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Report to MW Klunzinger, SJ Beatty and AJ Lymbery Centre for Fish & Fisheries Research Murdoch University July 2011 FRESHWATER MUSSEL RESPONSE TO DRYING IN THE LOWER HELENA PIPEHEAD DAM & MUSSEL TRANSLOCATION STRATEGY FOR CONSERVATION MANAGEMENT

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Page 1: IN THE LOWER HELENA PIPEHEAD DAM & MUSSEL … · in the lower helena . report to . mw klunzinger, sj beatty and aj lymbery centre for fish & fisheries research murdoch university

Report to

MW Klunzinger, SJ Beatty and AJ Lymbery

Centre for Fish & Fisheries Research Murdoch University

July 2011

FRESHWATER MUSSEL RESPONSE TO DRYING

IN THE LOWER HELENA PIPEHEAD DAM &

MUSSEL TRANSLOCATION STRATEGY FOR

CONSERVATION MANAGEMENT

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FRESHWATER MUSSEL RESPONSE TO DRYING IN THE LOWER HELENA PIPEHEAD DAM & MUSSEL TRANSLOCATION STRATEGY FOR CONSERVATION MANAGEMENT

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FRESHWATER MUSSEL RESPONSE TO DRYING IN

LOWER HELENA PIPEHEAD DAM & MUSSEL

TRANSLOCATION STRATEGY FOR

CONSERVATION MANAGMENT

Report to

Swan River Trust

Frontispiece: Lower Helena Pipehead Dam (top and middle); a Carter’s freshwater mussel left open to bird predation after drying

(bottom). Photos: Michael Klunzinger, Stephen Beatty and Suzanne Thompson.

This report should be referenced as: Klunzinger, M.W., Beatty, S.J. & Lymbery, A.J. (2011). Freshwater mussel response to drying in the Lower Helena Piphead Dam & mussel translocation strategy for conservation management. Centre for Fish & Fisheries Research, Murdoch University Report to Swan River Trust.

ACKNOWLEDGEMENTS:

THIS PROJECT WAS FUNDED BY SWAN RIVER

TRUST. WE WOULD LIKE TO THANK

SUZANNE THOMPSON (SWAN RIVER TRUST)

FOR CO-ORDINATING THE PROJECT. THANKS

TO MARK PAGANO (WA DEPT. OF FISHERIES)

FOR ORGANISING LEGISLATIVE EXEMPTION

PERMISSION AND WATER CORPORATION

FOR COOPERATIVE CONSULTATION AND

ACCESS TO SITES. WE ESPECIALLY THANK

JAMES KELEHER FOR ASSISTANCE IN THE

FIELD AND GORDON THOMSON FOR

ASSISTANCE WITH HISTOLOGY

Authors:

MW Klunzinger, SJ Beatty &

AJ Lymbery

Centre for Fish & Fisheries Research, Murdoch

University

July 2011

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FRESHWATER MUSSEL RESPONSE TO DRYING IN THE LOWER HELENA PIPEHEAD DAM & MUSSEL TRANSLOCATION STRATEGY FOR CONSERVATION MANAGEMENT

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Executive Summary

The Helena River is a major tributary of the Swan River. Carter’s Freshwater Mussel,

Westralunio carteri, is currently listed as ‘Vulnerable’ on the International Union for the

Conservation of Nature Red List of Threatened Species and as a ‘Priority 4’ species by the

Department of Environment and Conserveration (WA). The species is the only freshwater

mussel native to the south-west and may play an integral role in maintaining water clarity and

quality through its filter-feeding habit. Indeed, in other parts of the world, freshwater mussels

have been shown to benefit water supplies through the remediation of the effects of

eutrophication. The species has declined in recent years primarily from increased salinisation

of waterways throughout its historic range. Maintaining existing populations is becoming

increasingly important. Recent research is beginning to show that the survival of the species is

becoming increasingly challenged by not only salinity, but also drought, chemical and nutrient

pollution, habitat loss, and factors that threaten host fishes (which are required as obligate

hosts for the parasitic larval stage *‘glochidia’+).

Historically, the species was found throughout the freshwaters of the Swan-Avon

catchment, based on museum records and early reports (pre-1970s). Current information on

the species distribution within the Helena system is sparse, but recent reports by Wetland

Research and Management, Swan River Trust and Murdoch University have found the mussel

existing above and, although to a far lesser extent, below the dam.

This study aimed to collate previously existing data on the freshwater mussels in the

Helena River, update species information within the Lower Helena Pipehead Dam and

translocate part of an existing population in the dam which would be exposed to drying and

increased predation during a drop in water volume following maintenance works.

A total of 1205 live freshwater mussels were hand-collected from the areas that were

likely to be most exposed during dam works, translocated to Pools 4 and 5, located 250-300 m

downstream from the dam, and placed in habitats which were deemed appropriate, based on

surveys conducted elsewhere within the Swan Coastal Plain.

Population measures including size (age) classes, densities and reproductive status were

determined during this study. Estimates of mortality from drying and predation were also

determined within the dam, where mussels had been exposed and, for comparison, of mussels

which were still submerged but would become exposed during dam works. The existence of

smaller shells suggests some recruitment, but the large number of old mussels in the dam

suggests that this is an aging population with few recent cohorts. Mortality on banks in which

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mussels had been exposed to direct sunlight, warm temperatures and extreme dying was

relatively high (25.61%), whereas mortality of mussels that remained submerged within the

dam and pools below the dam was only 0.31%. A significantly greater number of large mussels

were found on exposed banks than submerged (71 mm compared to 61 mm average shell

length, respectively). The loss of larger mussels is of concern, particularly because larger

mussels presumably have a greater filtration capacity as well as reproductive potential, with

larger females able to produce more offspring and large adults able to filter a greater volume of

water than smaller/immature individuals. There were also a number of shell deformities

(2.72%). Shell deformities are sometimes caused by prolonged infection by parasites, but more

often it is suggested that deformities occur from long periods of exposure to contaminants such

as heavy metals or agricultural or industrial chemicals.

The population composition of freshwater fishes within the dam is currently unknown,

but the authors did notice a large Koi Carp (Cyprinus carpio), many Eastern Gambusia

(Gambusia holbrooki), some Swan River Goby (Pseudogobius olorum) and anecdotal reports of

Redfin Perch (Perca fluviatilis) from Water Corporation staff. Recent work by the authors has

shown that the larvae of Carter’s Freshwater Mussel do attach to a majority of native

freshwater fish species, but to a lesser extent, if at all, on feral fish species such as C. carpio and

G. holbrooki. It is unknown whether glochidia attach to and survive to the juvenile stage on

Redfin Perch, Koi Carp or Eastern Gambusia within the Helena River.

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Summary & Recommendations

A preliminary survey of permanent pools (4 and 5) downstream from the dam showed

that mussels were present in low densities and there was suitable habitat with enough

space to accommodate densities typical of other localities in south-western WA.

1205 live mussels which would be at risk of dehydration and predation during dam

works were translocated from the dam to these pools to improve their chances of

survival.

The study has demonstrated that a sizeable population of Carter’s Freshwater Mussel

exists within the Lower Helena Pipehead Dam and these would contribute to

maintaining water quality in the reservoir.

The study showed that mussels exposed to drying in the dam were vulnerable to not

only dehydration, but also a greater risk of predation by other animals. This resulted in

a loss of at least 25% of the mussels which were surveyed.

It is recommended that translocated mussels in pools 4 and 5 be monitored for health

and freshwater fishes within the system be examined for glochidia during October to

January and micro-habitats be sampled for the presence of newly-released juveniles

during the summer. This information will indicate the recruitment success within the

system and viability of the population.

The population should be tested for levels of heavy metals, pesticides or other

contaminants to help determine the cause of shell deformities should be determined

and. If contaminants are recorded, their source should be located.

The value of freshwater mussels as biological filters should be quantified in a dedicated

study to determine filtration capacity and benefit in terms of maintaining water quality

in water supply reservoirs.

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Contents

Executive summary ...................................................................................................................................... 3

Contents ........................................................................................................................................................ 6

1. Introduction ................................................................................................................................... 7

1.1 Freshwater mussels ......................................................................................................................... 7

1.2 Lower Helena Pipehead Dam .......................................................................................................... 8

1.3 Aims of the study ............................................................................................................................ 9

2. Methods ...................................................................................................................................... 10

2.1 Study sites ..................................................................................................................................... 10

2.2 Mussel sampling and collection .................................................................................................... 10

2.3 Water quality sampling ................................................................................................................. 11

2.4 Statistical analysis ......................................................................................................................... 11

3. Results ........................................................................................................................................ 17

3.1 Mussel population parameters ..................................................................................................... 17

3.1.1 Shell structure ....................................................................................................................... 17

3.1.2 Mortality .............................................................................................................................. 22

3.1.3 Population density ................................................................................................................ 25

3.1.4 Reproductive status .............................................................................................................. 26

3.1.5 Estimated age structure ........................................................................................................ 26

3.2 Water quality ................................................................................................................................ 28

4. Discussion ................................................................................................................................... 28

4.1 Mussel mortality ........................................................................................................................... 28

4.2 Shell deformities ............................................................................................................................ 30

4.3 Population structure and viability ................................................................................................. 31

Summary and recommendations ...................................................................................................... 32

References ....................................................................................................................................... 33

Appendices ...................................................................................................................................... 36

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1. Introduction

There have been a number of projects in the last decade that have involved

management of aquatic fauna during drainage works within reservoirs of south-western WA to

mitigate negative impacts on resident species (Beatty et al. 2003a, 2003b, 2003c; Morgan &

Beatty 2004; Beatty & Morgan 2006; Molony et al. 2003, 2005; Beatty & Morgan 2006; Beatty

et al. 2006). However, this project is the first that has focussed on the freshwater mussel.

1.1 Freshwater mussels

Freshwater mussels (Unionoidea) are benthic bivalve molluscs. They have important

functional roles in aquatic ecosystems through biological filtration of water, contributing to

clarity and quality. They oxygenate sediments through their burrowing habit, provide structure

to stream banks and sediments, provide refuge for other freshwater life including juvenile

crayfishes and provide food for other animals (Vaughn and Hakenkamp 2001). Freshwater

mussels are sensitive to environmental changes (Ponder and Walker 2003) and are considered

important bio-indicators of aquatic ecosystem condition.

Virtually all freshwater mussel larvae are obligate parasites, normally using a fish as a

host (Bauer and Wächtler 2001). Mussel larvae (glochidia) attach to the gills, body surfaces

and/or fins of fish, remaining as parasites for a period of weeks to months before undergoing

metamorphosis to emerge as juvenile mussels (Bauer and Wächtler 2001). Fish native and

endemic to a particular region where unionoids occur are the usual hosts for their glochidia and

most unionoids are host generalists, able to utilize a variety of native and endemic host fish

species (Haag & Warren 1997, 2003; O’Brien & Brim Box 1999; Rogers et al. 2001; Layzer et al.

2002; Strayer 2008a). Transformation success to the juvenile stage of native glochidia is

generally unsuccessful or less successful when they attach to feral or non-endemic host fishes

(Bauer & Vogel 1987; Rogers & Dimock 2003; Dodd et al. 2005).

Carter’s freshwater mussel, Westralunio carteri Iredale, 1934 is the only freshwater

mussel species endemic to south-western WA. It was historically found from the Moore River

to the south coast, west of Esperance and may have been found as far north as the Gascoyne

River prior to the late 1800s (Klunzinger et al. unpublished data). Population decline of W.

carteri in salt-affected systems such as the Avon River (Kendrick, 1976) has resulted in its

current listing as ‘Vulnerable’ by the International Union for the Conservation of Nature (IUCN),

meaning the species is facing a high risk of extinction in the wild in the medium-term future,

under international criteria (IUCN 1996) and as a Priority 4 species, which is defined as taxa in

need of monitoring, under inter-departmental fauna rankings by the Department of

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Environment and Conservation (DEC), Government of Western Australia. Currently, there is a

lack of published information regarding the precise distribution, biology and ecology of W.

carteri in south-western Australia.

1.2 Lower Helena Pipehead Dam

The Helena River is a tributary of the Swan River within the South West Coast Drainage Division, WA. Much of the water collected in the Pipehead Dam has historically been pumped back to the Mundaring Weir and utilised for drinking water. (Elliot 1983; Siemon 2001; Spillman 2003)

Recent works within the Pipehead Dam have resulted in the need to empty much of the water to allow access for engineers to the scour valve. The Swan River Trust was concerned for the freshwater mussels that would be stranded on the banks of the reservoir and contacted Murdoch for technical advice. Murdoch University Freshwater Fish Group was contacted for technical advice. It was proposed that a large number be moved downstream into a permanent pool to ensure survival of a proportion of the population. Communication between Water Corporation, Swan River Trust and the Department of Fisheries resulted in an emergency action plan agreed upon in which Murdoch researchers and Swan River Trust staff would conduct an ecological survey followed by a translocation of at-risk mussels.

1.3 Aims of the study A group of Carter’s Freshwater Mussels (Westralunio carteri) from Canebreak Pool, Margaret River,

where average mussel densities can be in excess of 100 mussels/m2. Photo: MW Klunzinger

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The aims of this study were to:

Determine the population structure and abundance of freshwater mussels in the pools

that were identified as potential refuges for mussels from the dam and determine

whether there would be adequate habitat and space for the translocated mussels.

Translocate a large number of mussels most at risk to exposure during the draining of

the dam.

Estimate mortality of mussels which had already been exposed to extreme drying and

predation within the Lower Helena Pipehead Dam.

Determine existing population structure, abundance and viability of live mussels within

the dam.

Provide recommendations that will aid in ensuring the long-term viability of this

population.

Lower Helena Pipehead Dam (top and bottom left); a dead Carter’s Freshwater Mussel after drying (bottom right). Photos:

Michael Klunzinger and Suzanne Thompson.

2. Methods

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2.1 Study sites

The study area included 520 m2 within the Helena Pipehead Dam (Figure 1) as well as

Pool 4 and Pool 5, approximately 250-300 m downstream of the dam wall. To estimate

mortality of mussels within the area of the dam that was dry, a series of transects (n=6) were

placed according to the diagram in Figure 2. Prior to translocation of mussels from the dam,

Pool 4 and Pool 5 were surveyed for the presence of mussels and densities and population

structure were determined.

2.2 Mussel sampling and collection

Within Pools 4 and 5, 1 m2 quadrats (n=11 and 14, respectively), constructed using 15

mm diameter round PVC tubing, were placed randomly within wadeable areas of each pool and

the number of freshwater mussels within each were recorded to determine density. All

mussels were identified to species level according to McMichael and Hiscock (1958) and Walker

(2004). Within each quadrat, dominant benthic substrate type was recorded (rock, gravel,

sand, mud, silt and/or detritus). For each mussel collected from the quadrats, the maximum

length (ML), maximum height (MH) and width (W) of the shell was measured with vernier

callipers to the nearest 0.02 mm.

Within the Lower Helena Pipehead Dam, a series of six transects were undertaken, each

50 m in length, oriented parallel to the water’s edge and 2.5-14 m from the water on the area

of the dam which had been exposed to drying (see Figure 2). For each transect, on either side

of the transect line, 10 x 1 m2 quadrats were placed randomly and the number of mussels

within each transect were counted and measured as above. The status of each mussel

measured (live or dead) was also recorded. Live mussels reacted to handling by closing their

shells tightly. Mussels were collected, by hand, from a 400 m2 submerged area within the dam

(see Figure 2), which was predicted to dry following water release, and translocated into Pools

4 and 5. Prior to release, a sub-sample of these mussels (n = 150) was measured as above. A

sub-sample (n = 28) was kept for reproductive status, age determination and future genetic

analysis. All other live mussels from quadrats were translocated to Pool 4 and Pool 5. Mussels

were released to a density of up to a maximum of 20 mussels/m2, which is well within the

density range of W. carteri found elsewhere in similar habitats for this species.

Reproductive status and gender of sub-sampled mussels was determined in the

laboratory. The sex of each mussel was determined by examining the gills. In the inner portion

of the inner demibranchs of females, the interlamellar septa (marsupia) are notably thickened

to provide support for developing embryos/glochidia. Males do not have this gill feature. The

visceral mass (minus the gills and mantle) was cut from the base of the shell hinge and the foot

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was removed by trimming with surgical scissors. The resulting ‘gonad’ tissue was fixed in

Bouin’s fluid for 24 hours in preparation for histology. These tissue samples were dehydrated

in graded ethanols, embedded in paraffin, sectioned (6 μm thick) and stained with

haematoxylin and eosin.

Shells of dissected mussels, as well as a range of various size classes of empty (dead)

shells collected from the dam, were embedded in FR251 epoxy resin, sliced longitudinally

through the beak of each shell, and sections examined microscopically and the number of

internal shell rings on each section was counted. In other species where growth has been used

to verify age, these lines normally represent annual growth or age. Measured shells were also

examined for ‘umbone’ or ‘beak’ wear as the degree to which the ‘pointy’ region of the shell

surface was worn away. Scores ranged from I - IV with I being the most and IV being the least

amount of wear.

2.3 Water quality sampling

Water quality (temperature (°C), pH, dissolved oxygen (% and ppm), NaCl concentration

(ppt), total dissolved solids (ppt) and conductivity (µS/cm)) was measured using an Oakton™

PCD650 waterproof portable multimeter at three locations at each site and a mean and

standard error (SE) determined.

2.4 Statistical Analysis

Mapping of mussel distributions was undertaken using NearMap™ online imagery

(http://www.nearmap.com/), under a personal license agreement. Graphical and statistical

analysis of mussel densities, length-frequencies, shell rings, reproductive status and umbone

scores were undertaken using Excel and SigmaPlot™11.0. Differences were tested for

significance by Analysis of Variance; correlations between shell length and the number of

internal shell rings were tested using Spearman’s Rank Order Analysis; and comparisons

between the proportions of shells with various umbone scores was tested by Chi-square

analysis with SigmaPlot™11.0.

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Figure 1 Location of the Lower Helena Pipehead Dam; diagram showing locations of transects

within the dam and opaque shaded area (outlines in red) of submerged mussels which were collected

for translocation. N.B. Pools 4 and 5 are not shown, but are located 250-300 m downstream from the

dam wall.

Transects

1-4 Transects

5-6

Lower Helena

Pipehead Dam

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Figure 2 Location of transects sampled for freshwater mussels along the dry portion of the

shoreline within the sampling area of the Lower Helena Pipehead Dam; (a) Transects 1-4; (b) Transects

5-6. N.B. diagrams are not drawn precisely to scale and images were taken on 20 April 2011 (4 weeks

prior to sampling), after which time, water levels had dropped and a greater area of bank was exposed.

a b

4

1

2

3

6 5

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Figure 3 Examples of submerged areas sampled for freshwater mussels in May 2011, including,

(a), (b), (c) Pool 5; (d), (e), (f) Pool 4, downstream from the Lower Helena Pipehead Dam. Photos:

Stephen Beatty

a

b

c f

e

d

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Figure 4 Morphological measurements for the Freshwater Mussels.

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Figure 5 Examples of submerged areas sampled for freshwater mussels in May 2011, within (a),

(b) the Lower Helena Pipehead Dam; (c) mussel collection nylon mesh dive bag; (d) upstream end of the

dam (extent of mussel population) facing downstream towards dam wall (north-west); (e) view from

Transect 1 facing upstream (south-east); (f) condition of dam water – turbid; (g) view from Transect 1

facing downstream towards dam wall (north-west); (h) view from Transect 1 facing tree line and access

track on east bank; (i), (j) view from Transect 5 (west bank) facing downstream towards dam wall (north-

west); (k) extent of area surveyed for mussels within the dam – view facing downstream towards dam

wall (north-west). Photos: Michael Klunzinger.

a b

d e

f g h

i j k

c

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Figure 6 Shell umbone wear scores given in Roman numerals; severity ranging from I (most) to

IV (least). N.B. The ‘umbone’ or ‘beak’ area of the shell is the point which lies directly below the

numeral in each image.

3. Results 3.1 Mussel population parameters

3.1.1 Shell structure

A total of 544 freshwater mussels were measured for MH, ML and W. Mean MHI

(=MH/ML) was 61.38%, which confers to W. carteri. Shell deformities were observed in 2.72%

of mussels collected (Figures 7 and 8), resulting in abnormal MHI (74.89%), causing

misidentification of the species.

IV

II

III

I

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Figure 7 A side-by-side comparison of a normal shell (left) and deformed mussel shell (right) of

Westralunio carteri, both from Lower Helena Pipehead Dam.

Figure 8 Examples of shell deformities (left) and normal shells (right) of Westralunio carteri collected from

Lower Helena Pipehead Dam.

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Overall shell length frequencies from each sampling area are given in Figure 9. Mean

shell length of submerged mussels (61.56 mm) was significantly smaller than mussels which

were exposed within transects (70.88 mm) (t = 10.335, df = 513, P < 0.001). Mussels sampled

from Pool 5 were smaller than those sampled from Pool 4 (55.48 vs. 68.79 mm, respectively) (t

= 3.664, df = 29, P < 0.001). Submerged mussels from the dam were larger than those from

Pool 5 (t = 2.360, df = 157, P = 0.020), but smaller than those from Pool 4 (t = -4.089, df = 168, P

< 0.001).

The proportion of mussels with each of the umbone wear scores is given in Figure 10.

Umbone scores from Transects 1 and 2 were not recorded, but a majority would have scored as

IV and the remainder as III. Excessive umbone wear (i.e. scores of I or II) was not observed in

Transects 3, 4, 5 or 6. Shell umbone wear was greater in Pool 4 or Pool 5 than within the dam

(χ2 = 37.833, df = 5, P < 0.001). Across all transects, the greatest proportion of shells had

umbone scores of IV and very few shells had scores of less than III (χ2 = 83.026, df =7, P <

0.001). Within Pool 4, there was no difference between the proportion of shells with umbone

scores of I or II (χ2 = 2.667, df = 1. P = 0.102), but there was a greater proportion of shells with

an umbone score of III, compared to the proportion of shells with an umbone score of IV (χ2 =

15.676, df =1, P < 0.001). Also, in Pool 4, very few shells were free of umbone wear, but there

was a greater proportion of shells with a score of III compared with scores of I and II (χ2 =

21.192, df = 3, P < 0.001). Within Pool 5, there was no difference between the proportion of

shells with umbone scores of I and II or III and IV (χ2 = 5.867, df = 3, P = 0.118).

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Transect 1

Shell Length (mm)

30 40 50 60 70 80 90

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Transect 2

Shell Length (mm)

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Transect 3

Shell Length (mm)

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Shell Length (mm)

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Dam

Shell Length (mm)

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Shell Length (mm)

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22Pool 5

Shell Length

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a

b

c

Figure 9 Shell length frequency histograms of Westralunio carteri; (a) Samples collected 2.5-5.0 m from shoreline (Transects 1,

2, 5); (b) Samples collected 7-14 m from shoreline (Transects 3, 4, 6) within the Lower Helena Pipehead dam; (c) submerged mussels

within the dam, Pool 4 and Pool 5.

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Figure 10 Proportion of individual Westralunio carteri with umbone scores (I – IV) within each

sampling site.

0

20

40

60

80

I II III IV

Pro

po

rtio

n o

fM

uss

els

(%)

Umbone Score

Transect 3

0

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I II III IV

Pro

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(%)

Umbone Score

Transect 4

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Pro

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(%

)

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Transect 5

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Pro

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(%

)

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Transect 6

0

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Pro

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(%)

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Pool 4

0

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Pro

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(%

)

Umbone Score

Pool 5

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Table 1 Mean density (±95% C.I.) of Westralunio carteri in Pool 4 and Pool 5, Helena River and in

the Lower Helena Pipehead Dam.

Site Locality Area

(m2)

No. live

mussels

Density of

live mussels

(number/m2)

No. dead

mussels

Density of

dead mussels

number/m2

(±95% C.I.)

Transect 1 Dam banks 20 32 1.60 (±0.60) 4 0.20 (±0.25)

Transect 2 Dam banks 20 51 2.55 (±1.08) 17 0.85 (±0.53)

Transect 3 Dam banks 20 41 2.05 (±0.84) 30 1.50 (±0.47)

Transect 4 Dam banks 20 89 4.45 (±1.69) 25 1.25 (0.45)

Transect 5 Dam banks 20 29 1.45 (±0.69) 9 0.45 (±0.39)

Transect 6 Dam banks 20 37 1.85 (±1.11) 9 0.45 (±0.42)

Submerged Dam 400 966 2.42 3 0.01

Submerged Pool 4 14 16 1.14 (±1.26) 1 0.05 (±0.15)

Submerged Pool 5 11 9 0.82 (±0.73) 1 0.05 (±0.20)

3.1.2 Mortality

Proportion mortality in each sampling area is given in Table 1. Mussel mortality was

greater in Transect 3 than any other transect (χ2 = 9.539, df = 5, P = 0.008). There was no other

differences in mussel mortality when comparing Transects 1, 2, 4, 5, and 6 (χ2 = 2.823, df = 2, P

= 0.244). Mortality of mussels within the submerged area of the dam was negligible compared

to the level of mortality observed on the shore within Transects 1-6 (χ2 = 294.434, df = 6, P <

0.001). Although mortality in Pool 4 and Pool 5 was greater than within the submerged area of

the dam (χ2 = 26.110, df = 2, P < 0.001), this was based on a much smaller sample size (n = 29

vs. n = 969, respectively). Side-by-side comparisons of length frequencies of live compared to

dead mussels for each sampling transects are given in Figures 11 and 12. Shell length of dead

mussels within Pools 4 and 5 were 69.52 mm (although this shell was deformed) and 39.84 mm,

respectively; dead shells within the submerged area of the dam were not measured.

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Transect 1 (LIVE)

Shell Length

30 40 50 60 70 80 90

Num

ber

of M

ussels

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18 Transect 1 (DEAD)

Shell Length (mm)

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Transect 2 (LIVE)

Shell Length (mm)

30 40 50 60 70 80 90

Num

ber

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18 Transect 2 (DEAD)

Shell Length (mm)

30 40 50 60 70 80 90

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ber

of M

ussels

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18

Transect 3 (LIVE)

Shell Length (mm)

30 40 50 60 70 80 90

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18 Transect 3(DEAD)

Shell Length (mm)

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ber

of M

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18

Figure 11 Shell length frequency histograms of Westralunio carteri sampled from Transects 1-3

with live mussels in the left figures and dead mussels in the right figures.

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Transect 4 (LIVE)

Shell Length (mm)

30 40 50 60 70 80 90

Nu

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18 Transect 4 (DEAD)

Shell Length (mm)

30 40 50 60 70 80 90

Num

ber

of M

ussels

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18

Transect 5 (LIVE)

Shell Length (mm)

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Nu

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18 Transect 5 (DEAD)

Shell Length (mm)

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ussels

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Transect 6 (LIVE)

Shell Length (mm)

30 40 50 60 70 80 90

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Shell Length (mm)

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18

Figure 12 Shell length frequency histograms of Westralunio carteri sampled from Transects 4-6

with live mussels in the left figures and dead mussels in the right figures.

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3.1.3 Population density

Population density, given as the number of mussels within each sampling area (m2) is

shown in Table 2. Differences in mean live mussel densities between sampling sites were

significant (F = 5.261, df = 8, 143, P < 0.001) as were differences in dead mussel densities (F =

6.884, df = 8, 140, P < 0.001). Pair wise multiple comparisons (Bonferroni t-tests) are given in

Appendix II.

Table 2 Mortality of Westralunio carteri in the Lower Helena Pipehead Dam, Pool 4 and Pool 5, Helena River.

Site Locality No. live mussels

No. dead mussels

Total no. mussels

Mortality % (95% C.I.)

Transect 1 Dam banks 32 4 36 11.11 (3.11-26.07)

Transect 2 Dam banks 51 17 68 25.00 (15.28-36.99)

Transect 3 Dam banks 41 30 71 42.25 (30.61-54.57)

Transect 4 Dam banks 89 25 114 21.93 (14.72-30.65)

Transect 5 Dam banks 29 9 38 23.68 (11.44-40.25)

Transect 6 Dam banks 37 9 46 19.57 (9.35-33.92)

Submerged Dam 966 3 969 0.31 (0.06-0.91)

Submerged Pool 4 16 1 17 5.88 (0.31-28.73)

Submerged Pool 5 9 1 10 10.00 (0.52-44.64)

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3.1.4 Reproductive status

Of the mussels dissected in the laboratory (n = 28), 15 were males and 13 were females,

as confirmed by histology (Figure 13). No eggs, embryos or glochidia were present in the gill

marsupia, but gonads contained eggs or sperm. In two of the mussels with deformed shells,

the gills were also deformed. Some orange deposits, which might indicate heavy metal

deposits, were found on the mantle tissues and gills of the mussels which were examined, but

without chemical analysis no conclusions can be made. These tissues are currently frozen and

stored at the Murdoch University Fish Health Unit.

Figure 13 Examples of histological sections of (a) male (20x); (b) female (20x) gonads of W. carteri.

3.1.5 Estimated age structure

Without validating the incremental growth of the mussel and confirming that growth lines are

deposited annually, it is impossible to be sure that shell rings truly represent age. However,

studies of different species of freshwater mussels in temperate climates elsewhere have shown

rings do represent annual growth, which is discussed below. In this study, the number of

internal shell rings significantly increases with shell length in normal shells (see Figure 14),

suggesting that larger mussels are older. In deformed shells however, the relationship between

the numbers of internal rings compared to shell length was not significant (P > 0.05).

(a) (b)

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Shell length (mm)

40 50 60 70 80 90

Nu

mb

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of

inte

rna

l sh

ell

rin

gs

0

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24

40 50 60 70 80 90

0

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18

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22

24

Figure 14 Regression plots of shell length vs. number of internal shell rings of Westralunio carteri

sampled from the Lower Helena Pipehead Dam. Regression plot (grey dashed with triangles) is

deformed shells and in black are normal shells. Linear regression analysis indicates that the number of

internal shell rings increases positively with shell length in normal shells (P<0.001), but not in deformed

shells (P = 0.267). Regression equations and r2 values are given for each plot.

Y = -11.087 + (0.335x) r2 = 0.822 F = 69.322, df = 16, P < 0.001

Y = -0.441 + (0.207x) r2 = 0.151 F = 1.425, df = 9, P = 0.267

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3.2 Water quality

Table 3 Physico-chemical parameters of water quality in the Lower Helena River.

Site Time Date Temperature (°C) Conductivity

(mS/cm)

NaCl

(ppt)

pH Dissolved

Oxygen (%)

Pool 4

1:26 PM 20 May 2011 15.36 0.9574 0.50 7.19 89.8

Pool 4

8:43 AM 24 May 2011 12.69 0.9259 0.48 8.09 91.7

Pool 5 1:35 PM 20 May 2011 15.32 0.9365 0.49 7.29 86.85

Pool 5 8:25 AM 24 May 2011 12.88 0.9230 0.48 7.92 90.1

Helena

Pipehead Dam

2:04 PM 20 May 2011 16.51 0.8846 0.46 7.82 107.8

Helena

Pipehead Dam

10:12 AM 24 May 2011 11.99 1.053 0.55 7.38 23.9

4. Discussion

4.1 Mussel mortality

From our surveys, it is evident that the primary cause of mortality within the dam was

exposure to drying and heat, with 25.2% of the 373 exposed mussels having died, compared to

just 0.31% of 969 submerged mussels within the dam. The precise time of death of these

exposed mussels is also uncertain. From the time series aerial photos in Plates 1-3 (Appendix I),

a significant drying event occurred in August 2010 which closely resembles the dam levels

observed during the drying in this study. Undoubtedly, mussel death may have occurred at this

time as well. Although there was some evidence of predation, this would require additional

investigation to quantify.

There have only been a few documented studies on freshwater mussels’ response to

drought. From the little amount of information available, some Australian hyriids, such as

Velesunio ambiguus and Velesunio angasi, may be well-adapted to aestivation for extended

periods, particularly those occurring in temporary floodplain billabongs and ephemeral streams

in remote inland areas (Walker 1981; Humphrey 1984; Sheldon and Walker 1989). The main

factor in survival is the mussel’s capability to utilize anaerobic respiration. Ch’ng-Tan (1968)

found that V. ambiguus survived for one year out of water. Walker et al. (2001), however,

notes that Australian hyriids are not widespread in ephemeral or salinised water. The ability of

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freshwater mussels to tolerate drought is enhanced by shells that fit tightly together and/or

encased in moist mud (Walker et al. 2001). Most mussels which had found their way into deep

cracks within drying mud in this study remained alive, but those which were exposed to direct

sunlight and mud that baked dry, had died. The lack of shade and habitat, such as leaves and

woody debris may have also exacerbated the problem.

In North America, where freshwater mussels were exposed to record drought

conditions, mortality rates ranged from 14-90%, depending on species and habitats available.

The presence of woody debris, shade, cooler groundwater inputs and the ability to burrow into

moist sediments assisted in mussel survival (Miller and Payne 1998; Golladay et al. 2004;

Gagnon et al. 2004; Haag 2008). Indeed, Velesunio angasi is known to survive the dry season by

sheltering in dense roots mats of riparian trees in the Alligator Rivers region of the Northern

Territory (Humphrey 1984). Storey and Edward (1989) suggest that W. carteri may survive in

intermittent streams by sealing their shells and awaiting the next flow event. This may be the

case in some habitats. In our surveys of this species throughout the south-west, we have rarely

found the mussel to exist in ephemeral systems and are generally restricted to areas that

receive flow throughout the year. In regulated systems surveyed in 2010-2011, we found that

the mussel is able to survive drying by burrowing into soft mud and sand that does not collapse

as mussels burrow, essentially creating ‘tubes’ which remain moist as water levels recede.

Where we have found live mussels in dry conditions, shells have always been tight fitting and

sealed with a mucus plug, as described by Storey and Edward (1989), but have always been in

the presence of woody debris, moist plant debris and/or been in the presence of moist

sediment, generally in shaded areas. In areas where mussels do not have these habitats for

protection, are exposed to direct light, high and dry ambient temperatures and sediments that

bake dry to a hard pan, they seldom survive and are also open to predation. Under laboratory

conditions, mussels left to dry in receding waters within sand substrate during the summer of

2010, had internal shell temperatures which reached a maximum of 60°C; cumulative

mortalities were 76% after 5 days and 88% after 10 days of drying. These mussels were

exposed to open sun without shade, simulating the most extreme of drought conditions.

Drought conditions and increased temperatures, combined with algal growth, are also

known to cause hypoxia and even anoxia in some systems which also impacts the survival of

freshwater mussels (Walker 1981; Humphrey 1984; Sheldon and Walker 1989; Miller and Payne

1998; Gagnon et al. 2004; Golladay et al. 2004; Haag 2008). Freshwater mussels are

particularly vulnerable to hypoxic stress during periods of gravidity when the gills have to

support brooding glochidia (Aldridge and McIvor 2003). It is also well known that elevated

water temperatures decreases dissolved oxygen concentrations. Other factors, such as

hypocalcemic conditions (low calcium), low pH and pollutants are also harmful to freshwater

mussels (Bauer and Wächtler 2001; Strayer 2008). Recently (summer 2011), a mass kill of W.

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carteri in the Lower Canning River presumably resulted from hypoxia or anoxia where dissolved

oxygen measured 30.87% saturation in the afternoon. This level was probably much lower

during night time, when algae (of which there was a large concentration) switches to aerobic

respiration, thus using much of the already limited amount of dissolved oxygen. This likely

resulted from lack of flow, excessive nutrient inputs and stagnation, which put the mussels

under hypoxic stress and resulted in death, similar to studies reported by Gagnon et al. and

Golladay et al. 2004. From the water quality measurements tested in this study, we observed

that within the dam, dissolved oxygen was supersaturated (107.8%) in the afternoon and very

low (23.9%) in the morning, which suggests high algal loads. In pools 4 and 5, however, this

was not the case and stability of oxygen in these pools increased their suitability as

translocation sites.

4.2 Shell deformities

There are few data available on the chronic effects of metal exposure on growth in

freshwater mussels (Naimo 1995). Lasee (1991), however, reported that juveniles of Lampsilis

cardium exposed to cadmium (Cd) concentrations as low as 0.01 mg/kg significantly reduced

anterior shell growth. Shell curling, such as the individual in Figure 7 has been reported in

marine mussels exposed to tributyltin (TBT - a compound found in marine anti-fouling paints)

(see Batley and Scammell 1991). Other shell deformities have also been reported for other

marine and estuarine bivalves exposed to heavy metals (Sunila and Lindström 1985; Nias et al.

1993; Yap et al. 2002). Another major consequence of environmental contamination includes

loss of reproductive function. For example, a study Humphrey (1995) indicated that

reproduction and glochidia release in the freshwater mussel Velesunio angasi was negatively

affected by water released from Ranger Uranium Mine to Magela Creek, Northern Territory.

We are aware of only one other study which reported similar shell deformities in freshwater

mussels; these were from areas prone to agricultural and household chemical use in southern

New York Strayer (2008b).

The differences in umbone scores in this study are more likely due to a prolonged

difference in both flow regime and sediment type. The sediments within the dam were

composed mainly of clay and soft mud, which undoubtedly resulted in higher umbone scores

than shells from Pool 4 and Pool 5. Pool 4 sediments were composed of a sandy base with

areas of deep detritus and sediments in Pool 4 had sandy bottoms with detritus. These pools

were also more likely to experience greater flow velocities than within the dam. The

combination of coarse sediments and greater water turbulence probably resulted in more shell

wear, as suggested by Hinch and Green (1988) as well as Roper and Hickey (1994). We also

observed blistering of the periostracum (outer shell covering) in most of the shells that had

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been exposed to light, heat and drying. Excessive wear on the umbone area weakens the shell

and could lead to an increased vulnerability to avian predation (Vestjens 1972).

4.3 Population structure and viability

The occurrence of mussels less than 40 mm in length suggests recruitment has occurred

within the last 5-10 years (based on unpublished data), although numbers of mussels in this size

class were relatively low. The transects were dominated by mussels in the 70+ mm size classes,

whereas most submerged mussels within the dam were less than 70 mm in length. This

suggests that larger, presumably older mussels may be less mobile in terms of receding water

levels; perhaps smaller mussels may normally occupy deeper waters or that predation may

have eliminated smaller mussels which were exposed to drying. Vestjens (1972) found that

Australian white ibis tend to be size selective on the freshwater mussels they predate. Without

baseline data on the usual distribution of mussels within the dam when at higher storage level,

it is difficult to make any clear conclusions.

Coker et al. (1921), Negus (1966), Ahlstedt (1979), Neves & Widlak (1987) and Neves et

al. (1980) established that mussels less than 20 mm in length are either difficult to find in

nature or that recruitment in freshwater mussel populations is sporadic or has been less

successful in recent years (however, we are confident that our manual sampling method would

have adequately recovered these smaller individuals). The loss of native host fishes required

for the mussels’ life cycle may have contributed to the apparent low level of recruitment in the

dam. From our studies (unpublished data), we have found that the occurrence of glochidia on

native fishes is much higher than on feral, introduced species such as Eastern Gambusia

(Gambusia holbrooki), One-spot Livebearer (Phalloceros caudimaculatus) and virtually nil in

Goldfish (Carassius auratus). Thus, in systems dominated by feral fishes, the lack or rarity of

smaller mussels may be indicative of poor recruitment as suggested by Aurajo and Ramos

(2000). In systems where a wide variety of native candidate host species occur, such as the

lower reaches of the Collie River and where feral species are virtually absent, such as Margaret

River, recruitment is more evident by the presence of a greater number of smaller mussels than

was observed in this study.

To date, accurately predicting the age of W. carteri at various lengths within various

systems has not been documented, but is one area of our research focus. The use of annual

growth rings as an estimate of age is supported, particularly in temperate climates such as

northern Europe (Bauer 1992; Schöne et al. 2004; Helama et al. 2006; Helama and Valovirta

2008) and has been validated in >20 North American species using mark-recapture methods

(Neves and Moyer 1988; Howard and Cuffey 2006; Haag and Commens-Carson 2008). Within

the Australasian region, maximum ages are reportedly 13-35 years (Grimmond 1968; Walker

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1981; Humphrey 1984; James 1985; Ogilvie 1993; Roper and Hickey 1994;). Although several

authors have reported the age in other unionoids ranging from 20 to 100 or more years old,

based on growth lines, there has been no consensus on the actual age of very large individuals

(i.e. maximum sizes) (see Anthony et al. 2001) and is still a debatable topic (Haag 2009).

Furthermore, growth interruption lines can arise from environmental factors, such as flood and

drought (e.g. Walker et al. 2001), environmental contamination or change in water chemistry

(Roper and Hickey 1994), which may not be annual events. So, without validating age with

growth we cannot accurately predict age-at-length within the Lower Helena Pipehead Dam,

however, using the number of growth lines in various size classes relative to one another is

probably a useful tool reflective of relative recruitment history.

Example of a Carter’s Freshwater Mussel (Westralunio

carteri) exposed to drying which has been hammered

open at the weak point of a worn umbone by an

unknown avian predator. Photo: Michael Klunzinger.

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Summary and recommendations

This was, to our knowledge, the first comprehensive ecological study of the response of

W. carteri to drying within a reservoir in WA.

The results revealed that considerable mortality of freshwater mussels had already

occurred from previous draining of Lower Helena Reservior.

We recommend flow releases from the dam should ensure water levels within Pool 4

and Pool 5 are maintained and follow-up monitoring of translocated mussels occur to

ensure they survive through future drying periods.

Sampling of the fish community should be conducted in November/December 2011

within the dam and within Pool 4 and Pool 5 and be examined for glochidia to

determine future recruitment potential within these sites.

Sampling microhabitats in March 2012 for juvenile mussels would enable recruitment

success to be determined.

We also recommend rigorous testing for possible source contaminants of deformed W.

carteri that were found in this study.

The likely benefits W. carteri provide in terms of providing water filtration should be

quantified.

Knowledge gained from this study will be useful in developing management plans for

the conservation of this species and during future surface water management projects.

As has occurred previously (Beatty et al. 2003a, 2003b, 2003c; Morgan & Beatty 2004;

Beatty & Morgan 2006; Molony et al. 2003, 2005; Beatty & Morgan 2006; Beatty et al.

2006), future drawdowns of water supply reservoirs should involve early development

of aquatic fauna management plans to mitigate negative impacts on resident

populations of native species, including Carter’s Freshwater Mussel.

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APPENDIX I

Temporal changes in the volume of water

within Lower Helena Pipehead Dam

July 2010 - April 2011

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Plate 1 Lower Helena Pipehead Dam water levels (a) 20 July 2010; (b) 1 August 2010; (c) 19

September 2010; (d) 23 October 2010. Photos were obtained from NearMap™ under a private user

license.

a b

c d

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Plate 2 Lower Helena Pipehead Dam water levels (e) 30 November 2010; (f) 13 December 2010;

(g) 8 January 2011; (h) 15 February 2011. Photos were obtained from NearMap™ under a private user

license.

e

h g

f

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Plate 3 Lower Helena Pipehead Dam water levels (i) 14 March 2011; (j) 20 April 2011. Photos

were obtained from NearMap™ under a private user license. N.B. Photos for May 2011 are not yet

available.

j i

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APPENDIX II

Significant pairwise multiple comparisons

(Bonferroni t-tests) of mussel densities

between sampling sites within Lower Helena

Pipehead Dam and Translocation Pools.

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Mussel Density

Comparisons

(T = Tansect)

Difference of Means t-statistic P - value

T3 dead vs. submerged

Dam dead

1.49 3.281 0.047

T3 dead vs. Pool 4 dead 1.43 4.945 <0.001

T3 dead vs. Tiger Snake

Pool dead

1.41 4.528 <0.001

T3 dead vs. T1 dead 1.30 4.958 <0.001

T3 dead vs. T5 dead 1.05 4.005 0.004

T3 dead vs. T6 dead 1.05 4.005 0.004

T4 dead vs. Pool 4 dead 1.18 4.079 0.003

T4 dead vs. Tiger Snake

Pool dead

1.16 3.724 0.010

T4 dead vs. T1 dead 1.05 4.005 0.004

T4 live vs. Tiger Snake

Pool live

3.63 4.949 <0.001

T4 live vs. Pool 4 live 3.31 4.855 <0.001

T4 live vs. T5 live 3.00 4.853 <0.001

T4 live vs. T1 live 2.85 4.610 <0.001

T4 live vs. T6 live 2.60 4.206 0.002

T4 live vs. T3 live 2.40 3.882 0.006