Effects of flow on lateral interactions of fish and shrimps with off

WORLD’S LARGE RIVERS CONFERENCE

Effects of flow on lateral interactions of fish and shrimpswith off-channel habitats in a large river-floodplain system

K. Gorski • K. J. Collier • D. P. Hamilton •

B. J. Hicks

Received: 31 January 2012 / Accepted: 3 October 2012 / Published online: 4 December 2012

� Springer Science+Business Media Dordrecht 2012

Abstract Off-channel habitats play a crucial role in

the life-cycles of many riverine fish species, but lateral

movements of fish into these habitats are poorly

understood. We tested how flow dynamics affects the

movement of fish and shrimps between the main river

channel and different types of off-channel habitats: a

riverine lake and a wetland. Our study site was the

lower Waikato River, North Island, New Zealand,

where there are numerous off-channel habitats. Fish

were sampled using directional fyke nets. Shortfin eel

(Anguilla australis) migrated mostly into the wetland

at night, particularly during high river flows. Common

bullies (Gobiomorphus cotidianus) were most abun-

dant during the day and in low-discharge conditions,

moving mostly into the riverine lake, whereas fresh-

water shrimp (Paratya curvirostris) moved mostly

into the wetland. High numbers of non-native larval

common carp (Cyprinus carpio) moved out of the

wetland with retreating flood water. This study

emphasises the importance of lateral connectivity

and flooding in functioning of this river system where

numerous native fish, but also exotic fish, used off-

channel habitats. Floodplain management strategies

should promote ‘controlled connectivity’ measures

that provide access for native species at key times

while limiting opportunities for introduced species to

utilise their favoured off-channel habitats.

Keywords Fish movement � Habitat connectivity �Floodplain � Riverine lake �Wetland �Waikato River

Introduction

Floodplain wetlands adjoining large lowland rivers are

amongst the most biologically productive freshwater

ecosystems (Tockner et al., 2009). Seasonal flow and

flood pulses increase connectivity between various

components of riverine systems, expanding habitat

availability for aquatic organisms (Junk et al., 1989;

Tockner et al., 2000). Currently, large river–floodplain

systems that retain a high degree of natural function-

ality are rare in temperate regions, and most are highly

modified by river regulation, e.g. construction of dams

and dykes, changes in land use, and introductions

of non-native species (Bayley, 1995; Tockner &

Stanford, 2002; Nilsson et al., 2005). Off-channel

Guest editors: H. Habersack, S. Muhar & H. Waidbacher /

Impact of human activities on biodiversity of large rivers

K. Gorski (&) � K. J. Collier � D. P. Hamilton �B. J. Hicks

Environmental Research Institute—Te Putahi Rangahau

Taiao, University of Waikato, P. O. Box 3105, Hamilton,

New Zealand

e-mail: [email protected]

K. J. Collier

Waikato Regional Council, P. O. Box 4010, Hamilton,

New Zealand

123

Hydrobiologia (2014) 729:161–174

DOI 10.1007/s10750-012-1352-1

habitats such as floodplain wetlands and lakes can be

crucial in supporting feeding, spawning and nursery

areas for many riverine fish species (Junk et al., 1989;

Baber et al., 2002; King et al., 2003; Jimenez-Segura

et al., 2010; Gorski et al., 2011a; Magana, 2012).

Consequently, in many river systems, fish community

structure and production are directly related to the

quality and quantity of connections between main

river channels and off-channel floodplain habitats

(Welcomme, 1979; Moses, 1987; De Graaf, 2003;

Gorski et al., 2011b).

In European rivers, cyprinids have been shown to

migrate from the main river channel into off-channel

habitats, including tributaries (Nunn et al., 2010),

floodplain water bodies (Molls, 1999; Hohausova

et al., 2003) and temporarily inundated grasslands

(Cucherousset et al., 2007), especially with increas-

ing river discharge and during floods. Similar

phenomena have been observed in large tropical

rivers with highly predictable seasonal floods, where

the lateral migration of fish closely followed the

dynamic ‘pulsing’ of water levels (Junk et al., 1989),

enabling fish to access superior feeding and nursery

habitats as water levels rose and advanced over

terrestrial riparian habitats during seasonal floods

(Wantzen et al., 2002; Castello, 2008). In the North

American temperate Kankakee River, for example,

fish species adapted to seasonal flooding have been

shown to repeatedly seek out floodplain habitats

during high flows, whilst being forced to the main

channel during low flows (Kwak, 1988). The

importance of off-channel habitats during low flow

conditions has been stressed in recent studies from

the Murray Darling Basin in Australia, where high

numbers of both native and non-native fish moved to

accessible off-channel habitats during low water

levels (Lyon et al., 2010; Conallin et al., 2011).

During winter, when low water temperatures limit

metabolism and swimming performance, many tem-

perate fish species prefer lentic backwater conditions

to conserve energy (Lucas & Baras, 2001). Off-

channel habitats can also provide refuges for fish from

high water velocities during high flows (Schwartz &

Herricks, 2005), and allow fish to restore energy

reserves after spawning (Fernandes, 1997; Gorski

et al., 2010). However, organically enriched backwa-

ter and floodplain habitats may often suffer hypoxia,

forcing fish to move back to flowing water of the main

river channel (Knights et al., 1995).

To make informed decisions about the rehabilita-

tion of riverine fish production, it is essential to

understand the ecological role of off-channel habitats

for both native and introduced species (Galat et al.,

1998; Buijse et al., 2002). Compared to large-scale

longitudinal migrations within river corridors, how-

ever, lateral fish movement and migrations in lowland

rivers remain one of the most poorly understood

dispersal mechanisms in temperate freshwater eco-

systems (Lucas & Baras, 2001; Nunn et al., 2010).

This is particularly so in New Zealand and other

southern hemisphere temperate countries where the

native fish fauna is dominated by diadromous species

(McDowall, 1990), and studies on lateral movement

are scarce.

The Waikato River is New Zealand’s longest river.

Its lower reaches were historically characterised by

extensive river–floodplain interactions (Collier et al.,

2010a), and the river once supported the most

productive whitebait (Galaxias spp.) and eel (Anguilla

spp.) fisheries in New Zealand (Stancliff et al., 1988;

Chapman, 1996). Complex riverine habitats associ-

ated with floodplain wetlands and riverine lakes

provided important habitat for a range of species in

the Waikato River, including many native fish that are

now considered to be endangered (Collier et al.,

2010b). Before European settlement, floodplain areas

in the lower Waikato River extended to around

364 km2, but at present only about 53% of that area

remains due to implementation of flood protection

schemes and changes in land-use (Collier et al.,

2010a). Twelve introduced fish species now co-occur

with native species, with common carp Cyprinus

carpio (Linnaeus, 1758) often dominating fish bio-

mass in the lower river (Hicks et al., 2010).

In this study, we investigated movement of larval

and juvenile fish between the main river channel and

two types of lateral habitats (riverine lake and

floodplain wetland) in the lower Waikato River. Our

specific objectives were to: (1) quantify the abundance

of fish moving into and out of a riverine-lake and

wetland habitats in different seasons compared to

movements within the main river channel, and (2)

investigate the role of selected environmental cues

associated with movement into and out of off-channel

habitats for different fish species. We hypothesised

that seasonal variations in flow could play a crucial

role in triggering lateral movement of both native and

introduced fish from the Waikato River into off-

162 Hydrobiologia (2014) 729:161–174

123

channel habitats (Hohausova et al., 2003; Nunn et al.,

2010). We further hypothesised that high flows could

trigger movement of fish, so that shortfin eel, Anguilla

australis (Richardson, 1841), might exploit off-chan-

nel habitats for feeding as well as exposed floodplain

areas with retreating water, as shown for Anguill

anguilla (Linnaeus, 1758) in Europe (Lasne et al.,

2008). Small-bodied fish species such as ınanga

Galaxias maculatus (Jenyns, 1842) as well as fresh-

water shrimps Paratya curvirostris (Heller, 1862)

might be expected to move into the off-channel

habitats to avoid high water velocities in the main

channel during floods (Schwartz & Herricks, 2005).

Methods

Study area

The Waikato River flows in a northerly direction for

around 442 km from its headwaters above Lake Taupo

to the Tasman Sea at Port Waikato (Collier et al.,

2010b). It drains a total catchment of 14,443 km2 and

has a mean annual discharge at the mouth of approx-

imately 450 m3 s-1 (Brown, 2010). The catchment

has been significantly altered from its natural state,

mostly for agriculture (62%) and exotic forestry

(19%), as well as some urban development (Collier

et al., 2010b), whilst the upper river is punctuated by

eight hydro dams. A dam 152 km upstream from the

river mouth effectively acts as a barrier to the natural

upstream movement of aquatic fauna. Soils in the

Waikato River catchment are dominated by sediments

with low infiltration rates, and therefore the river

system is highly responsive to rainfall, with large peak

flood flows after heavy rain (usually in winter and

spring) and low flows after periods of low rainfall

(summer and autumn) (Brown, 2010).

This study was conducted in the Waikato River’s

lower reaches, where it forms a low-gradient river

accommodating extensive floodplains which are char-

acterised by peat wetlands and several riverine lakes,

although these interactions are now highly regulated

by a flood protection scheme which was initiated in

the 1950 and 1960s (Chapman, 1996). Lake Whangape

(‘Lake’) and Opuatia wetland (‘Wetland’) were

selected as off-channel habitats to quantify fish move-

ment to and from the river main channel at different

discharges (Fig. 1). These habitats are amongst the few

remaining which have relatively undisturbed connec-

tivity with the river main channel in the lower Waikato.

Lake Whangape has a surface area of 14.5 km2 and

muddy sediments, and is a turbid, hyper-eutrophic lake

with high productivity dominated by phytoplankton;

aquatic macrophytes are absent (Hamilton et al., 2010).

A stone weir constructed in the lake for water level

control limits connectivity with the river main channel

in low flows, but during moderate to high flows the weir

is fully submerged, allowing unrestricted fish move-

ment. Opuatia wetland covers approximately 9.5 km2

and is one of the few remaining wetlands retaining

dominance of native restiad rushes (Clarkson et al.,

2004), supported by a peat bog surrounded by minera-

lised margins (Beard, 2010). Water originating from

the peat bog may periodically induce hypoxia in the

outflow of the wetland (Kuder & Kruge, 2001). Some

flood protection works have been carried out in this

wetland, including construction of bunds along the

boundaries of willow and pasture areas (Beard, 2010).

Fig. 1 The study area indicating sampling sites (circled fish)

and its location in New Zealand

Hydrobiologia (2014) 729:161–174 163

123

Data collection

Water level and temperature were recorded at 1-h

intervals at the inlet to each off-channel habitat using

automatic data loggers (3001 Levelogger� Junior;

Solinst Canada, Ontario, Canada). Daily discharge data

of the Waikato River at Rangiriri (approximately 5 km

upstream from the main channel sampling site (Fig. 1);

37�25055.8800S, 175�7041.5000E) were obtained for the

sampling period from Waikato Regional Council,

Hamilton, New Zealand. In addition, spot water

temperature, dissolved oxygen and specific conduc-

tance were recorded approximately 1 m below the

water surface on each sampling occasion using hand-

held devices (YSI model 55 handheld DO meter and

YSI 30 meter; Yellow Springs Instruments, OH, USA)

for conductance and temperature, respectively.

Fish were sampled bi-monthly between 29 September

2010 and 16 September 2011 to encompass seasonal

variations and different water levels and flows. Sam-

pling was conducted at the Lake Whangape inlet

(37�25050.7900S, 175�4042.9000E), which was character-

ised by water depth 1–2 m, width c. 30 m in low flow

conditions, muddy sediment and aquatic macrophytes

absent, and at Opuatia wetland inlet (37�24037.5000S,

175�3028.5500E), where thedepth was 1–2 m, the width c.

20 m in low flow conditions, muddy sediment and about

10% areal coverage with aquatic macrophytes (mostly

hornwort Ceratophyllum demersum L.) (Fig. 1).

To compare movement patterns into and out

of the lake and wetland with those in the main channel

during each sampling, fish were also collected from

the shore of a mid-channel island (37�2500.1600S,

175�4015.1600E; Fig. 1), where the depth was 1–2 m

and sediments were sandy, while aquatic macrophytes

absent. On each sampling occasion, a set of two-

directional double-wing fyke nets (2.5 m wing span)

was set in each location over the same 24-h period. As

this study concentrated on small and juvenile fish, fine-

mesh (2 mm) nets were used. The nets were set at

similar depths in each location, facing both upstream

(‘in’) and downstream (‘out’) to catch fish moving ‘in’

and ‘out’ of the off-channel habitats. Nets were

checked close to dawn and dusk to give an indication

of diurnal movement patterns. For each sample, fish

were identified to species-level based on morpholog-

ical features and pigmentation (Koblitskaya, 1981;

McDowall, 1990). Fish were measured for total length

(±1 mm) for at least 100 individuals if available to

derive length–frequency distributions. For the less

abundant fish, all individuals were measured. Catch per

unit effort (CPUE) was calculated for day and night

samples, expressed as number of fish caught per hour.

Data analysis

We used permutational multivariate analysis of variance

(PERMANOVA) (Anderson, 2001; McArdle & Ander-

son, 2001) to determine differences in CPUE between

the main channel and off-channel habitats for different

species. This test was chosen because we wished to

assess the differences between habitats and seasonal

dynamics as well as their interactions, and CPUE data

did not meet normality assumptions for parametric tests.

PERMANOVA provides analysis of the variance of

data for a set of explanatory factors on the basis of

dissimilarity measures, thereby allowing a wide range of

empirical data distributions. Models were run based on a

Bray–Curtis dissimilarity matrix for the whole fish

community, and separately for the four most abundant

native fish species and freshwater shrimp based on

similarity among dates and sites. Data were square root-

transformed prior to analysis to reduce the effect of

outliers. For the two most abundant species—the

common bully Gobiomorphus cotidianus (McDowall,

1975) and shortfin eel—we compared size distributions

of individuals between habitats and directions of

movement using a Kolmogorov–Smirnov test.

Finally, to identify the relationship between move-

ment patterns and habitat type, flow magnitude, water

temperature and measured water quality variables

(dissolved oxygen concentration and conductance),

we performed redundancy analyses (RDA) (based on

correlation matrices) (Jongman et al., 1995; ter Braak

& Smilauer, 2002) on square root-transformed CPUE

data of the five most abundant species fish moving into

or out of off-channel habitats (river samples were

excluded). Global Monte Carlo permutation tests

(1,000 permutations) were performed to determine

the significance of the ordination at a = 0.05.

Results

Flow and water quality parameters

The river flow during the sampling period was

characterised by high discharge in spring 2010

164 Hydrobiologia (2014) 729:161–174

123

(September–October) and winter 2011 (May–August),

with a short (1-week) flood peak in summer (February

2011) (Fig. 2, top panels). Water temperature was

highest in the lake inlet and the lowest in the wetland

inlet (Table 1) over the period of sampling. During

spring flooding, water temperature gradually

increased in both the lake and the wetland, and

reached 20–24�C during the summer months (Janu-

ary–February), gradually decreasing during winter

high flows (Fig. 2, top panels). Daily fluctuations in

water temperature were about threefold larger at the

lake inlet compared to the wetland inlet (Fig. 2, top

panels). On average, during the sampling period,

oxygen saturation was high in the lake inlet (*102%)

and main channel (*92%) and low in the wetland

inlet (*38%), whereas specific conductance was

higher in both lake and wetland inlets (191 and

206 lS cm-1, respectively) compared to the main

channel (148 lS cm-1) (Table 1).

Fish movement patterns

During the entire period of sampling, we caught a total

of 2,856 fish and 1,834 freshwater shrimp (Table 2).

Highest numbers of fish were captured at the inlet to

the lake (1,506) with about half the number in the

wetland (837) and fewer in the main channel (513).

Freshwater shrimp were most abundant at the inlet to

the wetland (1,029), followed by the main channel

(539) and lake (266). The majority of fish species, as

well as the freshwater shrimp, moved in both direc-

tions at all sampling locations, but the overall direction

of movement varied between different habitats.

Overall, the majority of fish moved downstream in

the main channel (77%) or into the lake (69%),

whereas for the wetland, similar numbers of fish

moved in both directions.

The frequency and direction of fish movements

varied both temporally (on both diel and seasonal

scales) and between different off-channel habitats.

Comparisons were made of abundance over time

between the different habitats and directions of

movement for the four most abundant native fish

species, as well as the abundant freshwater shrimp

(Fig. 2). Similar sampling depths and method allowed

abundance comparisons between different sites and

indicated species-specific behaviours. The movements

of shortfin eels were mostly nocturnal, whereas

ınanga, common bully and common smelt moved

mostly during the day (Fig. 2). In contrast, the

freshwater shrimp showed less diurnal variation in

directional movement, with similar numbers during

both day and night. High river discharge in spring

triggered movement of shortfin eel into the off-

channel habitats at night, with numbers of eels moving

into the wetland about fivefold higher than those

moving into the lake (Fig. 2; Table 3). In contrast,

common bullies were most abundant during low

discharge in summer and at the river inflow into the

lake, moving in both directions but with more than

70% moving into the lake (Fig. 2; Table 3). Inanga

moved into both the lake and wetland in similar

numbers, and their abundances at the inlets were about

twofold higher than in the main channel. Common

smelt was most abundant in the main channel. We

observed about twofold higher abundances of fresh-

water shrimp in the wetland compared to the main

channel, with significantly more individuals moving

out of the wetland after high flows (Fig. 2; Table 3).

Abundances of freshwater shrimp in the inlet to the

lake were lower than those of the main channel.

We also observed significant numbers of intro-

duced fish species at inlets to both off-channel

habitats. Gambusia was the third most abundant

species in the lake inlet (after native common bully

and shortfin eel), and moved in both directions. It was

also present at the other locations, but in very low

numbers (Table 2). We recorded numerous larvae of

non-native common carp moving out of the wetland

(Table 2). One individual each of brown bullhead

catfish and rudd was caught moving out of the wetland

(Table 2). There were no differences in size distribu-

tion of shortfin eel between habitats (Fig. 3), but we

found some indication of slightly higher numbers of

smaller common bully moving into the lake compared

with moving out of the lake (Fig. 3; Table 2), although

these differences were not statistically significant.

Results of the redundancy analyses showed that

habitat type, flow, water temperature, dissolved oxy-

gen and specific conductance explained much of the

variation in the abundance of fish moving in both

directions (51 and 59% for fish moving into and out

of the off-channel habitats, respectively; Table 4). We

observed consistent patterns in species—environment

associations for some common species (Fig. 4).

Common bully and gambusia were positively associ-

ated with lake habitat, higher dissolved oxygen

concentration and higher water temperature (summer

Hydrobiologia (2014) 729:161–174 165

123

Main channel

0

6

6

12

12

6

12

6

12

Lake

0

6

12

Wetland

0

6

12Anguilla australis

0

2

2 2 2

0

2

0

2

Galaxias maculatus

CP

UE

(n

o h

-1)

0

20

20

40

40

20

40

20

40

0

20

40

0

20

40

Gobiomorphus cotidianus

0

2

2

4

4

2

4

2

4

0

2

4

0

2

4

Retropinna retropinna

Oct Dec Feb Apr Jun Aug

0

10

10

20

20

10

20

10

20


0

10

20

NightDay


0

10

20

Paratya curvirostris

6

12

18

24

30

Wat

er le

vel (

m)

1

2

3

4

5

Daily water temperature (mean ± range)Daily discharge / Relative water level

Tem

per

atu

re (

°C)

6

12

18

24

30

1

2

3

4

5D

aily

dis

char

ge

(m3 s

1 )

0

400

800

1200

In

Out

In

Out

In

Out

In

Out

In

Out

In

Out

In

Out

In

Out

In

Out

In

Out

In

Out

In

Out

In

Out

In

Out

In

Out

-

Fig. 2 Waikato River discharge at Rangiriri, and water level

and temperature at Lake Whangape and Opuatia wetland

between September 2010 and September 2011 (top). Movement

dynamics of fish and freshwater shrimp (bottom) in the main

channel as well as at inlets to Lake Whangape and Opuatia

wetland between September 2010 and September 2011

166 Hydrobiologia (2014) 729:161–174

123

season) for both ‘in’ and ‘out’ movements. In contrast,

shortfin eel and common smelt moved mostly into the

wetland, especially with higher flows and lower water

temperatures (spring). Common carp moved mostly

out of the wetland habitat, mostly with higher flows

(spring).

Discussion

We observed significant numbers of fish moving to

and from both the riverine lake and wetland habitats

investigated in this southern hemisphere large river,

supporting the conclusion that lateral connectivity

between the main river channel and off-channel

habitats is a key feature of ecological integrity in

riverine ecosystems (Copp, 1989; Junk et al., 1989;

Hohausova et al., 2003). Moreover, the present study

highlights the importance of different off-channel

habitats (i.e. lakes, wetlands) for fish productivity as

different native species varied in their use of riverine,

lake or wetland habitats. In this study, we refer to

the exchange of biota between the main river channel

and off-channel habitats as ‘movements’ rather than

‘migrations’, defined as synchronised movements that

are large relative to the average home range for a

species and occur at specific stages of the life-cycle

(Lucas & Baras, 2001). Although it is possible that

some species interact with off-channel habitats during

migrations, most of the movements in the present

study likely occurred over a small scale in response to

environmental factors associated with inundation of

lateral habitats.

Flow triggers the movement of native fish

Movements of shortfin eels and ınanga into the

wetland were driven mostly by changes in river

discharge, especially spring floods, supporting our

hypothesis that seasonal variations in flow can play a

crucial role in triggering lateral movement of fish.

Similar behaviour has been observed in Europe for

cyprinids, which move from the main river channel

into tributary (Nunn et al., 2010) or floodplain (Grift

et al., 2001; Hohausova et al., 2003) habitats at high

flows. Directional movement of shortfin eels is likely

to be for opportunistic feeding (Chisnall, 1989;

Chisnall & Hayes, 1991; Lasne et al., 2008), while

ınanga could be searching for refugia from high flow

velocities (Schwartz & Herricks, 2005) or alterna-

tively seeking shelter in wetland habitats where

predation pressure by larger fish is likely to be lower

than in the main channel (McDowall, 1990).

Aquatic insects and large-bodied zooplankton that

do not normally grow in flowing waters (Baranyi et al.,

2002; Kim et al., 2002; Collier & Lill, 2008) can

flourish in wetlands and floodplain retention zones

where they may serve as a suitable food source for

ınanga and other small-bodied fish species (McDo-

wall, 1990; Rowe et al., 2002). In tropical river

systems, with highly predictable flood pulses, feeding

migrations during floods are widely documented

(Benech & Penaz, 1995; Fernandes, 1997; Wantzen

et al., 2002; Castello, 2008). These lateral movements

between the main channel and riverine lakes and

wetlands have also been shown to play an essential

role in exchange of organic carbon between off-

channel and main river food webs (Burford et al.,

2008; Hunt et al., 2012). Conversely, in the smaller

Manu River in Peru, with more unpredictable, short-

duration flood pulses, fish moving into the off-channel

lake had full gut contents more often than fish leaving

the lake, suggesting the main channel can also be a

preferred feeding habitat (Osorio et al., 2011).

Common bully movements occurred predomi-

nantly during low flow conditions and higher summer

temperatures, when mostly juvenile bullies were

moving into the lake. This indicates that the produc-

tive Lake Whangape, and potentially other riverine

lakes with intact connections to the main river, may be

important for common bully which has an excellent

ability to maintain feeding in waters with low water

clarity (Rowe, 1999). Lakes such as Whangape may

also serve as a juvenile nursery. Interestingly, in

contrast to the main river channel and wetland, we did

not observe larger ([7 cm) common bullies at the lake

inlet, suggesting that the more structurally complex

main channel and wetland habitats are more suitable

Table 1 Mean (SE) for water quality parameters measured at

the study sites during sampling period (n = 17)

Location Temperature

(�C)

Specific

conductance

(lS cm-1)

Dissolved

oxygen

(mg l-1)

Main channel 17 (1.3) 148 (4.3) 7.2 (0.4)

Lake 18 (1.5) 191 (12.3) 8.0 (0.5)

Wetland 16 (0.7) 206 (6.3) 3.8 (0.5)

Hydrobiologia (2014) 729:161–174 167

123

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au

ratu

s(L

inn

aeu

s1

75

8)a

Go

ldfi

sh–

–9

5.7

(0.6

)–

––

–4

5.5

(1.4

)4

5.6

(1.4

)

Cyp

rinus

carp

io(L

inn

aeu

s,1

75

8)a

Co

mm

on

carp

––

17

.8–

––

–3

1.1

41

1.2

(0.0

3)

Ga

mb

usi

aa

ffin

is(B

aird

&G

irar

d,

18

53

)aG

ambu

sia

22

.9(0

.6)

36

3.1

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)–

–1

2.1

77

2.9

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2.4

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)

Ga

laxi

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cula

tus

(Jen

yn

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84

2)

Inan

ga

10

5.7

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)1

85

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.2)

18

7.5

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4.4

(1.4

)2

15

.5(0

.3)

66

.5(0

.5)

Go

bio

mo

rph

us

coti

dia

nu

s(M

cDo

wal

l,1

97

5)

Co

mm

on

bu

lly

35

5(0

.7)

83

42

.4(0

.1)

12

34

.3(0

.2)

37

13

.1(0

.6)

29

63

(0.1

)3

05

4.7

(0.2

)

Ret

rop

inn

are

tro

pin

na

(Ric

har

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48)

Co

mm

on

smel

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.2(0

.2)

15

.71

58

.1(0

.4)

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(0.2

)2

66

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.3)

––

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rdin

ius

eryt

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phth

alm

us

(Lin

nae

us,

17

58)a

Ru

dd

––

––

––

––

––

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1.2

Pa

raty

acu

rvir

ost

ris

(Hel

ler,

18

62)

Fre

shw

ater

shri

mp

24

79

23

91

29

21

74

63

8

Av

erag

e(S

E)

tota

lle

ngth

(L)

of

spec

ies

caug

ht

isal

sosh

ow

na

Intr

odu

ced

spec

ies

168 Hydrobiologia (2014) 729:161–174

123

Table 3 PERMANOVA results conducted on CPUE for the fish community as well as four most abundant native fish and native

freshwater shrimp individually

Fish considered Source df SS Pseudo-F P

Community (all fish species) Location 2 7,556.4 5.5792 0.001

Date 5 8,860.6 2.6168 0.001

Location 9 date 10 10,482.0 1.5479 0.023

Residual 36 24,379.0

Anguilla australis Location 2 1,662.9 6.3287 0.004

Residual 36 4,729.7

Galaxias maculatus NS

Gobiomorphus cotidianus Location 2 4,649.0 13.914 0.001

Date 5 4,547.4 5.4441 0.001

Location 9 date 10 5,427.6 3.2489 0.002

Residual 36 6,014.1

Retropinna retropinna NS

Paratya curvirostris Location 2 958.0 4.0972 0.006

Date 5 2,687.6 4.5978 0.001

Location 9 date 10 8,588.8 7.3467 0.001

Residual 36 4,208.7

df Degrees of freedom, SS sums of squares, Pseudo-F distance-based pseudo F statistic, P probability values (obtained using 999

permutations of residuals under a reduced model)

Main channel

<10

10-1

920

-29

30-3

940

-49

50-5

960

-69

70-7

9

0

20

20

40

40

60

60

20

40

60

LakeAnguilla australis

<10

10-1

920

-29

30-3

940

-49

50-5

960

-69

70-7

9

Wetland

<10

10-1

920

-29

30-3

940

-49

50-5

960

-69

70-7

9

In

Out

In

Out

In

Out

Length class (cm)

<11-

1.92-

2.93-

3.94-

4.95-

5.96-

6.97-

7.98-

8.99-

9.9

10-1

0.9

11-1

1.9

12-1

2.9

Fre

qu

ency

(%

)

0

20

40

60

<11-

1.92-

2.93-

3.94-

4.95-

5.96-

6.97-

7.98-

8.99-

9.9

10-1

0.9

11-1

1.9

12-1

2.9 <1

1-1.

92-

2.93-

3.94-

4.95-

5.96-

6.97-

7.98-

8.99-

9.9

10-1

0.9

11-1

1.9

12-1

2.9

In

Out

In

Out

In

Out

Gobiomorphus cotidianus

Fig. 3 Length–frequency relationships of the two most abundant native species, shortfin eel (Anguilla australis) and common bully

(Gobiomorphus cotidianus), for each location and movement direction

Hydrobiologia (2014) 729:161–174 169

123

for adults of this species which can use aquatic

vegetation as a spawning substrate (McDowall, 1990).

Habitat complexity may also account to some

degree for the strong preference of freshwater shrimp

for wetland habitats with abundant vegetation

(Carpenter, 1982). Vegetated wetland habitats can

provide shelter from predation (Jordan et al., 1996;

Banha & Anastacio, 2011), as well as detritus and

periphyton as a source of food (Azim, 2005). Indeed,

shrimps moved in large numbers into and out of both

of the off-channel habitats, mainly during low flows

and high summer temperatures, suggesting dynamic

interactions between river and off-channel habitats for

this species, depending on environmental cues.

The role of temperature and water quality

As ectotherms, fish are generally more active at the

higher end of their preferred thermal optimum (Lucas

& Baras, 2001). This would explain the frequent

movements to off-channel habitats that we observed

during high discharge in spring, but not during winter

Table 4 Eigenvalues and cumulative percent variance explained

(in parentheses) from redundancy analyses (axes I–III) of habitat

type, flow magnitude and temperature explaining the abundance of

moving fish

Direction Pa Axes

I II III

In \0.01 0.450 0.036 0.015

(45) (48.6) (50.1)

Out \0.01 0.375 0.119 0.048

(37.5) (49.4) (54.3)

a Based on 999 permutations (test of significance of the first

canonical axis vs. all canonical axes gave the same results)

-1

0

1

-1 0 1

AX1

AX

2

G. cotidianus

G. maculatus

R. retropinna

A. australis G. affinis

Lake

Temperature

In

Wetland

Flow

Dissolved oxygen

Specificconductance

-1

0

1

-1 0 1

AX1

AX

2

G. cotidianus

G. maculatus

A. australis

G. affinis

LakeDissolved oxygen

Out

Wetland

Flow

C. carpio TemperatureSpecificconductance

Fig. 4 Redundancy

analyses of fish CPUE of

different species explained

by habitat type, flow

magnitude, temperature and

selected water quality

variables. The greater the

similarity in length and

direction of the vectors,

the stronger the association

between the abundance

of particular species and

associated environmental

characteristics

170 Hydrobiologia (2014) 729:161–174

123

when water temperatures remained low. Off-channel

habitats inundated during spring floods are often

warmer than the main river channel and thermally

heterogeneous (Gorski et al., 2010), suggesting inun-

dation may trigger movements in search of optimal

temperature regimes (Lucas & Baras, 2001). In

support of the role of thermal cues, adult barbel

Barbus barbus (Linnaeus, 1758) were shown to

progressively shift their diurnal pattern of feeding in

order to move to the foraging places at the time of the

day when water temperature is the closest to their

thermal optimum (Baras, 1995). In contrast to

Hohausova et al. (2003), who showed that water

quality parameters had little influence on fish move-

ment into off-channel habitats in the River Morava

(Czech Republic), we observed that variations in

specific conductance and dissolved oxygen between

different off-channel habitats could partly explain fish

movement into and out of these habitats. This finding

indicates that, after the governing roles of flow and

temperature are accounted for, other environmental

characteristics that define off-channel habitats (i.e.

water quality) may also play a role in initiating fish

movement. In support of this, North American

centrarchids overwintering in backwater lakes on the

upper Mississippi River were shown to move out of

lentic areas when oxygen levels dropped below

2 mg l-1 (Knights et al., 1995). Similarly, higher

numbers of common bullies moving into the lake

habitat in our study could, to some extent, reflect a

preference for turbid waters (Rowe, 1999). Factors

stimulating fish migration and movement are complex

(Lucas & Baras, 2001), with multiple cues working in

concert, and movement occurring when factors pro-

moting residency are outweighed by those stimulating

movement, irrespective of the physiological or envi-

ronmental nature of those cues (Ovidio et al., 1998).

Management implications

This study emphasises the importance of lateral

hydrologic connectivity and seasonal flooding of off-

channel habitats in the functioning of this southern

hemisphere temperate large river system, as demon-

strated by the numerous native fish using the lake and

wetland during the flood and at low flows. Therefore,

management strategies that promote connectivity

within lowland river–floodplain ecosystems and reha-

bilitate (semi-)natural flow dynamics are likely to

maximise habitat availability and feeding opportunities

for key life stages of native fish as well as providing

refugia at important times of year. Similar findings have

been shown for large European rivers (Lasne et al.,

2007), where numbers of native species increased with

increasing connectivity. Furthermore, body condition

of European eels Anguilla anguilla (Linnaeus, 1758)

has been shown to decrease with decreasing lateral

connectivity (Lasne et al., 2008), potentially because

inundated terrestrial habitats provided food of better

nutritional quality (Van Liefferinge et al., 2012). A

similar reliance on lateral connectivity with wetlands is

supported for shortfin eels in the lower Waikato River

(Chisnall & Hayes, 1991).

However, high numbers of non-native larval com-

mon carp moved out of the wetland with retreating

flood water, and the introduced gambusia was

recorded in both off-channel habitats through the

year. Maintenance of high flows and artificial floods

may potentially be important for preservation of native

fish communities in river systems in which most of the

introduced fish species are adapted to lentic conditions

(Bernardo et al., 2003). In the lower Waikato River,

however, flooded wetland habitats appear to serve as

superior spawning habitat for phytophilic common

carp whose larvae were highly abundant moving out

of the wetland with receding spring floods, as has

also been observed for this species in the Australian

Murray River (Stuart & Jones, 2006). Indeed, adult

common carp were recorded to move frequently to off-

channel habitats during spring spawning time in both

the Australian Murray River (Jones & Stuart, 2009)

and the lower Waikato River (Daniel et al., 2011).

Therefore, to be successful, management aimed at

enhancing ecological integrity must consider the

potential negative and ongoing implications of non-

native fish movement and reproduction in off-channel

habitats, which our study has shown can provide

important nurseries for invasive common carp. Para-

doxically, main channel–off-channel connections can

serve as natural movement bottlenecks for such

species and therefore may provide opportunities for

targeted control of non-native invasive fish species

during movement phases. Appropriate management

of floodplain ecosystems may therefore involve

implementing ‘controlled connectivity’ measures that

provide access for native species at key times while

limiting opportunities for introduced species to utilise

favoured off-channel habitats.

Hydrobiologia (2014) 729:161–174 171

123

Acknowledgments We thank Courtney Kellock, Warrick

Powrie, Ray Tana, Michael Pingram and Dudley Bell

(University of Waikato) for assistance during field sampling.

Funding for this work was provided by the Strategic Investment

Fund of the University of Waikato.

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