Recovery of the endangered trout cod Maccullochella macquariensis: What have we achieved in more than 25 years?
ABSTRACT Recovery of threatened species is often necessarily a long-term process. The present paper details the progress towards the recovery of trout cod, Maccullochella macquariensis, an iconic, long-lived fish species first listed as threatened in the 1980s. The objectives, actions and progress over three successive national recovery plans (spanning 18 years) are assessed, documenting changes to population distribution and abundance and updating ecological knowledge. Increased knowledge (especially breeding biology and hatchery techniques, movements, habitats and genetics) has greatly influenced recovery actions and the use of a population model was developed to assist with management options and stocking regimes. Key recovery actions include stocking of hatchery-produced fish to establish new populations, regulations on angling (including closures), education (particularly identification from the closely related Murray cod, M. peelii) and habitat rehabilitation (especially re-instatement of structural woody habitats). In particular, the establishment of new populations using hatchery stocking has been a successful action. The importance of a coordinated long-term approach is emphasised and, although there is uncertainty in ongoing resourcing of the recovery program, much has been achieved and there is cautious optimism for the future of this species
- SourceAvailable from: Brett A. Ingram[Show abstract] [Hide abstract]
ABSTRACT: Freshwater catfish (Tandanus tandanus) is a popular freshwater angling species in rivers of the Murray-Darling Basin (MDB). In recent decades the species has, however, undergone substantial declines in abundance and distribution. In Victoria, freshwater catfish is listed as threatened under the Flora and Fauna Guarantee Act 1988, occurring in small isolated populations only. In 2011, Native Fish Australia (NFA), (Wimmera) Inc. received funds from the Department of Primary Industries’ Recreational Fishing Grants Program to help restore Victoria’s recreational fishery by captive breeding and re-stocking freshwater catfish. The aim of this study was to observe spawning trials on freshwater catfish to produce juveniles for restocking into Victorian waters. Spawning trials focused on use of Ovaprim® to induce ovulation and spermiation in fish that where hand-stripped, and Ovaplant® (Syndel, Canada) to enhance maturation and ovulation in fish allowed to spawn naturally. Nineteen females and 26 males were used in spawning trials conducted in late October 2013, when water temperatures approached 20oC. Tetrauronema, a new myxozoan parasite of freshwater catfish, was observed in a milt sample from one fish. The implications of the presence of this parasite in freshwater catfish is unknown. Most females (70%) injected with Ovaprim were induced to spawn. Broodstock implanted with Ovaplant and transferred to a 0.15 ha pond containing patches of gravel subsequently spawned naturally within a month of being implanted. Small amounts (0.1-1.0 mL) of milt, which was generally watery in consistency, was stripped from males. There was a slight improvement in both milt consistency and sperm activity 2 days after injection with hormones. Relative fecundity of stripped females was low (920-3,370 eggs/kg). Although fertilisation rates (65-100%) were generally high, hatch rates (0-42%) were low and a high proportion (10-20%) of larvae were deformed. These results suggest that not all oocytes in the ovary were mature and consequently, responded to the hormone treatment, and/or fish were not fully conditioned for spawning. Spawning trials are likely to have occurred at the very beginning of the spawning season and so fish may not have been fully mature, which may affected fecundity, gamete quality, hatch rates and larval quality. Fry were reared to fertilised earthen pond, and on 7th February 2014, 1,900 fingerlings were harvested, 1,400 of these were released into Moodemere Lake (Rutherglen) and 500 into Crusoe Reservoir (Bendigo). Observations made during these trials will be used to guide future R&D to improve captive breeding of freshwater catfish. Recommendations include: • Increase broodstock numbers to increase productivity • Reduce broodstock densities in ponds to optimise growth, conditioning and maturation • Supplement the diet of broodstock with live prey (small yabbies) • Remove the fish infected with Tetrauronema to reduce the risk of spreading the disease to other stock • Check for the presence of this parasite in other broodstock during the next spawning season • Undertake induced spawning later in the season, once water temperatures exceed 21oC and before reach 25oC (i.e. early to mid-November) • Inject females and males to be hand-stripped each with a single doses of Ovaprim at 0.5 ml/kg • Repeat natural spawning trials to obtain further information on production levels for comparison with to induced spawning and hand-stripping methods • Rear fry into a pond by themselves using standard fry rearing methods employed for other native fish species.
Technical Report: Managing genetic viability of freshwater catfish in Victoria[Show abstract] [Hide abstract]
ABSTRACT: The freshwater catfish (Tandanus tandanus) is endemic to Australia, and can be found in slow moving or still waters throughout Victoria, New South Wales, South Australia and Queensland. The species was once widespread throughout the Murray-Darling River system, but their range has been altered dramatically since European settlement. Native Fish Australia, (Wimmera) Inc received funds from the Recreational Fishing Grants Program to restore Victoria’s recreational fishery by captive breeding and re-stocking freshwater catfish. To support of this project, this study was undertaken to: Describe the genetic structure of the freshwater catfish populations in Victoria Provide guidelines for genetic management of a freshwater catfish captive breeding program. Using DNA from 218 individuals, this study examined levels of genetic diversity, expected heterozygosity, effective population size and genetic structuring of the populations, using both mitochondrial and microsatellite data. Both mitochondrial and microsatellite data suggest that the Mallee, a naturally occurring population, is the most diverse population in Victoria. In contrast, the Avoca and Goulburn populations contained the least amount of diversity. The Loddon and Avoca sites were the only populations to contain no unique alleles. The Wimmera contained the highest effective population size (Ne) and the Loddon the lowest. Four genetic clusters were identified in the Victorian populations, which were Zone 1. Mallee, Wimmera, Little Murray, Gum Lagoon, Safe Lagoon Zone 2. Loddon and Avoca (Amphitheatre Res. and Centenary Res.) Zone 3. Phyland Lagoon and Turners Lagoon Zone 4. Goulburn (Lake Nagambie, Majors Creek and Tahbilk Lagoon). The genetic structuring observed in Victoria may be due to limited gene flow between populations along with influences of translocations. Results from the genetic analyses were used to develop guidelines for the genetic management of freshwater catfish broodstock used for stock-enhancement. The following points were recommended: 1. Avoid mixing fish between the four genetic zones identified in Victoria. 2. Use wild-born fish from Zone 1 and Zone 3 populations as broodstock. 3. Tag all broodstock for identification purposes. 4. Develop artificial spawning methods to improve genetic management of stock. 5. Spawn as many broodstock as possible each year. As a minimum spawn at least 10 fish (5 females and 5 males) each year. 6. Maintain as many broodstock as possible. 7. Spawn an equal number of female and male fish each year. 8. Undertake single-pair (one female and one male) matings only. 9. Mate different individuals each year (do repeat mating crosses that were undertaken in previous years). 10. Replace 10% of broodstock each year with new stock. 11. Maintain detailed and accurate breeding records. 12. Undertake stockings in accordance with the policies and protocols of Fisheries Victoria. 13. Spread the target number of fish to be stocked into each site over 4-5 consecutive years. 14. Mark (tag) all fingerlings before release. Following these recommendations will increase genetic diversity of the stocked fish while minimising loss in genetic diversity or changes in genetic structure of existing wild populations.
Recovery of the endangered trout cod,
Maccullochella macquariensis: what have
we achieved in more than 25 years?
John D. KoehnA,F, Mark LintermansB, Jarod P. LyonA, Brett A. IngramD,
Dean M. GilliganC, Charles R. ToddAand John W. DouglasE
AFreshwater Ecology, Arthur Rylah Institute for Environmental Research, Department of
Environment and Primary Industries, 123 Brown Street, Heidelberg, Vic. 3084, Australia.
BInstitute for Applied Ecology, University of Canberra, Canberra, ACT 2601, Australia.
CFisheries NSW, Department of Trade & Investment, Batemans Bay Fisheries Office,
PO Box 17, Batemans Bay, NSW 2536, Australia.
DDepartment of Environment and Primary Industries, Snobs Creek hatchery, Private Bag 20,
Alexandra, Vic. 3714, Australia.
EDepartment of Environment and Primary Industries, Fisheries, 5 Binns McCraes Road,
Alexandra, Vic. 3714, Australia.
FCorresponding author. Email: email@example.com
towards the recovery of trout cod, Maccullochella macquariensis, an iconic, long-lived fish species first listed as
threatened in the 1980s. The objectives, actions and progress over three successive national recovery plans (spanning
18 years) are assessed, documenting changes to population distribution and abundance and updating ecological
knowledge. Increased knowledge (especially breeding biology and hatchery techniques, movements, habitats and
genetics) has greatly influenced recovery actions and the use of a population model was developed to assist with
management options and stocking regimes. Key recovery actions include stocking of hatchery-produced fish to establish
new populations, regulations on angling (including closures), education (particularly identification from the closely
related Murray cod, M. peelii) and habitat rehabilitation (especially re-instatement of structural woody habitats). In
coordinated long-term approach is emphasised and, although there is uncertainty in ongoing resourcing of the recovery
program, much has been achieved and there is cautious optimism for the future of this species.
Additional keywords: Australia, conservation, endangered species, freshwater fish, recovery, rehabilitation.
Received 17 September 2012, accepted 4 June 2013, published online 6 September 2013
Freshwater fishes are imperilled world-wide (Sala et al. 2000),
with 30–60% of species considered threatened (e.g. Smith and
Darwall 2006; Jelks et al. 2008). Riverine fishes are susceptible
to a range of anthropogenic threats both directly and to their
habitats (Malmqvist and Rundle 2002; Dudgeon et al. 2006;
Cooke et al. 2012). Large, long-lived species can be especially
susceptible to exploitation (Hogan et al. 2004; Dudgeon et al.
no different; being subjected to a similar range of threats
(Cadwallader 1978; Lintermans 2013a) with serious population
declines as a result (Wager and Jackson 1993). The Murray–
Darling Basin (MDB) in south-eastern Australia (Fig. 1), for
example, has a naturally depauperate native fish fauna consist-
ing of only 44 species, 24 of which are considered threatened at
2004; Koehn and Lintermans 2012). Trout cod, Maccullochella
macquariensis, along with three closely related freshwater ‘cod’
species (genus Maccullochella: eastern freshwater cod M. ikei,
Mary River cod M. marienis and Murray cod M. peelii), are all
listed as threatened nationally (Lintermans et al. 2005).
Trout cod is a large, iconic Australian freshwater Per-
cichthyid fish, endemic to the rivers of the MDB. This species
has suffered major declines in range and
(Cadwallader and Gooley 1984; Douglas et al. 1994; Fig. 1)
and only one ‘natural’ population (remnant from pre-European
settlement) remains, although this is now accompanied by
several populations re-established via translocation and
Marine and Freshwater Research, 2013, 64, 822–837
Journal compilation ? CSIRO 2013www.publish.csiro.au/journals/mfr
stocking. Trout cod is listed as endangered under both national
accessed 22 July 2013) and State legislation (New South Wales
Department of Primary Industries 2006; ACT Government
2007; Department of Sustainability and Environment 2007;
Hammer et al. 2009). Prior to the first formal conservation
listing in 1980 (Burbidge and Jenkins 1984), concerns had been
At this time, there were no legislative obligations for recovery
plans or actions. A national recovery plan (the first for an
Australian fish) was published following its national conserva-
tion listing (Douglas et al. 1994). This recovery plan has now
been through three iterations over an 18-year period (Douglas
et al. 1994; Brown et al. 1998; Trout cod Recovery Team
2008a). Even though trout cod is a nationally listed species,
and other threatened species is largely undertaken by individual
The present paper (1) summarises recovery actions for
trout cod since its taxonomic description and subsequent
listing as a threatened species 40 years ago, with a focus on
the past 25 years, (2) provides an update on biology, ecology,
distribution and threats, (3) reviews the progression of three
consecutive recovery plans, assessing the success of on-ground
recovery actions, (4) illustrates actions with regional case
studies and (5) provides an assessment of the future for
1972), it had been suspected that this species was a taxon sep-
(see Cadwallader 1977). Lake (1971) had pre-empted this
description and, indeed, had considered the species as threat-
ened. Even after its first conservation listing in 1980, there was
limited understanding of trout cod biology, ecology, distribu-
tion, population status or threats and its biology was often
assumed to be ‘similar to’ that of the closely related Murray cod
(Koehn and O’Connor 1990).
Species description and ecology
recorded to 850mm in total length and 16kg,but is nowusually
preference for in-stream woody structure (Koehn and Nicol
1998; Growns et al. 2004; Nicol et al. 2004). Adults exhibit
high site fidelity, generally undertaking limited movements but
with occasional large-scale excursions (Koehn et al. 2008;
Ebner and Thiem 2009). The diet is dominated by shrimps and
crayfishes (75%) but includes other aquatic and terrestrial
macro-invertebrates and some fishes (Baumgartner 2005,
2007). Maximum life-span is thought to be 20–25 years (Todd
et al. 2004), with sexual maturity reached at 3–5 years of age.
Spawning occurs in spring at a water temperature of ,158C
(Koehn and Harrington 2006). Females produce 1200–11000
1 – Seven Creeks
Recent oral history records
Recent oral history records
2 – Ovens River
3 – Goulburn River
4 – Lakes Sambell and Kerferd
5 – Dartmouth Dam
6 – Lake Nagambie
7 – Burrinjuck Dam
8 – Cataract Dam
9 – Burrendong Dam
10 – Yarrawonga Weir
Historic and recent distribution of trout cod in Australia. (a) 1990, (b) 2012.
Recovery of the endangered trout cod
Marine and Freshwater Research
relatively large (2.5–3.6-mm diameter) adhesive eggs that are
attached to hard substrates and tended to and guarded by the
male. Larvae hatch after 5–10 days at 6–9mm, then disperse by
drifting in the water column (Koehn and Harrington 2006).
a sport fish (Berra 1974; Cadwallader 1977) and, as a top-order
predator, could be viewed as both a ‘keystone’ (Paine 1966,
1969) and ‘flagship’ species (Simberloff 1998).
Distribution and abundance
Because of the early taxonomic confusionand misidentification
Macquarie River (Berra and Weatherley 1972; Cadwallader
1977; Douglas et al. 1994; Fig. 1). Recently, historical records
documented for the Lachlan River (Trueman 2011). Trout cod
has been translocated into several waters, including Cataract
Dam (Nepean Catchment, New South Wales) before 1910
(Douglas et al. 1994), Seven Creeks (Goulburn Catchment,
Victoria) in the early 1920s and Lake Sambell (Ovens Catch-
ment, Victoria) in 1928 (Cadwallader and Gooley 1984; Fig. 1).
The species underwent an extensive decline in range and
abundance, apparently within only a few decades (Cadwallader
and Gooley 1984). Until at least 1950, trout cod was present in
the Murray River, as far downstream as Mildura (Lake 1971;
Cadwallader 1977), although they were considered rare down-
stream from Echuca (Cadwallader 1977). By the late 1970s, the
only remaining potentially sustainable breeding populations of
trout cod were the naturally occurring population in the Murray
River (New South Wales); occupying the reach from Yarra-
wonga Weir downstream to Cobram (Cadwallader and Gooley
1984; Ingram et al. 1990; Douglas et al. 1994) and the translo-
cated population in Seven Creeks (Cadwallader 1979; Morison
and Anderson 1987; Richardson and Ingram 1989). The trans-
located population in Lake Sambell apparently died outin a fish
kill in 1970 (Cadwallader and Gooley 1984) and the trout cod
and Murray cod translocated into Cataract Dam are known to
hybridise (Wajon 1983; Harris and Dixon 1988). The last
reported trout cod in the lower Murrumbidgee was collected at
Narrandera (Fig. 1) in 1969 (Gilligan 2005). The Australian
Capital Territory (ACT) population of trout cod in the upper
Murrumbidgee River disappearedonlyin the 1970s,(Lintermans
et al. 1988), as did the population in the Mitta Mitta River
following the construction of Dartmouth Dam, which was
completed in 1979 (Koehn et al. 1995; Ryan and Koehn 2001).
Recovery plans and actions
Following the brief description of recovery actions for trout cod
and many Australian freshwater fishes (Wager and Jackson
1993), there have been three consecutive national recovery
plans and four state/territory recovery plans (Douglas et al.
1994; Brown et al. 1998; ACT Government 1999, 2007;
Department of Sustainability and Environment 2003; New
South Wales Department of Primary Industries 2006; Trout cod
Recovery Team 2008a) dedicated to trout cod, spanning 18
years (Table 1). Some recovery actions have been underway
longer than this, but most concerted efforts have occurred over
the past 25 years. There have, however, been significant time
lags between recovery plan development, recognition and
was finalised in draft form in 2004.
Trout cod is subject to a range of threats that are often cited
as being responsible for the declines of many Australian native
fish species (Cadwallader 1978; Ingram and Douglas 1995;
Lintermans et al. 2005; Table 2). Recovery actions for trout
ing biological-knowledge requirements, habitat restoration,
of status and legislation. Some actions in the three national
recovery plans overlap in time with some actions continuing
across plans (e.g. recovery teams, monitoring; see Table S1,
available as Supplementary Material for this paper). Estimates
of the cost of implementing individual plans (undertaken at the
time of their preparation) are similar and reflect the on-going
nature of recovery actions. The current plan, however, has been
subject to more disparate funding opportunities, handled on an
individual state, catchment and site basis with no national funds
provided for implementation of the current recovery plan.
A summary of national recovery plan actions and review of
progress is given in Table 1 (see Table S1, available as
Supplementary Material for this paper), along with timelines
for significant ecological and management actions (Fig. 2).
Overall, a substantial number of actions have been completed
or partially completed (74% and 85% for Plans 1 and 2,
Table 1.Timeframes, objectives, actions, percentage actions completed and costs of the three national recovery plans for trout cod
C¼mostly (.80%) completed, N¼not (,20%) completed, PC¼partially completed (20–80%)
Attribute Recovery plan 1Recovery plan 2Recovery plan 3
Number of objectives
Number of actions
Douglas et al. (1994)
Many objectives, probably
Brown et al. (1998)
Trout cod Recovery Team (2008a)
Many smaller more site-specific actions;
lack of national funding
Marine and Freshwater Research
J. D. Koehn et al.
Formally recognised as a separate species
Last specimens reported from Upper Murray River
Considered seriously threatened
Seven Creeks closure
Last specimens reported from Murrumbidgee River
Disappear from Mitta Mitta River following construction
of Lake Dartmouth
Last specimens reported from the Macquarie River
Translocation of fish upstream in Seven Creeks and to a
dam in the Campaspe catchment
Seven Creeks closure extended
Hatchery techniques developed
Captive breeding and stocking programs commenceFive year ban on angling in the Trout Cod Protection
Area (Murray River between Yarrawonga and Cobram)
Translocation of fish upstream of bushfire affected
area in Seven Creeks
Flora and Fauna Guarantee Act Listing (Victoria)
Trout cod protected in NSW
First national recovery plan
Fishing re-opens in the Trout Cod Protection Area, but
only from Dec to Aug (take of trout cod remains
Development of population model
Change in stocking strategy
Trout cod protected in ACT and first ACT recovery plan
Trout Cod Protection Area extended downstream to
Seven Creeks closure extended
Second national recovery plan
EPBC Act national conservation listing (endangered)
First evidence of natural recruitment in the re-
established lower Murrumbidgee population.
Angling closure for 9 km of upper Murrumbidgee River
Third national recovery plan
Confirmation of historical records from the Lachlan
River (present up to at least c1968).
First NSW recovery plan
Second ACT recovery plan
Timeline of important events for the conservation of trout cod.
Table 2.Description of threats relevant to trout cod
Removal of woody habitatsRemoval of in-stream wood is widespread especially
mid-river, close to the deep, fast-flowing water
(preferred trout cod habitat).
Sand sediments in Seven Creeks and post-fire inputs.
As a result of poor water quality, including blackwater events.
Koehn and Nicol (1998), Growns et al. (2004),
Koehn et al. (2004a), Nicol et al. (2004)
Alteration of flow regimes
Saddlier et al. (2002), Lyon and O’Connor (2008)
Koster et al. (2004), King et al. (2012)
Reduced flooding and altered seasonality in the Murray River;
extraction from Seven Creeks.
Caused loss of the trout cod population in Mitta Mitta River.
Movements of adults appear limited but subadults exhibit
Risks to safe fish passage at Yarrawonga Weir.
Loss of drifting larvae into irrigation channels, through
pumps and passing over weirs.
Close (1990), Close (2002)
Cold water dam releases
Koehn et al. (1995), Todd et al. (2005)
Koehn et al. (2008), Ebner and Thiem (2009)
Thorncraft and Harris (1997), Stuart et al. (2010)
Koehn and Harrington (2006), Koehn et al. (2004b),
King and O’Connor (2007), Baumgartner et al.
Removal by fishing
Brown trout, redfin perch.
Impacts of carp, Cyprinus carpio, are unknown.
An aggressive predator susceptible to angling; captured
by commercial fishing operations in the past, some angler
take (either illegally or through misidentification).
Injury and post-release mortality of concern
(2–15% mortality estimated for Murray cod).
Cadwallader (1996), McDowall (2006)
Koehn et al. (2000)
Berra (1974), Rowland (1989), Trout cod Recovery
Team (2008a), Lintermans (2007)
Muoneke and Childress (1994), Bartholomew
and Bohnsack (2005), Douglas et al. (2010),
Hall et al. (2012)
Recovery of the endangered trout cod
Marine and Freshwater Research
respectively). In many cases, partial completion has meant that
the action has been undertaken only at some sites and a lack of
sampling consistency among sites has led to difficulties in
determining population status. Fewer actions have been com-
pleted for the current plan (only 15%), which is current until
been less certainty of funding. In particular, the lack of comple-
tion of communication actions is very evident.
Ecology and knowledge to inform recovery
Recovery plans have been supported by compendiums of
knowledge (Douglas et al. 1995; Trout cod Recovery Team
2008b) to inform particular recovery actions. The generation of
available as Supplementary Material for this paper) and,
importantly, new knowledge and ‘grey literature’ have been
incorporated. A search of the Scopus database using ‘trout cod’
or ‘Maccullochella macquariensis’ in the title or as a keyword
revealed a total of 29 publications, with the number of pub-
lications increasing substantially over the decades since trout
cod was recognised as a distinct species (Berra and Weatherley
1990s, and 18 in the 2000s.
The recognition of trout cod as distinct from Murray cod
highlighted its restricted distribution, which subsequently
7km of Seven Creeks were closed to angling and in 1993 this
was extended a further 4km downstream (Barnham 1995). In
area; TCPA) to protect the remnant wild population. In 1992,
the recreational Murray cod fishery within the TCPA was
re-opened, but with an ongoing annual ‘spawning’ seasonal
boundary of the TCPA was extended a further 33km in con-
sideration of the new spatial extent of the trout cod population.
cod state-wide, and New South Wales enacted similar restric-
in 1996 and, in 2000, a complete fishing closure was imposed
along 9km of the Murrumbidgee River, downstream of the
Angle Crossing stocking site (ACT Government 2007).
In an attempt to establish new trout cod populations, captive
breeding programs were established at both New South Wales
and Victorian government hatcheries (Narrandera and Snobs
Creek, respectively) (Ingram et al. 1990). Techniques to induce
(Rimmer 1987; Ingram and Rimmer 1992), and since then,
juvenile fish have been produced for release within the pre-
much of the information regarding the reproduction and early
life history of trout cod. Captive breeding methods have been
described by Ingram and Rimmer (1992), Douglas et al. (1994)
and Ingram (2009). Wild-caught broodstock have been sourced
mainly from the Murray River population. Fish are held in
earthen ponds but unlike other Maccullochella spp., trout cod
does not readily spawn unassisted in captivity. As a conse-
quence, during September–October (with water temperatures
rising to ,168C), mature broodstock are removed from the
ponds and injected with gonadotrophin after which eggs (up to
5600eggskg?1; mean 3300eggskg?1) and sperm are manually
stripped. The eggs are adhesive, 2.7–3.5mm in diameter and
take 5–10 days to hatch at 18–208C. Hatch success rates vary
from 0% to 98% (mean 50%) and the larvae (6.0–8.8mm in
length) begin exogenous feeding after 10 days. Following
commencement of feeding, fry are stocked into fertilised
earthen ponds for on-growing. Harvested fingerlings are then
held for a short period in the hatchery and in recent years, have
been chemically marked (oxytetracycline or calcein) before
release to discriminate stocked fish from those naturally
The first stocking of trout cod fingerlings occurred in 1986/
1987, with 1000 fish released into the upper Murray River
(Fig. 1). Since then, a total of 1.56 million trout cod individuals
has been released into 17 areas (32 sites) across eight river
catchments (Table 3). Most fish (.99%) have been released as
fingerlings (0.5–1.5g), although a small number (11430) were
released as yearlings (10–50g). As hatchery production
increased through the early 1990s, more fish became available,
resulting in higher numbers stocked at more sites (Table 3;
Douglas et al. 1994). A range of criteria was developed for site
selection (Douglas et al. 1994) with an early preference for
pristine habitats,withoutMurraycod (toavoidhybridisation) or
redfin perch (Perca fluviatilis) (to reduce predation), relative
isolation from population centres (to reduce angling pressure)
but accessible for monitoring (Douglas et al. 1994). These
factors, combined with the example of a successful historical
translocation to Seven Creeks, initially led to the stocking of
many smaller, upland streams (Table 3). Following a lack of
apparent long-term success at these sites, there was a change in
stocking philosophy from small creeks and few fish to larger
waters and many fish over the longer term (Table 3). These
subsequent stocking regimes for trout cod have been more
successful, with natural recruitment observed or inferred in
at least six of the areas. What might be described as ‘self-
sustaining’ populations now occur in the mid-Murrumbidgee
(Gilligan 2005; Ingram and Thurstan 2008), the lower Ovens
been variable and the time between initiating stocking and
observing recruitment has been 5–13 years (Lyon et al. 2012;
M. Lintermans, pers. comm.).
Currently, there are no genetically discrete populations of trout
cod (Moore et al. 2010). Stocking programs have often been
diversity or reduced viability (Allendorf 1991; Philipp et al.
1993). Using mitochondrial DNA, Bearlin and Tikel (2003)
Marine and Freshwater Research
J. D. Koehn et al.
Stocking-site details for trout cod
(incl. Ryans Ck)
Recruitment recorded in Loombah Weir (1998),
but no monitoring since then
Recent (2012) anecdotal report
(below L. Nagambie)
Comment on recent kill?
Upper Lachlan R.
(below Burrendong Dam)
Small size classes reported by anglers –
but none confirmed.
Upper Macquarie R.
(above Burrendong Dam)
No recruitment observed from initial stockings.
Too early to tell for most recent stockings.
Burrumbuttock Farm dams
Refuge population created at beginning
of conservation program. The population did not survive.
Cotter R. (Bendora Dam)
Recruitment observed in Bendora Dam
(below Burrinjuck Dam)
Tumut R. (Talbingo Res.)
Upper Murrumbidgee R.
(above Burrinjuck Dam)
2011 capture of potential natural recruit
at a single site
Upper Ovens R. (Buffalo Ck,
Buffalo R. and Rose R.)
Lower Ovens RiverA
Upper Murray R.
(above Hume Dam, incl.
Upper Mitta Mitta R.
(above L. Darmouth)
AIncludes some yearlings.
Recovery of the endangered trout cod
Marine and Freshwater Research
found the Murray River population (11 haplotypes) to be sig-
nificantly different from both the translocated Seven Creeks
population (2 haplotypes) and the stocked population in the
Ovens and King Rivers (6 haplotypes). A recent microsatellite-
marker study showed that although there were more alleles
detected in the stocked population of the Ovens River (total 55)
is likely to be the result of successful application of breeding-
program protocols, which were established early in the stocking
program and aimed to maximise the genetic diversity of the fish
produced. These included regular replacement of broodstock,
maintaining a sex ratio of 1:1, undertaking single-pair matings,
avoiding repeat matings of the same pairs of fish, and mixing
progeny fromall matingstogether beforerelease (Douglaset al.
1994). A low level of natural hybridisation between trout cod
and Murray cod is observed in the Murray River, which has not
appeared to affect the genetic status of either species (Douglas
et al. 1995).
The use of a population model
A stochastic population model for trout cod was developed to
assess the viability of the Murray River population and the
importance of management strategies, as well as to guide future
research and management actions (Todd et al. 2004). Model
development used input from key stakeholders to garner a
consensus (Burgman and Possingham 2000) and examined a
range of factors, including the sensitivity of management deci-
sions for trout cod, model structure, statistical distributions of
uncertainties, parameter values and the relative importance of
identified knowledge gaps. The model was influential in
changing the process and design of hatchery production and
stocking regimes (Lyon et al. 2012).
of the species. Important education issues for trout cod conser-
vation include the need to assist anglers to distinguish between
trout cod and the closely related Murray cod (Douglas et al.
1994), methods to release any trout cod by-catch to minimise
activities targeted at increasing public awareness, including
advisory signs along the trout cod management zone, infor-
identity-crisis, accessed 22 July 2013), magazine and internet
articles, information pamphlets, interpretation of scientific
results for the public and field-day displays including live fish,
trout cod information, and videos. Angler and other interest
groups have produced various educational websites, brochures
and trout cod promotional information (e.g. Native Fish
Australia; www.nativefish.asn.au, accessed 22 July 2013).
realised but has much potential.
The restoration of degraded aquatic habitats is a key driving
action in the recovery of many threatened fishes (Barrett 2004;
Murray–Darling Basin Commission 2004; Nagayama and
Nakamura 2010). Trout cod is strongly associated with struc-
tural woody habitats (Koehn and Nicol 1998; Growns et al.
2004; Nicol et al. 2007), and habitat restoration is seen as a key
recovery action. Techniques for restoring large woody habitats
have been developed and trialled in the trout cod reach of the
Murray River (Nicol et al. 2002, 2004). These trials found that
all species of large-bodied fish (Murray cod, golden perch,
Macquaria ambigua, and trout cod) utilised restored habitats.
Subsequent habitat-rehabilitation works have been undertaken
in the Murray River upstream of Yarrawonga weir (Lake
Mulwala), the Ovens River near Wangaratta, Tarcutta Creek
near Wagga Wagga and Wodonga Creek. In response to sedi-
ment (sand) accumulation in the upper Murrumbidgee River,
installation of rock groynes resulted in increased hydraulic and
habitat diversity by creating scour pools (Lintermans 2005). In
Seven Creeks, a 3-km reach at the lower extent of the trout-cod
range was restored with groynes, rock, structural wood and
riparian revegetation (Saddlier et al. 2002). Trout cod has been
recorded utilising these new habitats in a variety of locations
(Lintermans 2005; Stoessel 2008).
The Murray River trout cod population (Fig. 1) is vital to any
national conservation effort. Concern had been expressed
regarding the viability of this population and surveys were
et al. 1994). Data from population surveys (undertaken since
1994) has been used in the population model (Todd et al. 2004)
considered relatively stable (Fig. 3). The population has
expanded downstream to Torrumbarry, and is now common to
Barmah (Douglas et al. 2012). Despite the installation of a fish
lift in October 1999, upstream expansion of the population
remains limited by Yarrawonga Weir. Only a relatively small
number of trout cod individuals (254 to February 2012:
Goulburn–Murray Water, unpubl. data) has negotiated this
barrier (2% of fish passing though the fish lift), compared with
trout cod comprising 30–69% of the large-bodied fish popula-
tion in the river downstream (J. Lyon, unpubl. data) during that
time. The lift was designed for the passage of Murray cod and
other native species, with little consideration given to trout cod
and there is the potential for injury during fish passage (Stuart
et al. 2010).
Historically, trout cod occurred throughout the entire length of
the lower Murrumbidgee River and ,200km of the upper
being last recorded from the river in 1976. In an attempt to
restore trout cod populations,both theupperand mid-reaches of
the Murrumbidgee River have been stocked. In excess of
326000 trout cod fingerlings have been stocked across eight
sites in the upper Murrumbidgee River since 1988 (average
40775, range 18000–99500 fingerlings; Table 3). Regular
monitoring at some sites has demonstrated survival and growth
of stocked individuals, but no recruitment or establishment of
Marine and Freshwater Research
J. D. Koehn et al.
self-sustaining populations has occurred. Detection of stocked
individuals .3 years old at riverine sites has been unsuccessful,
except at one site where 3þ-year-old trout cod individuals are
being regularly recorded (ACT Government, unpubl. data).
Angle Crossing received the greatest stocking effort (99500
fingerlings over 9 years to 2005) and the collection of a single
small fish (,190mm TL) in 2011 (.5 years after stocking
has occurred (ACT Government, unpubl. data). However,
stocking of 8740 fingerlings over 2 years in Bendora Reservoir
resulted in recruitment from stocked fish more than a decade
after stocking (ACT Government 2007; Lintermans 2007) and
these recruits now dominate the population.
Trout cod was reintroduced into the mid-Murrumbidgee
downstream of Burrinjuck Dam (Fig. 1) between 1992 and
2001 (320800 fingerlings at six locations; Table 3). An average
of 11881 (range 2500–20000) fingerlings were stocked annu-
ally for three to seven consecutive years (Table 3). Population
monitoring suggests that since 2001, the average electrofishing
catch per unit effort (CPUE) has remained stable (from
0.13?0.04 in 2001 to 0.12?0.07 in 2012) and is similar to
that of the remnant population in the Murray River
(0.16?0.04). The re-introduced population has expanded at
least 81km downstream of the lowest release site and trout cod
individuals have been collected at 77% of sites sampled. The
in eight of the past nine years indicates natural recruitment. The
proportion of recruits over the past 5 years (23%) has been
similar to that observed during the same period in the Murray
River population (19%).
Trout cod became extinct in the lower reaches of the Ovens
River (Fig. 1) by the early 1980s. A 10-year stocking program
commenced in 1997 with .277000 fingerlings of trout cod
stocked in a 15-km reach of river downstream of Wangaratta
until 2007, after which more targeted surveys were undertaken.
cohorts, and although CPUE data are variable, the population
has expanded at least 50km upstream (in the Ovens, King and
Buffalo Rivers) and ,100km downstream to the Murray River.
as far as Wodonga (,200km from the original stocking sites)
and in the lower reaches of the Kiewa River (J. Lyon, pers.
The lower Goulburn River (downstream from Lake Nagambie)
was stocked with .58000 trout cod individuals between 1993
and 1997 (Table 3). Drifting trout cod larvae and some 1-year-
old juveniles were collected in November and December 2003
(Koster et al. 2009), indicating successful breeding. In January
2004, a fish kill involving many hundreds of fish, including at
least 20 trout cod individuals, was recorded. Subsequent annual
surveys from 2004 to 2007 failed to detect any trout cod, but
small numbers of adults have been recorded every year from
2007 to 2011, with larvae being collected in 2007/2008 and
2008/2009 (W. Koster, unpubl. data).
Seven Creeks is a small (5–7m wide) stream with rock, gravel
and sand substrates, and pools ,2m deep interspersed with
rapids and cascades. Some cascades, together with the larger
Goorum Falls, are natural barriers to upstream fish movement.
Population surveys conducted on an ad hoc basis (e.g. Morison
and Anderson 1987; Anderson 1991; Saddlier and Harrington
1997; Kearns et al. 2012) have indicated downstream dis-
placement of the trout cod population over time. To prevent the
main population being relocated into less favourable habitats in
the lower reaches, trout cod individuals have been collected
using electrofishing and relocated back upstream. In 1991, fire
affected the Seven Creeks Catchment and over 100 trout cod
individuals were relocated upstream before rains washed ash
into the stream (Anderson 1991). Despite the persistence of a
self-sustaining trout cod population, Seven Creeks remains
vulnerable to many threats. The populations restricted distri-
bution and small size makes it vulnerable to stochastic events
Trout cod per electrofishing second?1
Yarrawongaweir and Tocumwal with trend line (N¼9092). Note that 2011had high flows and reduced capture rates.
Trout cod catch per unit effort (CPUE) for fish over 200-mm length from the Murray River between
Recovery of the endangered trout cod
Marine and Freshwater Research
(Cyprinus carpio), and more particularly, the predatory redfin
suggests that urgent action is required (Kearns et al. 2012).
Historically, trout cod has occurred throughout the upper
reaches of the Macquarie River Catchment (type specimen
collected near Bathurst in 1824), but it was thought to have
have been stocked upstream (17900 at four sites in 1991–1993
and 92000 at 10 sites in 2009–2011) and downstream of
Burrendong Dam (232000 fingerlings in 1998–2004; Table 3).
Opportunistic scientific data and reports from recreational
fishers indicate that trout cod survives in both the upper and
lower catchments; however, there has been no dedicated mon-
itoring undertaken in the Macquarie to establish whether or not
self-sustaining populations have established.
The New South Wales Rivers Survey (Harris and Gehrke 1997)
sampled 27 sites within the likely historical range of trout cod
(1994–1996) and captured no individuals. From 2004 to 2012,
extensive sampling across the MDB by the Sustainable Rivers
wild population: 20); Upper Murray: 1; Kiewa: 1) (Davies et al.
2008; Murray–Darling Basin Authority, unpubl. data). The fish
fish, rather than evidence of self-sustaining populations. There
are many stocking locations (particularly smaller sites) where
the establishment of populations has not been successful,
although in some areas, small numbers of individuals have been
observed (e.g. Douglas and Brown 2000). This has included
major stocking sites such as the upper Murray River (.68000
fingerlings released over 8 years), the upper Mitta Mitta River
(above Lake Dartmouth) and a range of lesser sites, including
Talbingo Dam (Tumut River), Ryans, Koetong, Buffalo and
Rose rivers (Table 3).
Cataract Dam is believed to have been stocked with translo-
cated trout cod and Murray cod in 1914, and trout cod exhibits
high levels (32–50% of the population) of hybridisation with
Murray cod (Wajon 1983). The future of 1600 fingerlings
stocked into lakes Kerferd and Sambell (2008–2011) to estab-
lish a recreational fishery is uncertain and Lake Sambell has
recently (2011–2012) been partially drained for maintenance.
There is currently no stocking being undertaken solely for the
purpose of trout cod conservation in Victoria.
Assessment of recovery actions shows that much has been
achieved in the recovery of trout cod, through the implemen-
tation of three successive national recovery plans. Managing
fish across multiple jurisdictions and agencies in the MDB
has previously been identified as problematic (Koehn and
Lintermans 2012) and the present study has highlighted
the benefits of a dedicated, multi-jurisdictional recovery team
that can coordinate across state boundaries and agencies in
the implementation of a long-term, committed, coordinated
approach. Thediffering responsibilities
(e.g. management of water, habitat, fishery, fish production and
conservation) of the many state agencies has posed difficulties
for consistent communication, consensus and coordinated
actions. For example, the need for consistency among different
State legislation and regulations. However, there remains much
the progression of trout cod recovery actions will remain
Trout cod is typical of many large, long-lived freshwater
fishes that are subject to angling and other pressures (Hogan
provides lessons forother similar species, includingthoseof the
same genus (Lintermans et al. 2005). Clear, well designed and
written recoveryplans,togetherwithfundingand completionof
recovery actions, are key components to threatened species
recovery (Abbitt and Scott 2001; Crouse et al. 2002). Funding
for conservation is often directed towards ‘charismatic’ species
(Male and Bean 2005) and because trout cod is a large, iconic
species, with an interest by anglers, its recovery has been given
more attention and resources than that of other smaller Austra-
progress, as demonstrated in the present paper.
Trout cod has undergone serious declines in both range and
abundance and this causes an issue of ‘shifting baselines’ that
can be an impediment to recovery when recovery targets are set
too low (Lintermans et al. 2005; Humphries and Winemiller
2009). Managing public perception of when a fish has
‘recovered’ can be problematic when anglers see a surge in
abundance of the target species, but do not appreciate that they
are all stocked individuals, or what the historic levels of the
fishery originally were. Such misconceptions of ecology and
In this respect, the value of angler knowledge and historical
records (Trueman 2011) has been illustrated through clarifica-
tion of the true historical range and relative abundance of the
species (e.g. the presence of trout cod in the Lachlan River).
The expansion of the remnant population and establishment
of new populations reduce the extinction risk of a threatened
species and, despite some early failures, the long-term commit-
ment to re-establishment of trout cod throughout their former
range has been a key tenet to the recovery of this species. The
stocking-strategy and adaptive-management approach outlined
by Bearlin et al. (2002) and Todd et al. (2004) for trout cod has
been successful for some populations. Lyon et al. (2012)
postulated that an important facet of a long-term stocking
program for recovery of an endangered species is the increased
chance of a stocked cohort encountering favourable environ-
mental conditions that promote local survival, particularly
immediately following release. In the Ovens River, fish stocked
in 2003 and 2004 were more highly represented in sampling
programs than those from other years, supporting the conserva-
tive approach of releasing stock over a longer period of time
(Lyon et al. 2012). Further, stocking yearling trout cod had no
discernable impact on the final population size or structure
(Lyon et al. 2012). Ebner et al. (2007, 2009) found that
Marine and Freshwater Research
J. D. Koehn et al.
2-year-old and adult hatchery-reared trout cod individuals had
high mortality rates after release into the Murrumbidgee and
Cotter Rivers, probably because of behavioural deficits of such
on-grown fish (Brown and Day 2002; McDermid et al. 2010).
Consequently, releasing larger and more mature on-grown fish
via conventional hatchery practices seems unlikely to provide
major advances in trout cod recovery.
Apart from translocations, there are few options other than
stocking for population establishment. Translocation is widely
used in threatened species conservation (Seddon et al. 2007;
Lintermans 2013b) and, historically, has been used to establish
populations of trout cod and other freshwater angling species
(Cadwallader 1981; Cadwallader and Gooley 1984). Lake
Sambell, Seven Creeks and Cataract Dam trout cod populations
were all established through translocations; however, recently
this has not been pursued as an alternative to stocking. Impor-
tantly, there is only a single source population (Murray River)
potentially able to sustain the loss of translocated individuals.
This lack of alternate source populations may be alleviated as
reach sufficient abundances to sustain the harvest of fish for
It is recognised that there is a need to better engage recrea-
tional anglers in the management and conservation of fishes
(Granek et al. 2008). This aspect of trout cod recovery has not
been undertaken as well as for the other Maccullochella species
and increased community involvement and participation would
be valuable. The national recovery team has not been able to
secure an ongoing community representative, whereas eastern
freshwater cod and Mary River cod both have considerable
community involvement in their recovery programs (NSW
in the conservation of Murray cod is proving fruitful (Koehn
2010; Koehn and Todd 2012), helping real and perceived
conflicts between conservation and recreational-fishery objec-
tives toberesolved(Koehn2010).Because ofitsrecognitionby
anglers, trout cod is an ideal candidate to be used as a ‘flagship’
species (Simberloff 1998) that can be then used to convey
conservation messages for other species.
Angling is an important and popular pastime in Australia,
with Murray cod being a popular target species where it occurs
(Henry and Lyle 2003). While the take of trout cod is currently
prohibited, accidental captures are common, especially in highly
fished areassuch as the remnant Murray River population,where
the trout cod population may be affected by post-release mor-
long-lived species such as trout cod (Coggins et al. 2007). Short-
term release mortalities for Murray cod are considered to be
relatively low (Douglas et al. 2010; Hall et al. 2012) but such
impacts are not transferable between species (Bartholomew and
Bohnsack 2005), can be cumulative (with multiple captures),
change with conditions and fishing methods (Bartholomew and
Bohnsack 2005) and sublethal effects need to be considered
(Cooke and Suski 2005). The protection of larger, older fish in
the population is seen as important for fish species such as trout
cod (Birkeland and Dayton 2005) and the education of anglers in
(e.g.barblessorcirclehooks,lureonlyfishing) (Cookeand Suski
2005) to reduce any impacts should be seen as a priority.
The prospect of once again allowing fishing for trout cod is
popular with anglers and this has been recognised in the latest
recovery plan, with steps taken to establish trial fisheries in a
few small lakes in Victoria and New South Wales. There is,
however, a clear need to manage public expectations because
no timelines or measures for success have been provided for
these fisheries. The active involvement of anglers contributing
substantially to conservation measures for this species could
greatly improve both conservation and fishery outcomes, as
without a down-listing of conservation status, it is unlikely that
any take of trout cod from existing populations will be allowed
in the near future.
The long-term nature of the recovery of trout cod was
recognised by Brown et al. (1998) in the second recovery plan,
by setting a recovery objective of downgrading of trout cod
conservation status from endangered to vulnerable within 25
years (by 2023). Achieving this objective is, however, depen-
recovery actions, supported by adequate population monitoring
to ensure that the required criteria have been met. IUCN
threatened species criteria are highly sensitive to the number
of mature individuals and to uncertainty in parameter values;
support any down-listing (Nicol 2005). A lack of data to show
unequivocal changein populationsor alackof precisionin such
data will only delay down-listing (Nicol 2005). Although
establishing new populations reduces the overall risk to the
species, monitoring of population status has been variable;
a common failing in threatened species recovery efforts
(Campbell et al. 2002). Monitoring must not only be directed
at the focal species, but also at current and emerging threats
(e.g. in the present case, at climate change, blackwater events,
River population (Lyon et al. 2012) at other sites, together with
predictions made using an updated population model, would
assist such assessments.
Climate change has the potential to affect Australian fresh-
water fishes in multiple ways (Koehn et al. 2011). Changes to
reproduction (Pankhurst and Munday 2011), range contractions
(particularly in upland areas; Bond et al. 2011), changes to
habitats (e.g. water quality or reductions in refuge pools;
Balcombe et al. 2011; Pratchett et al. 2011), expansion of other
native or alien species (such as golden perch, which may
compete or predate; Bond et al. 2011) and the combination of
an impact on trout cod populations. Some potential impacts
relating to the reduced flows predicted for south-eastern
Australia as a result of climate change were highlighted during
the ‘millenium’ drought (Murphy and Timbal 2008). Trout cod
susceptible to drought impacts (Crook et al. 2010). Climate
change has not been adequately considered in the trout cod
recovery plans and this needs to be rectified for this and all
threatened freshwater fishes (see Koehn et al. 2011).
Recently, trout cod recovery has become less certain as a
recovery actions. The certainty of recovery-plan progression
and coordination is vital, because species for which there has
Recovery of the endangered trout cod
Marine and Freshwater Research
been increased spending on recovery actions are more likely to
decline (Male and Bean 2005; Kerkvliet and Langpap 2007).
Species with recovery coordinators have more recovery actions
implemented and species with a higher proportion of planned
recovery actions implemented are more likely to show positive
population trends (Lundquist et al. 2002). The continuation of a
National Recovery Team for this species is essential.
Recovering species with large geographical ranges, at low
population levels when listed and with multiple and difficult
threats, has been demonstrated to be problematic (Abbitt and
range of recovery actions has been implemented for trout cod,
the benefits of some actions such as legislation (e.g. closure to
fishing) and education (e.g. correct species identification),
habitat rehabilitation, environmental flows and fishways have
either not been monitored or their benefits to trout cod are
difficult to quantify (e.g. King et al. 2009). Many other actions
fish-related management actions (e.g. fishways, riparian plant-
ing, alien species control) that may help trout cod conservation.
In particular, these include actions such as those in the Native
2004; Koehn and Lintermans 2012) for other sympatric species
Koehn and Todd 2012).
There is of course, much still to be done before trout cod can
be removed from threatened species lists. The current national
recovery plan is due for revision in 2013 and assessments in
the present paper suggest that the following actions need
? re-instate a funded national recovery team,
? greater inclusion of climate-change and extreme-events
planning (fires, floods, droughts),
? need for additional recovery-success criteria (e.g. delisting
? greater surety of funding for recovery actions,
? expanded monitoring (species and threats),
? re-evaluation of the Yarrawonga Weir fish lift to pass trout
cod as well as provision of fish-passage facilities at barriers
within the ranges of re-establishing populations,
? habitat rehabilitation or enhancement where necessary and
? further research into the species flow requirements, so as to
advise effective environmental-flow delivery programs,
? resumption of conservation stocking in Victoria to establish
? engagement of anglers and other stakeholders to establish
trout cod as an icon fish species,
? education of anglers in low-impact fishing and catch-and-
? renewal and/or installation of trout cod information at all
priority trout cod sites,
? updating the trout cod population model, incorporating
new knowledge and model functions (see Koehn and Todd
? actions to prevent redfin perch predation impacts on the
Seven Creeks population and assessment for other sites.
The present paper has provided a valuable case study for the
time, effort, actions and difficulties of recovering a long-lived,
iconic native fish species across multiple jurisdictions. It has
highlighted the need for a long-term, committed, coordinated
approach; in the present case, over three consecutive national
recovery plans. Actions have been supported by considerable
new knowledge (breeding biology, movements, habitats,
genetics) and use of a population-modelling tool. Population
protection and regulations on angling (including closures), and
education materials/programs, have all played important roles,
and, in particular, the stocking of hatchery-produced fish has
established several new populations to reduce extinction risk.
Such stocking, however, needs to be planned, with adequate
stocking numbers being sustained over the appropriate long-
group in the recovery effort and the promotion of trout cod as a
flagship species. The establishment of additional breeding
populations and expansion of the largest natural population
(Murray River), together with other actions over the past
25 years, provides cautious optimism for the ultimate recovery
of this species.
The authors thank all those who have contributed significantly to national
Trout Cod Recovery plans, teams and recovery actions over the past 25
years, especially Simon Nicol, Alistair Brown, Jason Lieschke, Steve
Saddlier (DSE-ARI Vic), Paulo Lay, Glen Johnson (DSE Vic), Joy Sloan,
Anthony Forster, Geoff Gooley (DPI Fisheries Vic), Steve Thurston,
Matthew McLellan, John Pursey, Andrew Sanger Andrew Bruce, Craig
Copeland, Allan Lugg, John Pursey, Adam Vey (Fisheries NSW DPI),
Matthew Beitzel, Mark Jekabsons, Terry Rutzou (ACT Government, Con-
servation Planning and Research), Jim Barrett, David Hohnberg (Murray–
Kaminskas (ACT) and Ron Lewis. Thanks go to Wayne Koster for discus-
sion on the Goulburn River, Adrian Kitchingman for assistance with Fig. 1,
Luke Cruikshank and Goulburn–Murray Water for access to fishway data
and to Steve Saddlier and the two anonymous reviewers for comments on
drafts of this manuscript.
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