclined or able to migrate
under conditions of lower light intensity. This was most
obvious in study MCTDD where, in contrast to M.
splendida, some individuals were moving during complete
darkness, substantial increases in movement rates com-
menced earlier and substantial decreases in rates
commenced later. Corresponding with this observation.
Bishop (1993) noted that Ambassis spp. has a larger
relative eye diameter than M. splendida: 12% versus
7% SL, respectively. Glossamia aprion was another
species which appeared to move under conditions of
lower hght intensity in study MCTDD (Bishop unpubl.
data) and, similarly, this species also has a relative eye
diameter (10% SL) larger than M. splendida (Bishop
1993).
Representativeness of midday counts
One-hour, midday counts were found to be representative
of total ADH counts for each of the four species for
which representativeness could be determined. Given
the evidence that the movements of the examined species
were essentially diumal, it could reasonably be inferred
that the midday counts would be representative of total
diel movement for these species. Accordingly, daily
pattems in midday counts would be expected to depict
daily pattems in total diel movement, an initial validation
of the temporal standardization of the routine monitoring
procedure.
In respect to diurnal pattems of movements, it is
notable that Ambassis spp., the species which appeared to
be more able or inclined to move in conditions of lower
light intensity, had the greatest proportion of movement
in the early morning and the early evening. In contrast,
M. splendida and L. unicolor, the species which appear to
be less able or inchned to move in conditions of lower
light intensity, had the greatest proportion of movement
in the middle of the day.
The diurnal movement pattern of Af. splendida could
also reflect the downstream distribution of habitats where
the number of moving fish build up through dusk or
101
during the night. This is most likely to occur where deep
waterbodies are adjacent to the creek and be the result of
an increased reluctance by moving fish to leave deep
waters as light intensity decreases or, alternatively, be
due to night movements within the deep waters. The
closest deep waterbodies adjacent to the creek are in the
vicinity of the Magela-Djalkmara confluence, some
1500 m downstream of site PO. Assuming that (i) move-
ment from these waterbodies would commence at about
0630 h (see Fig. 8), and (ii) the upstream progress rate is
approximately 0.1 m s ' ' (see Table 3), then upstream-
moving fish from this source would be expected to
commence passing site PO by about 1100 h. This time
corresponds with a very significant rise in the proportion
of movement at this site (Wilcoxon W = 118, f < 0 . 0 0 1 ;
comparison of median proportions of 1000-1100 h vs
1100-1200 h).
Longitudinal changes in movements
Progress rates based on tracking
Considerable differences in observed upstream progress
rates were detected between species in the present study.
Limited additional data from the study area (Bishop
1993) indicate that juvenile Toxotes chatareus have pro-
gress rates comparable to the slow-moving species, and
sub-adult N. erebi have rates in excess of the fast-moving
species (11000m day"' vs 8530-9440m d a y ' ) .
Differences in progress rates were also apparent
between size classes of A. percoides, and it is very likely
that such differences occur within the other species, as it
is a common observation that swimming speeds are
positively related to body size (e.g. Bainbridge 1958;
Stahlberg & Peckmann 1987).
It is informative to examine some of the data on
progress rates in relation to information on mean water
velocities in the creek and critical swimming speeds of
the fish. Had M. splendida breasted the mean creek
velocity of 0.5 to 0.8 ms"' (derived from East et al. [1987]
for a discharge of lOm^s*'), their progress rates relative
to the water would have been 0.62-0.92ms'. These
rates are considerably greater than the critical swimming
speed of 0.32 m s"' (6.4 body lengths per second) derived
in the laboratory for this species by Bishop and Walden
(1991). It can therefore be inferred that M. splendida
follow minimum velocity paths in the creek to progress
as they do. By routinely monitoring moving fish along
the creek's margins, these minimum velocity paths are
effectively targeted.
Longitudinal continuity of movements
From the head of the floodplain to site PO, a high level
of longitudinal continuity of movements of M. splendida
102 K, A, B I S H O P ET AL.
and Ambassis spp, was indicated. This result gives support
to the hypothesis that fish movements at site PO were a
part of creek-long migrations rather than independent,
local activity.
The calculated progress rates from the analyses varied
between sites, and was always less than the observed
progress rates. It is likely that this was mainly caused by
the occurrence of intra- and inter-species interactions
along the creek which disrupt movement activity. Such
interactions were commonly seen along the length of the
creek, but were rare within the tracking sites from where
the observed progress rates were determined. Variation
in hydraulic conditions along the creek could also have
been partially responsible for the progress rate differences.
It would therefore appear that the observed progress
rates provide an indication of rates achievable under
optimal conditions (low interactions, ideal hydraulics),
while the calculated rates are more realistic in that they
reflect rates achievable under the range of conditions
along the creek. Consequently, the relative progress rates
given in Table 4 essentially provide an index of the
suitability of conditions for upstream movement along
the creek; values near unity reflect near ideal conditions.
Data on progress rates in the tropics are sparse and
most originate from studies on South American fishes.
Bay ley (1973) noted that characin, Prochilodus platensis,
took 3-5 weeks to progress 120 km up the River
Pilcomayo, and this is equivalent to an average progress
rate of 3430-5710m day"'. This is within and above the
higher end of range of calculated progress rates for the
slow-moving species in the present study, De Godoy
(1959) studied the curimbata, Prochilodus scrofa, in the
Mogi Guassu River and average progress rates of 10000
to 16 500 m day"' were apparent before spawning. This is
greater than the observed progress rates of the fast-
moving species in the present study.
In temperate Australia, Reynolds (1983) found the
average upstream progress rate of golden perch, Mac-
quaria ambigua, along the Murray-Darling River system
to be 1900-4100 m day ', Codwatch (1993) recorded
downstream progress rates of Murray cod, Maccullochella
peeli, in the upper Murray River system to be up to
4500 m day"', These rates are comparable to calculated
progress rates for the slow-moving species in the present
study. Reynolds indicated that the maximum progress
rates for golden perch were 5900-15 700m day"', which
is comparable with the observed progress rates of the
moderate- to fast-moving species in the present study.
These higher progress rates are still very much lower
than recorded for North American salmonids. For
example. Idler and Clemens (1959, in Weatherley 1972)
recorded a progress rate of 51 200 m day"' when following
the spawning movement of a wave of migrating sockeye
salmon, Oncorhynchus nerka, up the Frazer River, British
Columbia,
Sources and destinations of moving fish
The five species studied can be separated into two groups
based on longitudinal changes in their mean upstream
movement rates over the study period. Group A included
M. splendida and Ambassis spp,, the slow-moving species.
The decline in movement rate indicated the primary
source of these moving fish appeared to be the floodplain
downstream of site MX, and a secondary source existed
between site CA and PO, Group B included H. fuliginosus,
L. unicolor and A. percoides (the terapontids), the
moderate- to fast-moving species. The increase in move-
ment rate indicated the majority of fish passing site PO
appeared to arise diffusely from the creek upstream of
Mudginberri Billabong (more so for H. fuliginosus and
less so for/I, percoides).
The result for the terapontids (group B) ties in with
the notion that they are typically associated with upper-
reach escarpment areas and, in the Wet season, they
move downstream into the lowlands (Bishop et al. 1990),
The decreased upstream movement rates further down-
stream in the creek can be viewed as an indication of the
primary downstream distribution limit of these species.
Loss of moving fish
In comparison with group A species, only L, unicolor of
group B exhibited some loss of moving fish along the
creek. This occurred between the two most downstream
sites, and is likely to have been caused by some individuals
of this species refuging in Mudginberri Billabong over
the dry season. Being a major predator which uses
ambushing tactics on moving fish (Bishop & Forbes
1991), L. unicolor could be significantly advantaged by
such refuging, as it would be in a position to consume
pioneering migrants in the following early-Wet season.
The greatest losses of group A species (i,e. greatest
reduction in mean movement rates between sites)
occurred between the most downstream sites, the section
of the creek where slowest progress rates were recorded.
To explain this it is necessary to consider major mech-
anisms by which associations between reduced progress
rates and moving fish loss may be expected.
Mechanism Ll Ambushing by predators (see Bishop &
Forbes 1991). Moving fish may be consumed by predators
thus reducing numbers travelling. Their upstream pro-
gress rate would be also greatly slowed where many
ambush sites had to be negotiated.
Mechanism L2 Encountering sites where space is abun-
dant and water velocity gradients are poorly defined.
Moving fish either choose to occupy these sites, thereby
not continuing their journey, or their journey could be
delayed by the absence of well-defined water velocity
 F I S H M O V E M E N T D Y N A M I C S AND M O N I T O R I N G 103

gradients which would normally elicit a rheotaxic response
by the fish, keeping them moving upstream.
Mechanism L3 Encountering tributaries of the creek.
Moving fish either permanently enter the tributaries
thus reducing the number of fish moving up the creek, or
they may temporarily enter these tributaries thus slowing
the overall progress rate along the creek. Evidence that
fish moving up Magela Creek will be diverted into
backflow billabongs was found for M. splendida, L.
unicolor and Ambassis spp. in the present study.
The first two mechanisms are relevant to losses and
slowed progress rates of moving M. splendida and
Ambassis spp. between sites MX and MI. First, large
numbers of predators are typically a feature of corridor
waterbodies such as Mudginberri Billabong during the
Wet season (Bishop 1987). Second, Mudginberri
Billabong is characterized by an abundance of space and
less well-defined water velocity gradients (e.g. in com-
parison with the creek environment). An examination of
the daily pattern of movement of Af. splendida (Fig. 10)
lends support to the involvement of mechanisms Ll and
L2, in that the plot for site MI is more rounded (i.e. less
'spikey') than the other sites, which suggests a coalescing
of short-term pulses of moving fish. Such coalescing may
be expected when predators are encountered or when
large-volume sites gradually fill with moving fish and
then empty when a cue initiates en masse emigration. The
definition on these resultant medium-term pulses of
moving fish would be expected to be attenuated gradually
with increasing distance upstream.
Upstream of Mudginberri Billabong, in the creek
proper, mechanism L3 appears to be relevant for group
A species, as the section of the creek with the greatest
loss of moving fish (i.e. between sites MI and GD) has
the largest tributaries with the largest billabongs located
at their mouths. Correspondingly, the section of the
creek with the lowest loss of moving fish (i.e. between
sites GD and CA) has the smallest tributaries with the
smallest billabongs located at their mouths.
Information on the size of these tributaries' catchments
and their associated billabongs are given in Table 8. With
large catchments, and subsequently greater discharges
reaching the creek, there is a greater chance that fish
would be diverted from the creek to the tributaries.
Additionally, with larger catchment areas, there is a
greater chance that moving fish would leave the creek
permanently by progressing considerable distances up-
stream along the tributaries. With larger billabongs at
their mouths, there is also an increased chance that fish
will remain in the billabongs for longer periods (possibly
even permanently).
Rather than being considered as a disadvantage in
terms of slowing progress up the creek, and/or halting
progress for many individuals, the diversion of moving
fish from the creek to the billabongs may be a significant
advantage. First, food organisms (i.e. invertebrates) are
more abundant in the billabong habitats (Marchant 1982),
and so entry into this habitat would be beneficial as
sustenance could be obtained for continuing the upstream
journey. Second, diversion into billabongs and then into
tributaries enhances dispersal of the species as multiple
refuge colonies may be established and maintained.
The small loss of moving fish noted in the G D - C A
section of the creek can be attributed primarily to losses
occurring within the sandy creek habitat, as only very
small tributaries enter the creek here, and accordingly
the associated billabong habitat is limited (Table 8).
Mechanism Ll is likely to be important in the creek
habitat as large numbers of predators are typically
associated with this habitat in the Wet season, albeit less
so than in Mudginberri Billabong (Bishop 1987), and
ambushing of moving fish by predators is conspicuous
(Bishop & Forbes 1991). Another loss mechanism would
be related to mechanism L2 in terms of moving fish

Table 8. Features of tributaries and their billabongs which enter Magela Creek between the Ranger Uranium Mine and the floodplain

Tributary/billabong Estimated areas (km^) Ratio of billabong area

Section of creek name Catchment'' BiUabong' to catchment area

Within 1 km upstream of site PO Georgetown 23.6 0.19 0.008

Between sites CA and PO Djalkmara-Indium 1.5= 0.123 0.082
Coonjimba 1.2' 0.012 0.010
Between sites GD and CA Surshar 1.0 0.003 0.003
Between sites MI and GD Gulungul 90.7 0.29 0.003
Comdorl 92.8 0.76 0.008

"Estimated from 1:100000 topographic maps. ''All estimates are for the late-Wet season (Bishop et al. [1990] was used for Georgetown,
Djalkmara-Indium and Coonjimba Billabongs; Surshar Billabong was estimated to be one quaner the size of Coonjimba Billabong; a map in
Bishop & Walden (1991) was used to estimate Gulungul; Baker et al. [1984] was used for Corndorl [floodplain area excluded]). 'The values are
for catchments truncated by the construction by Ranger of retention ponds on the tributaries' upper reaches. Refer to the text for site codes.
104 K. A. B I S H O P ET AL.
colonizing unoccupied space in the sandy creek habitat.
Such space would be abundant as, at the beginning of
the Wet season, a new environment is opened up.
Gains of moving fisk
Evidence that group A species will enter the creek from
backflow billabongs was found in terms of three separate
mechanisms. The first two involved exiting by backflow
or crossflow channels (Gulungul and Djalkmara Billa-
bongs, respectively). The third was the use ofthe outflow
channel whereby the fish move down the channel and
upon reaching the confluence with the creek turn into
the creek and proceed upstream. This mechanism is of
considerable interest as it raises the possibility that
cognitive behaviour may be involved.
Extended observations in the outflow channel of
Gulungul Billabong in 1988 indicated that rapidly falling
base-flow water levels in Magela Creek are associated
with the commencement of emigration from the billabong
at the end ofthe Wet season. At this time the water level
in the billabong also falls rapidly and aquatic plant beds
collapse, thus greatly reducing the available space in the
system. Additionally, water quality would start to
deteriorate at this time and this may elicit an avoidance
response by the fish. It is recognized that the emigrants
observed could have been temporary immigrants from
the creek which could not proceed further into the
billabong because of restricted access (e.g. shallow waters
or collapsed aquatic plant beds). However, assuming the
observations adequately 'sampled' inputs and outputs
from the billabong, it is more likely that the moving fish
observed were primarily derived from billabongs as the
number of apparent immigrants at these times was always
much lower than the number of emigrants.
As indicated previously, it is apparent that the billa-
bongs can be independent sources of group A species
and, accordingly, the larger the billabong the greater will
be the input of billabong-derived moving fish to the
creek once an appropriate connection is made (i.e. as
permitted by sufficiently high water levels). This 'reser-
voir' of potential moving fish in the billabongs could be
drained if a large proportion of them travelled up the
tributary streams. Tributaries with larger catchment areas
and hence with larger discharges into the billabongs
would attract larger numbers of these moving fish and,
accordingly, the input to Magela Creek from such
billabongs would be lowered.
As a result, the relationship between billabong size
and the input of billabong-derived moving fish into the
creek, requires consideration ofthe size ofthe catchment
of the tributary on which the billabong is located. The
propensity for a billabong to provide an input of moving
fish to the creek can be expressed then in terms of the
ratio of the area of the billabong to the area of the
catchment of the tributary (the greater the ratio the
greater the propensity). These ratios are given for each
billabong/tributary in Table 8. It can be seen that the
Djalkmarra-Indium Billabong system has by far the
greatest ratio. It should be noted, however, that the
catchment area for this billabong complex, and that of
Coonjimba Billabong, has been greatly reduced since the
placement of retention ponds in the tributaries' upper
reaches by the Ranger Uranium Mine. Tying in well
with the hypothesis outlined previously, it is apparent
that the section of the creek which runs by the above-
mentioned billabongs is the only section in which a gain
in both of the group A species was apparent.
The above hypothesis could well be relevant to the
dynamics (including fate) of creek-derived moving fish
which are diverted into the billabong systems. For
example, with a larger ratio of billabong area to catchment
area, fewer moving fish will be diverted into the billa-
bongs, and similarly, fewer will progress up the tributary
streams. In other words, such billabongs and their
tributaries are less likely to 'absorb' creek-derived moving
fish and/or reduce upstream progress rates through
temporary colonization. These considerations may well
be complicated by the capacity of the billabongs them-
selves to absorb moving fish (positively related to billa-
bong size). However, competition for space between
immigrants and billabong-derived fish may buffer this
effect. Another complication may arise during very high
discharges if the billabongs become anabranches of the
creek, which is the case for many of the billabongs in the
study area. In such a situation, creek-derived discharge
leaving the billabongs would divert moving fish into the
billabongs, whereas normally the tributary discharge
would not. This effect may also be buffered as diverted
moving fish may readily emigrate through the area where
the creek is spilling into the billabong.
For each of the group B species, maximum and
minimum gains in movement rates occurred at differing
sections of the creek, making interpretations difficult.
The maximum gain for L. unicolor occurred in the
MIGD section. Significantly, this is the section which
receives Gulungul Creek, the creek which is known to
contain large populations of L. unicolor (Bishop unpubl.
data), and which therefore could be a major source of
immigrants. Relevant here is the observation that large
numbers of L. unicolor were observed leaving Gulungul
Billabong for Magela Creek during 1988 when base-flow
water levels were falling at their greatest rate. For A.
percoides, the maximum gain occurred within Mudgin-
berri Billabong. Sampling in this billabong has indicated
that large numbers of this species are present (Bishop et
al. 1990).
Origins offish passing site PO
In order to interpret impacts on movements near the
mine, it is necessary to examine the sources of group A
 F I S H M O V E M E N T D Y N A M I C S AND M O N I T O R I N G
species passing site PO in the t989 Wet season. The
component derived from the Coonjimba and Djalkmara-
Indium Billabong systems (the 'Bb component') can be
estimated as:
Bb component POmmr CAmmr + adjustment for
creek loss
where, POmmr is the mean movement rate at site PO,
and CAmmr is the mean movement rate at site CA.
It is assumed that an adjustment for net loss to the
billabong systems is urmeccessary because of the buffering
effect caused by the competition for space with billabong-
derived fish. The adjustment for net loss of moving fish
along the sandy creek habitat is assumed to be at the
same rate that occurred between sites GD and CA in
which there are no significant tributaries or billabongs.
This rate is given in Table 5 and when multiplied out for
the 3.5 km between sites CA and PO the values are
1610nh"' for M. splendida and 525h'' for Ambassis
spp.
The mean rates at sites PO and CA for M. splendida
were 9415 and 6901 n h ' respectively. Accordingly, using
the above expression and adjustment, the Bb component
was 4124nh^', which is equivalent to 44% of those
passing the site. FOT Ambassis spp., the mean rates at sites
PO and CA were 7219 and 10956, respectively. The Bb
component is then 4262 nh"' which is equivalent to 39%
of those passing site PO.
An imphcation of the substantial proportions of the
Bb component is that billabong-derived fish from the
Coonjimba and Djalkmara-Indium Billabong systems
play a significant role in providing recruits to upstream
dry-season refuges, a key ecological process for fish
within this seasonally flowing creek. It is notable that
these billabongs, being the closest downstream from the
Ranger Uranium Mine, are at greatest risk of degradation
arising from the accumulation of harmful substances
transported by water from the mine. Ranger presently
has a policy of releasing waste waters through the
Djalkmara-Indium Billabong system in an attempt to
effect some form of improvement to the quality of these
waters before they reach Magela Creek, a management
strategy considered problematic by the Commonwealth
Government's Office of the Supervising Scientist (OSS)
in 1989 (OSS 1989). Ranger plans to remove this billa-
bong system when they exploit their Ore Body No.3.
It would be difficult to obtain greater definition on the
origins of those fish passing site PO that do not arise
from the Coonjimba and Djalkmara-Indium Billabong
systems. The fundamental problem is to determine what
proportions of moving fish arise from the floodplain
downstream of site MX and the billabongs adjacent to
the creek. This problem is complicated by the possibility
that the ftoodplain-derived fish may temporarily enter
the biUabongs on their journey (i.e. they may have a
staged journey). The billabong-derived moving fish may
105
also have a staged journey, thus making it difficult to
determine which billabongs are important nursery areas
and, hence, which are potentially most sensitive to con-
taminants arising from waste-water releases from Ranger.
A first step in determining the relative contributions
would be to make movement counts at strategic sites in
the vicinity of the billabongs and at the head of the
floodplain. In such an investigation automatic counters
for moving fish (Hellawell 1978) could be used to
minimize labour costs. This could be followed up by the
examination of age structure and stomach contents of
samples of moving fish to obtain evidence of billabong or
creek origins.
CONCLUSIONS
The studies reported in this paper provide evidence to
support the view of Bishop and Forbes (1991) that
movements of fish observed in Magela Creek are indic-
ative of migratory activity. From a wider perspective,
they provide detailed new information on an Australian
tropical floodplain river, a region and a river type where
knowledge and concepts of movement dynamics are at an
early stage of development. At an apphed level the
studies give an initial validation for an efficient, direct-
observation technique to monitor these movements.
Outside South and Nonh America, direct-observation
techniques are seldom used to study fish movements,
and this probably variously reflects the lack of conducive
conditions, the small amount of time researchers spend
making straightforward observations and a certain rigidity
of thinking whereby researchers are not inclined to use
less-traditional techniques.
With the improved knowledge of movement dynamics,
particularly in relation to the sources and destinations of
moving fish, future monitoring of impacts from mining
in Magela Creek using this technique will be made
considerably more powerful as more detailed inter-
pretations of detected impacts will be possible. The
complexity of the dynamics between the creek and the
billabongs, a feature also of fish movements in the lower
reaches of South American rivers (Bonetto et al. 1981,
Quiros & Cuch 1986, both in Quiros 1989), makes it
necessary for more investigations in these areas.
Fish movements were selected for monitoring in
Magela Creek because their use was seen to be a unique
means of focusing on a survival strategy which was
directly significant to the structure and functioning of
the ecosystem, and also reflected conditions in a broad
area downstream of the mine. Unfortunately, monitoring
systems very rarely have these attributes, with the result
that the ecological significance of detected impacts is
usually exceedingly difficult to determine. Fish move-
ments have seldom been used for monitoring impacts,
even though they have been identified as valuable subjects
for impact assessment (Hellawell 1978; Nezdoliy 1984;
106 K. A. B I S H O P ET AL.
Petrere 1985). It is hoped that the development of
efficient, direct-observation techniques for monitoring
fish movements will facilitate the greater use of such
movements as the subject of monitoring.
Within Australia, monitoring of fish movements has
already been useful in detecting short-term avoidance
responses to the release of mine-site wastewaters (Bishop
& Walden 1990), and the long-term impact of river
regulation (Mallen-Cooper & Brand 1992). In North
America, studies on fish movements have demonstrated
avoidance responses to elevated levels of metals arising
from mining operations (Sprague et al. 1965; Saunders &
Sprague 1967).
ACKNOWLEDGEMENTS
The authors would like to thank the Northern Territory
Water and Sewage Division for data supplied on creek
flows. We thank Ms C, Camilleri for occasional assistance
in the field and Ms C. Taylor for editorial comments. For
discussions concerning studies on fish movements within
Australia we gratefully acknowledge Mr M, Mallen-
Cooper (New South Wales Fisheries), Mr R, Griffin
(Northern Territory Fisheries) and Mr A. Hogan (Queens-
land Fisheries),
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