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Predation by rainbow trout (Oncorhynchus mykiss)

on a Western Australian icon: Marron (Cherax
cainii)

Michelle Y. Tay , Alan J. Lymbery , Stephen J. Beatty & David L. Morgan

To cite this article: Michelle Y. Tay , Alan J. Lymbery , Stephen J. Beatty & David L. Morgan
(2007) Predation by rainbow trout (Oncorhynchus�mykiss) on a Western Australian icon: Marron
(Cherax�cainii) , , 41:2, 197-204, DOI: 10.1080/00288330709509908

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New Zealand Journal of Marine and Freshwater Research, 2007, Vol. 41: 197-204 197
0028-8330/07/4102-0197 © The Royal Society of New Zealand 2007

Short communication

Predation by rainbow trout (Oncorhynchus mykiss)

on a Western Australian icon: marron (Cherax cainii)
MICHELLE Y.TAY INTRODUCTION
ALAN J. LYMBERY The considerable predatory impact of introduced
STEPHEN J. BEATTY rainbow trout (Oncorhynchus mykiss Walbaum 1792)
DAVID L. MORGAN on native aquatic fauna is widely acknowledged
Fish Health Unit (Jenkins 1952; Tilzey 1976; Pidgeon 1981;
Centre for Fish and Fisheries Research Cadwallader & Eden 1982; Welcomme 1984; Crowl
Murdoch University et al. 1992; Fuller et al. 1999), resulting in this
South St, Murdoch species being included by the IUCN in the list of 100
WA 6150, Australia of the world's worst invasive alien species (Lowe
et al. 2000; Lintermans 2004). Rainbow trout were
email: [email protected]
first introduced into waters of southwest Western
Australia in 1902 and large-scale stocking of the
species began in the 1930s (Coy 1979), with c. 2.5
Abstract Considerable numbers of rainbow trout million O. mykiss stocked into public waterways for
(Oncorhynchus mykiss) are captively produced recreational fishing between 1999 and 2004 (Astbury
then stocked into many wild aquatic systems in 2004). Populations of O. mykiss in natural waterways
Western Australia despite a lack of understanding in Western Australia are restricted to the cooler
of their impact on the highly endemic native fish water bodies between Perth in the north and Albany
and freshwater crayfish of the region. The diet of in the south (Morgan et al. 2004). Although some
an introduced, yet self-sustaining population of O. populations are known to be self-maintaining, in most
mykiss was examined at Churchman Brook Reservoir instances they are maintained by continuous stocking
in the Canning River Catchment, Western Australia, (Morrissy et al. 2002; Morgan et al. 2004).
between September and November 2004. The large
endemic freshwater crayfish marron (Cherax cainii), The impact of trout stocking in southwestern
which supports an iconic recreational fishery in Australia, which contains a highly endemic freshwater
Western Australia, comprised 61% by volume of fish (80% endemic) (Morgan et al. 1998) and crayfish
all O. mykiss stomachs examined. The contribution (100% endemic) (Crandall et al. 1999) fauna, has
made by C. cainii to the assimilated biomass of O. received little attention to date. Oncorhynchus mykiss
mykiss was confirmed by stable isotope analyses, is known to be an opportunistic feeder; feeding on
which revealed that it dominated the assimilated diet a wide variety of prey, ranging from fish (including
of larger O. mykiss. The substantial contribution of other O. mykiss), aquatic and terrestrial insects,
C. cainii to the diet of wild O. mykiss suggests that, decapod crustaceans and sometimes amphibians and
at least in habitat limited water bodies, it is likely to birds (Jenkins 1952; Tilzey 1977; Leitritz & Lewis
have a considerable predatory impact on C. cainii 1980; Pusey & Morrison 1989). The few studies
populations and may compromise some elements of that have examined the diet of stocked O. mykiss
this iconic recreational fishery. in southwest Western Australia recorded predation
on the highly endemic fish and freshwater crayfish
of the region (Jenkins 1952; Pusey & Morrison
Keywords dietary analysis; trophic position; stable 1989; Morgan et al. 2004). Of particular interest
isotopes; marron; predatory impact; introduced is the extent of predation on the endemic decapod
species crayfish, the smooth marron (Cherax cainii, formerly
C. tenuimanus - although these names are currently
under review (Molony et al. 2006)), which supports
M06050; Online publication date 31 May 2007 an important recreational fishery in the region
Received 14 August 2006; accepted 24 January 2007 (Molony & Bird 2002). In recognition of the high
198 New Zealand Journal of Marine and Freshwater Research, 2007, Vol. 41

Fig. 1 Churchman Brook Reser-

voir in the southwest of Western
îN Australia.
! WESTERN
<i) AUSTRALIA
\

Churchman Brook Reservoir

—K
Indian Ocean

^VIA/

--/"N&-- Albany 119 Southern Ocean

social value of the recreational marron fishery, it classes, i.e., (1) <200 mm TL, (2) 200-300 mm TL,
has recently been recommended that the species (3) >300 mm TL. Each stomach was removed and its
be nominated as a State freshwater faunal emblem contents examined under a dissecting microscope. All
(Department of Fisheries 2006). The aim of this animal matter was identified using Davis & Christidis
study was to determine the diet and trophic position (1997) to the lowest possible taxon (usually order or
of a wild, self-sustaining reservoir population of family) and classified as being terrestrial or aquatic.
O. mykiss in southwestern Australia via stomach Stomach contents were analysed using the percentage
content and stable isotope analyses. frequency of occurrence and the percentage of
volumetric contribution of each dietary item. The
percentage frequency of occurrence is the proportion
MATERIALS AND METHODS of the number of stomachs containing a particular
prey type to the total number of stomachs examined,
Study site and sampling regime whereas the percentage of volumetric contribution
Churchman Brook Reservoir is located c. 30 km is the proportion of each prey type to the overall
southeast of Perth in the Canning River (Fig. 1). stomach contents of each fish, or all fish combined
The earth-fill embankment dam has a catchment (Hynes 1950; Hyslop 1980). These two different
area of c. 16 km2, with a reservoir capacity of 2.16 methods were applied as they may show different
gigalitres stretching 1.4 km in length (Department trends if large prey items dominate the stomachs of
of Water 2007). Rainbow trout were sampled on particular individuals. Oncorhynchus mykiss with
four separate occasions during spring, 2004; once in no dietary items present in stomach contents were
September and October, and twice in November. On excluded from these analyses. Wherever possible
each sampling occasion, four gill nets (50,75,100, (i.e., measurable), each C. cainii present in the guts
and 125 mm stretched mesh) were set overnight (5 of O. mykiss was measured to the nearest 1 mm
pm - 9 am) within the reservoir. To obtain adequate orbital carapace length (OCL).
numbers of juvenile O. mykiss and C. cainii, they Similarities in dietary composition among
were collected from c. 100 m of stream immediately individual O. mykiss were estimated from square-
above the reservoir using a back-pack electrofisher root transformed percentage volumetric data
(Smith Root Model 12-A). using the Bray Curtis similarity coefficient (Bray
& Curtis 1957; PRIMER v5, Clarke & Gorley
Stomach contents analysis 2001). Patterns of similarity among different size
The total length (TL) of each fish was measured to classes of O. mykiss were assessed with non-metric
the nearest 1 mm and wet weight measured to the multidimensional scaling (MDS) of the Bray Curtis
nearest 10 mg. Fish were categorised into three size similarity matrix. Stress values, or the distortion
Tay et al.—Predation by rainbow trout on marron 199

between the original similarity rankings and similarity and conventional standards (Pee Dee Belemite
13
rankings in the ordination plot, were compared at carbonate for δ C and atmospheric nitrogen for
15
different dimensionalities of ordination to determine δ N). The precision of the analytical equipment was
13 15
the most suitable plot dimensions. A stress value ± 0.1‰ for δ C and ± 0.3‰ for δ N.
<0.2 indicates reasonable representation of rank 13
The mean and standard error of the δ C and δ N
15

similarities by the ordination plot (Clarke 1993). The ratios for each sample group (i.e., size categories,
significance of differences in dietary composition type of species, and sampling time) were calculated.
between size classes was tested by analysis of To infer the relative trophic positions of C. cainii
similarities (ANOSIM, PRIMER v5, Clarke & and O. mykiss, we assumed the mean δ N ratio
15

Gorley 2001). Where multiple statistical tests of fractionation rate of O. mykiss to be 3.36‰, which
the same hypothesis were applied, a Bonferroni was the difference in the signatures of O. mykiss fed
correction was used to maintain an experiment-wide solely on pelleted food over a period of 9 months
Type I error rate of 5% (Bland & Altman 1995). at an aquaculture facility (Tay 2004). The equation
The contribution of individual dietary items to the T
L = (δ 1 5 N O.mykiss- δ 1 5 N CcalJ 13.36 (adapted
similarity within size classes or the dissimilarity from Post 2002), was used to determine the trophic
among size classes was determined by averaging difference between predator and its prey, where T L
the Bray-Curtis similarity (or dissimilarity) term is the trophic level difference between O. mykiss and
for each item over all pairwise individual O. mykiss C. cainii, δ15NO. m ykiss is the mean stable δ 15 N ratio of
combinations, using SIMPER (PRIMER v5). To O. mykiss and O.mykssi „ . Nis the mean δ 15 N ratio of C.
determine the strength of the relationship between cainii.
size of O. mykiss and the ingestion of particular
dietary items, percentage volumetric data were
arcsine transformed before simple linear regression
analysis (Zar 1998). RESULTS
The size range for 123 O. mykiss captured in
Stable isotope analysis Churchman Brook Reservoir from September 2004
Stable isotope analysis (δ 1 3 C and δ 1 5 N) was to November 2004 was 23-392 mm TL: 16 were
also undertaken to further determine the trophic <200 mm TL, 48 were between 200-300 mm TL,
relationships of O. mykiss in the reservoir. Four and 59 were >300 mm TL. One-hundred-and-five
replicates of both small (224-305 mm TL) and large of these fish had distinguishable dietary items in
(334-380 mm TL) O. mykiss, each consisting of their stomach. Of the 21 food categories ingested
one randomly selected individual, were collected by O. mykiss, C. cainii comprised the bulk of the
in October and November 2004. A sample of dorsal diet (c. 60% by volume, c. 70% by occurrence)
tissue (wet weight of 2-5 g) from each O. mykiss (Table 1). Both whole juvenile marron (OCL 7-23
was removed and dried at 70°C for 48 h. Similarly, mm, n = 9 measurable OCLs) and the appendages
four replicates of both juvenile (<30 mm OCL) of adult marron were found in trout stomachs. The
and adult (>30 mm OCL) C. cainii were analysed next most important aquatic invertebrates ingested
in September, with three replicates per size class were dytiscids that were found in c. 30% of all O.
analysed in October for δ 13 C and δ 15 N. Between mykiss examined and contributed c. 10% of diet by
2 and 5 g wet weight of abdominal muscle tissue volume. Other aquatic invertebrates consumed were
(without carapace or digestive tract) was removed anisopteran larvae and amphipods, while the main
from each individual and placed in 1 mol/litre of terrestrial invertebrates ingested included formicids,
HCL for 24 h to remove inorganic carbonates. dipterans and diplopods (Table 1). Overall, terrestrial
Samples were then rinsed in distilled water and oven fauna comprised c. 19%, and aquatic invertebrates,
dried at 70°C for 48 h. Before being finely ground excluding C. cainii, comprised c. 18% by volume
to a powder with a mortar and pestle, all samples of O. mykiss diets.
were re-weighed to a dry weight of between 2 and Ordination analysis of dietary composition
3 mg. A tracermass iron ratio mass spectrometer showed marked differences between size classes
(Europa PDZ, Cambridge, United Kingdom) fitted (Fig. 2), which were confirmed by ANOSIM (R =
with a Roboprep combustion system was used for 0.38, P < 0.001). Pairwise tests found significant
oxidising samples. The ratios of carbon δ 13 C:δ 12 C differences in dietary composition between all
and nitrogen δ15N:δ14N were expressed as the relative size classes (1 and 2, R = 0.45, P < 0.001; 1 and
parts per thousand differences between the sample 3, R = 0.86, P < 0.001; 2 and 3, R = 0.10, P <
200 New Zealand Journal of Marine and Freshwater Research, 2007, Vol. 41

50
Fig. 2 Multidimensional scaling (MDS) ordination of ° 150 200 250
the similarity matrix of dietary composition (calculated Total length (mm)
from % points data) of the different size classes of Onco-
rhynchus mykiss from Churchman Brook Reservoir. Size Fig. 3 Relationship between total length of Oncorhyn-
groups are: • , <200 mm TL; O, 200-300 mm TL; • , chus mykiss (mm) and % volumetric contribution of
>300 mm TL. Cherax cainii in the diet.

Table 1 Relative contribution by volume (% points) and percentage occurrence

of food items of Oncorhynchus mykiss (n = 105) captured at Churchman Brook
Reservoir, Western Australia. (* denotes a terrestrial food item.)

Prey type % contribution (± 1SE) % occurrence

Cherax cainii 60.94(19.13) 67.62
Dytiscidae 9.99 (6.66) 30.48
Formicidae* 6.05 (7.86) 7.62
Diptera* 4.43 (4.95) 10.48
Diplopoda* 4.16(5.88) 6.67
Anisoptera larvae 3.38 (4.67) 12.38
Amphipoda 2.34 (2.55) 17.14
Tabanidae* 2.22 (3.63) 3.81
Vegetation 1.93(3.12) 5.71
Unidentified animal matter 1.42(2.93) 3.81
Oncorhynchus mykiss 0.78 (2.45) 0.95
Chironomidae larvae 0.68(1.18) 3.81
Terrestrial insects* 0.62(1.01) 3.81
Chironomidae pupae 0.42(1.16) 1.90
Noteridae 0.19(0.61) 0.95
Apoidea* 0.16(0.49) 0.95
Culicidae larvae 0.10(0.33) 0.95
Coleoptera 0.09 (0.25) 1.90
Tabanidae larvae 0.05(0.16) 0.95
Zygoptera larvae 0.04(0.12) 0.95
Arachnida* 0.01 (0.03) 0.95

0.010), although the differences were most marked volume to the diets of fish of size class 1, 67.6%
between those fish <200 mm TL (size class 1) and by frequency of occurrence and 59.8% by volume
those 200 mm TL (size classes 2 and 3). SIMPER to the diets of fish of size class 2, and 70.2% by
analysis revealed that over 70% of the differences frequency of occurrence and 84.4% by volume to
in diet between fish <200 mm TL and those 200 the diets of fish of size class 3. Regression analysis
mm TL were explained by a preponderance of confirmed a significant relationship between fish
terrestrial fauna in small fish and a preponderance size and percentage volumetric contribution of C.
of C. cainii in larger fish. Cherax cainii contributed cainii (Fig. 3; r2 = 0.29; n = 104, F1 m = MAI, P
14.3% by frequency of occurrence and 5.7% by < 0.001).
Tay et al.—Predation by rainbow trout on marron 201

studies in Western Australia that found terrestrial

c.
0
V c.
ainii small (Sept)
ainii large (Sept) invertebrates to be more important components
D
0
c
c
ainii small (Oct)
ainii large (Oct) of O. mykiss diets than aquatic invertebrates. For
A 0.
0 0
•nyWss small (Oct)
mykiss large (Oct) example, Jenkins (1952) found that terrestrial insects
• 0.
V 0
vykiss small (Nov)
mykiss large (Nov)
T T
had 18 times the contribution to O. mykiss diets
nty y—H
I 5 l than aquatic invertebrates at several rivers in the
Nitrogen fractionation
dis anee between
southwest. Similarly, Pusey & Morrison (1989)
Or nykiss and its food
found that 73% of O. mykiss diets at Wungong
i Dam consisted of terrestrial insects. The difference
\ .i. i i i between these studies and the results of the current
Ï^ i study may be attributed to differences in aquatic
habitats. Churchman Brook Reservoir is typical of
impoundment reservoirs of the region; it is largely
devoid of riparian and aquatic vegetation (M. Tay
pers. obs.). As a result, tree stumps rather than
Fig. 4 Mean values (± 1SE) of stable carbon δ 1
3C (‰) fringing aquatic vegetation surround the banks of
isotope ratios and stable nitrogen δ15N (‰) isotope ratios Churchman Brook Reservoir, thus possibly limiting
of Oncorhynchus mykiss (n = 4 per size class and sam- the availability of terrestrial insects for O. mykiss in
pling occasion) and Cherax cainii (n = 4 and 3 per size this ecosystem.
class in September and October, respectively) tissue in
spring 2004 from Churchman Brook Reservoir, Western The preference for C. cainii by O. mykiss at
Australia. δ15N (3.36‰) and δ13C (1.29‰) fractionation Churchman Brook Reservoir increased markedly
(enrichment) rates of O. mykiss and its food are displayed with increasing fish size. This finding is similar
as determined by Tay (2004). to a study by Pidgeon (1981) in two streams in
New South Wales that demonstrated that freshwater
crayfish increase in proportions in the diets of larger
The mean δ13C and δ15N signatures of C. cainii O. mykiss. The dominance of C. cainii in O. mykiss
analysed in September were generally more diets in Churchman Brook Reservoir suggests that O.
enriched than C. cainii signatures in October mykiss is a specialist and/or opportunistic predator in
(Fig. 4). Similarly, O. mykiss in October had more this system; selecting the most accessible prey and
enriched mean δ 13 C and δ 15 N signatures than O. larger prey that provide a large proportion of total
mykiss in November, except for large O. mykiss in energy intake (Tilzey 1977). Foraging for larger
November that had the most enriched δ 15 N signature sized prey results in lower energy cost compared
(11.27‰) (Fig. 4). Both large and small O. mykiss with foraging for many smaller sized invertebrates
had significantly enriched carbon (δ13C) and nitrogen with the equivalent amount of energy; resulting
(δ15N) signatures compared with large and small C. in a faster growth rate (Kerr 1971). As a result,
cainii; with the exception of δ 13 C signatures of large freshwater crayfish could make an important
O. mykiss in November (-21.78‰), when compared contribution to the growth rate of O. mykiss and
to large C. cainii in September (-21.59‰) (Fig. the availability of C. cainii as a food source in the
4). Oncorhynchus mykiss were approximately 0.99 habitat-limited Churchman Brook Reservoir has
trophic levels higher than C. cainii in Churchman probably contributed to the successful establishment
Brook Reservoir. of O. mykiss in this system. It is likely that O. mykiss
and C. cainii have coexisted in Churchman Brook
for at least 18 years (Brett Molony pers. comm.
Department of Fisheries, Government of Western
DISCUSSION Australia). It is possible these species continue to
The analyses of stomach contents and stable carbon co-exist despite the heavy predation recorded here
and nitrogen isotopes of a self-sustaining population owing to the life-history strategy of C. cainii. These
of O. mykiss at Churchman Brook have demonstrated life-history traits, such as having a high fecundity
that O. mykiss consume and assimilate significant and rapid growth to a large size (Beatty et al. 2003,
quantities of C. cainii. Other aquatic invertebrates 2005), would result in a proportion of individuals
and terrestrial fauna were also important components rapidly reaching a mature size large enough to
of O. mykiss diets in this system, especially for preclude predation by O. mykiss, therefore allowing
smaller fish. Our results contrast with previous on-going recruitment to the breeding population.
202 New Zealand Journal of Marine and Freshwater Research, 2007, Vol. 41

The highly modified environment of Churchman a known predator of C. cainii in reservoirs (Morgan
Brook Reservoir may limit the extent to which we et al. 2002; Molony et al. 2004).
can generalise our results to more natural systems. This study demonstrates that C. cainii appears to
For example, there are considerable differences be a very important component of the diet of a feral
between the habitat complexity of natural systems O. mykiss population for at least part of the year.
and this impoundment reservoir, with natural rivers This level of prédation is of great concern given
generally having more refuge structures available the current status of C. cainii and the other endemic
owing to the greater amount of large wooden debris decapod crustacean populations in the southwest
input from the riparian zone. This greater habitat of Western Australia. Further assessments of the
complexity in more natural lotie systems may result predatory impact of O. mykiss in other freshwater
in lower prédation rates by O. mykiss on C. cainii in systems of the region need to be undertaken to better
those systems than was recorded in the present study. understand the ecological impacts caused by the
However, previous studies have also documented stocking regime of this species.
O. mykiss prédation on C. cainii and other decapod
crustaceans in Western Australia. Pusey & Morrison
(1989) showed that at Wungong Dam, C. tenuimanus ACKNOWLEDGMENTS
(now known as C. cainii) was the most important
prey by weight (c. 80%), occurring in over a third Thanks to Peter Chalmers for field assistance, the Water
of all O. mykiss stomachs examined. Jenkins Corporation of Western Australia for site access, Lydia
Bednarekfor assistance with stable isotope analyses, and
(1952) also found that C. tenuimanus made up a Drs Rob Doupé and Howard Gill for assistance invarious
considerable proportion of O. mykiss diets, occurring stages of the study.
in seven out of 10 O. mykiss caught in several rivers
in southwestern Australia. Morgan et al. (2004)
demonstrated that 23% of 92 O. mykiss collected REFERENCES
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