Thé
Victorian

Naturalist

Volume 119 (1) February 2002

Published by The Field Naturalists Club of Victoria since 1884

From the Editors

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The

Victorian
Naturalist

Volume 119 (1) 2002 February

Executive Editor: Merilyn Grey
Editors: Alistair Evans and Anne Morton

Research Reports The Fish of Kororoit Creek — Stressed Relicts and City Slickers,
Dy Pau GA CLO SAM UR set ee recap savin uh d Everett rer AP e UIS 4

The Mammal Fauna of Remnant Native Grasslands of the
Western Basalt Plains and Northern Plains of Victoria,

"yy eter lie ts (Cle a7 Bee ae Ay n TU HE RT RTRS 14
How Do Nectar Foraging Butterflies Select Flowers?
O MSSE De VIII Ts, OPE certs ieee etas zi been: 21
Contributions Hooded Plover Thinornis rubricollis Chick Attacked by
Conspecifics, by V Teoh and MA Weston .............. eene 27
Invertebrates of Mount McKay — a Brief Survey, by EJ Grey ........ 29
Naturalist in the The Biodiversity Blitz, by Ken Green... DO
Mountains
Honours Australian Natural History Medallion 2001
— Alan Bridson Cribb, by Jan Endersby................ sess. 38
Tributes Graham Martin Pizzey, by Sheila Houghton ............... nenne 39
Book Reviews Wyperfeld: Australia's First Mallee National Park,
by Geoff Durham, reviewed by Sara Maroske............... eee 40

How to Identify Wildflowers of the Grampians,
by Ken Woodcock, reviewed by Margaret Corrick............... sess. 44

New Guidelines for Authors at the end of this issue

ISSN 0042-5184

ne ae
Cover: Dr Alan Cribb, the 2001 Australian Natural History Medallion recipient, with his
wife Dr Joan Cribb, (ANH Medallion recipient in 1994), Photo: Wendy Clark. Empathy

Photographics.

Web address: http://www.vicnet.net.au/-fncv/
email: fnev@vicnet.net.au

Research Reports

The Fish of Kororoit Creek — Stressed Relicts and City Slickers

Paul G Close!

Abstract

Kororoit Creek is a metropolitan watershed in which human disturbances may have reduced the
health of aquatic fauna populations. Identifying changes in the distribution, diversity and abundance
of aquatic fauna is hindered by a paucity of biological information, especially with regard to fish.
Intensive surveys conducted during January 2000 quantified the species richness and abundance of
fish and decapod crustacea at six sites in Kororoit Creek. A total of nine freshwater and one estuar-
ine fish species was collected, of which five were native species representing approximately 50% of
the native taxa expected to occur in the catchment. The relative abundance of exotic species tended
to be high in the lower catchment and very low in the mid- to upper catchment. This study identifies
a number of anthropogenic factors affecting aquatic fauna populations in Kororoit Creek. (The

Victorian Naturalist 119 (1), 2002, 4-13)

Introduction

With increasing urbanisation and indus-
trialisation, there is increasing pressure on
the ecological integrity or ‘health’ of
aquatic ecosystems (e.g. Klein 1979;
Growns er al. 1998; Sonneman et al. 2001;
Walsh ef al. 2001). Until recently, few
investigations into the ecology of urban
watersheds and their aquatic fauna have
been reported, making assessment of
human disturbances on these environments
difficult. In particular, there is little infor-
mation regarding the diversity, abundance
and distribution of fish species in urban
systems. Furthermore, historical fish data
are often compromised by survey method-
ology and design. For example, whilst the
health of Kororoit Creek has been
described based on macroinvertebrate pop-
ulations (Papas er al. 2000) and physical
and chemical characteristics such as
instream and riparian habitat and water
quality (Mitchell 1990; Melbourne Water
1997), previous studies of fish in the catch-
ment are limited by age (McKenzie and
O'Connor 1989) and geographical extent
(McGuckin 1999; DNRE 2000).

Nevertheless, there is increasing recogni-
tion of the intrinsic value of urban streams
and their associated flora and fauna. Basic
biological data, such as species richness,
abundance and distribution can provide
important information for the conservation
of values such as biological diversity,
research and education, and aestheties. In
an attempt to determine the health of

! Arthur Rylah Institute for Environmental Research,
PO Box 137, Heidelberg, Victoria 3084

4

streams in the Melbourne area and identify
priority management areas, Melbourne
Water Corporation has established a
stream health assessment program includ-
ing ongoing monitoring programs such as
the Melbourne Water Water Quality
Monitoring Network, Biological
Monitoring Program and the Tributary
Investigation Program. These Programs
combine a range of stream health indica-
tors (e.g. fish and macroinvertebrate
assemblages and water quality) at various
temporal and spatial scales. In 1999,
Melbourne Water commissioned the
Arthur Rylah Institute for Environmental
Research to assess the current state of fish
assemblages in Kororoit Creek as part of
the Tributary Investigation Program.

This paper reports the species richness,
abundance and distribution of fish and
decapod crustacea in Kororoit Creek and
identifies anthropogenic disturbances
potentially impacting on the integrity of
aquatic fauna populations. This provides
valuable base-line information with which
future monitoring events may be compared.

Methods
Study area

Kororoit Creek catchment covers an area
of approximately 250 km" (Fig. 1). Stream
flow is intermittent, with zero discharge
occasionally recorded at the hydrographic
station near Deer Park (station number
5509) (McGuckin 1999). Although stream
flow is intermittent and ephemeral in the
upper reaches, Kororoit Creek is dominat-
ed by long and relatively deep pools,
which may provide refugia for aquatic

The Victorian Naturalist

Research Reports

Fig. 1. Location of sites in Kororoit Creek where aquatic fauna surveys were undertaken.

fauna during periods of low flow if
instream habitat becomes discontinuous.
The catchment relief is relatively low, with
the headwaters rising just south of
Gisborne at an altitude of approximately
460 m above sea level (m ASL) in basalt
country of the Great Dividing Range.

Vol. 119 (1) 2002

Kororoit Creek is approximately 80 km in
length and flows in a south-easterly direc-
tion through rural farmlands in the head-
waters and mid-catchment and
urban/industrial areas in the mid-catch-
ment and coastal lowlands. Kororoit Creek
flows into Port Phillip Bay near Altona.

Research Reports

Quantification of site attributes

Five study reaches were selected along
the length of Kororoit Creek (Fig. 1), based
primarily on altitude and distance to river
mouth. Within each study reach, a single
survey site was established to represent the
available aquatic habitat attributes, includ-
ing flow type and cover elements. At site 3,
an additional survey reach was established
to include a reach of riffle habitat.
Although riffle habitat is scarce in the mid-
reaches of Kororoit Creek, riffles often sup-
port different assemblages of fish species
(see Matthews 1998 for a recent review).

For each survey reach, the mean width
and mean maximum depth (m) was calcu-
lated from at least five measurements of
each dimension. Spot measurements of
water quality parameters were recorded
within each survey reach. Water tempera-
ture (°C) and electrical conductivity
(uS/em at 25°C) were measured with a
WTW LF 320 meter, dissolved oxygen
(mg/L) with a WTW OXI 320 meter and
pH with a WTW pH 320 meter.

The relative abundance (% of wetted
area) of hydraulic units, habitat attributes
and cover elements was estimated for each
survey reach. Hydraulic units at each site
were defined using the following cate-
gories: cascade, rapid, riffle, glide, run,
pool or backwater (after Anderson and
Morrison 1989; Anderson er al. 1989).
Substrata were described in terms of the
percentage composition of various particle
size groups, namely boulder (particle size
2256 mm), cobble (64-255 mm particle
size), pebble (16-63 mm particle size),
gravel (2-15 mm particle size), sand (0.1-2
mm particle size) and silt/clay (particle
size «0.1 mm). The relative abundance of
dominant cover elements, including sub-
stratum, woody debris (logs, branches),
leaf litter (including bark), bank overhang,
vegetation overhang and aquatic vegeta-
tion, was also estimated and recorded.

Aquatic fauna survey

Surveys were conducted during January
2000. AIl sites (except site 2) were sur-
veyed using a consistent array of several
different net types including single-winged
fyke nets, fine mesh larval nets (200 um
mesh), and bait traps. A combination of a
single cyalume light stick and a small

quantity of cat food was placed in each
bait trap to attract fish. The wetted time
(fishing time) for each net was recorded.
Site 2 and the riffle habitat at site 3 were
surveyed using a two-pass bank-mounted
(Smith-Root® 7.5GPP electrofishing unit)
electrofishing technique with fine mesh
nets blocking the downstream and
upstream margins of the survey reach,
Voltage settings ranged from 170 to 340 V
to produce approximately 10 amps. For
these sites, the length of the survey reach
(m) and the electrofishing time (min; total
time during which electrical current is
applied to the water) taken to complete
each pass were recorded.

All fish and decapod crustacea collected
were identified and counted. Nomenclature
for fish species follows Allen (1989) and
Paxton ef al. (1989), and Horwitz (1990)
for decapod crustacea.

Analyses
Catch per unit effort, fish biomass and
density

Estimates of catch per unit effort (CPUE)
were calculated for both netting surveys
and electrofishing surveys. For netting sur-
veys, CPUE was calculated for each net-
ting technique and expressed in terms of
the number of fish collected per hour. The
wetted time for individual nets was
summed to give a total wetted time and
was used in the calculation of total CPUE
for each site. CPUE for electrofishing sur-
veys was also expressed in terms of the
number of fish collected per hour using the
summed time of two electrofishing passes.
Estimates of biomass (g/m^) and density
(fish/m^) were only calculated for elec-
trofishing surveys.

Results
Site descriptions

The five survey sites were distributed
evenly throughout the catchment in terms
of altitude and distance from river mouth
(Fig. 1; Table 1). The uppermost site (site
5) was located near the headwaters of
Kororoit Creek at an altitude of 155 m
ASL and 40 km from the river mouth.
Sites 4 and 3 were located in the mid-
catchment at altitudes of 100 and 50 m
ASL and 32 and 15 km to river mouth,
respectively. Site 2 was located on the

The Victorian Naturalist

Research Reports

Table 1. Characteristics of the stream, reach and instream habitat for each of the study sites located
on Kororoit Creek. Substrata and instream cover are defined in the text.

Habitat Survey
Characteristics

Site 5

Site 4

Site 3 Site 3 Site 2 Site 1
* *

Stream Characteristics
Elevation (m ASL)

Distance to river mouth (km)
Discharge ML/day

Reach Characteristics

Mean stream width (m)

Mean depth (m)

Water Quality

Temperature (?C)

Electrical conductivity (uS/cm)
Dissolved oxygen (mg/L) 2
pH 7.1
Substratum (Composition %)

Sheet rock -
Boulder
Cobble
Pebble -
Gravel
Sand
Silt
Clay -
Instream Cover (% of wetted area)
Substrate (Rock)

Logs/Log jams

Branches/Branch piles

Leaf litter

Bank overhang

Vegetation overhang

Urban rubbish

Aquatic vegetation

Flow Type (%)

Rapid/cascade

Run -
Riffle =
Glide
Pool
Backwater -
Reservoir/lake -
Tidal t

Un Un o3

tas IT

Un

a

* denotes sites surveyed using electrofishing techniques.

coastal lowlands 6 km from the river
mouth at an altitude of 15 m ASL. Site 1
was located on the coastal margin, 3 km
from the river mouth at an altitude of 5 m
ASL. This site represented the upper limit
of estuarine influence.

Except for site 2, instream substrata were
dominated (>50%) by finer particles, name-
ly sand and silt/clay, with the remaining
substrata comprised of boulders and/or cob-
bles (Table 1). The substratum at site 2
comprised approximately equal proportions
of silt (40%) and coarser particles of cob-
bles and boulders (50% combined).
Instream habitat that may provide cover for

Vol. 119 (1) 2002

aquatic fauna varied between sites. At the
majority of sites, cover was predominantly
emergent aquatic vegetation (Typha sp. and
Phragmites australis) and rocky substratum
(Table 1). At sites 3, 4 and 5 cover was pre-
sent in the form of overhanging banks and
overhanging terrestrial vegetation. Instream
woody debris (logs and branches) only con-
tributed small amounts of cover and was
present at all sites except site 2 (Table 1).
Stream discharge was recorded for each
survey site on the day of survey except for
sites 4 and 1 where a suitable discharge
transect could not be established.
Discharge estimates varied between sites

7

Research Reports

"— L——————— — —Ó QM
Table 2. Freshwater native and exotic fish species previously recorded from Kororoit Creek, and

their current conservation status.

Species Common Name Conservation Reference
Status

Native

Anguilla australis* Short-finned Eel CW McKenzie and O'Connor (1989)
MeGuckin (1999)

Galaxias maculatus* Common Galaxias CW McKenzie and O'Connor (1989)
McGuckin (1999)

Nannoperca australis Southern Pvgmy Perch CW DNRE (2001)'

Philvpnodon grandiceps Flat-headed Gudgeon CW McKenzie and O'Connor (1989)
MeGuckin (1999)

Pseudaphritis urvillii* ^ Tupong CW DNRE (2001)!
McGuckin (1999)

Retropinna semoni Australian Smelt CW McKenzie and O'Connor (1989)

Macquaria australasica
Exotic

Macquarie Perch

Carassius auratus Goldfish
Gambusia holbrooki Eastern Gambusia
Tinea tinea Tench

FFG listed

DNRE (2001)

* Freshwater migratory; ^ Translocated population; ' Recorded by Museum of Victoria; C =
Common, W = Widespread ° Unpublished survey results (Cadwallader 1981); Conservation status

follows DNRE (2000).

and ranged from zero near the headwaters
(site 5) to 3.8 ML/day at site 2 in the mid-
catchment (Table 1). Water quality para-
meters also varied between sites (Table 1).
Dissolved oxygen was generally low, rang-
ing from 2.2 mg/L at the uppermost site
(site 5) to 9.0 mg/L at site 2 (Table 1).
Electrical conductivity was high at all sites
(71500 uS/cm) except the uppermost site
(site 5) (Table 1). At all sites pH was alka-
line and tended to increase slightly in alka-
linity closer to the river mouth.

Historical data

Few fish surveys have been conducted in
Kororoit Creek, with most records of
species richness and distribution from
McKenzie and O'Connor (1989) and
MeGuckin (1999), Ten species of fish have
previously been recorded in the system
(DNRE 2001) (Table 2). Seven species are
native, and of these, three are considered
diadromous (i.e. migrate between freshwa-
ter and estuarine marine habitats at particu-
lar stages of their life; Table 2). The
Macquarie Perch Maequaria australasica
is native to Australia but is not endemic to
Kororoit Creek. While 50 Macquarie Perch
were stocked in 1912 at Sunshine, this
species has not subsequently been recorded
(DNRE 2001). A recent collection of
Tupong Pseudaphritis urvillii in Kororoit
Creek (McGuckin 1999) is the first record

8

of this species since 1894 by the Museum
of Victoria (DNRE 2001), The only record
of Southern Pygmy Perch Nannoperca
australis is ftom 1934 and the specimen is
held at the Museum of Victoria.

Three exotic fish species have previously
been recorded from Kororoit Creek (Table
2). Tench Tinca tinea were recently col-
lected in Kororoit Creek for the first time
since they were stocked in 1914
(MeGuckin 1999; DNRE 2001).

Survey results
Species distribution

Aquatic fauna surveys were conducted
over a period of 8 days between 17 and 25
January 2000. A total of nine species of
freshwater fish was recorded, of which five
(55%) were native (Table 3). Three of
these native species are diadromous. One
estuarine species, Tamar River Goby
Afurcagobius tamarensis, was collected at
the most downstream site (site 1). Two
genera of decapod crustacea were collected
in Kororoit Creek. Although no specimens
were collected, an additional genus of
decapod crustacean, Engaeus, was also
recorded by the presence of their charac-
teristic burrows.

Four exotic fish species were collected in
Kororoit Creek (Table 3) including Carp
Cyprinus carpio, another new species
record for Kororoit Creek.

The Victorian Naturalist

Research Reports

DJ" RR
Table 3. Relative abundance (%) of fish species and total number of decapod erustacea collected at

each study site.

Species Site5 Site4 Site3 Site3 Site3 Site2 Site I
N N N EF EF+N EF N

Teleosts

Anguillidae

Anguilla australis (Richardson) - 10.2 3.7 5.9 4.2 10.0 40.9

Galaxiidae

Galaxias truttaceus (Valenciennes) -- -- - 2.8 F4 -

Galaxias maculatus (McDowall) 4 : 4K4 78.8 23.5 47,5 10.6

Retropinnidae

Retropinna semoni (McDowall) - L4 1.4 3,1 1.9 32 7.0

Cyprinidae

Carassius auratus (Linnaeus) * - E r4 4.9 1.5 12,3 r4

Cyprinus carpio (Linnaeus) * - - - - - 13717 xS

Tinca tinca (McDowell) * : B wa 3 Kk - v

Poeciliidae

Gambusia holbrooki (Girard) * - F4 r4 2.1 F4 12.8 4,2

Gobiidae

Philypnodon grandiceps (McCulloch) - 87.8 93,2 24 66.8 r4 17.6

Afurcagobius tamarensis (Johnston) E - = - - - 3.5

Total Number of Fish 0 147 1036 — 425 1461 219 142

Decapod Crustacea

Parastacidae

Cherax destructor 149 | E | l - -

Atyidae

Paratya australiensis - Present Present Present Present Present Present

Number of Species 1 6 8 9 10 8 10

Habitat survey characteristics are defined in Table 1. / indicate the presence of the species at low
abundances (<1%). * denotes exotic species. N, netting survey. EF, electrofishing survey.

Species composition and richness varied
throughout the system. At most sites a
range of aquatic fauna species was record-

ed, except at site 5 in the upper reaches of

the system where only one species,
Common Yabby Cherax destructor, was
collected. At the site immediately down-
stream (site 4), a total of six species was
recorded. Aquatic fauna assemblages were
most diverse throughout the middle and
lower reaches of the system (sites 1, 2 and
3). Species richness ranged from eight to
ten species at each of these sites.
Short-finned Eel Anguilla australis were
collected at all sites except the uppermost

site (site 5). The relative abundance of

Short-finned Eel was greatest (approxi-
mately 4096) at the most downstream site
(site 1). At all other sites, the relative
abundance ranged from 3.7% at site 3 to
10.2% at site 4.

Spotted Galaxias Galaxias truttaceas
were collected only at site 3 (12 individu-
als) using bank-mounted electrofishing

Vol. 119 (1) 2002

techniques. This is the first documented
record of this species in Kororoit Creek.

Common Galaxias Galaxias maculatus
were collected at three sites located in the
mid to lower catchment (sites 1, 2 and 3)
with relative abundance estimates ranging
from <1 to 78.8%. The highest relative
abundance was recorded at sites surveyed
using electrofishing techniques (sites 2 and
3). Estimates of relative abundance from
netting surveys were relatively low (1194).

Australian Smelt Aetropinna semoni
were collected at all sites except the upper-
most site (site 5). The greatest relative
abundance (7%) was recorded at the most
downstream site (site 1). Relative abun-
dance estimates were similar (approxi-
mately 3%) in the two reaches (sites 2 and
3) surveyed using bank-mounted elec-
trofishing techniques.

Flat-headed Gudgeon Philypnodon
grandiceps were recorded at all sites
except the uppermost site (site 5). Relative
abundance was high (780946) at sites sur-

Research Reports

veyed with netting equipment (sites 4 and
3) and lowest at the electrofishing survey
sites 2 and 3, Young-of-the-year fish were
numerically dominant in samples from
sites 3 and 4 (Close 2000).

Tamar River Goby were restricted to
estuarine habitats and only three individu-
als were collected from the most down-
stream site (site 1). Tamar River Goby
comprised 3.5% of the total number of fish
collected at this site.

Of the exotic fish species collected,
Eastern Gambusia Gambusia holbrooki
was the most widespread, occurring at all
sites where fish were caught. Estimates of
relative abundance were generally low
(<5%), except at site 2 where they con-
tributed 12.8% of the total catch. Goldfish
Carassius auratus exhibited a similar dis-
tribution and relative abundance, being col-
lected at three sites (sites 1, 2 and 3) with
the highest relative abundance (12.3%)
recorded at site 2. Tench were only collect-
ed at two sites (sites 3 and 1) and were pre-
sent in low abundance. Similarly, Carp
were only collected from the two most
downstream sites (sites | and 2).

Catch per unit effort, fish biomass and
density

The results of netting and electrofishing
surveys were different in terms of both
species richness and species abundance,
and catch per unit effort (CPUE) (c.f.
Tables 4 and 5). CPUE for netting surveys
at four sites ranged from 0.34 to 2.66
fish/hr (Table 4). In comparison, CPUE for
electrofishing surveys conducted at sites 2
and 3 was 114.3 and 269.1 fish/hr respec-
tively (Table 5). At site 3, where both net-
ting and electrofishing survey techniques
were employed, both these techniques
recorded similar species except for Spotted
Galaxias and the Common Yabby that were
only collected by electrofishing (Table 5),

The efficiency of the different netting
techniques used in this survey varied
between sites, although larval nets proved to
be the most efficient netting technique at all
sites (Table 4). Flat-headed Gudgeon domi-
nated species abundance and CPUE for lar-
val nets. At site 5, only one species, the
Common Yabby, was recorded in larval
nets. The only large-bodied native fish
recorded in Kororoit Creek was the Short-

10

finned Eel. Other large-bodied fish present
in the catchment were exotic species, name-
ly Carp, Tench and Goldfish. Fyke nets rep-
resented the only efficient technique to cap-
ture these larger-bodied species.

Discussion
Species distribution and abundance

The composition and species richness of
aquatic fauna assemblages varied between
survey sites within Kororoit Creek. In gen-
eral, species richness was greatest in the
mid- to lower catchment (sites 3, 2 and 1).
Species richness decreased with increasing
distance from the creek mouth to a mini-
mum at the uppermost site (site 5) where
only one species, Common Yabby, was
collected. The relative abundance of exotic
species in aquatic fauna populations
ranged from zero to 38.8% and generally
increased with increasing distance from
source.

The collection of Spotted Galaxias and
Carp represent the first documented
records of these two species in Kororoit
Creek. In a recent survey of the system by
McGuckin (1999), Tupong was collected
in a fyke net survey near Clarkes Road,
Rackbank. Tupong was the only species
recently recorded in the catchment that
was not collected in this study. While there
are historical records of Southern Pygmy
Perch and Macquarie Perch, both from the
early 1900s, neither species was collected
in the present study.

The low abundance of the migratory
Tupong recorded by McGuckin (1999) and
the absence of Tupong and Southern
Pygmy Perch in the present survey is of
conservation concern, as the reasons for
their poor representation in fish assem-
blages is unknown. In Victoria, Tupong
are considered diadromous, with adult fish
migrating downstream to estuaries to
spawn during autumn and winter ( Andrews
1996). Upstream migration of juvenile
Tupong from estuaries into riverine reach-
es has also been described during spring
and summer. Only one instream barrier to
fish movement is documented on Kororoit
Creek (MeGuckin 1999). Although this
barrier may allow fish passage at some
high flows (MeGuckin 1999), its presence,
along with other possible barriers, may
contribute to the low abundance of Tupong

The Victorian Naturalist

Research Reports

Table 4. Results of netting surveys including the number and type of netting equipment used at each
site, pooled wetted time, and abundance (in parentheses) of each species collected. Catch per unit
effort (CPUE) for each net type is expressed in terms of number of fish caught per hour of wetted net
time.

Type and Pooled Total Number of CPUE Total
Number Wetted Species Fish/Decapod fish/hr CPUE
of Nets Time (hrs) Crustacea fish/hr
Site 1
Fyke - 5 96.7 Aa (58), Cc (7), Tt (1) 66 0.7
Mesh 3" - 1 19.3 Ca (1), Cc (4) 5 0.3
Mesh 4” - 1 19.3 Co) 2 0.1 0.82
Larval - 2 38.7 At (2), Cc (4), Gm (15), 62 1.6
Gh (6), Pg (26), Rs (10)
Bait Trap - 10 193 At (3), Pg (4) 7 0.05
Site 3
Fyke - 8 156 — Aa (37), Gm (1), Pg (1), Tt (4) 43 03
Larval - 2 39 Aa (1), Pg (961), Gm (5), 987 25.3 2.66
Gh (3), Rs (14), Tt (3)
Bait Trap - 10 195 Ca (1), Gm (2), Pg (4) 7 0.05
Site 4
Fyke - 9 184.5 Aa (13), Pg (4) 17 0.1
Larval - 2 41.0 Aa (2), Cd (1), Gh (1), 116 2.8 0.34
Pg (110), Rs (2)
Bait Trap- 10. — 205 Pg (15) 15 0.1
Site 5
Fyke -4 69 Cd (24) 24 0.3
Larval - 2 34.5 Cd (71) 71 2.1 0.54
Bait Trap - 10 has Cd (54) 54 0.3

cae ae lf LE M) NN EE e
Abbreviations denote species: Aa, Short-finned Eel Anguilla australis; At, Tamar River Goby
Afurcagobius tamarensis; Ca, Goldfish Carassius auratus; Cc, Carp Cyprinus carpio, Gh, Eastern
Gambusia Gambusia holbrooki; Cd, Common Yabby Cherax destructor; Gm, Common Galaxias
Galaxias maculatus; Gt, Spotted Galaxias Galaxias truttaceus; Pg, Flat-headed Gudgeon
Philypnodon grandiceps; Rs, Australian Smelt Retropinna semoni; Tt, Tench Tinea tinca.

DG Ag came MON" CU CLAU DM i
Table 5. Results of electrofishing surveys at sites 2 and 3, including electrofishing time, area sam-
pled, species diversity, abundance and biomass. Estimates of fish biomass, density and catch per unit
effort (CPUE) are also provided.

Sample Area Species Biomass Total Density CPUE
time Sampled Biomass g/m* Number fish/m* fish/hour
(hrs) (m?) (g) of Fish

Site 2
1.92 546 Aa (2774) 7.08 Aa (22) 0.40 114.3
Ca (406.2) Ca (27)
Cc(323.7) Cc (30)
Gh (15.5) Gh (28)
Gm (661.7) Gm (104)
Pg (4.6) Pg (1)
Rs (8.1) Rs (7)
Site 3
1.58 362 Aa (1773) 9.0 ^a (25) 1.2 269.1
Ca (470.3) Ca 21)
Cd (7.7) Cd (1)
Gh (5.3) Gh (9)
Gm (960.8) Gm (335)
Gt (50.7) Gt (12)
Pg (2.9) Pg (10)
Rs (1.3) Rs (13)

o
Abbreviations denote species: Aa, Short-finned Eel Anguilla australis, Ca, Goldfish Carassius aura-
tus; Ce, Carp Cyprinus carpio; Gh, Eastern Gambusia Gambusia holbrooki; Cd, Common Yabby
Cherax destructor; Gm, Common Galaxias Galaxias maculatus; Gt, Spotted Galaxias Galaxias trut-
taceus; Pg, Flat-headed Gudgeon Philypnodon grandiceps; Rs, Australian Smelt Retropinna semoni.

Vol. 119 (1) 2002 11

Research Reports

Table 6. The proportion of species recorded (bold) at each site expressed as a percentage of the total
number of expected species (non-bold) and the relative abundance (%) of exotic fish at each site.

Site Expected native % of % exotic
species expected abundance

I Aa, Gau, Gm, Gt, Mm, Pa, Pg, Pm, Pu, Rs 50 20.4

2 Aa, Gau, Gm, Gt, Mm, Na, Pa, Pg, Pm, Pu, Rs 45.5 38.8

3 Aa, Cd, Gau, Gm, Gt, Mm, Na, Pa, Pg, Pu, Rs 54.5 27

4 Aa, Cd, Gau, Gm, Gt, Mm, Na, Pa, Pg, Pu, Rs 45.5 0.6

5 Aa, Cd, Gm, Na, Pa, Pg, Rs 14.3 0

* denotes migratory species. Abbreviations denote species: *Aa, Short-finned Eel Anguilla australis;
Cd, Common Yabby Cherax destructor; *Gau, Pouched Lamprey Geotria australis; *Gm, Common
Galaxias Galaxias maculatus; Gt, Spotted Galaxias Galaxias truttaceus; *Mm, Short-headed
Lamprey Mordacia mordax; Na, Southern Pygmy Perch Nannoperca australis; Pa, Freshwater

Shrimp Paratya australiensis; Pg, Flat-headed Gudgeon Philypnodon grandiceps;

*Pm, Australian

Grayling Prototroctes maraena; Rs, Australian Smelt Retropinna semoni; *Pu, Tupong

Pseudaphritis urvillii.

by inhibiting recruitment of young fish
migrating upstream from estuarine areas.

Southern Pygmy Perch has not been
recorded in Kororoit Creek since 1934,
although it has been collected in nearby
tributaries of the Werribee River (DNRE
2001). Kororoit Creek is considered to
have a significant amount of potentially
suitable habitat (abundant submerged/
aquatic vegetation) for Southern Pygmy
Perch (pers, obs.) and its absence in sam-
ples from recent surveys is currently unex-
plained.

Impacts on the Kororoit Creek System

Numerous factors potentially affect the
environmental condition of Kororoit
Creek, and more specifically, populations
of aquatic fauna. In general, these impacts
relate to the whole catchment, although
there are additional disturbances that are
site specific. Impacts on aquatic fauna pop-
ulations in Kororoit Creek include urbani-
sation and industrialisation, mainly in the
mid- to lower reaches of the catehment;
clearing of native riparian vegetation and
the introduction of exotic plant species;
bed and bank instability, mainly in agricul-
tural areas in the upper catchment; barriers
to fish migration; introduction of exotic
species; and alteration of the natural flow
regime.

As with many other aquatic ecosystems in
Victoria (Koehn and O'Connor 1990),
habitat loss or alteration is considered a pri-
mary factor that has an impact on the health
of aquatic fauna populations in Kororoit
Creek (Close 2000). To assess the condi-

12

tion of fish populations, a list of taxa
expected to occur at each site on Kororoit
Creek was compiled using available histori-
cal survey data (DNRE 2001) and refined
using knowledge of distribution and habitat
requirements of Victorian fish species. The
richness of freshwater fish species at sites
located on the coastal margin (site 1) and
coastal lowlands (site 2) was approximately
50% of the total expected species richness,
and the relative abundance of exotic
species ranged from 20.4 to 38.8% (Table
6). Similarly, approximately 50% of the
species expected to occur at sites in the
mid- to upper catchment (sites 3 and 4)
were recorded in this study, although the
relative abundance of exotic species was
very low (2.7 and 0.6%, respectively), The
species richness recorded from the upper
catchment (site 5) in this study represented
only 14.3% of what was expected to occur.
No exotic species were recorded at this site.

The composition of fish assemblages
may vary temporally and respond to varia-
tions (natural or anthropogenic) in stream
discharge, water quality, habitat structure
and the abundance of food as a result of
variations in spawning, recruitment and
recolonisation success (see Matthews 1998
for a recent review). Nevertheless, the
species composition of fish assemblages
recorded in this survey appears altered
from that which would be expected to
occur naturally in Kororoit Creek: native
species richness appears reduced and exot-
ic species have been introduced. Those
species not recorded in the current survey
which are expected to occur in the catch-

The Victorian Naturalist

ment are Pouched Lamprey Georria aus-
tralis, Short-headed Lamprey Mordacia
mordax. Southern Pygmy Perch
Nannoperca australis, Tupong Pseuda-
phritis urvillii and Australian Grayling
Prototroctes maraena. Whilst ongoing
monitoring will provide a better under-
standing of the dynamics of the Kororoit
Creek fish assemblage, the data discussed
here, coupled with the identification of
potentially numerous anthropogenic
impacts on the health of Kororoit Creek,
suggest that the current fish assemblage
may represent stressed and relict native
species as well as exotic species or ‘city
slickers? which may be more resilient and
resistant to disturbed environments.

Acknowledgements

Thanks to Melbourne Water Corporation for
commissioning this study. Thanks also to Peter
Fairbrother for fieldwork assistance and Rhys
Coleman (Melbourne Water), the Arthur Rylah
Institute internal review panel and two anony-
mous referees for comments on an earlier ver-
sion of the manuscript.

References

Allen GR (1989) Freshwater Fishes of Australia. (TFH
Publications: New Jersey)

Anderson JR and Morrison AK (1989) Environmental
flow studies for the Wimmera River, Victoria, Part
B. Fish habitat assessment. Arthur Rylah Institute for
Environmental Research, Technical Report Series No
74. Department of Conservation, Forests and Lands,
Victoria.

Anderson JR, Hill IM and Morrison AK (1989)
Environmental flow studies for the Wimmera River,
Victoria. Part E. Technical appendices. Arthur Rylah
Institute for Environmental Research, Technical
Report Series No 77. Department of Conservation,
Forests and Lands, Victoria.

Andrews AP (1996) Family Bovichtidae Congolli. In
Freshwater Fishes of South-eastern Australia, pp
198-200. Ed RM McDowall. (Reed Books:
Chatswood)

Close PG (2000) An Assessment of the Aquatic Fauna
of Kororoit Creek, Victoria Consultancy report to
Melbourne. Water. Department of Natural Resources
and Environment, Victoria.

DNRE (2000) Threatened Vertebrate Fauna in
Victoria. Department of Natural Resources and
Environment, Victoria,

Research Reports

DNRE (2001) Victorian Aquatic Fauna Database. Fish
survey data for Kororoit Creek, Laverton Creek,
Skeleton Creek, and the Werribee River including
tributaries. Department of Natural Resources and
Environment, Victoria.

Growns IO, Pollard DA and Gherke PC (1998)
Changes in river fish assemblages associated with
vegetated and degraded banks, upstream of and with-
in nutrient-entiched zones. Fisheries Management
and Ecology 5, 55-69.

Horwitz P (1990) A taxonomie revision of the species
in the freshwater craylish genus Engaeus Erichson
(Decapoda: Paratacidae). Invertebrate Taxonomy 4,
427-614.

Klein RD (1979) Urbanization and stream quality
impairment, Water Resources Bulletin 15, 948-963.
Koehn JD and O'Connor WG (1990) Threats to
Victorian native freshwater fish. The Victorian

Naturalist 107, 5-12.

Matthews WJ (1998) Patterns in Freshwater Fish
Ecology. (Chapman and Hall: New York)

MeGuckin J (1999) Fish of Kororoit Creek, Prepared
for Biosis Research Pty Ltd.

McKenzie JA and O'Connor WG (1989) The fish
fauna and habitats of Kororoit Creek, Truganina
Swamp, Laverton Creek and Cherry Lake. Arthur
Rylah Institute for Environmental Research,
Technical Report Series No 97, Department of
Conservation, Forests and Lands, Victoria.

Melbourne Water (1997) Health of Waterways Within
the Port Phillip & Western Port Catchements
Annual Stream Health Monitoring Report, 1997. Eds
R Coleman, M Batty and V Pelttigrove, Melbourne
Water Corporation, Melbourne,

Mitchell P (1990) The environmental condition of
Victorian streams. A report for the Department of
Water Resources Victoria.

Papas PJ, Nicol M and Crowther D (2000) Melbourne
Water Biological Monitoring Program - 1999/2000.
Consultancy report to Melbourne Water. Department
of Natural Resources and Environment, Victoria.

Paxton JR. Hoese DF Allen GR and Hanley JR (1989)
Zoological Catalogue of Australia. Volume 7. Pisces.
Petromyzontidae to Carangidae. (Australian
Government Publishing Service: Canberra)

Sonneman JA, Walsh CJ, Breen PF and Sharpe AK
(2001) Effects of urbanisation on streams of the
Melbourne region Victoria, Australia. Il. Benthic
diatom communities. Freshwarer Biology 46, 535-
551,

Walsh CJ, Sharpe AK, Breen PF and Sonneman JA
(2001) Effects of urbanisation on streams of the
Melbourne region Victoria Australia, L Benthic
macroinvertebrate communities, Freshwater Biology
46, 553-565,

Vale Jack Hyett

Members of The Field Naturalists Club of Victoria will be saddened to
learn of the death of Jack Hyett in July 2001. Jack was an honorary life
member of the Club and an Australian Natural History Medallionist. A
tribute to Jack will appear in a later issue,

Vol. 119 (1) 2002

Research Reports

The Mammal Fauna of Remnant Native Grasslands of the
Western Basalt Plains and Northern Plains of Victoria

Susan A Hadden'

Abstraet

Twenty-four native grassland remnants in the Western Basalt and Northern Plains of Victoria,
Australia, were investigated to determine the composition of the mammal fauna and evaluate their
habitat use. These sites included remnants that had differing levels of grazing pressure, structural
attributes and floristic diversity. Mammals were assessed by pitfall trapping and systematic search-
ing. Grassland characteristics, representing structure and diversity, that might affect the mammal
species richness of the grassland were recorded. Five mammal species were recorded in the Western
Basalt Plains and four in the Northern Plains. Sites with light grazing, open ground cover and high
native plant richness favoured native mammal species. (The Victorian Naturalist 119 (1), 2002, 14-20)

Introduction

In many regions of Australia, only dis-
junct remnants of the original forest,
woodland and grassland vegetation remain
within an expanse of cleared agricultural
land (Kitchener ef al. 1980; Bennett 1990),
Remnants are under continuing pressures
from a variety of pastoral activities, includ-
ing grazing, fire and pasture ‘improve-
ment’. Their ability to cope with changes
in management is of great interest.
Because most vertebrate fauna have not
successfully adapted to modified environ-
ments, remaining patches of natural vege-
tation are of importance to the regional
conservation of the indigenous fauna
(Bennett 1987a; Hadden and Westbrooke
1996), With increasing loss and fragmenta-
tion of the natural habitat, determining
both the consequences of habitat fragmen-
tation for faunal communities and the char-
acteristics of remnants that favour fauna
conservation — in particular, the role of dis-
turbance — is essential.

Native grasslands have been identified as
one of the most threatened ecosystems in
Victoria (Groves 1979; DCE 1990, 1992;
Lunt 1991; McDougall and Kirkpatrick
1994; Foreman 1996). Prior to European
settlement they covered approximately
30% of Victoria, occurring across the
western and northern plains, generally con-
fined to the lowlands, below 500 m alti-
tude. The western and northern plains lie
within the two natural regions of the
Volcanic Plains and Riverina of Victoria
(Conn 1993). Grasslands may appear to be

! Centre for Environmental Management, University
of Ballarat, PO Box 663, Ballarat, Vietoria 3353
Email; s.hadden@ballarat.edu.au

14

simple-structured communities (Lunt
1991); however, they are complex at the
small scale (Patton 1935). Their complexi-
ty arises from their diverse floristic and
faunal composition and the interaction
between these elements (Lunt 1991;
Morgan 1994). They have been largely
cleared for agriculture because the fertile
soils were suited to pasture improvement
and cropping, with the structure of remain-
ing grassland remnants being severely
reduced by cultivation.

The aim of this research was to document
the occurrence and abundance of ground-
dwelling mammal species in native grass-
lands and relate these to the importance of
vegetation structure.

Methods
Study sites

Twenty-four grassland remnants were
surveyed for ground-dwelling mammals.
These sites included remnants that had dif-
ferent levels of grazing pressure, structural
attributes and floristic diversity.

The Western Basalt Plains comprise a
series of lava flows, which are interrupted
in places by outcropping of other rock
types. The flat to undulating basalt plains
on which the community occurs cover
approximately 21,000 km' of Victoria. The
area is bounded by Melbourne in the east,
Hamilton in the west, Beaufort to the north
and Colac to the south. The average annual
rainfall ranges between 500 and 800 mm,
most falling between April and November.
The soils of the Western Basalt Plains are
described as duplex clays, with a shallow
clay and humus profile overlying another

The Victorian Naturalist

less permeable clay layer, restricting root
development and drainage (Conn 1993).
The Riverine Plain of south-eastern
Australia is the eastern portion of the
Cainozoic sedimentary Murray River
Basin dominated by Quaternary alluvial
sediments, and occupies a large part of
both Victoria and New South Wales
(Butler et al. 1973). The Victorian portion
is typically referred to as the Northern
Plain, occupying approximately 25,000
km: extending from Wangaratta and
Wodonga in the east to Charlton in the
west (Conn 1993), It consists of a relative-
ly homogeneous flat to undulating quater-
nary alluvium land-form, generally receiv-
ing less than 400 mm of rainfall per
annum. The grasslands and grassy wood-
lands occur on relatively dry and high
ground, away from zones of seasonal inun-
dation. The soils are typically poorly
drained with a firm clay surface texture
(McDougall et al. 1994; Foreman 1996).

Survey methods
Pitfall drifi-fence system

The traps consisted of thirteen 20 litre
plastic buckets buried in the ground so that
the lip was flush with the ground level to
form a pit. The traps were arranged in a
grid system of 3 * 2 (repeated 2.5 times),
20 m apart. A drift-fence extended through
the centre of each trap and extended for 2
m either side. The lower 2 cm of the fence
was buried in the ground to prevent ani-
mals from crossing under the fence. Holes
were drilled in the bottom of each pit to
prevent them from filling with water when
it rained. Traps were opened for four
nights during each survey period and were
checked every morning and afternoon,
resulting in 52 trap-nights for each site per
survey period. All vertebrates trapped were
identified and released at the point of cap-
ture, with care being taken to minimise any
distress to the animals. Mark-recapture
(ear-tagging) methods were employed to
obtain estimates of abundance.

Systematic and opportunistic searching
Systematic searching involved searching
for mammals under logs, behind bark,
through ground cover and in water. Any
fauna seen or hand-caught were identified

Vol. 119 (1) 2002

Research Reports

and released. Twelve hours of searching
were undertaken at each site.

Assessment was undertaken from
January 1995 to February 1996. Six sur-
veys, approximately 8 weeks apart, were
conducted at each site for each grassland
region, involving 7488 trap nights and 288
hours of systematic searching.

Habitat analysis

The habitat variables considered as
potentially being important in affecting the
richness and abundance of mammal
species in remnant native grassland were:

* cover of cool season perennial grass;

e cover of warm season perennial grass;

s cover of native herbs;

* cover of exotic grasses;

* cover of exotic herbs;

e cover of bare ground;

e cover of dry litter material;

e floristic composition (total, natives only
and exotics only);

egrazing pressure (sheep per ha and
months grazed); and

e invertebrate richness.

The percentage cover of the seven func-
tional groups described above was record-
ed using the point quadrat method
(Goldsmith et al. 1986). A 10 m tape mea-
sure was laid out through the trapping grid
and at every 10 cm point, a steel pin was
lowered into the vegetation, and the func-
tional group contacted by this pin record-
ed. Only one functional group was record-
ed per point, giving 100 point touches per
grassland site. This can be interpreted as
the percentage of cover for each functional
group. Measurements of cover were
recorded seasonally, in conjunction with
the trapping session. The botanical compo-
sition of the grassland site was determined
from the number of vascular plants record-
ed from a 5 * 5 m quadrat within the trap-
ping grid, during spring. All vascular
plants were identified and recorded.

The number of sheep per hectare and the
number of months the paddocks are grazed
per year defined the grazing pressure,
Information about these variables was
obtained directly from the relcvant
landowners.

Invertebrate richness was estimated
based on the number of invertebrate orders

15

Research Reports

at each site. Invertebrates were sampled
using small plastic cups filled with alco-
hol/glycerol solution. Ten cups were
placed randomly in the ground throughout
the trapping site and sampled during each
trapping session.

Species were categorised as common or
uncommon, based on records held in the
Atlas of Victorian Wildlife (NRE). Faunal
species were then given a classification as
being rare and/or endangered based on the
Atlas of Vietorian Wildlife records, Species
known from less than 50, 5? latitute/longi-
tude grids, were categorised as uncommon.

Some variables were combined to compare
particular trends in the data. The habitat
variables of grazing, cover and native plant
richness were classified into groups for an
initial review of the data, The variable of
grazing was split into lightly and heavily
grazed sites and was based on the DSE lev-
els and number of months grazed - those
which were grazed all year round with a
DSE rating of 3/ha and over were classed as
heavily grazed. Those sites that were rested
for at least three months of the year and with
a DSE of less than 3/ha were lightly grazed
sites. The variable of cover was split into
open ground cover sites and dense ground
cover sites, This was based on the percent-
age cover of tussocks, with sites having
greater than 50% tussock cover classed as
dense ground cover sites and those sites with
less than 50% tussock cover as open ground
cover sites. The variable of native plant rich-
ness was split into floristically rich sites and
floristically poor sites and was based on the
number of native plant species. Those sites
which had more than 30 native plant species
were classed as floristically rich and those
with fewer than 30 native plants were
classed as floristically poor.

Statistical analysis

Descriptive statistics were used to list
and describe the mammal composition of
the native grassland sites. They were used
to outline the characteristics of the fauna
and to investigate general similarities and
differences between the grassland sites.

Results

Two native mammal species were identi-
fied within the 24 remnants of native
grassland, with both represented in the
Western Basalt Plains and one in the

16

Northern Plains. The Fat-tailed Dunnart
Sminthopsis crassicaudata was found in
both grassland regions, whilst the
Common Dunnart S. murina was only
encountered on the Western Basalt Plains
with just a single capture at one site, Three
exotic species, the Brown Hare Lepus
capensis, House Mouse Mus musculus and
European Rabbit Oryctolagus cuniculus,
were recorded in both regions. Tables |
and 2 record the abundance of faunal
species recorded within each remnant, for
each region. Because the European Rabbit
and Brown Hare were sighted only and not
trapped, accurate abundance levels could
not be determined; their presence at a site
is indicated by +. All further results and
discussion are based on the native species.

Site occurrence of native mammals

Fat-tailed Dunnarts were recorded at
every site in the Northern Plains (100%)
and seven of the 12 sites in the Western
Basalt Plains (58%) (Tables 1 and 2). The
single record of the Common Dunnart was
at a site in the Western Basalt Plains where
Fat-tailed Dunnarts were also recorded.
Five of the 12 sites in the Western Basalt
Plains revealed no native mammals.

Site occurrence of the House Mouse

The introduced House Mouse occurred at
five sites on the Western Basalt Plains
(42%) and eight sites on the Northern
Plains (67%).

Native mammals in relation to site
characteristics and previous information
The most common marsupial, Fat-tailed
Dunnart, was abundant at a number of
floristically rich sites, but was also found
in high numbers on degraded grassland
remnants and in exotic pastures. This
species has adapted to altered grassland
conditions, Found living under stone walls
and rock piles, the species was observed to
use wolf spider holes on the Northern
Plains. It is considered to be a common
and widespread species with few concerns
about its conservation status. However, at
a number of grassland sites around
Melbourne, the species has disappeared
from areas where it was once in high abun-
dance (Colin Hocking pers. comm.). Very
few recaptures were made in this study,
and the species may warrant careful survey

'The Victorian Naturalist

Research Reports

Table 1. Mammal species recorded from grassland remnants of the Western Basalt Plains, with
numbers of captures during 7488 trap nights. + present but abundance levels not determined; * intro-

duced species.

Species

Site

Family Dasyuridae

Sminthopsis
crassicaudata [Q5 9 3

Sminthopsis murina - - - -
Family Muridae

Mus musculus* - - = 2
Family Leporidae

Lepus capensis* - t B +

Oryctolagus cuniculus* — - = + 1

pes ML es ee eo -
E b 1 À : € d i
eae eee 2 b n^
t + E + + + +
E 4 + E 4 n n

Table 2. Mammal species recorded from grassland remnants of the Northern Plains, with numbers of
captures during 7488 trap nights. + present but abundance levels not determined; * introduced

species.

Species

Family Dasyuridae

Sminthopsis

crassicaudata EL 7 S 7

Family Muridae

Mus musculus * - - - -
Family Leporidae

Lepus capensis* - - - -

Oryctolagus cuniculus* — - - + -

over the coming years to assess fluctua-
tions in population numbers and status.

The second species, Common Dunnart, is
considered uncommon throughout Victoria
despite its name (Morton 1995). All other
records for this species are in woodlands
(Menkhorst 1995; Morton 1995) so this
record from grassland suggests that the
species may occupy habitats outside its
previously known range.

Mammal abundance and grazing pressure

The relative abundance of the Fat-tailed
Dunnart was higher on lightly grazed sites
than on heavily grazed sites. The differ-
ence was more pronounced on the Western
Basalt Plains (p = 0.002) and not signifi-
cant on the Northern Plains (p — 0.241).
'The opposite was observed for the House
Mouse which was more abundant on heav-

ily grazed sites. There were no records of
the species on the lightly grazed sites of

the Western Basalt Plains. This difference
was significant for the Western Basalt

Vol. 119 (1) 2002

Site
5 6 vi 8 9 10) de 2
6 3 5 6 7 4 10 8
3 2 4 3 2 3 2 1
n 4 H + i i 4 +
E 4 - ; z : = x

Plains (p = 0.001) but not for the Northern
Plains (p = 0.156).
Mammal abundance and ground cover
The Fat-tailed Dunnart was more abun-
dant in the open vegetation cover sites than
in the dense vegetation cover sites. The
difference was more pronounced in the
Western Basalt Plains, but not significant
in either region (p = 0.196 and 0.278
respectively). The House Mouse showed a
different response to ground cover in the
two regions. On the Western Basalt Plains,
sites with a dense ground cover supported
significantly higher numbers of House
Mouse (p = 0.003), whereas on the
Northern Plains, sites with open ground
cover supported higher numbers of House
Mouse (p = 0.141).

Mammal abundance and plant species
richness

The Fat-tailed Dunnart was more com-
mon at the richer native plant sites than at
sites which were floristically poor. The dif-

17

Research Reports

ferences observed on the Western Basalt
Plains were significant (p — 0.002), but not
on the Northern Plains. There were higher
numbers of House Mouse on floristically
poor sites than on those which were floris-
tically rich. The differences observed on
the Western Basalt Plains were significant
(p = 0.003), but not on the Northern Plains.

Discussion
The mammal fauna

Few mammal species were recorded on
native grassland sites on either the Western
Basalt Plains or the Northern Plains, with
just one native mammal species, Fat-tailed
Dunnart, occurring widely in both regions.
Fat-tailed Dunnarts were found to be wide-
spread and common in both native grass-
lands and exotic pastures. Other mammal
species, including introduced Brown Rat
Rattus norvegicus and native Swamp Rat R.
lutreolus (J Seebeck pers. comm.), have
been recorded in grasslands, particularly on
the Western Basalt Plains, but were not
encountered during this survey. The Fat-
tailed Dunnart has a broad geographical dis-
tribution, occurring across southern
Australia in a variety of vegetation habitats,
including open woodland, low shrublands of
saltbush and bluebush, tussock grasslands
and gibber plains (Morton 1995). It inhabits
only lowland areas in the western half of
Victoria (Menkhorst 1995). Morton (1995)
and Menkhorst (1995) state that unlike most
native mammals in Victoria, the Fat-tailed
Dunnart appears not to have been greatly
disadvantaged by early agricultural develop-
ments, and populations survive on improved
pasture throughout the Volcanic Plains,
Otway Plains, Wimmera and Riverina.
Morton (1978a) states that the species is pre-
dominately an animal of sparse grassland
and open shrubland where there is a signifi-
cant component of bare ground. From this
information on the species, it was expected
that captures of Fat-tailed Dunnart would be
mainly confined to grassland sites with bare
ground a significant component.

The Common Dunnart was found at one
site from the Western Basalt Plains and
despite its vernacular name, is not common
in Victoria. It is most commonly found in
woodland, open forest and heathland but
has also been recorded near rainforest veg-
etation (Fox 1995). Sites of Common

18

Dunnart are characterised by sparse shrub
and ground cover (<50%), but often with
dense leaf and bark litter (Menkhorst
1995). This species has not previously
been recorded from grassland vegetation
and this record provides a major range
extension for the species. The site near
Colae is close to woodland vegetation,
which may provide an explanation for this
record.

Factors influencing abundance of the
Fat-tailed Dunnart

The abundance of the Fat-tailed Dunnart
appeared to be influenced by a number of
habitat characteristics and it was more
common on sites that were lightly grazed.
This suggests that high levels of grazing
may affect the species’ abundance, by
reducing the availability and quality of
ground cover. Sites which had an open
ground vegetation cover tended to support
greater numbers of Fat-tailed Dunnarts than
those which were structurally dense.
Because the species is an active forager of
insects and other invertebrates, open spaces
for hunting prey may be extremely impor-
tant, Dense vegetation cover at ground
level would severely limit the ability of the
species to move. However, the need for
cover is also important for protection from
raptors, foxes and cats and for shelter from
adverse weather conditions (Birney ef al.
1976; Abramsky et al. 1979; Grant er al.
1982; Alder 1985; Clark et a/. 1989;
Bowles and Copsey 1992). Although bur-
rows in the soil (Northern Plains) and under
rock (Western Basalt Plains) are vital for
Fat-tailed Dunnarts, some form of tussock
cover is required. Selection of sites with
open ground cover but with tussock cover
minimises the detection by aerial predators.

Potential predators of Fat-tailed Dunnarts,
and other native mammals, include intro-
duced Dog Canis familiaris; Fox Vulpes
vulpes; Cat Felis catus; native Brown
Goshawk Accipiter fasciatus; Grey
Goshawk 4. novaehollandiae; Wedge-tailed
Eagle Aquila audax and Whistling Kite
Haliastur sphenurus (Bennett 1987b). On
grasslands, tussock cover is most effectively
provided by scattered tussock grasses Poa
spp. on the Western Basalt Plains and wal-
laby grasses Austrodanthonia spp. on the
Northern Plains.

The Victorian Naturalist

Cover is also important to the Fat-tailed
Dunnart for diurnal shelter, and may ame-
liorate local weather conditions. The Fat-
tailed Dunnart constructs nests within tus-
socks or other dried plant material beneath
rocks and within cracks in the soil. Sites
with a greater richness of native plants sup-
ported more Fat-tailed Dunnarts. Although
the species was found at every site on the
Northern Plains and over half of the
Western Basalt Plains sites, as well as on
all exotic pasture sites, relative abundance
did have a slight correlation with native
species richness. The more floristically rich
sites, which are most likely to be more
intact, will presumably support greater
numbers of animals. As invertebrates are
still found in high numbers on more floris-
tically depauperate sites, food resources are
still adequate to support Fat-tailed Dunnarts
but may not be able to do so in the long
term. Such sites may only provide àn
extension of marginal habitat from a core
grassland area — an area which is used for
foraging, but not as a permanent living site.
Fat-tailed Dunnarts have been found to
have a preference for loose scattered rocks
in grassland remnants (Mathieson 1987).
Rock piles in grassland areas did provide
some habitat; however, those on agricultur-
al land did not (Mathieson 1987). Areas
with numerous, scattered rocks provide
more potential nesting sites. Because the
species tends not to be territorial, the pres-
ence of numerous scattered rocks may help
in the migration of individuals between
areas for both breeding and exploitation of
available food resources (Morton 1978a, b,
€). In this study, the Fat-tailed Dunnart was
found living under stone walls and rock
piles, and was observed using wolf spider
holes on the Northern Plains. Micro-habitat
use by this species at grasslands reflects the
need for three important requirements:

l. open ground cover for actively forag-
ing for invertebrate food items;

2. tussock cover for diurnal shelter and
predator avoidance; and

3. floristic diversity.

Factors influencing abundance of the
House Mouse

The relationship between habitat variables
and the introduced House Mouse contrast-
ed with the Fat-tailed Dunnart. House Mice

Vol. 119 (1) 2002

Research Reports

are highly fecund and adaptable, and have
been described as the ultimate mammalian
*weed' (Menkhorst 1995). They have
exploited a wide range of habitat types
across Australia. In natural vegetation,
House Mice rely heavily on their ability to
construct burrows or to shelter beneath
rocks or logs, and their distribution is there-
fore largely determined by characteristics
of the soil (Menkhorst 1995). The species
has excellent dispersal and short-term
colonising abilities and usually occupies
unfilled niches (Braithwaite and Gullan
1978), It is a primary coloniser after distur-
bance but once vegetation is suitable for
native mammal species, populations of
House Mice rapidly decline. Numbers of
House Mice on both the Western Basalt
Plains and Northern Plains were higher on
heavily grazed sites and on sites which
were floristically poor. This supports the
findings of Hadden and Westbrooke
(1996), who found numbers of exotic
species higher on more degraded, heavily
grazed and weedy Buloke woodland sites.
The Northern Plains sites with an open
ground cover supported greater numbers of
mice than sites with closed ground cover.
On the Western Basalt Plains, sites that had
a dense vegetation cover supported more
individuals. It may be that, on the more
intact grassland sites, the House Mouse is
unable to compete effectively with the Fat-
tailed Dunnart, while on more degraded
and depauperate sites, the Fat-tailed
Dunnart is less favoured, thus enabling the
House Mouse to be the dominant species.
Fox and Pople (1984) have stated that
House Mice are apparently unable to com-
pete successfully with small native mam-
mals. Abramsky et al. (1979) reported that
native mammals remained on natural short-
grass prairie and only the exotic mammals
occurred in completely modified areas.

Management implications
Conservation of native mammals

The Fat-tailed Dunnart was located across
a range of grassland types of the Western
Basalt Plains and Northern Plains.
However, very few recaptures were made. It
is not known if populations of Fat-tailed
Dunnarts are secure on these grassland rem-
nants. The Common Dunnart was recorded
at one site on the Western Basalt Plains.

19

Research Reports

Given that this species is uncommon and
has not previously been recorded in grass-
land vegetation, survey work to attempt to
locate more individuals is warranted.

Acknowledgements

The author would like to thank the editorial
assistance of Martin Westbrooke, John Seebeck
and Andrew Bennett. Thankyou to the land-hold-
ers and their wonderful grasslands on which
many a day was spent searching ~ they are beau-
tiful places: Ian and Trish Taylor, Kevin and
Jenny Blake, Cam and Cathy Nelson, Claire and
Jamie Dennis, Janet and Lochy Gordon, Angus
and Sally Ramsay, lan and Jill Buchanan, Jack
and Jean Wishart, Dorothy Davies, George
McGillivray, Wendy Rolfe, Don Glasson, Laurie
Boyd and Greg Rankin. Fauna survey was ĉon-
ducted under the Department of Natural
Resources and Environment Research Permits
RP-94-173 and RP-95-199. The research also
received approval from the University of Ballarat
Animal Ethies Experimentation Committee.

References

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530-536.

Alder GH (1985) Habitat selection and species interac-
tions; an experimental analysis with small mammal
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Bennett AF (19872) Conservation of mammals within a
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Bennett AF (1990) ZJubitat corridors: their role in
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Birney EC, Grant WE and Baird DD (1976) Importance
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by small mammals in a Victorian heathland,
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Butler BE, Blackburn G, Bowler JM, Lawrence CR,
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(1989) Small mammals in tall-grass prairie: patterns
associated with grazing and burning. Prairie
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Cann BJ (1993) Natural regions and vegetation of
Victoria, In Flora of Victoria Volume 1 Introduction,
pp 79-158. Eds DB Foreman and NG Walsh. (Inkata
Press: Melbourne)

DCF (1990) Remnant nutive grasslands und grassy
woodlands of the Melhourne area. Department of
Conservation and Environment Victoria

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DCE (1992) Draft conservation program for native
grasslands and grassy woodlands in Victoria.
Department of Conservation and Environment
Victoria.

Foreman PW (1996) Ecology of native grasslands on
Victoria's Northern Riverine Plain, (MSc thesis, La
Trobe University, Bundoora)

Fox BJ (1995) Common Dunnart Sminthopsis murina.
In The Mammals of Australia, pp 150-151. Ed R
Strahan (Australian Museum/Reed Books: Sydney)

Fox BJ and Pople AR (1984) Experimental confirma-
tion of interspecific competition between native and
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323-334.

Goldsmith FB, Harrison CM and Morton AJ (1986)
Description and analysis of vegetation, In Methods in
Plant Ecology, pp 437-525. Eds PD Moore and SB
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Grant WE, Birney EC, French NR and Swift DM (1982)
Structure and productivity of grassland small mammal
communities related to grazing-induced changes in
vegetative cover. Journal of Mammalogy 63, 248-260,

Groves RH (1979) The status and future of Australian
grasslands. New Zealand Journal of Ecology 2, 76-81.

Ifadden SA and Westbrooke ME (1996) Habitat rela-
tionships of the herpetofauna of remnant buloke
woodlands of the Wimmera Plains Vietoria. Wildlife
Research 23, 363-372.

Kitchener DJ, Chapman A, Dell J, Muir BG and Palmer
M (1980) Lizard assemblage and reserve size and
structure in the Western Australian wheatbelt - some
implications for conservation. Biological
Conservation 17, 25-62.

Lunt ID (1991) Management of remnant lowland grass-
land and grassy woodland for nature conservation: a
review. The Victorian Naturalist 98. 56-66,

Mathieson G (1987) The distribution and abundance of
the fat-tailed dunnart on the western Plains of
Victoria. Ballarat College of Advanced Education, Mt
Helen, Ballarat.

McDougall K and Kirkpatrick JB (1994) Conservation
of lowland native grassland in south-eastern
Australia, (Worldwide Fund for Nature Australia:
Canberra)

McDougall K, Appleby M and Barlow T (1994)
Western Busalt Plains Lake Omeo Murray Valley
Riverine Plains and the Wimmera. In Conservation of
lowland native grassland in south-eastern Australia,

p 44-112, Eds K McDougall and JB Kirkpatrick,
(Worldwide Fund for Nature Australia: Canberra)

Menkhorst PW (1995) Mammals of Victoria
Disiribution, Ecology and Conservation. (Oxford
University Press: Melbourne)

Morgan JW (1994) The ecology of grasses and grass-
land in lowland Victoria. The Victorian Naturalist
111, 87-93.

Morton SR (1978a) An ecological study of Sminthopsis
crussicuudata (Marsupiala; Dasyuridae) 1,
Distribution study areas and methods. dustralian
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Morton SR (1978b) An ecological study of Sminthopsis
crassicaudata (Marsupiala: Dasyuridae) HL, Behaviour
and social orgainsation, Australian Wildlife Research
5, 163-182.

Morton SR (1978c) An ecological study of Sminthopsis
crassicaudalu (Marsupiala: Dasyuridae) Ill.
Reproduction and life history, Australian Wildlife
Research 5, 183-211.

Morton SR (1995) Fat-tailed Dunnart Sminthopsis cras-
vicaudala, In The Mammals of Australia, pp 130-131.
Ed R Strahan. (Australian Museum/Reed Books: New
South Wales)

Patton RT (1935) Ecological studies in Victoria Part 1V

basall plains association, Proceedings of the Royal
Soview of Victoria 48, 172-191,

The Victorian Naturalist

Research Reports
How Do Nectar Foraging Butterflies Select Flowers?

Melissa B Nunn'

Abstract

Nectar foraging tendencies of two species of butterfly was experimentally investigated using charac-
teristics of flowers that may influence selection of nectar resources. Potentially influential character-
istics of nectar flowers were simulated in various combinations in a controlled laboratory situation.
Wild-caught butterflies were captured for use in the experiments. The two species preferred different
combinations of factors, potentially reflecting their natural foodplants. Common Grass-blue Zizina
labradus visited small white flowers most frequently, a combination of factors displayed by known
foodplants. Australian Painted Lady Vanessa kershawi preferred to feed on yellow, medium or large
flower combinations that match the flowers of Capeweed Arctotheca calendula and some other
observed foodplants. The position of the experimental flower on the ‘stem’ was not a significant
contributor to flower selection; however, flower size did contribute, There was a positive relation-
ship between the size of the butterfly and that of the flower chosen. In the wild, this may reflect not
only the flower's ability to support the weight of the butterfly, but also the amount of nectar supplied

by the flower, (The Victorian Naturalist 119 (1), 2002, 21-27)

Introduction

Butterflies are the most studied of the
invertebrates, yet there is still much to dis-
cover about their ecology, behaviour and
general biology. The focus of research is
generally on rare or exotic species, and lit-
tle information is known on the more com-
mon species (Kitching and Scheermeyer
1999),

Definitive studies of ecology and behav-
iour are lacking for common species such
as Yellow Admiral Vanessa itea,
Australian Painted Lady Vanessa kershawi
and Common Grass-blue Zizina labradus,
which occur frequently where this study
was conducted.

Butterflies feed solely on a liquid diet
consisting of plant nectar and sometimes
water or other fluids (Braby 2000). The
preferred source of nectar for butterflies is
nectar-bearing flowers. The nectar that is
produced by plants contains two of the
most vital ingredients for obligatory feed-
ers such as butterflies: carbohydrates and
protein (Brackenbury 1995). Butterflies,
with their specialised narrow tubular pro-
boscis, are able to access carbohydrates
and proteins contained in nectar by capil-
lary action (CSIRO 1991).

Vanessa kershawi generally appears in
late August and early September in the
Ballarat region and is usually the first adult
butterfly to be seen each season (G
Ambrose pers. comm. 2000). Zizina

Centre for Environmental Management, University of
Ballarat, Ballarat, Victoria 3353

Vol. 119 (1) 2002

labradus is more often found in gardens,
lawns, pastures, and weedy areas than in
bushland (Coupar and Coupar 1992). They
are common in many suburban areas,
including Ballarat.

Insects locate nectar-bearing flowers by
means of visual cues rather than by smell
or touch. The attraction is so powerful that
butterflies will be attracted to flowers that
are placed in an artificial setting. These
visual cues attract the butterfly to the
mouth of the corolla. Nectar guides (mark-
ings on flowers) provide a way of guiding
the proboscis of the butterfly toward the
nectaries (organs that secrete nectar),
Nectar guides may reflect strongly in the
ultraviolet section of the light spectrum in
which butterflies are sensitive
(Brackenbury 1995).

Ford (1985) has suggested that smaller
nectarivorous bird species are more mobile
than larger species and perch on flowers on
the end of branches without breaking them.
The smaller bird species are also more
acrobatic and are able to hover above the
flowers on the tips of the branches and
feed. Butterflies may mirror this behav-
iour, whereby smaller butterflies may pre-
fer flowers on the tips of stems.

It is documented that sometimes different
butterfly species prefer different flowers
(Clyne 2000; Braby 2000). However, over
100 species of butterflies feeding on
Lantana Lantana camara have been
observed, which suggests that butterflies
may also be opportunistic. The aims of this

21

Research Reports

study were to use artificial flowers to dis-
cover which characteristics of flowers
attract two butterfly species, and to com-
pare the findings to known foodplants and
their characteristics. It was expected that
the butterflies in the study would also pre-
fer a particular group of flowers.

Methods
Preparation

Two butterfly species, Vanessa kershawi
and Zizina labradus, were used in the
examination of three important flower
characteristics. These were chosen based
on the literature (Brakenbury 1995; Coupar
and Coupar 1992). These characteristics
were size, colour and the position on the
plant where flowers grow. Artificial plastic
flowers were used and the sizes (small,
medium and large) were scaled to known
nectar source species from the daisy family
Asteraceae. The artificial flowers were 2.5,
4.0 and 6.0 cm in diameter respectively.
The paint colours chosen for the study
included three that were common in the
Ballarat region in September, namely
white, yellow and purple. Two positions
were chosen to represent flowers that grow
close to the main trunk of the plant and
flowers that grow on the tips of the plant.

A cage was constructed by modifying an

old plywood finch cage. The sides and roof

were cut out and replaced by fibreglass fly-
wire, with the exception of one side that
was draped with muslin. This enabled
access to the inside of the cage and was
knotted when not in use. The frame of the
cage was painted white so that it provided
a light coloured backdrop against which
butterfly activity could be seen clearly,
The artificial flowers were drawn on ice-
cream container lids in the shape of flow-
ers from the Asteraceae family. These
were then coloured with acrylic paints.
Acrylic paints were used because they dry
quickly, are odourless and non-toxic. They
also do not run into the nectar solution on
contact. Four millimetre plastic hose was
sliced to produce thin plastic rings that
acted as nectar cups. These were glued to
the centre of the flowers with non-toxic
glue. Three timber frames were construct-
ed to hold the flowers and to simulate posi-
tions on a plant where the flowers might
grow. The design of the timber frame used

22

in the study is shown in Fig. 1. Small nails
were used to hold the flowers to the frames
and the glue used on the nectar cups was
used to seal the centres of the flowers
against leakage of nectar. Yellow cotton
wool was then packed tightly into the nec-
tar cups to absorb the nectar, preventing
drowning of the butterflies whilst still
allowing aecess.

Nectar

Artificial nectar was produced using a
recipe used at Melbourne Zoo's Butterfly
House. The nectar used in this study and in
a study conducted by Harris (1993) con-
tained approximately 15 percent sugars.
The recipe (see below) was halved to make
one litre of nectar. This formula makes
enough nectar to feed about 800 butterflies
for 24 hours.

The recipe used by the Melbourne Zoo's
butterfly house comprises 500 ml of puri-
fied water (boiled), with 130 g of sucrose
and 95 g of glucose or dextrose added and
stirred until they were dissolved. 200 g of
liquefied fructose (approximately 130 g
when crystallised) were added and stirred
in until dissolved. 500 ml of cold purified
water and half a teaspoon of honey were
added and the solution and stirred. Cold
purified water was added to take the quan-
tity up to two litres.

The different types of sugars are from
various plant sources, each supplying dif-
ferent molecular structures and nutrients.

Fig. 1. The design of the timber frame on which
the artificial flowers were placed during the
study. The base and main trunk were made of
25 mm * 19 mm pine painted green. The three
branches were made from dowel 10 mm in
diameter. The dowel represents the branch of a
plant and the flower placed closer to the centre
of the dowel represents flowers close to the
trunk,

'The Victorian Naturalist

The addition of honey is thought to be the
vital ingredient that provides the butterflies
with the essential nutrients that can sustain
them over a long period. Honey contains

Vitamin C, protein and small quantities of

nitrogenous compounds, minerals, organic
acids and lipids (Winston 1987). The but-
terflies at the Melbourne Zoo spend their
entire lives on this formula.

Capture of butterflies

Butterflies were captured on sunny days in
early September when they first began to
emerge. Optimal capture numbers were on
days that had followed one or two fine,
warm days. The capture sites included gar-
dens in town and an open area on the
Ballarat University campus at Mount Helen
that has both native and exotic vegetation.
An insect net was used to capture the butter-
flies. Undamaged specimens were retained
and put into a container with a muslin insert
in the lid. The butterflies were not fed for 48
hr before any sampling to encourage
hunger. Sixty-seven individuals were cap-
tured during the study, 35 of which were F.
kershawi and 31 Z. labradus.

Foraging observations

Observations were undertaken on four
occasions, 48 hours after capture. The
study cage was placed in a glasshouse that
provided a warm and humid environment,
which encouraged butterfly activity. The
glasshouse was connected to a thermostat,
which was kept at around 25°C. Each
observation period took one hour, which
was the optimum time discovered from a
pilot study. One hour was enough time for
each butterfly to feed three or four times
and was brief enough to allow the nectar to
stay moist.

Data analysis

Data were analysed using the software
package SPSS 10 for Windows. Univariate
ANOVA test was used to test for signifi-
cance between the categories of data
(Coakes and Steed 2001; Kitchens 1998).
The ANOVA statistic is used to examine
the difference between the means of forag-
ing frequencies for the two species, and to
determine whether they are significantly
different. Profile plots and ANOVA were
used to see if there were any interactions
between the characteristics studied,

Vol. 119 (1) 2002

Research Reports

Limitations

Other variables or characteristics that
may influence the frequency of visits such
as nectar volume, shape of flower and
ultraviolet reflectivity were not tested. The
brief nature of the study limited the num-
ber of variables that could be tested. It was
decided only to test what were thought to
be the most influential factors including
size, colour and position, allowing these
variables to be thoroughly examined in the
time allocated.

All taxonomic names in this paper follow
that of Braby (2000) for butterflies and
Ross (2000) for vascular plants.

Results

The flower characteristics preferred by
each butterfly species are shown in Figs 2,
3 and 4. Vanessa kershawi visited yellow
flowers more often than white or purple
flowers. Response to colour itself, when
tested for both species, showed no signifi-
cant difference of frequencies compared to
the other factors (F — 5.907, p — 0.064).
This indicates that there was no particular
colour preference overall for both species.
There was, however, a colour-species
interaction evident, which indicates that
each species tended to prefer a particular
colour (F = 9.559, p = 0.030).

The majority of flowers visited by E. ker-
shawi were large or medium in size. As
shown in Fig. 4, V. kershawi also tended to
visit flowers that were positioned close to
the trunk of the plant more often than those
positioned on the tips of the branches.
However, there was no significant differ-
ence found between the use of the two
positions on the plant (F = 1.113, p
0.402).

Zizina labradus significantly favoured
white flowers (F = 9.559, p = 0.030; Fig.
3), with the purple and yellow flowers
being less frequently visited. There was
also a significant preference for small
flowers (F = 8.72, p = 0.041), with the
medium and large flowers much less often
visited (Fig. 2). There appeared to be a
slight preference for flowers positioned on
the tips of the branches, but the difference
was not significant (F — 3.692, p — 0.195).

Vanessa kershawi tended to visit medium
sized yellow flowers that were positioned
on the trunk more frequently than any

23

Research Reports

20

Frequency of visits/hour.

—
| aessa
kershawi

O Zizina labradus

small medium

large

Flower characteristic

20

15

Frequency of visits/hour.

. 2. Frequency of visits for Vanessa kershawi and Zizina labradus with respect to flower size.

yellow

purple

| WB anessa |
| | kershawi
| | O Zizina labradus |
| n=
| H
(er -—

white

Flower characteristic

Fig. 3. Frequency of visits for Vanessa kershawi and Zizina labradus with respect to colour of

flower.

other combination of characteristics (Table
1). Flowers that were not visited at all
included large purple trunk, small white
trunk and medium white branch flowers.
Zizina labradus tended to visit small, white
flowers (Table 2). Use of the two small
white combinations were significantly dif-
ferent from the other combinations. The
flowers that were not visited at all were the
purple, medium and large flowers posi-
tioned on the branch.

The characteristics that had the strongest
association with visiting frequency were
the colour and size of the flower. This sug-
gests that each species has a preference for
particular characteristics in these groups of

24

factors. This is also evident from Figs 2 and
3. The position of the flower on the plant
had little influence on visit frequency.

Discussion

Vanessa kershawi visited yellow flowers
most often. These flowers were mostly
medium or large. It is evident that flowers
with combinations of traits that V. kershawi
was observed to feed on in this study may
match such species as Capeweed

Arctotheca calendula, Daisies Euryops sp.

and Everlasting Daisies Bracteantha sp.
Low (2000) details some butterfly species
found to have fed on such species of food-
plants. Foliage of these plants is also eaten

The Victorian Naturalist

16
d

8 14
3 12
un i0
-

o 6
Le]

3 4
g 2
= 9 eee

trunk

Research Reports

Wl anessa kershawi

O Zizina labradus

branch

Flower characteristic

Fig. 4 Frequency of visits for Vanessa kershawi and Zizina labradus with respect to flower position.

by the butterfly's larval stages (Braby
2000). Smooth Cats Ear Hypochoeris
glabra and Hairy Hawkbit Leontodon
taraxacoides are also abundant introduced
yellow flowered daisies in the region. They
flower over many months and are common-
ly visited by V. kershawi for nectar (G
Arnbrose pers. comm. 2000). Capeweed is a
weed of agricultural areas and is more often
encountered by butterflies (Low 2000) near
Ballarat than their native larval hostplants,
members of the family Asteraceae.

Zizina labradus showed a significant
preference for white flowers. This agrees
with the reports by Clyne (2000) and
Common and Waterhouse (1972) of butter-
flies feeding on flowers with similar char-
acteristics. Such flowers include Lawn
Daisy Bellis perennis, Clover Trifolium

subterraneum and native peas. The size of

these flowers is similar to that of the small
flower in this study, which could explain
the preference for small flowers. Small,
white flowers were the most visited of all
the combinations, and were significantly
different from the other combinations,
although not from each other. This indi-
cates that there is no preference by Zizina
labradus to the positioning of flowers on
the trunk or branch.

Vanessa kershawi visited large and medi-
um flowers with a slight but not significant
preference for the trunk. G Ambrose,
(2000 pers. comm.) suggests that flowers
that are pollinated by small butterflies may

Vol. 119 (1) 2002

not be able to support the weight of larger
butterflies. Therefore the larger of these
two butterflies may find the larger flowers
offer a more stable support when feeding.

Main (1981) suggests that the larger the
flower, generally the more nectar it pro-
duces, and that bigger flowers can support
larger birds both by the quantity of nectar
and by the strength of the flower. This can
also be applied to butterflies. Larger flow-
ers usually have a larger nectar reward
than smaller flowers, making them more
attractive to larger species of nectar forag-
ing fauna (Paton 1985). However, Dunn
(1999) lists several very large butterflies
feeding on small flowers of Lantana, and
highlights a low yield of nectar per flower.
The larger butterfly would need more ener-
gy to sustain itself; it would therefore tend
to visit the flowers that were more likely to
contain larger rewards in a field situation.

The body size of V. kershawi was much
larger than the small flowers in the study.
The results suggested that the bigger the
flower, the more likely it was to be visited
on the trunk. Under normal circumstances,
a larger flower with sturdier support would
offer these butterflies a more stable feed-
ing platform (Main 1981).

There may be two possible explanations
for the feeding patterns observed in this
study. Thompson (1999) suggests that but-
terflies tend to be specialised in their
choice of nectar and host plant. This sug-
gests that they may be predisposed to

25

Research Reports

Table 1. The number of visits per hour made by
Vanessa kershawi to each flower combination.

Table 2. The number of visits per hour made by
Zizina labradus to each flower combination.

Colour Size Trunk Branch Colour Size Trunk Branch
yellow 5 LS 2.25 yellow 5 2.25 2.5
m §.25 2.25 m 1 0.75
| 4.25 2.75 | 0.75 0.25
purple S L25 0.25 purple 8 l 2,25
m 0.25 0.75 m 0.75 0
| 0 l | 0.75 0
white S 0 0.5 white 5 Su 59/5
m | 0 m | 1.5
| 1.25 0.25 l 0.25 1.25

select the flower types or characteristics
that optimise their foraging efficiency. As
the preferences for natural hostplants
match the preferences of the flowers in the
study, it may be likely that these recorded
preferences are adaptive for the butterfly
species used in this study.

When foraging for nectar, butterflies
sometimes exhibit constancy (preference
for a particular species previously visited)
and often ignore rewarding flowers of dif-
ferent species (Goulsen e/ al, 1997),
Nectar rewards were constant for all flow-
ers during this experiment and so did not
contribute to the observed differences in
foraging frequencies. Darwin (1859)
described how the learning of handling
skills by a butterfly for one flower inter-
feres with the learning of handling skills
for another. Handling skills are skills that a
butterfly has had to learn to feed from a
flower. For example, a butterfly may have
to perform mid-flight ‘acrobatics’ to feed
from a particular tubular flower, or must
land on a certain area of another flower to
avoid falling off. The butterflies possibly
visited flowers similar to those with which
they were familiar and did not need to
learn such handling skills, in the short time
of the study.

The reflectivity of the paint on the flow-
ers Was not tested in this study. The effect
of ultraviolet reflectivity of the flowers on
frequency of visits is not known. However,
it is known that visual cues are used to
atiract butterflies to flowers (Brackenbury
1995). Colouration of the artificial flowers
was simple, without any ultraviolet pattern-

ing, Nectar guides were not provided nor

were patterns that may be visible on flowers
under ultraviolet light, Zizina labradus visits

26

purple flowers in the wild, including those
of Cullen, Hardenbergia and Hovea spp.,
which may or may not have ultraviolet pat-
terning (Clarke and Lee 1987).

The methods used were novel and impro-
vised because there has been limited study
in this area. There has been no benchmark
method for this particular area of research
and methods were chosen according to
their expected suitability. Other variables
may contribute to the selection of nectar
plants, such as the time of day, size and
type of nectar reward, shape of flower,
ultraviolet reflectivity and possibly pollen
volume. Further study should endeavour to
answer these questions.

Conclusion

Vanessa kershawi and Zizina labradus
prefer flowers that exhibit a different com-
bination of characteristics. The characteris-
tics preferred by V. kershawi include yel-
low, medium or large flowers and Z.
labradus, white, small flowers, These
combinations of characteristics tend to
match those of the butterflies’ known
foodplants.

Acknowledgements

Vhanks to Graeme Ambrose who supervised this
project, and gave me feedback on the develop-
ment of this manuscript, and to the anonymous
reviewer who read and commented on the manu-
script.

References

Braby MF (2000) The butterflies of Australia: Their
Identification, Biology and Distribution, Vol 1.
(CSIRO Publishing: Melbourne)

Brackenbury J (1995) Insects and Flowers; A
Biological Partnership. (Blandford: United Kingdom)

Clarke I and Lee H (1987) Name That Flower: The
Identification of Flowering Plants, (Melbourne
University Press; Burwood)

The Victorian Naturalist

Clyne D (2000) Attracting Butterflies to Your Garden.
(Reed New Holland: Sydney)

Coakes SJ and Steed LG (2001) SPSS Analysis Without
Anguish, (John Wiley and Sons: Milton)

Coupar P and Coupar M (1992), Flying Colaurs
Common Caterpillars, Butterflies, and Moths of
South-eastern Australia, (New South Wales
University Press: Kensington)

CSIRO (1991) The Insects of Australia, A Texthook for

Students and Research Workers, Division of

Entomology, CSIRO. (Melboume University Press:
Carlton South)

Darwin C (1859) The Origin of Species, (New Collier:
New York)

Ford HA (1985) A synthesis of the foraging ecology
and behaviour of birds in eucalypt forests and wood-
lands. In Birds of Eucalypt Forests and Woodland:
Ecology, Conservation and Management, pp 249-
254, Eds A Keast, H Recher, H Ford and D Saunders.
(Surrey Beatty and Sons Pty Ltd/RAOU: Chipping
Norton)

Goulson D, Stout JC and Hawson SA (1997) Can
flower constancy in nectaring butterflies be explained
by Darwin's interference hypothesis? Oecologica
112, 225-231.

Harris J (1993) The life history of the Australian
Admiral butterfly, Vanessa itea, in the Ballarat dis-
trict. (Student report, Biological Resource
Management, Ballarat University College, Ballarat)

Kitchens LJ (1998) Exploring Statistics: 4 Modern
Intraduction to Data Analvsis und Inference.
Duxbury Press: California)

Contributions

Kitching RL and Scheermyer E (1999) Foreword. In
Biology of Australian Butterflies, pp xv-xvi. Eds RI
Kitehing, E Scheermeyer, RE Jones and NE Pierce.
(CSIRO Publishing: Collingwood)

Low T (2000) Flutterby, butterfly. Nature Australia
Winter 2000, 24-25.

Main AR (1981) Plants as animal food. In The Biology
of Australian Plants, pp 342-360. Eds JS Pate and AJ
McComb. (University of Western Australia Press:
Nedlands, Western Australia)

Paton DC (1985) Food supply, population structure,
and behaviour of New Holland Honeyeaters
Phylidonyriss novaehollandiae in woodland near
Horsham, Victoria. In Birds of Eucalypt Forests and
Woodland: Ecology, Conservation and Management,
pp 124-137. Eds A Keast, H Recher, H Ford and D
Saunders, (Surrey Beatty and Sons Pty Limited/
RAQU: Chipping Norton)

Ross JH (2000) (ed) 4 Census of the Vaseular Plants of
Victoria, 5 ed. (Royal Botanic Gardens of Victoria:
Melbourne)

l'hompson JN (1999) What we know and do not know
about coevolution: insect herbivores and plants as a
test case, In Herbivores: Between Plants and
Predators, pp 730, Eds H Of, VK Brown, and RH
Drent, (Blackwefl Science Ltd: Oxford)

Winston ML (1987) The Biology of the Honey Bee.
(Harvard University Press: Cambridge)

Hooded Plover Thinornis rubricollis Chick Attacked by
Conspecifics

V Teoh' and MA Weston

We report an observation of adult
Hooded Plovers Thinornis rubricollis
attacking a flightless chick at Mornington
Peninsula National Park. The chick was
not seen again, and apparently did not sur-
vive to fly, although it is not known
whether the attacks contributed to its pre-
sumed death. Causes of mortality in
Hooded Plover chicks are poorly known,
although mortality rates are high (Weston
1998a, 2000). The eastern population of
this species is threatened, and is classified
as Vulnerable (Garnett and Crowley 2000).
Consequently, any observations of the dan-

! Mornington Peninsula National Park, Parks Victoria,
Hinton Street, Rosebud, Victoria 3939

* Threatened Bird Network, Birds Australia National
Office, 415 Riversdale Road, Hawthorn East, Victoria
3123 Email; m.weston@)birdsaustralia.com.au

Vol. 119 (1) 2002

gers that chicks may encounter are note-
worthy.

On 30 December 2000 one of us (VT)
was conducting a foot patrol along the
ocean beaches in the Mornington
Peninsula National Park, between Sorrento
and Portsea, Victoria. At the Franklin
Road access track (38?20' S, 144?43' E), at
Portsea, a pair of adults attending a young
chick was observed (the chick was
between 17 and 24 days old). This area is
used by flocks in winter, and by small
flocks during the breeding season (B
Dowling pers. comm.). A group of six
adults was close to the pair and chick. The
pair attending the chick was aggressive
towards the nearby flock, and a number of
short aerial chases were observed. In the

27

Contributions

confusion, the chick was temporarily sepa-
rated from its defending parents. Adults
from the flock ran directly at the chick, and
the chick ran away from them. Adults from
the flock also began ‘dive-bombing’ the
chick by flying low and directly at it.
About four such ‘diye-bombs’ occurred,
and the chick was struck twice. The chick
fell over on each occasion, and each strike
caused feathers to be dislodged from the
chick. Each time the chick was struck the
attacking adult landed and chased it on
[oot until a parent intervened. During these
attacks the chick piped continuously.

It seemed that the attacks would continue,
so when three adults from the flock aggres-
sively approached the chick, VT intervened
in order to give the chick a chance to find
shelter (these birds were part of an inten-
sively managed population; see Dowling
and Weston 1999), The chick managed to
hide. It was evident that the aggression
continued between the parents and the
flock after observations ceased. Although
the parents were seen on two subsequent
visits (on 5 January 2001 and 18 January
2001), the chick apparently did not survive
to fledge as it was not seen again.

No other reports of such aggression to
chicks are known to us (Marchant and
Higgins 1993). Weston (2000) observed
broods for more than 270 hours and never
recorded any such attacks. In an intensive
9-year study in the National Park, no such
attacks have been recorded until now
(Dowling and Weston 1999; B Dowling
and M Urquhart pers. comms). The aggres-
sive defence by the parents is common;
brood-rearing parents repel intruding con-
specifics vigorously, with one observation
of a parent *dive-bombing' and striking an
intruding adult (Weston 1998b).

The cause of this aggression is unknown,
but we speculate that displacement of the
flock occurred, such that territorial aggres-
sion escalated to the point where the chick
was attacked. The pair had been in the area
since 23 November 2000, and in early
December a l'emporary Beach Closure had
been implemented, so the territory seemed
spatially stable. Observations of colour-
banded birds have shown that birds in
flocks consist of non-breeders and some-
times include off-duty, and presumably
territorial, breeding birds (MAW pers.

28

obs.). The observations occurred on a busy
Saturday, and so recreationists (or VT)
may have displaced the birds, and caused
them to concentrate into one area,
Disturbance by humans displaces some
birds, including those with broods (Hockin
et al. 1992; Brown and Morris 1995; Hill
et al. 1997; Lord et al. 1997). Displace-
ment of breeding Black Skimmers
Rynchops niger to nearby less-disturbed
areas has caused nesting density to
increase, and this may have caused an
increased number of territoríal disputes
which could compromise the survival of
chicks (Safina and Burger 1983). A similar
displacement, albeit over a shorter time-
span, may have lead to the extreme aggres-
sion reported here.

Acknowledgements

Thanks to B Dowling and M Urquhart (Parks
Victoria), J Peter (Birds Australia), Anne
Morton (Assistant Editor) and an anonymous
referee for their comments on, and contributions
lo, the publication of this note. The Threatened
Bird Network is funded by the Natural Heritage
Trust.

References

Brown KM and Morris RD (1995) Investigator distur-
bance, chick movement, and aggressive behaviour in
Ring-billed Gulls. Wilson Bulletin 107, 140-52.

Dowling B and Weston MA (1999) Managing a breed-
ing population of the Hooded Plover Thinornis rubri-
collis in a high-use recreational environment. Bird
Conservation International 9, 255-70,

Garnett S and Crowley G (2000) The Action Plan for
Australian Birds. (Environment Australia Canberra)
Hill D, Hockin D, Price D, Tucker G, Morris R and
Treweek J (1997) Bird disturbance: improving the
quality and utility of disturbance research. Journal of

Applied Ecology 34, 275-88.

Hockin D, Ounsted M, Gorman M, Hill D, Keller V
and Barker MA (1992) Examination of the effects of
disturbance on birds with reference to its importance
in ecological assessments. Journal of Environmental
Management 36, 253-86.

Lord A, Waas JR and Innes J (1997) Effects of human
activity on the behaviour of northern New Zealand
Dotterel Charadrius obscurus aquilonius chicks.
Biological Conservation 82, 15-20.

Marchant S and Higgins PJ (1993) Handbook of
Australian, New Zealand and Antaretic Birds.
Volume 2. Raptors to Lapwings. (Oxford University
Press: Melbourne)

Safina C and Burger J (1983) Effects of human distur-
bance on reproductive success in the Black Skimmer,
Condor 85, 164-71.

Weston MA (19982) Nankeen Kestrel takes Hooded
Plover chick, Australian Bird Watcher V7, 266-7.

Weston MA (1998b) Some undescribed aggressive
behaviours, displays and calls of the Hooded Plover
in Western Australia. West Australian Naturalist 22,
105-14.

Weston MA (2000) The effect of human disturbance on
the breeding biology of Hooded Plovers.
(Unpublished PhD Thesis, University of Melbourne)

'The Victorian Naturalist

Contributions

Invertebrates of Mount McKay — a Brief Survey

EJ Grey!

Abstract

This note reports the results of a brief, small-scale survey of terrestrial invertebrates near Mount
MeKay in north-east Victoria conducted in October 1999. Survey areas, in this sub-alpine zone,
included Open Heathland, Tussock Grassland and small Snow Gum Eucalyptus pauciflora patches.
The invertebrates were collected in pitfall traps and by hand and the greatest numbers were taken in
the more open sites that were exposed to sunshine. (The Victorian Naturalist 119 (1), 2002, 29-35)

Introduction

A combined Field Naturalists Club of

Victoria and Victorian National Parks
Association expedition to Mount McKay
was held over the period 29 October-
2 November 1999. The purpose was to
increase appreciation and knowledge of the
ecology, geology, flora and fauna of the
area in view of its proposed excision from
the Alpine National Park, a decision that

has since been rescinded (Parliament of

Victoria 2001).

A small-scale survey of terrestrial inver-
tebrate fauna was undertaken as part of the
expedition. Two sites were sampled by pit-
fall trapping, and direct hand collecting
was carried out over several areas around
Mount McKay.

At the time of the expedition, inverte-
brate activity would have just started to
increase after the winter months. From this
point of view a later time, perhaps January,
would have produced greater numbers
(Green 1997).

A short search of the literature indicated
there were few studies of ground dwelling
invertebrates of the Australian high coun-
try. The most comprehensive general dis-
cussion of invertebrates in these areas is by
Green and Osborne (1994), while Green
(1997) details survey results from pitfall
trapping over a lengthy period in the
Snowy Mountains, particularly emphasis-
ing the seasonal and altitudinal variations.
Osborne et al. (1978) listed invertebrates
collected from the snow surface in
Kosciusko National Park and Green (1982)
reported the results of pitfall trapping
beneath the snow, from the same area.

A survey of ‘sub-alpine woodland’ con-
ducted in 1973/1974 in Victoria listed 51
invertebrate families from 7 orders (Land

8 Woona Court, Yallambie, Victoria 3085

Vol. 119 (1) 2002

Conservation Council 1977), and
Swarbrick (1946) gives notes on some of
the 50 species of insects collected by hand
at Mt Buffalo over a period of two weeks.

Methods
Mount McKay forms part of the Bogong

High Plains in the Victorian High Country

and lies approximately 2 km south of Falls

Creek Village. Vegetation varies from

Open Heathland through Closed Heathland

to Tussock Grassland with small patches

of Snow Gums. Rocky outcrops are also a

feature.

The two sites chosen for pitfall trapping
each comprised three distinct vegetation
types:

Il. Snow Gum Eucalyptus pauciflora
patches with exposed granite outcrops.
Open Heath species also occurred with
grassy, open spaces between shrubs.

2. Open Heath with Phebalium squamulo-
sum subsp. alpinum (in flower) and
Mountain Beard Heath Leucopogon
hookeri. Other species most likely pre-
sent but not identified would have includ-
ed Grevillea australis, Alpine Hovea
Hovea montana, and Alpine Orites Orites
lancifolia (Bogong High Plains,
Vegetation Map and Guide to Alpine
Flora, Pretty Valley Sheet);

3. Tussock Grassland with Soft Snow Grass
Poa hiemata and Ranunculus sp. (in
flower) (Bogong High Plains Vegetation
Map and Guide to Alpine Flora; Willis er
al. 1975).

The two sites were located near access
roads to the east of Mount Mckay and both
had elevations c. 1500 m ASL. The first site
established (site A in this article) was locat-
ed at 36"52738" S 147?14'45" E, and the
second (site B) at 36?52'54" S 147*15'01"E.
At each site, the three distinct habitat types

29

Contributions

were a short distance apart, separated by
some 20-30 m and with little difference in
altitude.

Direct hand collecting was done at a
number of areas around the vicinity of
Mount McKay. The results section records
where the specimens were found.

At the pitfall trapping sites A and B,
three lines of six traps were set with one
line in each habitat type. Each line was
numbered: 1, Snow Gum patch; 2, Open
Heathland; 3, Tussock Grassland. Plastic
coffee cups of 200 ml capacity, height 80
mm and top diameter 72 mm were used for
the traps. Each trap was dug in flush with
the ground and 30 ml of preserving fluid
was placed in each. The preserving fluid
was a 50/50 mixture of ethylene glycol and
70% ethanol. The pitfall traps were open
for four days from 28 October (the day
they were dug in) to 1 November 1999,
The weather during this period was vari-
able with cool, sunny days, one night with
severe frost, and some rain. A maximum/
minimum thermometer and a rain gauge
were set up at Site A, The thermometer
was suspended from a Snow Gum branch,
and the rain gauge was attached to a star
picket. Over the sampling period, the
recorded shade temperature range was —3°
to *13*C and 40 mm of rain fell. The
weather observations for Site A were
assumed to apply to Site B as the sites
were less than one kilometre apart, had
westerly aspects and were roughly at the
same elevation.

For comparison, weather data for the
months of October and November 1999
were obtained from the Bureau of
Meteorology for Mt Hotham, the nearest
recording site (Table 1). Mt Hotham is sit-
uated c. 15 km south-west of Mt McKay
and has an elevation of 1849 m, which is
approximately 350 m higher than the site
at Mt McKay.

Hand collection was carried out at ran-
dom and no attempt was made to regulate
the time spent in this activity, or the num-
ber of people involved. Invertebrates were
collected from vegetation, under rocks and
from streams, and each person kept a note
of the habitat for each sample.

At a later date, identification of ants was
to genus (species or species group where
possible); beetles and spiders to family

30

——— €—— a — -
Table 1. Mt Hotham Weather Station, Data for
October and November 1999.

Oct Noy

1999 1999

Mean daily maximum 8.0 8.1
temperature "C

Mean daily minimum 1.8 1.4
temperature °C

Highest temperature °C 15.6 15.4

Lowest temperature °C -54 -6.5

Mean monthly rainfall mm 292 137.6

level, and all other invertebrates to order
(sometimes class or phylum), The keys
used for identification included: ants
(Andersen 1991; Greenslade 1979;
Shattuck 1999); beetles (Moore 1980); spi-
ders (Davies 1986); and all other inverte-
brates (Harvey and Yen 1989),

Results

The material trapped in each of the six
traps per line was pooled to give one sam-
ple for that site, e.g. A1 contains all mater-
ial from the six traps in the Snow Gum
patch at Site A. Table 2 is a list of the total
numbers of invertebrates per pitfall trap
line.

The most numerous groups were found in
"Other orders’, and while Flies (Diptera)
made up most of the numbers, Springtails
(Collembola) were very abundant at site
Bl with an estimated number of 10,000
individuals (see note in Table 6).

The numbers of Ants (Hymenoptera)
caught in the pitfall traps are shown in
Table 3. The ants were identified to genus
(indicated as sp.), to species group (as
shown by species in inverted commas) or,
in some cases, to species.

A total of eight genera from four sub-
families was recorded from the pitfall
traps. The largest number were from the
Dolichoderinae (92%) with one species
group Anonychomyrma ‘itinerans’ making
up 96% of the sub-family.

The Dolichoderinae are generalist preda-
tors and favoured the Open Heath and
Tussock Grassland habitats. These ants are
characteristic of open and sunny places
(Andersen 1991). However, the large num-
bers found in the Tussock Grassland (site
A3) may be associated with a large nest (or
nests) or a foraging trail. Shattuck (1999)
notes that nest sizes for Anonychomyrma

The Victorian Naturalist

Contributions

e T———— A-——————————— — «€ €

Table 2. Total numbers of invertebrates collected in pitfall traps.

Habitat Snow Gum Patch Open Heath Tussock Grassland
Site Al Bl A2 B2 A3 B3
Ants (Hymenoptera) 18 7 52 40 259 5
Beetles (Coleoptera) 19 13 40 33 23 15
Spiders (Araneae) 5 4 17 3 20 15
Other Orders 285 10,293 367 500 267 324

Table 3. Total ants (Hymenoptera:Formicidae) in the pitfall traps.

Habitat Snow Gum Patch
Site A1 B1

Open Heath Tussock Grassland
A2 B2 A3 B3

Myrmicinae
Aphaenogaster sp. -
Monomorium sp. -
Pheidole sp. 6 -
Ponerinae
Amblyopone australis - -

Dolichoderinae
Anonychomyrma ‘itinerans’ - -
Iridomyrmex sp. 12 -
Formicinae
Plagiolepsis sp. - |
Prolasius sp. - l

un

D
si N
1
D

LE TOT o aaa

‘itinerans’ range from 500 to tens of thou-
sands of workers.

The maximum temperature of 13°C
recorded during the survey is at the lower
level for ant functioning with 10°C being
the limit for any functioning at all
(Hólldobler and Wilson 1990, 1994).
Therefore, the two Dolichoderinae genera
must have adapted to foraging in these
cool conditions, although the temperatures
at the ground surface and certainly within
the nest would have been higher than the
shade temperatures measured. On the other
hand, specialist cool climate ants, in this
case Monomorium, Plagiolepsis and
Prolasius, were not notably abundant in
this survey (Andersen 1990).

Table 4 lists the beetles (Coleoptera) found
in the pitfall traps. Identification was to fam-
ily level. However, the key used (Moore
1980) was not applicable for beetles smaller
than 3 mm, and so were classed as *undeter-
mined' specimens.

The beetle families found in this survey
can be divided into two main groups. The
first group is those adults and larvae that
rely on living prey or carrion for food.
These are the predators — the Ground
Beetles (Carabidae), Pselaphidae and Rove
Beetles (Staphylinidae) — and are the most

Vol. 119 (1) 2002

numerous group, making up 69% of the
total number of beetles collected. The
Ground Beetles were the most abundant
family and most were found in the Open
Heath and Tussock Grassland. These bee-
tles (and their larvae) feed on smaller and
weaker insects as well as on larvae of other
invertebrates. They use their strong jaws
for seizing, tearing and crushing their vic-
tims (Hawkeswood 1987).

The second group is the herbivores,
which encompass those that, as adults or
larvae, feed on living, as well as dead or
decaying plant material — Longicorn
Beetles (Cerambycidae), Leaf Beetles
(Chrysomelidae) and Weevils (Curculion-
idae).

Click Beetles (Elateridae) can be included
in both groups since larvae of some mem-
bers of this family are plant feeders while
others eat insect larvae (Mathews 1985).

Pitfall trapping favours the ground
dwelling families, hence larger numbers of
Ground Beetles, Rove Beetles and Weevils
(Green and Osborne 1994).

The greatest spider abundance came from
the Open Heath and Tussock Grassland
habitats. Most of the spiders caught in the
pitfall traps were from ground-dwelling
families and this reflects the method used

31

Contributions

Table 4. Total beetles (Coleoptera) in the pitfall traps.

Habitat Snow Gum Patch Open Heath Tussock Grassland
Site Al Bl A2 B2 B3
Family

Carabidae (Ground Beetles) 5 6 31 26 17 5
Cerambycidae (Longhorn Beetles) 4 - | - - -
Chrysomelidae (Leaf Beetles) - 1 - - - 3
Curculionidae (Weevils) - 3 6 6 l 3
Elateridae (Click Beetles) l - j! - -
Pselaphidae - - - - - 2
Staphylinidae (Rove Beetles) 5 2 - - - -
Undetermined 5 - 2 - 5 2

Table 5. Total spiders (Araneae) in the pitfall traps.

Habitat Snow Gum Patch Open Heath Tussock Grassland

Site B1 A2 B2 A3 B3

Family

Mygalomorphae Hexathelidae - - - 1 * -
Nemesiidae 2 - - - - -

Araneomorphae Gnaphosidae = - 10 - - l
Hadrotarsidae - - - - -
Lycosidae (Wolf Spider) - 2 7 D l 6
Miturgidae 2 - - 18 8

for sampling. Spiders that spend their lives
above ground level — the web builders and
other foliage-dwelling species — are unlike-
ly to be found in pitfall traps. Table 5 lists,
in families, the spiders found.

Three families, the Gnaphosidae,
Miturgidae and Lycosidae, comprised the
bulk of the specimens. All of these fami-
lies are active, open-range hunters preying
on a variety of invertebrate fauna including
crickets, locusts and other spiders (Brunet
1996).

The single member of the Hexathelidae
comes from the family that includes the
Sydney Funnel Web spider, but our speci-
men is not this species.

Table 6 gives an indication of the variety
of invertebrate fauna found by pitfall trap-
ping. Only adults from all the orders were
found in the traps, and larvae were notice-
ably absent. Most of the material was iden-
tified to order level, although one cen-
tipede could only be identified to class.

The flies (Diptera) and mites (Acarina)
were the most abundant orders, although at
one site in the Snow Gum Patch habitat an
estimated 10,000 springtails (Collembola)
were collected. Mites and Collembola have
both been recorded in large numbers in
alpine areas (Green and Osborne 1994;

32

McKeown 1942) and possibly the rain at
the time of the survey influenced the num-
bers of Collembola from site B1.

The large number of flies (Diptera) are
difficult to explain, although the cold to
cool weather may have inhibited their fly-
ing and kept them on the ground.

Although opportunistic and unregulated,
hand collecting (from foliage, under litter
and rocks, and in one case from a stream),
discovered members of the orders shown
in Table 7.

The Wolf Spider (Lycosidae) was teased
from its burrow with a grass stem, the bur-
row sealed with a knife and the spider cap-
tured on the ground surface.

Further observations

One interesting observation was the num-
ber of mating pairs of the Mountain
Spotted Grasshopper Monistria concinna
that were seen on roads and in the bush
while inspecting pit-fall traps and during
hand collecting. These are large grasshop-
pers with bright yellow spots on the body
and a pale streak extending from the body
to under the eye. The female is nearly dou-
ble the length of the male and both are
‘brachypterous’, i.e. lacking functional
wings (Green and Osborne 1994), The

'The Victorian Naturalist

Contributions

Table 6. Total ‘other invertebrates’ in the pitfall traps.

Habitat

Site Al

Snow Gum Patch
B1

Tussock Grassland
B3

Open Heath
A2 B2 A3

Order/Phylum

Acarina (Mites)

Amphipoda (Amphipods)
Phylum - Annelida (Worms)
Blattodea (Cockroaches)
Class - Chilopoda (Centipedes)
Lithobiida (Centipedes)
Collembola (Springtails)
Diplura (Diplurans)

Diptera (Flies)

Haplotaxida (Earthworms)
Hemiptera (Bugs)
Hymenoptera (Wasps, Bees)
Isopoda (Slaters)

Opilionida (Harvestmen)
Orthoptera (Grasshoppers)
Platyhelminthes (Flatworms)
Polydesmida (Millipedes)
Polyzoniida (Millipedes)
Sphaerotheriida (Millipedes)
Symphyla (Symphylans)
Undetermined

3

ON

to
©
t =u a NULU Bee iii anma

1a

4

ae

rDNYeMNwohw *—,. = i

(34

to

cee et 45:98
1 Wh

1
tt i

tm
o
j| NNS:

'
'
[

D
— le j
1

Ha

* estimated by counting a fraction of the sample and multiplying to arrive at an approximate total.

Mountain Spotted Grasshopper has the
ability to overwinter at any stage of devel-
opment from the third instar to adult by
entering diapause. This is the reason for
such an early appearance of adult
Mountain Spotted Grasshoppers and their
mating at this time.

Another part of our work took us into
sphagnum bogs in Pretty Valley at the
headwaters of Pretty Valley Creek. Here
we noticed two types of terrestrial inverte-
brates living in a wet environment. The
first was the presence in the sphagnum
beds of a number of spider holes, which
were 30-40 mm diameter and closed with a
dome-shaped web. These were taken to be
the retreats of Wolf Spiders (Lycosidae),
and Main (1984) notes that *some species
occur in bogs'. The only other likely spider
family to be found in this situation would
be the Water Spiders (Pisauridae) and, in
particular, the genus Dolomedes. However,
these spiders do not build a retreat or bur-
row (Main 1984).

The second was the surprising presence
of many ant nests in the bogs. These con-
sisted of mounds constructed of vegetation
fragments, largely sphagnum moss, that
were raised above the water level. Mounds

Vol. 119 (1) 2002

are usually built as an aid to regulating the
colony temperature and moderating
extremes, an advantage in this situation
(Hólldobler and Wilson 1990). These ants
were members of the genus /ridomvrmex
(sub-family Dolichoderinae), a large genus
noted for occupying both *the driest and
the wettest of snow country environments’
(Green and Osborne 1994).

It is interesting to note that the ant
species from the Dolichoderinae found in
the pitfall traps did not appear to have built
mounds as none was sighted and they are
normally quite visible.

Discussion

In spite of the cool weather and early (for
invertebrates) season, a variety of inverte-
brates from 25 orders were recorded in this
brief survey. Thus, the indication is that a
rich and diverse invertebrate fauna inhabits
this area. Further detailed surveys over
longer periods and including summer
would be necessary before firm conclu-
sions can be made about the invertebrate
fauna of Mt McKay.

The results of a survey of invertebrates
from a low sub-alpine (1680 m ASL) site
in the Snowy Mountains were given in

33

Contributions

Table 7. Specimens caught from hand collection. Habitat key: UR, under rock; OV, on vegetation;
RS, at road-side; IC, in cascade; OH, open heath; FB, from burrow.

Order Family Genus/Species Habitat/Location
Group/Species UR OV RS IC OH FB
Amphipoda = = t I - -
Araneae Araneidae - 1 - - - =
Gnaphosidae 1 - - - z
Lycosidae T E - ^ ^ I
Micopholcommatidae l - - - - -
Miturgidae ^ 2 à g a T
Nicodamidae | 2 = ^ ^
Symphytognathidae - l - - - -
Zodariidae [: 3 1 L s -
Blattodea 4 2 1 é 5 À
Coleoptera Carabidae 6 = z - z :
Chrysomeledae - 1 - z s E
Curculionidae l - - - = <
Tenebrionidae | z : 3 = z
Diptera E l z 4 - H
Ephemeroptera (larva) E E 1 x Z
Hymenoptera Formicidae Amblyopone 10 - - - - -
Hypoponera l - - - - 3
Anonychomyrma
'itinerans ' 9 - = z 13 -
Myrmecia pilosula - - - - 7 2
Isopoda l * " E E T
Odonata (nymphal case) 4 1 L > = x
Orthoptera t 6 " 2 = "
Hemiptera = 3 2 = - 2
Plecoptera (larva) 3 z 2 1 es "
Polydesmida 2 1 2 = = £

Green (1997). Table 8 compares the
results, for selected groups, from this study
with those from the work at Mt McKay. It
is interesting that the two sets of results are
quite comparable, despite the difference in
total numbers, and emphasise the influence
of Collembola on invertebrate fauna com-
position in these areas.

One interesting question that further
work on ants might answer is why in this
survey there was a virtual absence of cold
climate specialists species (Monomorium
spp., Notoncus spp. and Prolasius spp.).
From previous studies (Andersen 1990)
these would have been expected to make
up a substantial proportion of the ant
fauna.

Another point relating to ants which
requires further work is to understand how
the ant communities are interrelated at Mt
McKay. Other work has indicated that
Dolichoderinae species, which include
Anonychomyrma and Iridomyrmex, play a
less dominant role and are less abundant in
cooler climate communities (Andersen
1990). In these communities, cold climate

34

Table 8. Comparative results from two inverte-
brate studies shown as percentage contribution
by numbers for selected groups.

% Contribution

Low sub- Mt McKay
alpine area (1999)
(Green 1997)

Coleoptera 1.3 d
Hymenoptera 2.1 3.0
Araneae 1.8 0.5
Collembola 80.1 79.4
Diptera 9.6 12.9
Acarina 1.1 Les
Total 1,068 12,616
species and opportunists such as

Rhytidoponera spp. and Paratrechina spp.
are the most common ants. The results
from this survey do not bear this out and
possibly a different community organisa-
tion exists at Mt McKay where
Dolichoderinae were found to be the most
abundant family.

The results of this limited survey give
some indication that the Mt McKay habitat

The Victorian Naturalist

is relatively undisturbed because of the
absence of opportunistic ant species that
are characteristic of disturbed areas
(Andersen 1990). However, additional
work on this point is also needed,

Acknowledgements

The collecting was done under Flora and Fauna
Permit Number 10000715. Thanks are due to
members of the Field Naturalists Club of
Victoria and Victorian National Parks
Association who helped with the fieldwork and
subsequent identification, and to lan Bell for
providing weather bureau data. Special thanks
go to Erich and Elsbeth Sacco and Patricia Grey
for their assistance in setting up the pit-fall lines.
An early draft of this article was much improved
by the comments of an anonymous referee.

References

Andersen AN (1990) The use of ant communities ta
evaluate change in Australian terrestrial ecosystems:
a review and a recipe. Proceedings of the Ecological
Society of Australia 16. 347-357.

Andersen AN (1991) The Ants of Southern Australia: A
Guide ta the Bassian Fauna. (Commonwealth
Scientific and Industrial Research Organisation: East
Melbourne)

Bogong High Plains Vegetation Map and Guide to
Alpine Flora, Pretty Valley Sheet. (ND) (Victorian
Conservation Trust)

Brunet B (1996) Spiderwatch: A Guide to Australian
Spiders, (Reed Books: Australia)

Davies VT (1986) Australian Spiders: Collection,
Preservation and Identification. (Queensland
Museum Booklet 14)

Green K (1982) Notes on winter-active invertebrates
beneath the snow. The Victorian Naturalist 99, 144-
146.

Green K (1997) Inter-annual, seasonal and altitudinal
differences in invertebrate activity in the Snowy
Mountains. The Victorian Naturalist 114, 222-229.

Contributions

Green K and Osborne W (1994) Wildlife of the
Australian Snow Country. (Reed Books: Chatswood)

Greenslade PJM (1979) 4 Guide to Ants of South
Australia. (South Australian Museum: Adelaide)

Harvey MS and Yen AL (1989) Worms 10 Wasps: An
Illustrated Guide to Australia's Terrestrial
Invertebrates. (Oxford University Press; South
Melbourne)

Hawkeswood T (1987) Beetles of Australia. (Angus
and Robertson: Sydney)

Hólldobler B and Wilson EO (1990) The Ants. (The
Belknap Press of Harvard University Press:
Massachusetts)

Hólldobler B and Wilson EO (1994) Journey to the
Ants. (The Belknap Press of Harvard University
Press: Massachusetts)

Land Conservation Council (1977). Report on the
Alpine Study Area. (Land Conservation Council:
Melbourne)

Main BY (1984) Spiders. (William Collins Pty Ltd:
Sydney)

Mathews EG (1985) 4 Guide to the Genera of Beetles
of South Australia. Part 4. (South Australian
Museum: Adelaide)

McKeown KC (1942) Australian Insects, (Royal
Zoological Society of New South Wales: Sydney)

Moore BP (1980) A Guide to the Beetles of South-
Eastern Australia. (Australian Entomological Press:
Sydney)

Osborne W, Preece M, Green K and Green M (1978)
Gungarton: A Winter Fauna Survey above 1500
metres. The Victorian Naturalist 95, 226-235.

Parliament of Victoria (2001) Land (Further
Revocation of Reservations) Act 2001.

Shattuck SO (1999) Australian Ants; Their Biology and
Identification. (Commonwealth Scientific and
Industrial Research Organisation; Collingwood)

Swarbrick E (1946) Notes on insect life at Mt Buffalo,
The Victorian Naturalist 63, 19-23.

Willis JH, Fuhrer BA and Rotherham ER (1975) Field
Guide to the Flowers and Plants of Victoria. (AM
and AW Reed Pty Ltd: Sydney)

Vale Alexander Clifford Beauglehole OAM

Club members will be saddened to learn of the death of Cliff Beauglehole
in January 2002. Cliff made enormous contributions to our knowledge of
the distribution and conservation of Victorian plants. He was also a recipi-
ent of the Australian Natural History Medallion. A tribute to Cliff will

appear in a later issue.

For assistance with the preparation of this issue, thanks to Karen Dobson (label printing),
Dorothy Mahler (administrative assistance) and Phil Bock (web page).

Vol. 119 (1) 2002

ta
Un

Naturalist in the Mountains

The Biodiversity Blitz

There is something, very satisfying about
seeing a group of experts in their own
fields enthused by being together with
like-minded people, their only task to go
out and explore. This was the scene at
Kosciuszko National Park on the weekend
of 12-13 January when a ‘Biodiversity
Blitz’ was held over 24 hours. It was the
first major event held in Australia to cele-
brate the International Year of Mountains.
The concept of a Biodiversity Blitz origi-
nated in Germany, where it is a common
event, and spread to Switzerland in 2000,
where it was sponsored by the German
journal Gea. The task is to document every
species of living thing within a defined
area over a 24 hour period. The Swiss
event “Alp Blix’ was held near Chur and
was organised by Jürg Müller from the
Natural History Museum at Chur (well
worth a visit!). I arrived at Chur after the

event but the continuing enthusiasm of

Jürg and the results published in full colour
in ‘Geo’ sparked my interest.
Because we needed accommodation and

because it is the International Year of

Mountains, the blitz was set up to look at
the biodiversity between our accommoda-
tion at Thredbo at 1400 m altitude and the
summit of Mt Kosciuzsko at 2228 m - a
total of 27 square kilometres, rather big but
at least it had something for everybody.
Getting so many experts together in one

place for one weekend involved a lot of

organisation, not made easier as the date
approached by the fires that started in

NSW around Christmas and took myself

and most of the local National Parks and
Wildlife Service staff out of the running
for a good part of the time. There were
logistical problems too; how to find a punt
and then get it onto Australia’s highest
water body Lake Cootapatamba.
Victoria's Department of Natural
Resources and Environment solved the
first problem and Brian Timms, who has
worked on the lakes for many years, pro-
vided the muscle for the latter.

In all, 71 people participated, with
experts in birds, bats (and other beasts),
reptiles and frogs, fish, insects, collembola,

36

aquatic fauna and flora, soil invertebrates,
snowbank invertebrates, spiders, velvet
worms, vascular plants, horticultural
plants, fungi, mycorrhizal fungi, lichens,
mosses, liverworts and snow algae. The
public was invited to participate and vol-
unteers were on hand to help with sam-
pling and also to enjoy the ‘show and tell’
held in the Thredbo Community Hall at the
completion of the 24 hours.

Timing was always going to be a prob-
lem; there is, for example, a predictable
sequence of insect emergence in the moun-
tains and many adults are gone long before
others emerge. We were never going to be
in the right place at the right time for
everything - the annual grasshoppers
Kosciuscola had only reached their first
instar whereas the Mountain spotted
Grasshoppers Monistria concinna were
well represented by all instars. Many of the
herbs were in flower but we were about
three weeks too early for the grasses.

Following the early spring thaw in the
mountains, the following cool stretch
right up to snowfalls on Boxing Day
meant that snowbanks were still large and,
with the sun shining on them, snow algae
were blooming and warm conditions and
strong winds just before the blitz dumped
many insects on the snow surface ready to
be collected. Warm nights also aided the
entomologists with their insect traps.

Some species were easy to tick off
(Crimson Rosellas and Snow Gums were
never going to be a problem!). Ticking off
the Platypus was a simpler job than expect-
ed with a visit to the pool just below the

1 A Y

Lake

Biologists sorting the catch. from
Cootapatamba,

The Victorian Naturalist

Brian Timms leads out with the Victorian punt
above Lake Cootapatamba.

Thredbo Community Centre which result-
ed in a sighting within two minutes. To
complete the monotremes, an Echidna
wandered across the village green through
groups of holiday makers. Even when sur-
rounded by people it didn't go into its
defensive dig-in mode but carried on its
way. Among the rarer species, Mountain
Pygmy-possums and a Broad-toothed Rat
were caught on the summit of Mt
Kosciuszko. Just down from the summit in
the cross-drains beneath the walking track,
the Mountain Galaxias Galaxias olidus
showed which was Australia's highest fish.
One intrepid researcher was given the task
of documenting the feral fish using a dry
fly (Brown Trout were confirmed).

Bats presented à problem. It was easy
enough to set harp traps in flightways
through the trees but above the treeline
anabat detectors had to be used. The
Lesser Long-eared Bat, Chocolate Wattled
Bat and Southern Forest Bat were all
caught and the White-striped Freetail Bat
and Gould's Wattled Bat were identified
from their calls.

Dead male alpine funnel webs were
brought in by the public, their bodies are a
common sight in January (nearly as com-
monly seen as the males at night in
December as they wander across the grass-
land in search of females). It took a little
more effort to find live funnel webs, but a
live female was keen to demonstrate the
difference between modern and primitive
spiders. Primitive spiders have a vertical

Naturalist in the Mountains

bite and have to strike like snakes whereas
modern spiders bite horizontally.

Ted Edwards, who spent ten or so days in
the area, collected an estimated 400 moth
species including one last caught in 1922.
Another moth was previously collected
only once from Kosciuszko, the remaining
specimens known only from Tasmania and
Mt Buffalo. One new species was found,
and another moth of interest was a larva of
a Bogong Moth which turned up in a soil
sample. Other insects of interest were para-
sitic flies found remote from their hosts.

Among rarities, with a little adjustment
of the boundaries, we were able to include
one of the last ten known ponds in NSW
containing the endangered Alpine Treefrog
Litoria verreuxii alpina, but it has been a
long time since a Corroborree Frog has
been seen anywhere near the search area.
Thanks to the work of Alec Costin er al.
(their Kosciuszko Alpine Flora was
reviewed in volume 118 of The Victorian
Naturalist), there were no surprises among
the native vascular plants but we await the
identification of mosses, lichens and liver-
worts where there may be some new
species.

A report on the event with species lists
will be published in time for the
International Year of the Mountains
Conference in November this year.
Published by the Cooperative Research
Centre for Sustainable Tourism (Griffith
University) and the Australian Institute of
Alpine Studies, it will contain the methods
and the locations and where possible, a
complete species list as well as any inter-
esting findings to come out of the blitz.

Despite the complexity of the organisa-
tion it was good once again to participate
in an effort to document simply what is out
there as I creep towards my 25" anniver-
sary of naturalising in the mountains.

Ken Green
National Parks and Wildlife Service
PO Box 2228, Jindabyne, NSW 2627

Editors’ Note: To help celebrate the United Nations International Year of Mountains,
Ken Green will be our ‘Naturalist in the Mountains’ during 2002. We look forward to
publishing Ken's contributions about our mountain environments throughout the year,

Vol. 119 (1) 2002

37

Honours

Australian Natural History Medallion 2001
Alan Bridson Cribb

The winner of the 2001 Australian
Natural History Medallion was Dr Alan
Cribb, nominated by The Queensland
Naturalists' Club for an outstanding con-
tribution to natural history over a period of
fifty years, culminating in his publication

of the field guide Seaweeds of

Queensland. The nomination was support-
ed by The Royal Geographical Society of
Queensland for Dr Cribb's contribution to
four major expeditions, being involved in
research project selection, overseeing the
quality of submitted reports and refereeing
submitted papers. He attended the expedi-
tions to Cape York in the Wet Season and
to the Lawn Hill National Park.

Alan Bridson Cribb graduated with a
BSc from the University of Queensland in
1948 and subsequently gained a Masters
degree and PhD from the same university.
From the year of his graduation until 1988
he held positions at the University of
Queensland except for a short time with
CSIR Fisheries early in his career. He was
in turn, demonstrator, lecturer, senior lec-
turer and finally Head of the Botany
Department.

Dr Cribb is an authority in marine and
freshwater algae concentrating on system-
atics of the Queensland flora, marine and
terrestrial vegetation of the southern part
of the Great Barrier Reef, and marine
fungi. Voucher specimens from his
research form the bulk of the reference
collection at the Queensland Herbarium.
In association with his responsibilities for
teaching the principles of botany and biol-
ogy of terrestrial plants, Alan Cribb has
contributed significantly to the documen-
tation of the distribution of Queensland
flora by the provision of numerous check-
lists of plants for the National Parks and
Wildlife Service. Many of these record
new species or extensions of range. His
work on the Great Barrier Reef led to a
scheme of classification for the intertidal
reef and several papers on the algal ecolo-
gy of the area.

38

Dr Cribb has recorded his considerable
research activities in a total of 149 papers
and articles, 134 as sole author, and pub-
lished six books, four jointly with his
wife, Joan. His two books are Marine
Algae of the Southern Great Barrier Reef
Part I. Rhodophyta and the Seaweeds of
Queensland, He has described a number
of new species (24 algae, 14 marine fungi,
one flowering plant). Alan and his wife
kindled the current interest in native food
which had hitherto been overlooked by the
scientific community and general public.

In 1997 the Queensland Natural History
Award was presented to Dr Cribb for his
many contributions toward awareness and
knowledge of natural history in
Queensland. In addition to his academic
achievements he introduced a course at the
University of Queensland especially to
provide an appreciation of botany for non-
scientists. This theme was always present
in his talks to natural history groups and to
the public through radio and newspaper
articles, He has led many field excursions
for the Queensland Naturalists’ Club and
other groups and is a much appreciated
exhibitor at Club meetings. In retirement
since 1988, he has continued to promote
public awareness of the contribution of
plants to human welfare and their impor-
tance and role in biodiversity.

Dr Cribb has held membership of a num-
ber of professional and natural history
societies within the State, nationally and
internationally. This included the positions
of Editor and President for the Queensland
Naturalists’ Club and twenty years on the
council of the Great Barrier Reef
Committee. He was elected Fellow of the
Phycological Society (India).

In the area of conservation, Alan Cribb
has put his expertise to good use by
preparing submissions to relevant authori-
ties. These have included reports and evi-
dence to the Royal Commission on
Petroleum Drilling on the Great Barrier
Reef, the zoning of the Capricornia sec-

'The Victorian Naturalist

tion of the Great Barrier Reef Marine
Park, and the revegetation of North
Stradbroke Island after sand mining.
There are many people, not only mem-
bers of the QNC but also the public in
general, who are indebted to Dr Alan
Cribb for new knowledge and understand-
ing of natural history. For his contribu-

Tribute

tions to botany, marine biology and con-
servation, he is a worthy recipient of the
2001 Australian Natural History
Medallion.

Ian Endersby
56 Looker Road
Montmorency, Victoria 3094

Graham Martin Pizzey

4 July 1930 — 12 November 2001

i

A note of sadness was struck at the pre-
sentation of the Australian Natural History
Medallion on 12 November 2001, when
Tom May, after welcoming some previous
Medallionists, announced that Graham
Pizzey had died that day. Graham Pizzey
was awarded the Australian Natural
History Medallion in 1986, in recognition
of his long involvement with natural histo-
ry and conservation, and his work in shar-
ing this knowledge, through words and
photographs, with the general public. He
contributed a regular natural history col-
umn to The Age from 1954 to 1965 and to
The Herald from 1965 to 1983, The impor-
tance of ecology and the need for conser-
vation were reflected in the address he
gave to the Club on receiving the
Medallion. Too many people were
unaware of the natural world, being
focussed on material things, and the gap
between them and those that appreciated it
was wide, and he emphasised the need to
develop a ‘land ethic’? whereby people saw
themselves as belonging to the land rather
than its being something to be exploited.

Graham Pizzey played a significant part
in the campaign to protect the Little Desert
in the late 1960s, and in the conservation
movement which culminated in the estab-
lishment of the Land Conservation Council
in 1971, which led to a great increase in
protected land across the State. He was a
member of the National Parks Authority
from 1967-1971. In 1981 he was appointed

Vol. 119 (1) 2002

warden of the wildlife sanctuary at
Coolart, on Westernport Bay, greatly
appreciating the opportunity of involving
the public in the development of this
important wetlands site. Following this, in
1984, he and his wife, Sue, began a search
which led ultimately to their acquiring a
piece of natural bushland near Dunkeld, in
the Grampians, where, as he described in
articles in The Age, they could live and
observe nature, and provide an opportunity
for paying guests to share in this experience.

Although the move to Dunkeld meant
that Graham no longer participated in
many of the organisations with which he
had been involved, he continued as hon-
orary ornithologist to the Museum of
Victoria, and up until the time of his death
he was working on the protection of the
Brolga in western Victoria.

Birds were Graham Pizzey's major inter-
est and his authoritative book, 4 Field
Guide to the Birds of Australia, meticu-
lously researched and illustrated over 15
years, was published in 1980. It ran to 14
editions; the latest, completely revised,
appeared in 1997 and became a bestseller.

Graham Pizzey was a member of the
Field Naturalists Club of Victoria from
1962 to1994,

Sheila Houghton
12 Scenic Court, Gisborne
Victoria 3437

39

Book Review

Wyperfeld: Australia's First Mallee National Park

by Geoff Durham

Publisher: Friends of Wyperfeld National Park Inc., Elsternwick, Victoria, 2001.
Paperback, viii + 200 pp (including maps, black & white, and colour photographs,
and eleven appendices). ISBN 0 646 40101 7. RRP $25.00.

On the cover of this book is a picture of
the setting sun silhouetting a kangaroo and
her joey. Red light catches the hair around
her body and the grass at her feet, broken up
only by the long shadows cast by tree trunks
behind her. It is a scene recalled with plea-
sure by campers in any of the Mallee
national parks of northern Victoria. The
solitude, and remoteness, the ancient land-
scape, the regular sightings of large fauna,
the colourful sunsets followed by brilliant
starry skies on clear nights are all attractions
to travel to these wonderful places.

Wyperfeld National Park was, as the title
of the book says, Australia's first Mallee
national park. It started life as a 9,600 acre
(3,885 ha) reservation in 1909 and has been
added to a dozen times subsequently, with
the latest addition in 1992, making it, at
357,097 ha, the third largest national park
in Victoria (behind the Alpine and Murray-
Sunset National Parks). Wyperfeld lies in
thé north-western part of Victoria in an area
known as the Mallee and is dominated by
Mallee vegetation. The original section of
the Park was in the parish of Wyperfeld but
the origin of that name is obscure.

My father, lan Maroske, was a member
of the Committee of Management of
Wyperfeld National Park (later Advisory
Committee) from 1959 to 1993 when he
retired as Chairman. He joined after
curiosity about the large blanks on maps of
north-western Victoria led him to explore
the area. In his more than thirty years of
service he visited the Park numerous times
and his delight in what he saw never
dimmed, nor his wish to share that plea-
sure with others. His papers on the Park
are now held at the State Library of
Victoria, and include information for a
book that he was never able to publish.

If he had lived to see the use that Geoff
Durham has made of these and other mate-
rials he would have been very gratified.

40

Wyperfeld: Australia’s First Mallee
National Park is as handsome, accessible,
concise and informative a manual as any-
one could wish to see, and one that no visi-
tor to the Park will wish to be without.
Durham has presented an absorbing story
of the Park in 15 chapters accompanied by
numerous photos, maps and diagrams, all
carefully laid out by Leon Costermans.
There are eleven appendices at the end of
the book (including lists of plants and ani-
mals and an explanation of place names), a
bibliography and a comprehensive index.

The first part of the book deals with the
history of the Park, It includes information
on diverse topics such as the land, its peo-
ple and wild life, which Durham weaves
together skilfully into a sustained narra-
tive. The Wotjobaluk were dispossessed of
their land when the first settlers arrived in
the area in 1848. Within 50 years
Wotjobaluk culture was shattered and the
land also changed irrevocably. The birds,
however (including the Mallee Fowl),
were still sufficiently abundant at the end
of the century to attract the notice of field
naturalists and these visitors were the dri-
ving force in getting the first piece of land
reserved as a park in 1909.

Durham's chapters on natural history are
particularly good. In chapter four he
describes the three seasons of the Park, dry,
cold and breeding, and the ways in which
organisms have adapted to live through
them. Settlement has more often than not
disrupted these relationships. Exotic pests,
large bush fires, reduced water flows and
so on have all created management prob-
lems. For example, Cypress Pines burnt out
around Lake Brimin in 1946 showed no
signs of regenerating naturally due to over-
grazing by rabbits as well as kangaroos.
They have been replanted by hand and net-
ted to protect them, the nets only recently
having been removed.

'The Victorian Naturalist

In part two of the book, four distinctive
regions of the Park are described in detail:
the flood plain, Pine Plains, the eastern
interface and the western wilderness. The
flood plain is the termination of the
Wimmera River and splits the Park in two
from south to north. Floods are now rare in
the Park and one of my cherished memo-
ries is of swimming in Outlet Creek in
1976 (the last time that water reached the
Park). The northern most point of the flood
plain is Wirrengren Plain which is part of
Pine Plains. The land to the east of the
flood plain is a strong-hold of the Mallee
Fowl and to the west is a wilderness which
almost doubled the size of the Park when it
was incorporated in 1991.

Part three of the book consists of general
visitor information. It details facilities at
Wonga Camp Ground and Casuarina
Camp Ground in the Park, and in nearby
settlements. The established walks and dri-
ves in the Park are sufficiently numerous
and various to suit most kinds of visitors.
Maps and some notes (especially for the
drives) are provided in the book and com-
plement the information sheets produced
by Parks Victoria. Additional information
on the Park can be found using the bibliog-
raphy at the back of the book, or making a
trip to the Information Centre at Wonga
Camp Ground.

Book Review

I have many wonderful memories of
Wyperfeld National Park. As a child I
recall the approach through seemingly
interminable cleared land to the south. At
last the rustic gateway (no longer extant)
would appear in front of the car and we
would pass through into the bush with evi-
dence of European settlement rapidly
receding. We followed a ribbon of Black
Box along the course of Outlet Creek to
Wonga Camp Ground on the lookout for
kangaroos and emus. There we were often
met by the Park's first Ranger Rudd
Campbell who always had some pepper-
mints for my brother and sister and me.

Wyperfeld is not a place of dramatic vis-
tas like Mt Buffalo or Wilsons Promontory,
but its landscape is attractive. This wonder-
ful book will make an important contribu-
tion to increasing appreciation of the Park.
It is, as it says of itself, *an invitation, an
introduction and a practical guide with
background information to enhance every
visit’. It was published by the Friends of
Wyperfeld National Park and will make the
Park many new friends. | will have it, open,
in my hand on my next visit to the Park,
which I hope will be very soon.

Sara Maroske

1 Wastell Street
Northcote, Victoria 3070

Footnote: The book's editor, Leon Costermans, wishes to record some corrections to minor errors

which occurred in the production process.

Page 43: The caption for photo 6 should read *6 Copper Stag beetle (Lamprima varians)

Where Stipa is given in the text as the genus for Spear-grass species, this is synonymous with the
more recent Austrostipa S.W.L. Jacobs and J. Everett (1996), as given in the plant list on page 152.

Page 111: The photo caption should read *An overnight bushwalking group at Wonga Lake’,

(Round Lake is not ringed by Red-gums!).

Page 136: Under Walks, delete the word ‘Track’ in *... south-west to Outlet Creek Track ...".

Page 139: In the paragraph under Day visitors, replace the last sentence with ‘From here, the gravel
road is usually reasonable for two-wheel drive vehicles for 10 km northwards’.

Page 156: Under Epacridaceae, the common name for Brachyloma ericoides subsp. ericoides should
read Brush Heath. (Pink Beard-heath’ is Leucopogon ericoides which is not recorded north of the

Little Desert).

Page 171: After Silver Gull (Larus novaehollandiae), delete the code +A’.

Page 190: In the Bibliography, add: Cogger, H.G. 2000, Reptiles & Amphibians of Australia, 6th
edn, Reed New Holland, Sydney. This is referred to on page 168.

In the index, several page references are in error by one page, due to last-minute layout adjustments.

Vol. 119 (1) 2002

July 2001

41

Guidelines for Authors — The Victorian Naturalist

Submission of all Manuscripts

Authors may submit material in the form of

research reports, contributions, naturalist notes,
letters to the editor and book reviews. A
Research Report is à succinct and original sci-
entific paper written in the traditional format
including abstract, introduction, methods, results
and discussion. A Contribution may consist of
reports, comments, observations, survey results,
bibliographies or other material relating to nat-
ural history. The scope of a contribution is broad
and little defined to encourage material on a
wide range of topics and in a range of styles.
This allows inclusion of material that makes a
contribution to our knowledge of natural history
but for which the traditional format of scientific
papers is not appropriate. Research reports and
contributions must be accompanied by an
abstract of not more than 200 words. The
abstract should state the scope of the work, give
the principle findings and be complete enough
for use by abstracting services. Research reports
and contributions will be refereed by external
referees. Naturalist Notes are generally short,
personal accounts of observations made in the
field by anyone with an interest in natural histo-
ry. These may also include reports on excur-
sions and talks, where appropriate, or comment
on matters relating to natural history. Letters to
the Editor must be no longer than 500 words.
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quent delay of publication.

Taxonomic Names

Cite references used for taxonomic names.
References used by The Victorian Naturalist are
listed at the end of these guidelines.

Abbreviations

The following abbreviations should be used in
the manuseript (with italics where indicated): er
al.; pers. obs.; unpubl. data; and pers. comm.
which are cited in the text as (RG Brown 1994
pers. comm. 3 May). Use *subsp.' for subspecies.

Units

The International System of Units (SI units)
should be used for exact measurement of physi-
cal quantities.

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All illustrations (including photographs) are
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tographs should be of high quality/high contrast
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They may be colour slides, colour or black-and-
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may be computer generated or in black Indian Ink
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All figures and tables should be referred to in the
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etc.) and put on a separate page at the end of the
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tively (Table 1, Table 2, ete.) and have an explana-
tory caption at the top.

Please consult the editors if additional details are
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arrangements.

Journal Style

Authors are advised to note the layout of head-
ings, tables and illustrations as given in recent
issues of the Journal. Single spaces are used
afler full stops, and single quotation marks are
used throughout,

In all papers, at the first reference of a species,
please use both the common name and binomial,
However, where many species are mentioned, à
list (an appendix at the end), with both common
and binomial names, may be preferred. Lists
must be in taxonomic order using the order in

which taxa appear in the references recommend-
ed below.

The journal uses capitalised common names
for species followed by the binomial in italics
without brackets, e.g. Kangaroo Grass Themeda
triandra.

References

References in the text should cite author and
year, e.g. Brown (1990), (Brown 1990), (Brown
1990, 1991), (Brown 1995 unpubl. ), (Brown and
Green 1990), (Brown and Green 1990; Blue
1990; Red 1990). If there are more than two
authors for a paper use Brown ef al. (1990).
These should be included under References. in
alphabetical order, at the end of the text (see
below). The use of unpublished data is only
accepted if the data is available on request for
viewing. Pers, obs. and pers. comm. should not
be included in the list of references. Journal
titles should be quoted in full.

Leigh J, Boden R and Briggs J (1984) Extinct
and Endangered Plants of Australia.
(Macmillan: South Melbourne)

Lunney D (1995) Bush Rat. In The Mammals of
Australia, pp 651-653. Ed R Strahan.
(Australian Museum/Reed New Holland:
Sydney)

Phillips A and Watson R (1991) Xanthorrhoea:
consequences of ‘horticultural fashion’. The
Victorian Naturalist 108, 130-133.

Smith AB (1995) Flowering plants in north-
eastern Victoria. (Unpublished PhD thesis,
University of Melbourne)

Wolf L and Chippendale GM (1981) The natural
distribution of Eucalyptus in Australia.
Australian National Parks and Wildlife
Service, Special Publications No 6, Canberra.

Other methods of referencing may be acceptable
in manuscripts other than research reports, and
the editors should be consulted. For those using
the bibliographic software ‘EndNote’, a style
guide for The Victorian Naturalist will be avail-
able shortly on our website. For further informa-
tion on style, write to the editors, or consult the
latest issue of The Victorian Naturalist or Stvle
Manual for Authors, Editors and Printers
(Australian Government Publishing Service:
Canberra).

Manuscript Corrections

Authors can verify the final copy of their man-
uscript before it goes to the printer. A copy of
their article as ‘ready for the printer’ will be sent
and only minor changes may be made at this
stage.

Complimentary Copies

After publication of an article in the journal,
five complimentary copies of that issue are sent
to the author(s) for each paper. Authors of
Naturalist Notes and Book Reviews will receive
two complimentary copies of the journal.

Additional copies of The Victorian Naturalist:
25 copies, $50.00 (+ postage); 50 copies, $90.00
(+ postage), including GST. Additional copies
must be ordered before printing.

Checking species names is the responsibility of authors. The books we would like used as references
for articles in The Victorian Naturalist are listed below. Authors should refer to the source used for
species names in their manuscripts. In every case, the latest edition of the book should be used,

Mammals — Menkhorst PW (ed) (1995)
Mammals of Victoria: Distribution, Ecology
and Conservation. (Oxford University Press:
South Melbourne)

Reptiles and Amphibians ~ Cogger H (2000)
Reptiles and Amphibians of Australia, 6 ed.
(Reed Books: Chatswood, NSW)

Insects — CSIRO (1991) The Insects of Australia:
a textbook for students and research workers.
Vol I and TH. (MUP: Melbourne)

Birds - Christidis L and Boles W (1994) The
Taxonomy and Species of Birds of Australia
and iis Territories. Royal Australian
Ornithologists Union Monograph 2. (RAOU:
Melbourne)

Plants - Ross JH (ed) (2000) 4 Census of the
Vascular Plants of Victoria, 6 ed. (Royal
Botanic Gardens of Victoria: Melbourne)

The Editor

Please submit manuscripts and enquiries to:

The Victorian Naturalist
Locked Bag 3, P.O.
Blackburn, Victoria 3130

Phone/Fax (03) 9877 9860. Email fnev@vicnet.net.au

Web address: http://www. vicnet.net.au/~fnev/

How to Identify Wildflowers of the Grampians

by Ken Woodcock

Publisher: Tke Community Association of Halls Gap Inc. RRP $10.50

This is an unusual little book illustrated
with 150 very small colour drawings and
minimal text. There are six illustrations on
each page and species are grouped accord-
ing to flower colour. This presents some
problems in grouping the blue, violet,
mauve and pink species, where the subtle
colour differences are notoriously difficult
to describe or reproduce. The descriptions
are brief with one or two important fea-
tures arrowed on the drawing. Both the
common and botanical names are given
but any technical or botanical terms are
entirely avoided. This makes it easy for
beginners in wildflower study and should
enable recognition at least to genus.
However problems may occur in differen-
tiating the species in genera such as
Hibbertia or in the pea flowers, where sub-
tle differences in leaf characters can be
critical. For example no mention is made
of the leaves being in very obvious oppos-
ing pairs (decussate) in Eutaxia. This char-
acter is easy to see and distinguishes
Eutaxia from all other yellow or orange
flowered peas in the Grampians. A note or
sketeh of the legume shape would also
have been helpful in sorting out the peas.

Keeping abreast of changing nomencla-
ture is a problem for both authors and nat-
uralists. This should not deter beginners
from learning about native plants; even the
most knowledgeable of naturalists are
rarely completely up to date. An addendum
paragraph after the index lists some recent
changes; however Leionema bilobum is
still the valid name for what was formerly
Phebalium bilobum. The illustration of
Utricularia is presumably U. dichotoma,
not U. uniflora that is an East Gippsland
species,

This very elementary and relatively
cheap little book could be useful to those
totally unfamiliar with wildflowers and
would probably enable them to recognise
many of the common flowers, at least to
genus. Anyone seeking more information
would need to refer to one of the several
other more detailed and comprehensive
books dealing with Grampians plants.

Margaret Corrick
7 Glenluss Street
Balwyn, Victoria 3103

The Field Naturalists Club of Victoria Inc.

Reg No A0033611X

Established 1880
In which is incorporated the Microscopical Society of Victoria

OBJECTIVES: To stimulate interest in natural history
and to preserve and protect Australian flora and fauna.
Membership is open to any person interested in natural history and includes
beginners as well as experienced naturalists.
Registered Office: FNCV, 1 Gardenia Street, Blackburn, Victoria 3130, Australia.
Postal Address: FNCV, Locked Bag 3, PO Blackburn, Victoria 3130, Australia.
Phone/Fax (03) 9877 9860; International Phone/Fax 61 3 9877 9860.

Patron
John Landy, MBE, The Governor of Victoria

OPI Ve
Victorian
Naturalist

Volume 119 (2) April 2002

Published by The Field Naturalists Club of Victoria since 1884

Birds of Box Hill

by Tess Kloot with illustrations by Nicolas Day

Publisher: Victorian Ornithological Research Group, 2000. RRP $18.00 (* $2 postage
per copy) (Order from VORG, 133 Graydons Road, Moorooduc, Victoria 3933)

The Birds of Box Hill derives from a
study undertaken by the Victorian
Ornithological Research Group (VORG)
between 1988 and 1991, and it is refresh-
ing to see amateur research organizations
publishing the fruit of their work in a man-
ner accessible to the public. This book cer-
tainly has a lot in it that will appeal, espe-
cially to the residents of Box Hill and other
eastern suburbs of Melbourne. The VORG
survey arose from a desire to compare the
avifauna of Box Hill in the late 1980s with
that recorded by one Robert Hall, a local
naturalist, in the 1890s, when Box Hill was
a sparsely populated area. The book not
only presents a guide to the birds found in
the Box Hill area but also presents the

reader with an idea of the abundance of

each species and will be useful for genera-
tions to come as a base-line of the avifauna
of an established suburb. When Robert
Hall was conducting his original surveys
the area was mainly a collection of farms
with much remnant vegetation - times have
certainly changed.

The first part of the book is basically a
field guide to the birds of Box Hill, with
concise, pertinent text and excellent black
and white illustrations from Nicolas Day.
However the use of colour would have been
far more desirable in an identification guide
aimed at novices. In all a respectable 78
species were recorded on a regular basis and
are illustrated, with eight being introduced.
The thirty-seven species that were infre-
quently recorded during the VORG survey
(seen four times or less) include some very
interesting and exciting sightings such as
Swift Parrot, Baillon’s Crake, Rufous
Songlark and some obvious escapees such
as Budgerigar and Zebra Finch. These
species are not illustrated which I think is
entirely appropriate if the field guide por-
tion of this book is aimed at novices.

The chapter on Selected Birdwatching
Areas of Box Hill gives a good guide to

these areas although this chapter and those
on Parks and Gardens of Box Hill and
Birds Recorded in Selected Areas could
have been combined. I really enjoyed read-
ing the chapters on the history of Box Hill
and on Robert Hall's work. Lots of inter-
esting snippets and facts can be gleaned
from this book. I particularly liked the list
of birds recorded at the end of the 19" cen-
tury which includes their common name of
the time, for example Sordid Woodswallow
(Dusky Woodswallow), Funeral Cockatoo
(Yellow-tailed Black Cockatoo), and Rose
Hill Parrot (Eastern Rosella).

I have one major quibble with the book
given that there are chapters on Attracting
Birds to your Garden, Robert Hall (1867-
1949) One Hundred Years On, and The
History of Box Hill (which includes the
origin of the suburb's name) there is very
little prominence given to the vegetation of
the area and its change over the past centu-
ry. | am none the wiser after reading this
book about the original flora of this area.
Although given the loss of woodland bird
species (White-winged Chough, Diamond
Firetail, Regent Honeyeater, Spotted
Quail-thrush, Brown Treecreeper and
Speckled Warbler for example) in the past
hundred years it is presumed that a large
part of the area was open eucalypt (box)
woodland with a grassy understorey.

All up a terrific little book that will hope-
fully appeal to residents of Melbourne’s
middle eastern suburbs and those of us
who have a deeper interest in the changes
in Victorian avifauna wrought over the last
hundred years. If it inspires only some Box
Hill residents to take more interest in their
natural surrounding, which it will, it
should be considered a great success.

Stuart Dashper
13 Ryan Street

East Brunswick
Victoria 3057

The

Victorian
Naturalist

Volume 119 (2) 2002 S April

Executive Editor: Merilyn Grey
Editors: Alistair Evans and Anne Morton

Index to Volume 118, 2001 is in the centre of this issue

Research Reports A Herpetofauna Survey of the Victorian Alpine Region,
with a Review of Threats to These Species, by Nick Clemann.....48

The Mammals of Parker River Inlet, Otway National Park,

DY MAIV ESRO ORANA PUMEBPÓPelt IE Ao eiae eee oh 60
Contribution Habitat of the Endangered Hibbertia procumbens (Labill.) DC
(Dilleniaceae) from the Central Coast of New South Wales,
MEP BH auo Eee TU Ie in Pro oie a eria rur does 69
Naturalist Notes Wingan Wilderness, by Ron Fletcher................sceeeenenee 74
Naturalist in the
Mountains The Changing Mountains, by Ken Green .............. eee 76
Tributes Jack Hyett OAM, by Cecily Falkingham ................. ee 78
Alexander Clifford Beauglehole, by Margaret Corrick
Book Reviews Birds of Box Hill, by Tess Kloot with illustrations by
Nicolas Day, reviewed by Stuart Dashper ................. eene 46

Wild Solutions: How Biodiversity is Money in the Bank,
by Andrew Beattie and Paul Ehrlich, reviewed by TR New ......... 58

A Field Companion to Australian Fungi, by Bruce Fuhrer,
PEVIGVIEE DV EHV) FORK A IE rea aan aridis ccc ES 79

Wildflowers of Sydney and Adjoining Areas, by Alan Fairley,
reviewed by Cecily Falkingham................. eter 83

ISSN 0042-5184

oo
Cover: Tall Everlasting Helichrysum elatum photographed by Ron Fletcher at Wingan Inlet. See
Naturalist Note on page 74.

Web address: http://www.vicnet.net.au/~fnev/
email: fncv@vicnet.net.au

Research Reports

A Herpetofauna Survey of the Victorian Alpine Region, with a
Review of Threats to These Species

Nick Clemann!

Abstract

A wide-ranging survey of the reptiles and frogs of the Victorian alpine region was conducted in sum-
mer and early autumn 2001. Eight habitat ‘types’ that were easily discernible in the field were identi-
fied, and randomly-positioned sites chosen in these habitats. Surveys of these sites were conducted
using ‘active search’ techniques, and results bolstered with incidental records, Three frog and eleven
reptile taxa were recorded across all areas and habitat types. Notable records included a range exten-
sion for the threatened Alpine Bog Skink Pseudemoia cryodroma, and the collection of specimens of
undescribed lizards from the genus Egernia from the Bogong High Plains, Davies Plain and Mt
Bogong, Threats to Victorian alpine herpetofauna are discussed, as is the dramatic decline in many
areas of the Alpine Tree Frog Litoria verreauxii alpina. (The Victorian Naturalist 119 (2), 2002, 48-58)

Introduction

The herpetofauna (reptiles and frogs) of
the alpine region of Victoria is largely
endemic to alpine areas, the general ecolo-
gy of these taxa is poorly understood, and
they face a suite of threatening processes,
The aim of this project was to refine our
understanding of the habitat associations of
these taxa, and the processes that threaten
them. To this end, a broad-scale field sur-
vey and a detailed literature search were
carried out.

The alpine region of Victoria embraces
the highest plateaux in the State, and
extends from the Mt Cobberas area near the
border with New South Wales in the north-
east, along the top of the Great Dividing
Range, to the Mt Baw Baw area in the
south-west. This region can be crudely
defined as commencing at about 1200 m
altitude, and extending to the highest point
in Victoria, the summit of Mt Bogong at
1986 m. It receives a consistent snow cover
during winter months.

The alpine region contains several species
of threatened herpetofauna. This fauna
(with threatened categories according to
NRE (2000) in parentheses) includes:
Alpine Tree Frog Litoria verreauxii alpina
(listed as Vulnerable nationally on
Australian and New Zealand Environment
and Conservation Council (ANZECC 1999)
and Endangered Species Protection Act
1992 lists, and Critically Endangered in
Victoria), the Baw Baw Frog Philoria frosti
(Vulnerable on the ANZECC list, Critically
Endangered in Victoria), Alpine Water
' Arthur Rylah Institute for Environmental Research,

Department of Natural Resources and Environment,
PO Box 137, Heidelberg, Victoria 3084

48

Skink Eulamprus kosciuskoi (Critically
Endangered), Alpine She-oak Skink
Cyclodomorphus praealtus (Endangered),
Alpine Bog Skink Pseudemoia cryodroma
(Vulnerable, and endemic to Victoria) and
Glossy Grass Skink Pseudemoia rawlinsoni
(Lower Risk — Near Threatened). With the
exception of the Glossy Grass Skink, these
species are restricted to alpine areas. In
addition to these species, two undescribed
taxa of the scincid genus Egernia occur in
the region. Draft Action Statements have
been developed for the Alpine Water Skink
(Meredith et al. in press) and Alpine She-
oak Skink (Clemann in press).

The biology and ecology of these species
are poorly known, as are the processes that
threaten them. Plausibly, these threats
include grazing and trampling of habitat by
cattle, feral horses and deer, predation by
introduced carnivores such as cats and
foxes, habitat modification or destruction
due to development (e.g. ski resort infra-
structure, walking and vehicular tracks).
recreational activities or the invasion of
weeds, illegal collection, inappropriate fire
regimes and global warming due to the
effect of greenhouse gases. This latter
process is of particular concern for alpine
fauna; under an enhanced greenhouse sce-
nario, considerable contraction of alpine
habitat may occur, with little or no oppor-
tunities for emigration to other suitable
locations. Whilst the Alpine Tree Frog is
known to have suffered major declines
throughout much of its range (Gillespie et
al. 1995; Hunter ef al. 1997), the current
status of alpine reptiles is poorly known.
However, some of these species may also

The Victorian Naturalist

have declined due to one or more of these
processes.

The alpine region of south-eastern
Australia contains other threatened alpine
vertebrate species: Southern Corroboree
Frog Pseudophryne corroboree (Endan-
gered in NSW), Northern Corroboree Frog
P, pengilleyi (Vulnerable in New South
Wales), and the Mountain Pygmy Possum
Burramys parvus (Vulnerable on the
ANZECC list, Endangered in Victoria).
Several vegetation communities, namely the
Alpine Bog Community, Alpine Snowpatch
Community and Fen (Bog pool) Commun-
ity, are also listed on Schedule 2 of the
Victorian Flora and Fauna Guarantee Act
1988 (FFG). Also listed on the FFG as a
potentially threatening process is soil ero-
sion and vegetation damage and disturbance
caused by cattle grazing.

Methods
Study areas

Study areas were selected to represent
alpine areas across north-eastern Victoria
that were relatively accessible and have a
continuing history of human presence, and
thus were likely to contain a considerable
degree of anthropogenically-generated dis-
turbance. Examples of the processes that
typify such disturbances include mining,
recreational pressure, resort development,
stock and feral herbivore grazing and tram-
pling, and track and road construction.
Logistic and time constraints restricted sur-
veys to plateaux with relatively easy
access. As such the following areas were
chosen for surveys: Mt Buller/Mt Stirling,
the Dargo High Plains, Davies Plain,
Dinner Plains, Falls Creek/Bogong High
Plains, the Howitt Plains, Lake Mountain,
Mt Bogong and Mt Buffalo. Several of
these *areas' are generic terms for series of
alpine plains. For example “Howitt Plains’
incorporates areas such as Bryces, Snowy
and Bennison Plains; ‘Dinner Plains’
incorporates areas such as Flourbag,
Horsehair and Emu Plains; and ‘Dargo
Plains’ incorporates areas such as Gow,
Lankey, Treasure, Halfway and One Pole
Plains.
Habitat

Eight broad habitat types that were easily
identified in the field (bog, cleared and/or
disturbed, tussock grassland, rocky grass-

Vol. 119 (2) 2002

Research Reports

land, heathland, montane dry woodland,
Snow Gum Eucalyptus pauciflora wood-
land and Rocky Outerop Shrubland/Herb-
land Mosaic — this last habitat type is spe-
cific to the Mt Buffalo area) were chosen
for general herpetofauna surveys, and an
effort made (where possible) to sample
two sites in each habitat type in each area.
Although most of these sites showed vary-
ing degrees of disturbance, sites classified
as *cleared/disturbed’ were usually grossly
modified. These sites included areas such
as cattle yards, ski runs, and areas used for
rock or gravel extraction or surrounding
human developments such as huts and
carparks.

These eight habitat types often were not
strictly discrete, and considerable variation
and overlap between habitat types was evi-
dent. For example, ‘heathland’ sites were
often found to surround bogs. This meant
that, whilst surveying a ‘bog’ site, this site
would grade into heath (and vice versa),
often with dense Epacris sp. in the inter-
grade zone. ‘Grasslands’ were often a com-
bination of tussock grass, herbfields and
very low (usually < 30 cm high) heath.
Snow Gum woodland, whilst always domi-
nated by an overstorey of Snow Gum, var-
ied markedly in understorey and ground-
storey composition. For example, some
Snow Gum woodland sites contained
extremely dense, head-high heath, others
contained a low, tussock grass groundcover,
and many were intermediate between these
extremes. Similarly, some Snow Gum sites
contained numerous, large rock outcrops,
whilst others were devoid of rocks.

Site selection

Within each survey area (‘Study areas’
above) a series of random grid points was
generated and plotted onto maps. Once in
the field, these grid points were considered
potential sites, and were visited (with
emphasis on those sites within a reason-
able walking distance from roads and
tracks) and systematically surveyed until
two sites had been sampled in each habitat
type in each area. All other grid points
were then disregarded. Using this system-
atic approach, grid points that occurred in
habitats that had already been adequately
surveyed were also disregarded. Wherever
possible, all sites were located more than

49

Research Reports

one km from any other site. Not all habitat
types were detected in each area, In some
areas it was only possible to locate enough
habitat to survey one site in some habitat
types. Also, in some areas it was necessary
to select habitat in a non-random manner,
because there were only one or two patch-
es of this localised habitat in some general
survey areas,

Survey techniques

Within each site a time-constrained (20
minute) ‘active search’ limited to the
appropriate habitat type was conducted
between the hours of approximately 0800-
1830 daylight savings time. This technique
involved noting all basking or active rep-
tiles, and any frog calling activity. Rocks
and logs were rolled, soft logs were
cleaved, loose bark was peeled, and litter
and gravel sifted to detect sheltering ani-
mals. At each site the habitat type and any
obvious disturbance were noted, and gen-
eral weather variables such as temperature.
cloud cover and wind velocity were
recorded,

This application of a single survey tech-
nique for a variety of taxa has obvious lim-
itations, including failure to detect particu-
larly cryptic species, and a high likelihood

of underestimating the abundance of

Species that are difficult to detect during
active searching. For example, species that
prefer dense tussock grass habitat are usu-
ally very difficult to detect or capture (for
confident identification) using this tech-
nique. Thus, the following results provide
only a coarse index of abundance.

Results

Three frog taxa (Alpine Tree Frog,
Common Froglet Crinia signifera and
Victorian Smooth Froglet Geocrinía victo-
riana) and 12 reptile taxa were recorded
either during surveys or incidentally (Table
1). Several threatened taxa were also
recorded, and these included the Alpine
Iree Frog on the Dargo Plains and Mt
Bogong, Alpine Water Skink on Davies
Plain and the Bogong High Plains north-
east of Falls Creek, and Alpine Bog Skink
in the Buller/Stirling, Davies Plain, Howitt
Plains, possibly Mt Buffalo (see below)
and Lake Mountain areas. This latter local-
ity represents an extension of the known
range of this species.

50

Several specimens of Pseudemoia could
not be confidently assigned to a particular
species. In particular, there was confusion
regarding intermediate colouration and mor-
phology between specimens of P. eryodro-
ma and P. pagenstecheri. Voucher speci-
mens of these taxa from these areas were
sent to the South Australian Museum
(whose staff described these taxa) for resolu-
tion. Despite this, some specimens from Mt
Buffalo could not be confidently identified.

Specimens of undescribed lizards from
the genus Egernia were collected on the
Bogong High Plains (Ruined Castle south
of Falls Creek), Davies Plain and Mt
Bogong. Additionally, a gravid female had
been collected on Mt Buffalo in December
1997, and one of this lizard's offspring has
been sent to the South Australian Museum
for taxonomic study, along with the
Fgernia and Pseudemoia specimens col-
lected during the present study.

The Alpine She-oak Skink was not
recorded during this study, despite surveys
and incidental searches in localities from
which this species had previously been
recorded (Lankey Plain on the Dargo High
Plains, Ruined Castle and Basalt Hill on
the Bogong High Plains and several locali-
ties in the vicinity of Mt Hotham) (Schulz
et al. 1995; Heinze 1997: Schulz and
Mansergh 1997; Atlas of Victorian
Wildlife database).

Two taxa were recorded from only a sin-
gle area. The Victorian Smooth Froglet
was detected in the Mt Buller/Mt Stirling
area, and the Black Rock Skink Egernia
saxatilis intermedia was detected on Mt
Buffalo (Table 1). The Alpine Water Skink
was detected only on Davies and the
Bogong High Plains. Conversely, the
Southern Water Skink Eulamprus tympa-
num was detected at eight of the nine
areas, and Tussock Skinks Pseudemoia
pagenstecheri were recorded at seven,

Davies Plain yielded the highest number
of taxa (not including taxa not identified to
species level) during this study, with 10
species, whilst nine were recorded on the
Howitt Plains (Table 1). All of the other
areas returned between five and eight
species,

The mean number of reptile species
recorded for each habitat type was four
(4.4 when Rocky Outerop Shrubland/

The Victorian Naturalist

Research Reports

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51

Vol. 119 (2) 2002

Research Reports

Herbland Mosaic, found only on Mt
Buffalo is disregarded). Both the Southern
Grass Skink Pseudemoia entrecasteauxii
and Tussock Skink were recorded in six
habitat types, and the Southern Water
Skink in five (Table 2). Species recorded
in only one habitat type include Coventry's
Skink Niveoscincus coventryi, Black Rock
Skink and Egernia sp. (although these
undescribed taxa were also incidentally
recorded in Snow Gum woodland). The
Alpine Water Skink was recorded in two
habitat types (bog and heathland), although
they show considerable overlap.

Bog habitat yielded the highest number
of reptile species (six) across all areas,
whilst heathland and rocky grassland
returned five species (Table 2). Other habi-
tat types yielded three or four species, with
the exception of Rocky Outcrop Shrub-
land/Herbland Mosaic (one species from
two siles on Mt Buffalo).

lhe largest reptile occurring in the alpine
region, the Highland Copperhead Austre-
laps ramsayi, was recorded at five of the
eight study areas. This species occasional-
ly reaches considerable densities for a top-
order predator, with up to four adults
recorded in areas of approximately
1600 m` during this study. These high den-
sities were noted where there was also a
high abundance of scincid lizards, particu-
larly Southern Water Skinks.

Discussion

During this study the Alpine Tree Frog
was not detected at several localities such
as Lake Mountain and the Howitt, Davies
and Bogong High Plains where it was once
known to be abundant and where many
specimens of this subspecies in the
Museum Victoria collection originate
(Atlas of Victorian Wildlife database).
Most of these specimens were collected in
the 1960s and '70s. The last records of this
frog from Lake Mountain and Davies Plain
were from 1993 and 1994, respectively
(Atlas of Victorian Wildlife database).
Whilst intensive frog surveys have been

conducted during the breeding season of

the Alpine Tree Frog on Mt Baw Baw
(where this taxa was once abundant) for
the last six years, no specimens have been
detected, suggesting that this frog no
longer occurs in this area,

52

The apparent declines detected in the pre-
sent study accord with those detected by
other authors. The population status of this
taxon was reviewed in 1995 (Gillespie er
al. 1995), at which time it appeared that it
had suffered major population declines in
the Australian Capital Territory and
Kosciusko National Park in New South
Wales. Subsequent surveys funded by the
Alps Liaison Committee confirmed major
declines in this region of its range, but did
not adequately assess the taxon’s status in
Victoria. Other targeted frog surveys con-
ducted on Mt Baw Baw (G Hollis unpubl.
data) and Lake Mountain (B Malone pers.
comm.) indicate that the taxon has disap-
peared from these parts of its former range.
Osborne et al. (1999) suggest that the
Alpine Tree Frog is in immediate danger
of extinction in New South Wales and is
highly threatened in Victoria.

Current knowledge of the status of the
Alpine Tree Frog and the nature of declines
has been reviewed recently (Osborne er al.
1999). It is clear that these declines have
occurred concurrently with those in all
other endemic alpine frog taxa (Philoria
frosti, Pseudophryne corroboree, P.
pengillevi, Limnodynastes dumerilii fryi).
However, it is still unclear what the causes
of these declines are. Possible causes
include changes in climate and increased
ultraviolet radiation. The disease
Chytridiomycosis, caused by a fungal
pathogen, has recently been implicated in
the decline of numerous frog species,
including taxa closely related to the Alpine
Tree Frog (Berger er al. 1999), and may yet
be found to be contributing to the decline
of the taxon. Post-European settlement
impacts, such as grazing and fire regimes
have also been implicated (Gillespie er al.
1995), Recent evidence from North
America (Kiesecker ef al. 2001) suggests
that synergistic affects of several processes
may cause the decline of some amphibian
species. In particular, Kiesecker er al.
(2001) have shown a pattern whereby cli-
mate change has reduced water levels at
some oviposition sites, causing high mor-
tality of embryos by increasing their expo-
sure to UV-B radiation (which is increasing
due to ozone depletion) and, consequently,
increasing embryo vulnerability to infec-
tion by pathogens.

The Victorian Naturalist

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53
2

wiih iiai
l.

Vo

Research Reports

ilf: Lars: om

Vig, L Alteration of vegetation structural complexity by cattle grazing and trampling on the Howitt

Plains, Heavy gazing and trampling inside the fenced area has profoundly modified the structural

integrity of the tussock grassland

* Alpine Water Skink (Meredith er al in
press); and
* Alpine Bop Community (NRE in prep.).
Similarly, soil erosion and vegetation dam-
ape assoctited with grazing are the primary
processes currently threatening the Alpine
Bog Community, Alpine Snowpatch
Community and l'en. (Bog pool)
Community, all of which have been listed
as threatened on Schedule 2 of the PEG,
Ihe Alpine Water Skink is restricted to
alpine sphagnum mossbed-heath associa-
tions (that forms "bog! and some “heath.
land? habitat as defined in the present
study) along drainage lines and small
streams, This habitat type is particularly
susceptible to damage by cattle seeking
water and grazing on the palatable sedge
Carex gaudichaudiana (McDougall 1982,
1989; NRE in prep). Severe habitat degra-
dation caused by grazing and trampling
has been noted at several sites where this
lizard occurs (Meredith er a/ in press;
pers, obs. Pig. 3). Whilst cattle typically
only graze on the sedge in late summer
(van Rees 1984), Me Dougall (1989) found
that this short exposure to cattle was suffi-
vient to severely damage bog margins,

54

l'videnee of extensive trampling of this
habitat by cattle is obvious in all areas
where cattle graze in the Victorian alps
(Wahren ef al, 1999; pers, obs.); This dam-
ape may eventually exclude the Alpine
Water Skink from these habitats. Other
authors (Coventry and Robertson. 1980;
Mansergh 1982) have also highlighted this
threat to the habitat of the Alpine Water
Skink,

The Alpine Water Skink occurs in the
stream/bog system along Nelse Creek,
north-east of Falls Creek. Within these
bogs the lizards use the cornice-like sphag-
num overhanging the water as basking
sites, When approached, they frequently
drop from this overhang into the water and
retreat beneath the overhang to avoid cap-
ture, This sphagnum margin is usually
destroyed by cattle seeking water and Tod-
der (NRE in prep.: pers. obs.; Pig. 4).
McDougall (1989: 8) notes that “the grow-
ing edge of the layer bordering pools is
most susceptible to dislodgment and
destruction by trampling’, Trampling is
likely to destroy a vital habitat component
lor these lizards, The Alpine Water Skink
has been recorded on the Bogong High

The Victorian Naturalist

Plains both north of Rocky Valley Storage,
where grazing has been excluded since
1991, and south of this dam where grazing
continues. During the present study we
detected this species in bog habitat in this
northern area, but did not detect it in the
currently grazed areas to the south. Further
work is needed to compare this species’
persistence and abundance in these areas.
Such work will further illuminate the
Alpine Water Skink’s response to grazing.
and its response to removal of grazing.

Less obvious than the highly visible dam-
age stock often cause to bogs, is alteration
to the hydrology of stream and bog sys-
tems caused by trampling. This impact can
result in channelling and resultant lowering
of the water table (LCC 1982). Bog sys-
tems and the peat soils derived from them
are dependent on a high water table. As
these peat soils dry out they become sus-
ceptible to erosion; thus grazing within
these zones causes high levels of damage
leading in the past to these areas being pro-
gressively withdrawn from grazing (LCC
1982). Of the broad habitat types surveyed
for this study, bog was the most species
rich for reptiles, and is probably the most
threatened habitat in the region. The
impacts to resident herpetofauna of this
habitat degradation have not been quanti-
lied, but may be severe.

The Alpine She-oak Skink was not
recorded during this study. This cryptic
skink, which may be common at some
localities, is often difficult to detect, and
detection may require the right weather
conditions (P Robertson pers. comm.).
Schulz and Mansergh (1997) note the
uncertainty associated with detection of
this species; searches by these authors in
the same places in different years have
yielded considerably different results. This
species has been recorded at only three
broad localities within Victoria, two of
which are in alpine resorts (in the vicinity
of Mt Hotham and Falls Creek Alpine
Resorts), where its habitat is subject to
ongoing disturbance due to recreational
activities and slashing of native vegetation
during grooming of ski runs (Schulz et al.
1995; Schulz and Mansergh 1997;
Clemann in press; D Heinze pers. comm.).
Slashing may be sufficient to destroy the
structural integrity of Alpine She-oak

Vol. 119 (2) 2002

Research Reports

Fig. 2. Alteration of vegetation structural com-
plexity by cattle grazing and trampling in the
Mt Stirling area. The area in the top of the pho-
tograph is a cattle exclusion zone. The structure
of the vegetation in the foreground (outside of
the cattle exclusion zone) has been profoundly
modilied by cattle grazing.

Skink habitat (D Heinze pers. comm.).
Habitat of this species at Basalt Hill near
Falls Creek is also subject to cattle grazing
in summer, as is the third broad locality at
which it has been detected (Lankeys
Plain). Cattle graze on species of tussock
grass that are important components of
Alpine She-oak Skink habitat (van Rees
1984).

The Alpine Bog Skink occurs in grass-
land, Snow Gum woodland and heath-
lands, yet is most abundant in bogs and
boggy creeks (Hutchinson and Donnellan
1992; pers. obs.). Its occurrence in these
habitats is often marginal, overlapping
with the Tussock Skink and Southern
Grass Skink. Despite this sympatry, the
Alpine Bog Skink tends to occur in wetter
microhabitats than the Tussock Skink, and
in more open areas than the Southern
Grass Skink (Hutchinson and Donnellan
1992: pers. obs.). The occurrence of the
Alpine Bog Skink on Davies Plain, close
to the New South Wales border, suggests
that it may yet be detected in similar habi-
tat over the border.

The Southern Grass Skink and the
Tussock Skink were frequently recorded
during this study; each was recorded in six
habitat types, within six and seven of the
broad study areas, respectively. This
reflects their widespread and abundant
nature in the alpine region. The taxonomy
of lizards in this genus has only recently
been clarified by using eletrophoretic,
karyotopic and morphological data
(Hutchinson and Donnellan 1992),

55

Research Reports

A^ : AEN c s
Fig. 3. Damage in bog habitat on Davies Plain

caused by cattle trampling. Nearby undamaged
habitat is occupied by the Alpine Water Skink.

Unfortunately, considerable variation and
overlap in colouration and morphology
exists between alpine taxa within this
genus (particularly P. pagenstecheri and P.
cryodroma), confounding the identification
of some specimens in the field. Tentative
identification of some specimens collected
during this study required confirmation by
Dr Mark Hutchinson (South Australian
Museum), In the absence of further elec-
trophoretic testing, the identification of
some specimens (notably those from Mt
Buffalo) remains uncertain.

Davies Plain was the most species rich
(10 taxa) of the broad areas surveyed in
this study. Threatened species recorded in
this area included the Alpine Water Skink
and Alpine Bog Skink (historically, the
Alpine Tree Frog also occurred on this
plain). The unidentified Egernia species
recorded on this plain (and throughout the
alpine areas) are significant fauna, and are
likely to be candidates for future listing as
threatened taxa (P Robertson pers. comm.).
Thus, Davies Plain is of particular value
for threatened alpine herpetofauna. Despite
this, the area remains subject to cattle graz-
ing, and is one of the strongholds for feral
horses in the Victorian alps. Many horses
were observed on this plain during two
days of fieldwork. Whilst horses on the
Bogong High Plains represent an isolated
population, those on Davies Plain are
members of a population extending from
Thredbo in New South Wales to the
Buchan River in Victoria (M Walters pers.
comm.). Both cattle and horses are causing
considerable damage to bog habitat on
Davies Plain, suggesting that their man-

56

LÀ

FTT .- AX p p -
Fig. 4. Damage caused by cattle to the margin
of a wet area that was once a bog/stream com-
plex on the Bogong High Plains.

agement should be an urgent priority in
this area.

There is a growing body of evidence that
cattle grazing has a significant detrimental
effect on alpine habitats (references cited
throughout this discussion) and, conse-
quently, a deleterious effect on terrestrial
alpine fauna. In particular, grazing has a
devastating impact on wet vegetation asso-
ciations in alpine areas that are important
habitat for the Alpine Bog Skink, and vital
for the Alpine Water Skink, and Wahren et
al. (1999; 165) note that ‘continued graz-
ing by cattle is not compatible with the
conservation objectives for this vegetation
type’. This damage has long been recog-
nised (Costin 1958), and it is now clear
that grazing in alpine areas places at con-
siderable risk the conservation value of the
region. The only way of ensuring the con-
tinued existence of several terrestrial verte-
brates and the integrity of some plant
species and communities may be to control
densities of cattle and horses, or remove
them from certain areas. Williams et al.
(1997) summarised the damage caused by
cattle grazing on the Bogong High Plains
(including the facilitation of weed inva-
sion), and recommended the removal of
cattle from these plains. The fact that cattle
have been successively excluded from
parts of the high plains (e.g. northern
Bogong High Plains, Mts Bogong, Hotham
and Feathertop) for conservation purposes
is indicative of the damage stock have
caused in these areas, The revegetation and
recovery of areas after cattle or other stock
have been removed is an extremely slow
process owing to the harsh nature of the

The Victorian Naturalist

alpine climate and the low mineral content
of soils (Downes 1961; Bridle and
Kirkpatrick 1999). In fact, some alpine
wetlands may take decades or even hun-
dreds of years to recover from the impact
of livestock incursion (Wahren er al.
1996). The last grazing leases were with-
drawn from Kosciuszko National Park in
1969, but rehabilitation of habitat damaged
by cattle is still needed today (Scherrer and
Pickering 2001). Consequently, grazing
pressure should be eliminated from these
areas sooner rather than later. Implicit in
the need to remove grazing from some
areas is the need for habitat rehabilitation.
A major consideration of such rehabilita-
tion is that the costs of these activities are
usually borne by organisations different
from those responsible for the damage.
Thus the benefits of grazing go to private
individuals, whilst the cost of reparation is
borne by the public (Scherrer and
Pickering 2001).

The impact of introduced predators such
as foxes and cats on alpine herpetofauna
has not been quantified; however, observa-
tions of foxes in some alpine areas has
shown that they prey on a range of reptile
species, including the Alpine Water Skink
(R Martin pers. comm.). Such impacts are
likely to be particularly devastating for
species with very small geographic ranges
(such as the Alpine She-oak Skink), those
with a high degree of habitat specificity,
particularly those occupying narrow, often
linear habitats (such as the Alpine Water
Skink), and those whose populations are
diminished or stressed due to other threat-
ening processes. Similarly, domestic pets
may be a threat to the Alpine She-oak
Skink in the vicinity of the Mt Hotham and
Falls Creek villages.

Climate change (i.e. *global warming")
may pose a considerable threat to alpine
organisms. Whilst some taxa may be able
to migrate to higher altitudes to maintain
their preferred climate, others will not, and
species already at the top of their possible
altitudinal distribution are likely to be
replaced by other species moving up
(Hughes and Westoby 1994). As climate
conditions change, generalist species may
have a greater capacity to occupy, reinvade
or outcompete specialist congeners
(Bennett ef al. 1991).

Vol. 119 (2) 2002

Research Reports

Acknowledgements

John Silins. Simon Scott and Cassie Wright pro-
vided enthusiastic and helpful field assistance,
and made field work considerably more enjoy-
able. Throughout the project, staff from Parks
Victoria and NRE regional staff provided vital
local knowledge and facilitated access to all
areas. In particular | thank Jeff Carboon and
Ron Riley from Parks Victoria. Similarly, staff
from the Lake Mountain Alpine Reserve assist-
ed with local knowledge and the provision of
access to this area. Mark Hutchinson (SA
Museum) confirmed the identities of' several
specimens collected during this project, and pro-
vided useful dialogue about the distribution and
ecology of various species. John Coventry
(Museum Victoria) and Peter Robertson
(Wildlife Profiles Pty Ltd) also provided useful
feedback during the project. Michelle Walters
(Australian National University) provided valu-
able information on feral horses. Marcus Whitby
provided the Egernia specimen from Mt
Buffalo, Geoff Brown and David Choquenot
(ARI) supervised this project and assisted with
critiques of the manuscript, lan Mansergh,
Graeme Newell and Tim Clancy (all NRE) also
provided valuable comments on the manuscript.
| thank Professor Richard Shine (University of
Sydney) for reviewing the manuscript and sug-
gesting improvements,

References

ANZECC (1999) Threatened Australian Fauna.
Australian and New Zealand Enyironment and
Conservation Council.

Bennett S, Brereton R, Mansergh Il, Berwick S,
Sandilord K and Wellington C (1991) The potential
effect of the enhanced greenhouse climate change on
selected Victorian fauna. Arthur Rylah Institute
Technical Report Series No 123,

Berger L, Speare R and Hyatt A (1999) Chytrid fungi
and amphibian declines: overview, implications and
future directions In Declines and Disappearances of
Australian Frogs, pp 23-33. Ed A Campbell.
(Biodiversity Group, Environment Australia:
Canberra)

Bridle KL and Kirkpatrick JB (1999) Comparative
effects of stock and wild vertebrate herbivore grazing
on treeless subalpine vegetation, Eastern Central
Plateau, Tasmania. Australian Journal of Botany 47,
817-834,

Brown G W (2001) The influence of habitat distur-
bance on reptiles in a Box-Ironbark eucalypt forest of
south-eastern Australia, Biodiversity and
Conservation 10, 161-176,

Brown G and Bennett A (1995) Reptiles in rural envi-
ronments. The distribution, habitat requirements and
conservation status of the reptile fauna of the
Murray-Darling Basin Area in Victoria. Report to the
Murray-Darling Basin Commission, Department of
Conservation and Natural Resources, Heidelberg,
Victoria.

Clemann N (in press) Action Statement for the Alpine
She-oak Skink Cyelodomorphus praealtus.
Department of Natural Resources and Environment,
Melbourne.

Costin AB (1958) The grazing factor and the mainte-
nance of catchment values in the Australian Alps.
CSIRO Australia Division of Plant Industry
Technical Paper No 10

57

Book Review

Coventry AJ and. Robertson P (1980) New records of
seineid lizards trom Victoria. The Vietortan
Naturalist 97, 190-193,

Downes RG (1961) The Victorian High Plains — the
environment and its use, Proceedings of the Royal
Noviety of Victoria 75, 339-347,

Gillespie GR, Osborne WS and McElhinney NA
(1995) The conservation status of frogs in the
Australian alps, a review. A report to the Australian
Alps Liaison Committee, Department ol
Conservation and Natural Resources, Heidelberg,
Victoria and Applied Ecology Research Group,
University of Canberra, Belconnen, ACT.

Hadden SA and Westhrooke ME (1996) Habitat rela-
tionships of the herpetofauna of remnant Buloke
woodlands of the Wimmera Plains, Vietoria, Wildlife
Research 23, 363-372

Heinze D (1997) Notes on a new survey technique in
locating the Alpine She-oak Skink Cvelodomorpliis
praealtus in the Mt Hotham area, Victoria. Zhe
Victorian Naturalist Wd, 176-177

Hughes L and Westoby M (1994) Climate change and
conservation policies in Australia, coping with
change that is far away and not yet certain, Z'uci/ic
Conservation Biology V, 308318,

Hunter D, Osborne WS and Smith MJ (1997)
Distribution and abundance of the Alpine Tree Frog
(Litoria verreauxit alpina) in the Australian alps
national parks, report on the first seasons survey
(1996-97), Heritage Unit Snowy Mountains Region,
NSW National Parks and Wildlife Service

Hutehinson MN and Donnellan SC (1992) Taxonomy
and genetic variation in the Australian lizards of the
penus Pseudemoia (Scincidae; Lygosominae).
Journal of Natural History 26, 215-264

Kiesecker IM, Blaustein AR and Belden LK (2001)
Complex causes of amphibian population declines.
Nature 410, 681-084,

LCC (1982) Report on the Alpine Study Area. Land
Conservation Council, Victoria

Manserph | (1982) Notes on the range extension of the
Alpine Water Skink (Sphenomorphus koseiuskor) in
Victoria. The Victorian Naturalist 99, 123-124

MeDougall KL (1982) The alpine vegetation of the
Bopgong High Plains. Environmental Studies
Publication No 357, Ministry for Conservation,
Victoria

McDougall KI. (0989) The effect of excluding cattle
Itom a mossbed on the Bogong High Plains, Victoria
Arthur Rylah Institute for Environmental. Research,
Vechnical Report Series No 95

Meredith C, Hudson $, Robertson P and Clemann N (in
press) Action Statement for the Alpine Water Skink
Eulamprus kosciuskoi. Department of Natural
Resources and Environment, East Melbourne,

NRE (2000) Threatened vertebrate fauna in Victoria
2000, a systematic list of vertebrate fauna considered
extinct, at risk of extinction or in major decline in
Victoria. Department of Natural Resources and
Environment, Fast Melbourne.

NRE (in prep.) Alpine Hog Community. Flora and
Fauna Guarantee Action Statement, Parks, Flora and
Fauna Division, Department of Natural Resources
and Environment, East Melbourne,

Osborne W, Hunter D and Hollis G (1999) Population
declines and range contraction in Australian alpine
frogs. In Declines and Disappearances of Australian
Frogs, pp. 145-157. Ed A Campbell. (Biodiversity
Group, Environment Australia; Canberra)

Scherrer P and. Pickering CM (2001) Effeets of praz-
ing, tourism and climate change on the Alpine
Vepetation of Kosciuszko National Park, The
Victorian Naturalist 118, 93-99,

Schulz M, Alexander J and Mansergh 1 (1995) Notes
on the Alpine She-oak Skink Cre/odomorphus
praealtus in the MU Hotham Area, alpine Victoria
with a deseription of a potential new survey tech-
nique. The Victorian Naturalist 112, 219-220,

Schulz M and Mansergh 1 (1997) New location for the
Alpine She-ouk Skink Cyelodomorphus praealtus in
Victoria, The Victorian Naturalist 114, 178-179

van Rees H (1984) Behaviour and diet of free-ranging
cattle on the Bogong High Plains Victoria.
Department of Conservation, Forests and Lands,
Victoria. Report No 409

Wahren CH, Williams KJ and Papst WA (1996) The
ecology of wetlands and snow patehes on the Bogong
High Plains. Report to the Australian Heritage
Commission and the Victorian Department of
Natural Resources and Unvironment. National Estate
Grants Program project No 710.

Wahren CH, Williams RJ and Papst WA (1999) Alpine
and subalpine wetland vegetation on the Bogong
High Plains, south-eastern Victoria. duszralian
Journal of Botany AT, 105-188.

Williams RJ, Papst WA and Wahren CH (1997) The
impact of cattle grazing on alpine and subalpine plant
communities of the Bogong High Plains,
Unpublished report to the Department of Natural
Resources and Environment.

Wild Solutions: How Biodiversity is Money in the Bank

by Andrew Beattie and Paul Ehrlich

Publisher: Melbourne University Press, 2001. 239 pp. ISBN 0 522 84986 5. $49.95,

Loss of biodiversity — the wealth of nat-
ural variety on our planet - is one of the
greatest concerns of modern times. In a
world driven largely by pragmatic deci-
sions, evaluating and communicating the
richness and. variety of living organisms is
an important basis for conservation. Wild

58

Solutions is written by two eloquent con-
servation ecologists to help convey the
potentially tangible values of organisms
that are most commonly regarded as pas-
sengers in wider conservation endeayour,
namely the invertebrates, microorganisms
and ‘lower’ plants. Beattie and Ehrlich

The Victorian Naturalist

demonstrate the immense importance of
these organisms, and their incredible spec-
trum of largely unexplored benefits to (and
uses for) humanity, together with their
variety and vital roles in ecological sus-
tainability, Collectively, invertebrates have
‘solved’ many of the problems of survival
in complex or extreme enyironments that
still challenge human ingenuity, and study-
ing how they have done this may provide
valuable clues for human wellbeing. The
breadth of examples used is impressive,
and the simplified language renders the
book accessible to general readers whose
knowledge of invertebrates may be limit-
ed. Christine Turnbull’s clear line draw-
ings of many of the organisms discussed
are also helpful and appealing.

Indeed, from some points of view, the
efforts to write in yery general terms are
occasionally frustrating, in that highly
complex scientific stories are presented
very simplistically and the sense of wonder
we should feel at their intricacy tends to
‘get lost! The information presented here
is too important to be taken for granted.
Nevertheless, the reader will surely appre-
ciate the authors’ attempts to summarise
such a vast compass in this form. Their key
message is that we must be induced to
value and develop the concept of ‘natural
capital’ before its value is destroyed, and
that natural capital is the very organisms
that we usually overlook or deride. The
loss of biodiversity can be attributed large-
ly to three categories of human activity:
overpopulation, overdevelopment and
overexploitation. One important message
(p. 224) is that *we can never predict what
species or populations are going to be use-
ful or even desperately important in the
future', and consequently the arguments
sometimes advanced on redundancy of
many taxa (so that these are then function-
ally dispensable in terms of pragmatic
human values) are very poorly founded in
present knowledge. Curtailing losses of
taxa from human activities may well be the
key to our future survival.

The book consists of 13 main chapters,
and a list of further reading, which usefully

Vol. 119 (2) 2002

Book Review

augments the main content. Chapter I
commences with an apt quotation from
Antonie van Leewenhoek (1632-1723)
‘All we have yet discovered is but a trifle
in comparison with what lies hid in the
great treasury of nature’, a sentiment
acknowledged by many modern commen-
lators and sobering in relation to the
assumptions we tend to make in our deal-
ings with life on earth. The history of dis-
covery of invertebrates and microorgan-
isms continually shows the uncertainties
we face in trying to define their richness
even within an order of magnitude. More
specific themes are then developed, cover-
ing most major ecosystems, and stressing
the uniqueness and intricacies of the inter-
actions between taxa and the ways in
which study of living organisms provides
lessons for human technological enterprise
in virtually any field of endeavour
including pharmaceutical/medical benefits,
robotics, chemical engineering (bio-
mimicry), biological monitoring, recycling
materials, and the wide variety of other
ecosystem services. Use of bacteria to help
clear up diesel spills in soils or feeding on
old car tyres and noxious agricultural
chemicals; of bull ant secretions as antibi-
otics; of spider silks as models for light-
weight body armour; of velvetworm ‘spit’
as rapid-setting glue and correspondingly
rapid solvents for it; and of tardigrades as
producers of antifreeze are amongst the
numerous tantalising issues discussed. The
potential list of such benefits is both end-
less and fascinating.

Much of this book induces an increasing.
sense of wonder at the possibilities lor
people to ‘capitalise’ on such values of
natural biodiversity, and the urgency to do
so. The authors convey powerfully that we
lose biodiversity at our peril, and the book
is a valuable contribution to the conserva-
tion debate, Read it, and spread its mes-
sages as widely as you can!

TR New

Department of Zoology
La Trobe University, Victoria 3086

Research Reports
The Mammals of Parker River Inlet, Otway National Park

ME Westbrooke! and PT Prevett'

Abstract

Studies of small mammals at Parker River Inlet from 1985-95 have demonstrated the presence of ten
species of small mammals, Trapping, spotlighting and incidental sightings have shown that a further
18 species of mammals occur within an area of 200 ha. Pseudomys fumeus and Antechinus minimus,
recorded in this survey, are rare and restricted in Victoria, The species richness of the area is related
to the heterogeneity of the vegetation, with twelve vegetation types being identified in the study
area. The occurrence of the more abundant small mammals, Rattus fuscipes, Rattus lutreolus, Mus
musculus and Antechinus swainsonii, is correlated with vegetation structure and floristies. Causes of
fluctuations of the populations of the exotic species M. musculus and Rattus rattus are considered.
Management recommendations, which reflect the high significance of faunal habitat in the area, are

made with the aim of minimising human impact. (7he Victorian Naturalist 119 (2), 2002, 60-68)

Introduction

Botanically, the Otways, in particular the
Cape Otway/Parker River area, is consid-
ered significant as an isolated western
extremity of a vegetation type much more
extensive in eastern Victoria and Tasmania
(Carr 1971). Preliminary surveys in 1982
and 1983 at Parker River Inlet, located in
the southwest corner of the main block of
Otway National Park (Fig. 1), revealed the
presence of six species of small mammals,
and sufficient interest was generated to
warrant a more extensive, long-term study.

At the mouth of Parker River, sandstone
cliffs give way to dune limestone cliffs,
outcrops and stable dunes (Fig. 2). The
lower course of Parker River. is aligned
along a fault. The estuarine sector has
developed where the fault zone crosses the
coast, allowing the stream to cut a wider
valley in the fractured rock. The estuarine
sector is short and largely filled by a tidal
delta of in-washed sand (Rosengren 1984).
Parker River enters the estuarv over a
series of cascades and small falls. Elevated
areas receive abundant rains, falling main-
ly between autumn and early summer,
Rainfall at Cape Otway averages 890 mm
per annum, but rises to twice this figure on
the Parker Ridge. The combination of
these elements has resulted in both floristic
and structural diversity of vegetation with
a great variety of associations being repre-
sented within a small area around Parker
River Inlet. Data presented relates to an
area of 200 ha located either side of the
lower reaches of Parker River.

' Centre for Environmental Management, University
of Ballarat, PO Box 663, Ballarat, Victoria 3353

60

Methods

Small mammal trapping

A trapping grid was established at Parker
River Inlet. It comprised 100 trap locations
20 m apart encompassing an area of
approximately 5 ha of diverse topography
and vegetation. One trap was placed at
each site. Small folding mammal traps 33

10 x 10 cm (Elliott Scientific Equip-
ment, Upwey, Victoria) were baited with a
mixture of peanut butter, rolled oats and
honey. Sites were trapped for three consec-
utive nights on 14 occasions from April
1985 to October 1988, a total of 4200 trap
nights. Traps were cleared and reset each
morning. Animals caught were identified,
sexed, weighed to the nearest gram and
marked with numbered fingerling tags
(National Band and Tag Co. No. 1) on the
ear.

Recording of other mammal species

From April 1985 to March 1995 inciden-
tal observations were made of other mam-
mals occurring in the vicinity of the Inlet
within the area shown on Fig. 1. In March
1989 a Harp Trap was set at the bridge
over Parker River, 1 km from the inlet, for
three consecutive nights to trap insectivo-
rous bats.

Vegetation assessment

At each trap site on the small mammal
grid, the structure and floristies of the veg-
etation were assessed and related to the
occurrence of the more frequently trapped
species, The vegetation of the surrounding
area was also surveyed.

'The Victorian Naturalist

NSW

VIC

Melbourne

NATIONAL

eo

Crayfis Seal Point

Point Franklin

Research Reports

Blanket Bay
PARK

Point Lewis

A Trap Grid

Study Area

Pz

0 0.5 1
— m
kilometres

Fig. 1. Location of study site at Parker River, Otway National Park, Victoria.

Results
Vegetation
Eleven vegetation associations were identi-

fied in the area around the lower reaches of

Parker River (Fig. 3). Seven of these were
represented within the trapping grid and
are marked *. Taxonomy is according to
Ross (2000).

*Eucalyptus globulus tall open-forest

(Gully)
Tall open forests dominated by Blue

Gum Eucalyptus globulus, with a range of

associated shrub species including Hazel
Pomaderris Pomaderris aspera, Prickly
Currant-bush Coprosma quadrifida and
Privet Mock-olive Notelaea ligustrina. The

Vol. 119 (2) 2002

ground layer was dominated by ferns
including Common Maidenhair Adiantum
aethiopicum, Hard Water-fern Blechnum
procerum, Common Ground-fern Culcita
dubia and Spreading Fan-fern Sticheris
lobatus.

Eucalyptus obliqua open-forest

Open forest dominated by Messmate Æ.
obliqua, to 20 m.with an understorey of
shrubs to 2m. Shrub species include Heath
Tea-tree Leptospermum myrsinoides,
Lance Beard-heath Leucopogon lanceola-
tus, Prickly Moses Acacia verticillata and
Large-leaf ^ Bush-pea Pultenaea
daphnoides.

61

Research Reports

PT es Ad r A

Fig. 2. Parker River Inlet, Otway National Park. Vi

*E, obliqua/E. globulus open forest (Open
Forest)

Open forest dominated by £. obliqua and
E. globulus. An open understorey of Poa
spp. with Austral Bracken Preridium escu-
lentum, Spiny-headed Mat-rush Lomandra
longifolia subsp. longifolia, Snowy Daisy-
bush Olearia lirata, A. verticillata and
Sweet Bursaria Bursaria spinosa.

*'Otway messmate' low open-forest (L.
Open Forest)

Low open forest of the hybrid Otway
Messmate with an understorey dominated
by Tussock Grass Poa labillardieri, L.
longifolia subsp. longifolia, and P. escu-
lentum, Scattered shrubs include Coast
Beard-heath Leucopogon parviflorus and
Silver Banksia Banksia marginata.

Eucalyptus viminalis woodland

On undulating stabilised dunes between
the lighthouse road and Blanket Bay Road
occurs a woodland of Manna Gum Æ. vimi-
nalis with a sparse shrub layer of A. verti-
cillata and B. spinosa, with a ground layer
dominated by P. /abillardieri and L. longi-
folia.

Leptospermum continentale/Melaleuca
squarrosa tall closed-shrubland

In more or less sheltered coastal gullies
to the west of Parker River a dense closed

62

ctoria.

shrubland of Scented Paperbark Melaleuca
squarrosa and Prickly Tea-tree Lepto-
spermum continentale occurs. The ground
layer is sparse or absent.

Closed heathland

On flat poorly drained areas along the
Blanket Bay Road a closed heath of Scrub
Sheoak A/locasuarina paludosa, Dwarf
Sheoak A. pusilla, Prickly Tea-tree L. con-
tinentale and B. marginata occurs. Other
associated species include Austral Grass-
tree Xanthorrhoea australis, Common
Heath Epacris impressa and members of
the Restionaceae. There is an occasional
overstorey of Bog Gum Eucalyptus kitso-
niana.

*Low open heathland (Heath)

Areas on the dune limestone cliffs are
dominated by 7. parviflorus. This varies
from small stunted shrubs less than | m
tall in exposed sites to 3 m in more shel-
tered situations. Leucopogon parviflorus
was generally an overstorey to either P.
labillardieri or Buffalo Grass Stenotaph-
rum secundatum but, where associated
with L. continentale, Manuka Leptosper-
mum scoparium and B. marginata, an
almost impenetrable thicket existed,
excluding any ground layer. Scattered
stands of Drooping Sheoak Allocasuarina
verticillata also occur.

The Victorian Naturalist

, À Point Franklin
0 0.5 1

kilometres

hs f

Research Reports

L.continentale/

= Leucopogon parviflorus
Low Open Shrubland

Parker River Inlet

Eucalyptus globulus

Tall Open Forest
2 L.continentale /

Allocasuarina spp

E.obliqua
C
Tall Open Forest Closed Heath
Disturbed
E.obliqua Low Heath
t
RUP Coastal Dune
Grassland / Shrubland
E.obliqua /

E.viminalis Open Forest Grazed
is Grassland / Herbland
Otway Messmate

Low Open Forest MES: australis

Grassland
E viminalis
Woodland k E. kitsoniana
Restricted occurence
“Leptospermum continentale Closed Heath
— / Melaleuca squarrosa
Tall Closed Shrubland

m Study Site
Ses

Fig. 3. Vegetation communities of Parker River, Otway National Park, Victoria.

*Coastal dune complex (Dune)

Marram Grass Ammophila arenaria and
Hairy Spinifex Spinifex hirsutus provide
the major stabilizing vegetation on the sea-
ward dunes. Associated species include P.
esculentum and Knobby Club-rush /solepis
nodosa. Sea Rocket Cakile edentula is
apparently the first species to colonise the
sand of the open beach but this may be the
result of large-scale sand movements.

*Phragmites australis grassland (Phrag)

Common Reed Phragmites australis
dominates a stretch alongside Parker River
between the waterfall and the beach. Poa
labillardieri occurs with it except where
inundated with water.

Vol. 119 (2) 2002

*Tussock grassland (Grass)

Poa labillardieri, forms dense stands to |
m in height with S. secundatum. Poa labil-
lardieri was the dominant species in low-
lying areas, whereas S. secundatum occurs
as the dominant species on level areas
between the dune system and the first dune
limestone cliff. In this situation the Buffalo
Grass has been grazed by roaming cattle to
produce a low, even lawn, interspersed
with other grass species such as Hare's
Tail Lagurus ovatus, Slender Tussock-
grass Poa tenera, Cocksfoot Dactylis
glomerata, Yorkshire Fog Holcus lanatus
and Tall Fescue Festuca arundinacea.

63

Research Reports

Captures/ 100 trap nights
14

42 |

R. fuscipes R. lutreolus R. rattus M. musculus A. swainsonii
Species
E 1985 B 1986 1987 B 1988

Fig. 4. Mammal captures/100 trap nights, Parker River Inlet, Victoria.

Small mammals

Nine species of small mammal were
recorded: Swamp Rat Rattus lutreolus,
Black Rat Rattus rattus, Bush Rat Rattus
Juscipes, Smoky Mouse Pseudomys
Jumeus, House Mouse Mus musculus,
Agile Antechinus Antechinus agilis,
Swamp Antechinus Antechinus minimus,
Dusky Antechinus Antechinus swainsonii
and White-footed Dunnart Sminthopsis
leucopus. A single dead specimen of
Southern Brown Bandicoot /soodon obesu-
lus was also found. Relative abundance of
the more frequently occurring species over
the four years based on captures per 100
trap nights is shown in Fig. 4. The occur-
rence of the small mammal species in the
seven vegetation types represented on the
trapping grid is shown in Table 1.

Other mammals
During the period of the study incidental
observations of ten larger mammal species

64

were made together with five bat species
trapped in March 1989 (Table 2).

Discussion

Small mammals frequently captured at
Parker River Inlet were R. fuscipes, R.
lutreolus, M. musculus and A. swainsonii.
Pseudomys fumeus, although only trapped
twice is a significant record for the area
since it has been infrequently recorded in
the Otways. Antechinus minimus is listed
as rare in Victoria because it is found only
in small isolated populations. There are
only two previous records of Sminthopsis
leucopus from the Cape Otway area. The
appearance and later disappearance of a
population of R. rattus is of interest and
may be related to changes in use in the
area.

Rattus fuscipes is a widespread and com-
mon species, being found in sub-alpine
woodland, coastal scrub, eucalypt forest
and rainforest. It is nocturnal, preferring

The Victorian Naturalist

Research Reports

=i 4 IL tcc —— — —
Table 1. Relationship of small mammals to vegetation types. o, not recorded; +, occasional; #, fre-
quent. * denotes exotic species. Taxonomy according to Menkhorst (1995).

Grass Phrag Dune Heath L.Open Open Gully
Forest Forest
Rattus fuscipes E = + # # # E
Rattus lutreolus # # 4 # + + +
*Rattus rattus + + + + o o o
Pseudomys fumeus o o o o ir 4 o
*Mus musculus # # # # E ! o
Antechinus swainsonii — o o + + [ E
Antechinus agilis o o o + + ! +
Antechinus minimus + + o o o + [U
Sminthopsis leucopus o o o o 4 + o

ce ee (AE
Table 2. Species of larger mammals and bats recorded adjacent to Parker River Inlet. * denotes exot-

ic species.

eS

Species

Evidence

Location

Ornithorhynchus anatinus

Tachyglossus aculeatus

Wallabia bicolor

Phascolarctos cinereus

Trichosurus vulpecula

Pseudocheirus peregrinus

Petaurus australis
Petaurus breviceps
Acrobates pygmaeus
Isoodon obesulus
*Vulpes vulpes

Diurnal observation
Diurnal observation
Diurnal observation
Diurnal observation
Nocturnal observation
Nocturnal observation
Nocturnal observation
Nocturnal observation
Nocturnal observation
Carcass

Scats, prints

Parker River
Open Forest
Widespread
Woodland

Open Forest
Open Forest
Tall Open Forest
Tall Open Forest
Tall Open Forest
Open Forest
Dunes

Vespadelus darlingtoni Harp trap Tall Open Forest
Vespadelus regulus Harp trap Tall Open-forest
Chalinolobus morio Harp trap Tall Open Forest
Chalinolobus gouldii Harp trap Tall Open Forest
Nyctophilus geoffroyi Harp trap Tall Open Forest

Falsistrellus tasmaniensis
Hydromys chrysogaster
Arctocephalus pusillus doriferus

areas where there is dense ground cover.
Although non-fossorial, it excavates bur-
rows under stumps, logs, bushes and
clumps of grass (Warneke 1971). In spite
of having broad habitat preferences (Hall
and Lee 1982), several studies have shown
it to be associated with dense cover and
structually complex vegetation types
(Leonard 1970; Warneke 1971;
Braithwaite er al, 1978). Rattus fuscipes
has been shown to be adversely affected by
fire (Fox 1982) and, in the Otways, it was
absent for three years of post-fire succes-
sion (Rogers 1991). The results of this
study associate R. fuscipes with dune com-
plex vegetation with dense ground cover.
Thus it appears that friable soils, which
provide favourable conditions for burrow-
ing and support a rich microfauna of

Vol. 119 (2) 2002

Harp trap
Cage trap
Diurnal observation

Tall Open Forest
Parker River
Parker River Inlet

arthropods and other animals, are a major
habitat requirement.

Rattus lutreolus has been described as a
rodent of heath, grass or sedge. Catling
(1986) pointed out that, whilst R. fuscipes
is primarily a forest species, R. lutreolus is
considered to be a wet heath species that
inhabits areas of thick vegetative cover.
Catling (1986) identified dietary dilTer-
ences as the reason that R. lutreolus pre-
cedes R. fuscipes post-fire. Rattus lutreolus
is predominantly a herbivore, feeding on
sedges, mosses, fungi, some herbs, seeds
and insects (Seebeck 1995a), whilst A.

fuscipes is omnivorous, eating insects and

plant material (Seebeck 1995b). R. lutreo-
lus was trapped most frequently in the
heath and tussock grassland, sites that had
a dense ground layer and a generally

65

Research Reports

sparse overstorey. This species was fre-
quently encountered in the coastal dune
complex, often adjacent to the beach.

Mus musculus is an introduced species
existing in large populations where appro-
priate food is available, and is an oppor-
lunistic and omnivorous species (Watts
and Aslin 1981). Norton (1987) pointed
out that at several heathland sites, the
species showed clear habitat preferences
for similar floristic groups. In dry heath-
land in north-easten Tasmania captures of
House Mice were positively correlated
with floristic richness and negatively cor-
related with vegetation cover less than 50
cm high. Floristically diverse habitats
could provide a range of food resources
throughout the year, support year-round
breeding and result in a rapid population
increase. Rapid population growth is well
documented for this species (Newsome
1969). Mus musculus was most commonly
trapped within the heath and low open-for-
est, this preference perhaps reflecting suit-
ability of the soil for burrowing, or related
to plant diversity and the greater range of
food sources throughout the year. Data
support a significant decline in the popula-
tion of this species over the four years of
the study at Parker River. This may be a
result of decline post fire, as has been
reported by Newsome (1995). He suggest-
ed a decline 3-4 years post-fire together
with a return of populations of native
rodents. The last fire in this locality, imme-
diately to the north-west of the study area,
was at least ten years prior to this study.

Antechinus swainsonii is a small ground-
dwelling marsupial. According to Dickman
(1995), the preferred habitats of A. swain-
sonii have a dense understorey of fern or
shrub. Although Dickman (1995) believed
the species to be a soil-fossicking insecti-
vore, Hall (1980) regarded A. swainsonii
as a generalist and an opportunistic feeder.
During the winter, nests are excavated in
creek banks or just below the soil surface,
often under the cover of decaying logs or
grass (Dickman 1995). Antechinus swain-
sonii was trapped only in the gully and on
the medium slopes and ridge within the
open forest communities.

Whilst A, minimus is listed as rare in
Victoria because it is found only in small
isolated populations which are thus vulner-

66

able to threats such as wildfire and feral
predators (Menkhorst 1995), recent studies
have shown populations along the Otway
coast (Westbrooke et al. 1995) and Port
Campbell National Park (Twyford 1997).
Whilst there are only two previous records
of S. leucopus from the Cape Otway area,
this may be the result of low capture rates
for this species rather than rarity. Its con-
servation status nonetheless remains
unclear (Menkhorst 1995).

Pseudomys fumeus is uncommon or rare
throughout its Victorian range (Menkhorst
1995), and was described by Brazenor
(1934) from two specimens taken at
Olangolah near Beech Forest in the Otway
Ranges of south-western Victoria, Another
specimen was collected in the same area in
1937. One specimen was trapped at the
mouth of Parker River in 1980 and two
were captured at Blanket Bay to the east in
1985. Cockburn (1981) suggested that the
habitat preferences of P. fumeus reflected
seleetion for those areas providing a year-
round source of high quality food, rich in
nitrogen and, further, suggested that post-
fire succession producing an understorey
rich in peas and wattles is essential for its
continued survival. In this study the two
specimens were trapped in low open-forest
vegetation, a trapping rate of approximately
one per 500 trap nights for this communi-
ty. Low trapping rates are common for this
species and a number of reasons for this
have been suggested. Menkhorst and
Seebeck (1981) noted that population den-
sities of P. fumeus were generally low,
possibly due to its preference for drier hill-
sides with a heath understorey. Even at
such sites the species may never develop
high population densities. Pseudomys

fumeus is considered to require careful

monitoring, principally because of succes-
sional changes in vegetation communities
and therefore change in habitat suitability
to the species (Menkhorst 1995).

Rattus rattus was introduced to Australia
from Europe. It is generally considered to
be a vagrant in undisturbed Australian veg-
etation and is more usually associated with
disturbed situations in coastal Australia.
Settlement on Parker Hill and camping at
Parker River Inlet may have provided the
required disturbance for the small popula-
tion of R. rattus that occurred there. A

The Victorian Naturalist

large percentage of the captures were on
the periphery of the flat grassland used by
campers and grazed by livestock. These
habitats of dense Buffalo Grass, heath or
tussock grass may have provided suitable
nest sites. Rattus rattus has not been cap-

tured at Parker River Inlet after 1986.

Factors that may have influenced this are:

(1) reduced use of the inlet for camping
since the establishment of the National
Park and the closure of the inlet access
track to vehicles;

(ii) continued euthanasia of specimens
trapped on a regular basis from what may
have been a population not well estab-
lished.

Other species recorded in this study are
unremarkable in themselves but the very

high species richness of mammals, 25

native and three exotic, within an area of

200 ha, makes the area highly significant.
Four further native species, Cercartetus
nanus, Potorous tridactylus, Dasyurus
maculatus and Miniopterus schreibersii
have been recorded within 2 km of the
inlet.

Management implications

This study at Parker River Inlet has
demonstrated the association of different
mammal species with particular vegetation
types. Consequently, concern is expressed
for the management of the area.

Clearing of woodland for grazing has
occurred from Point Franklin to Cape
Otway. Pastures are unimproved and were
used for grazing cattle, which wandered
freely over large areas of pasture, wood-
land and dune. These cattle have altered
the vegetation of the cliffs by trampling,
initiating and aggravating erosion, and
facilitating the spread of exotic weeds such
as Stenotaphrum secundatum and Lagurus
ovatus. Despite being established as a
National Park in July 1981, cattle were not
excluded until the purchase of a block of
private land to the west of the inlet, in
1985. At the inlet of Parker River, live-
stock had reduced areas of the coastal
heath and tussock grasslands to lawns of
exotic grass species. These open spaces on
level ground and close to the sea attracted
many campers. Native plant species such
as Spinifex hirsutus are susceptible to dam-
age by trampling. Spinifex hirsutus,

Vol. 119 (2) 2002

Research Reports

spreading by rhizomes, quickly colonises
recent sand accretions to the dunes.
However, the rhizomes are broken easily
under the loose sand when trampled, and
many areas have become devoid of this
valuable stabilizing species (Anon 1981).
The stability of coastal dunes in southern
Australia has been maintained by hand-
planting of the vegetatively propagated
Ammophila arenaria, introduced to
Australia from Western Europe and this is
well established. Cattle selectively graze
the native S. hirsutus in preference to the
exotic A, arenaria (pers. abs.). A number
of other exotic species occur in the dis-
turbed communities of the inlet and at least
two have become established within the
period of this study, i.e. Cape Gooseberry
Physalis peruviana and Sea Spurge
Euphorbia paralias.

The preferred habitat of P. fumeus, ridge-
top sclerophyll forest with a diverse under-
storey of heath dominated by legumes
(Menkhorst and Seebeck 1981), is regener-
ated by fire and Cockburn (1995) suggests
that it is dependent upon post-fire succes-
sion for its continued survival. He also
considered the species to be in danger of
extinction. This study has confirmed the
presence of P. fumeus within the Otway
National Park and appropriate manage-
ment procedures should be considered
along with the needs of other small mam-
mal species.

A notable absentee from the area is the
Broad-toothed Rat Mastacomys fuscus.
According to Menkhorst (1995), the
species has been found in dense Poa grass-
land beside Parker River and on dunes
within 50 m of the sea, No specimens were
recorded at Parker River Inlet despite trap-
ping between 1985 and 1989 in locations
where it might be expected.

Mus musculus becomes abundant and
ubiquitous about 18 months after a bush-
fire, at a time when native species of
rodent have become rare. The outbreak can
last up to 3-4 years, with M. musculus dis-
appearing as populations of native rodents
begin increasing again (Newsome 1995).
A controlled burn was carried out in open
forest at Blanket Bay, 4 km east of this
study site in January 1990. This has pro-
vided the opportunity to assess changes in
the small mammal populations following

67

Research Reports

fire at a site similar to parts of the trapping
grid used for this study (Westbrooke and
Prevett 2000).

It is apparent that major disturbances
through the activity of man or roaming
livestock are detrimental to both the plant
and animal communities at Parker River
Inlet. Such degradation of habitats may
favour the exotic mammal species R. ratius
and M. musculus (Watts 1995; Newsome
1995),

Acknowledgments

We would like to acknowledge the assistance
and cooperation of staff of DNRE, Colac
Region, and in particular Alan Rampel, Senior
Ranger, Otway National Park.

References

Anon (1981) 4 study of the land in the catchment of the
Otway Range and adjacent plains. (Soil
Conservation Authority: Kew)

Barnett JL, How RS and Humphreys WE (1976) The
use of habitat components by small mammals in east-
ern. Australia. Australian Journal of Ecology 3, 277-
85.

Braithwaite RW, Cockburn A and Lee AK (1978)
Resource partitioning by small mammals in lowland
heath communities of south-eastern Australia.
Australian Journal of Ecology 3, 423-45.

Braithwaite RW and Lee AK (1979) The ecology of

Rattus lutreolus in a Victorian heathland population,
Austrulian Wildlife Research 7, 173-79.

Brazenor CW (1950) The Mammals of Victoria.
(Brown, Prior and Anderson: Melbourne)

Carr GW (1971) Vegetation of the Parker River, Cape
Otway Region. Geelong Naturalist 8, 66-73.

Catling PC (1986) Ratus Iutreolus, colonizer of heath-
land after fire in the absence of Pseudomys species?
Australian Wildlife Research 13, 127-39.

Cockburn A (1981) Population regulation and disper-
sion of the Smoky Mouse, Pseudomys fumeus, |
Dietary determinants of microhabitat preference.
Australian Journal of Ecology 6, 231-54.

Cockburn A (1995) Smoky Mouse Pseudomys fumeus.
In The
Australian Mammals.
Robertson; Sydney)

Dickman CR (1995) Dusky Antechinus Antechinus
swainsonii, In The Australian Museum Complete
Book of Australian Mammals. Ed R Strahan. (Angus
and Robertson: Sydney)

Emerson er al. (1975) Ecological distribution of the
vertebrate animals of the volcanic plains — Otway
Range area of Victoria. Victoria Fisheries and
Wildlife Division, Wildlife Paper No 6.

Fox BJ (1982) Fire and mammalian secondary succes-
sion in an Australian coastal heath, Ecology 63,
1332-41.

Ed R Strahan. (Angus and

Hall 5 (1980) The diets of two coexisting species of

dnivchinus (Marsupialia: Dasyuridae). Australian

Wildlife Research 7, 365-78.

Mall S and Lee AJ (1982) Habitat use by two species of

AAnrechinus and Rattus Juscipes in tall open forest in
Southern Victoria In Carnivorous Marsupials, pp
209-20. Ed M Archer. (Royal Zoological Society of
New South Wales: Sydney)

68

Australian Museum Complete Book of

Hockings M (1981) Habitat distribution and species
diversity of small mammals in south-east Queensland
in relation lo vegetation structure, Australian Wildlife
Research 8, 99-108,

Leonard B (1970) Effects of control burning on small
mammal poru alons; 2" Fire Ecology Symposium
Papers, pp 1-14. Forests Commission of Victoria:
Melbourne.

Menkhorst PW (1995) Mammals of Victoria: distribu-
tion, ecology and conservation. (Oxford University
Press: Melbourne)

Menkhorst PW and Seebeck JI (1981) The distribu-
tion, habitat and status of Pseudomys fumeus,
Brazenor (Rodentia: Muridae). Australian. Wildlife
Research 8, 87-96

Newsome AE (1995) House Mouse Mus musculus. In
The Australian Museum Complete Book of Australian
Mammals. Ed R Strahan. (Angus and Robertson:
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Newsome AE (1969) A population of house mice tem
porarily inhabiting a South Australian wheathield.
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Norton TW (1987) The ecology of small mammals in
northeastern Tasmania II. Pseudomys novaehollandi-
ue and the introduced Mus musculus. Australian
Wildlife Research 14, 534-41.

Robinson AC (1975) Some aspects of the population
ecology of the bush rat, Rattus /uscipes,
(Waterhouse). In Barnett ef ul. (1978) The use of
habitat components by small mammals in eastern
Australia. dustralian Journal of Ecology 3, 277-285.

Rogers LM (1991) The effects of post-fire succession
on small mammals of the Otway National Park.
Undergraduate project report, Ballarat University
College, Mt Helen.

Rosengren N (1984) Sites of geological and geomor-
phological significance in the Shire of Otway.
Department of Geography: University of Melbourne.

Ross JH (ed) (2000) A Census of Vascular Plants of
Victoria, 5 ed, (Royal Botanic Gardens: Melbourne)

Seebeck JH (19954) Swamp Rat, In Mammals of
Victoria: distribution, ecology and conservation. Ed
PW Menkhorst. (Oxford University Press:
Melbourne)

Seebeck JIL (1995b) Bush Rat, In Mammals of
Victoria. distribution, ecology and conservation, Ed
PW Menkhorst, (Oxford University Press:
Melbourne)

lwyford KL (1997) Habitat relationships of small
mammals at Port Campbell National Park, Victoria,
Australian Mammology 20, 89-98.

Warneke RM (1971) Field study of the Australian Bush
Rat, Ruitus fuscipes, (Rodentia: Muridae). Wildlife
Contributions Victoria 14, 1-115.

Watts CHS (1995) Black Rat. In The Australian
Museum Complete Book of Australian Mammals. Ed
R Strahan, (Angus and Robertson; Sydney)

Watts CHS and Aslin HJ (1981) The Rodents of
Australia. (Angus and Robertson: Sydney)

Westbrooke ME and Prevett PT (2000) Post fire suc-
cession of small mammals in coastal woodland at
Blanket Bay, Otway National Park, Victoria,
Australia. 13th Conference on Fire and Meteorology,
Lorne, October 27-31, 1996.

Westhrooke ME, Burns D, Gibson M, Kerr M and
Prevett P (1995) The Fauna and Flora of Coastal
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The Victorian Naturalist

Contributions

Habitat of the Endangered Hibbertia procumbens (Labill.) DC
(Dilleniaceae) from the Central Coast of New South Wales

Stephen AJ Bell'

Abstract

Hibbertia procumbens (Labill.) DC (Dilleniaceae) is currently listed as an endangered species under
the NSW Threatened Species Conservation Act 1995. This species is extremely rare in New South
Wales, with only two records now known for the whole of the State; it is more common in Victoria,
where populations are concentrated in the coastal districts, and in Tasmania, Future taxonomic revi-
sion may possibly differentiate the New South Wales specimens from those in the more southern
States. A new population of Hibhertia procumbens is described here for Bumble Hill on the Central
Coast of New South Wales. Habitat details are provided for this and the only other known New
South Wales location at Strickland State Forest. A conservation risk code for the New South Wales
populations is suggested, following the system of Briggs and Leigh (1996), Additional searches for
Hibbertia procumbens in New South Wales should be made within appropriate habitat both in and
outside conservation reserves in the northern Sydney Basin region, to ascertain the conservation sta-

tus of the species and to assist taxonomic revision. (The Victorian Naturalisi 119 (2), 2002, 69-74)

Introduction

Hibbertia procumbens (Labill.) DC
(Dilleniaceae) is a very rare, summer-flow-
ering, prostrate shrub growing in scrubby
heath on sandy soils in New South Wales,
Victoria (Harden and Everett 1990), and
Tasmania (H Toelken, pers. comm. ).
Currently, H. procumbens is listed as

endangered in Schedule | of the NSW

Threatened Species Conservation Act
1995, although nationally it is not consid-
ered rare (Briggs and Leigh 1996). In New
South Wales, Harden and Everett (1990)
note that this species has been recorded

only from the Mangrove Mountain area of

the Central Coast botanical subdivision. In
Victoria, H. procumbens is locally com-
mon in coastal heathland and heathy
woodland, with a discontinuous distribu-
tion from the Portland area, Otway
Ranges, Anglesea, Western Port (including
French and Phillip Islands), Cape Paterson,
Wilsons Promontory, near Traralgon and
at a few sites east of Orbost (N Walsh,
pers. comm.). The species is also reported
to be locally common in Tasmania (H
Toelken, pers. comm.).

There is some taxonomic uncertainty
regarding the correct placement of the
New South Wales specimens within this
group. The disjunct occurrence in New
South Wales of what is currently described
as H. procumbens from populations in
Victoria and Tasmania may suggest that a

Fasteoast Flora Survey, PO Box 216, Kotara Fair
NSW 2289

Vol. 119 (2) 2002

different taxon is involved. In any case, the
New South Wales specimens are recog-
nised as problematic, and will require taxo-
nomic resolution in the future (H Toelken,
pers. comm.). Under current legislation in
New South Wales, H. procumbens is con-
sidered endangered, in recognition of the
paucity of records and its apparently high-
ly restricted range.

This paper reports on a new population
of H. procumbens located during vegeta-
tion surveys undertaken in Wyong Shire
on the Central Coast of New South Wales
(Bell in prog.). The site near Bumble Hill
on the edge of the Somersby Plateau was
recognised as being highly significant for
the State, and the opportunity is taken here
to document the habitat attributes in this
area. The only other known population in
New South Wales was also re-visited in
order to compare and describe the habitat
there.

Previously known records in New South
Wales

Previously, /7. procumbens was known
from only one collection in New South
Wales, made in Strickland State Forest at
Mangrove Mountain near Gosford in
October 1991 (P Hind, pers. comm.). This
is despite the relatively extensive vegeta-
tion survey work that has been carried out
in the Sydney region over recent decades.
Although present within this State Forest,
there is currently no known representation

69

Contributions

of this species within formal conservation
reserves, The existing site in Strickland
State Forest is approximately 15 km south-
east of the new location at Bumble Hill,

Hibbertia procumbens was also collected
in December 1998 during searches for the
endangered herb Prostanthera junonis on
behalf of the NSW National Parks and
Wildlife Service (S Douglas, pers. comm.).
Identification was not possible at this time
due to a lack of fertile material, and a cut-
ling was propagated to provide flowering
material for later identification. It was only
while assessing the status of threatened
flora in the Gosford City Council area that
the identity of the unknown //. was con-
firmed (Douglas and Burcher in prep.).
This was verified in August 2001 when
fertile specimens were observed in the
field.

Habitat at Bumble Hill

The Bumble Hill record was made in
November 2000, within a Banksia ericifo-
lia var. ericifoliu-Angophora hispida
serub/heath on skeletal sandy soils derived
from Triassic Hawkesbury sandstone (see
Appendix | for common names and
authorities of all plant species discussed in
the text). Soils have been mapped at this
location as part of the Lambert (la) soil
landscape, which consists of a loose, stony,
brown, sandy-loam topsoil with apedal,
single-grained structure and a porous
sandy fabric, over a yellowish-brown
clayey-sand subsoil with apedal massive-
to-weakly pedal structure, and a porous
earthy fabrie (Murphy 1993). The site is
located on a mid-to-upper exposed slope
(aspect 355°), on a gently sloping sandstone
bench (8°), and at an elevation of 90 m
ASL. Structurally, the vegetation is com-
posed of a tall shrub layer 2-3 m in height
and 20-40% cover, dominated by
Angophora hispida, Banksia ericifolia var.
ericifolia and Leptospermum trinervium.
Below this at a cover of 50-60% is a 1 m
laver of shrubs dominated by Banksia eri-
cifolia var. ericifolia, Hakea teretifolia and
Banksia oblongifolia. Ground layer vegeta-
tion merges with the shrub laver, and is
dominated by Lepyrodia scariosa.
Ptilothrix deusta and Actinotus minor. The
site appeared to have been burnt approxi-

»

mately 3-5 years prior to survey. Initially,

70

a population of around a dozen plants was
estimated for this site, but subsequent
searches for the species in adjacent areas
during October 2001 revealed a much larg-
er population of between 100-150 plants.
Other associated species within the
scrub/heath included (in decreasing order
of relative abundance) the shrubs
Pultenaea ferruginea var. ferruginea,
Petrophile pulchella, Baeckea diosmifolia,
Acacia linifolia, Phyllota phylicoides,
Bossiaea heterophylla, Lambertia formosa,
Hibbertia aspera, Platysace linearifolia,
Hakea dactyloides, Acacia sp. B, Grevillea
buxifolia subsp. buxifolia, Acacia suave-
olens, Epacris pulchella, Leucopogon
esquamatus, Dillwynia sericea and
Bossiaea scolopendria; the herbs
Mitrasacme polymorpha, Patersonia
sericea, Stackhousia nuda. Dampiera
stricta, Phyllanthus hirtellus, H. riparia
and Scaevola ramosissima; and the
graminoids Anisopogon avenaceus,
Cyathochaeta diandra, Xanthorrhoea
resinifera, Lomandra obliqua, Schoenus
lepidosperma subsp. pachylepis. Xyris
complanata, Aristida vagans and
Lomandra glauca. The small fern
Lindsaea linearis and thè terrestrial orchid
Cryptostylis erecta were also present.

Habitat at Strickland State Forest

Efforts were made to revisit the location
of the Strickland State Forest record in
July and August 2001. Notes accompany-
ing the original collection in 1991 included
a description of the habitat as ‘Banksia eri-
elfolia-Allocasuarina distyla open scrub,
on skeletal sandy soil with rocks’. The site
reference co-ordinates provided did not
correspond with the deseription of the area
(inaccurate by approximately 5 km), which
indicated that the site was ‘along track to
Banksia Picnic Area, off Dog Trap Road.
Strickland State Forest’, Consequently, a
search of the general area around the
Banksia Picnic Area managed to locate
suitable habitat and plants of H. procum-
bens. While detailed counts were not made
at this site, it was estimated that a popula-
tion of perhaps 100-200, possibly up to
300 plants, occurs here. The Strickland
State Forest population is probably the
larger of the two known New South Wales
sites.

The Victorian Naturalist

Like Bumble Hill, the Strickland State
Forest population occurs within a serub/
heath of Banksia ericifolia var. ericifolia
and some Angophora hispida, on skeletal
sandy soils derived from the Triassic
Hawkesbury sandstone series. Murphy
(1993) has mapped the soils at this site as
forming part of the Sydney Town (st) soil
landscape, although the description of both
the soils and the vegetation provided by
Murphy (1993) suggest that the area in
question is in fact a small unmapped
occurrence of the Lambert (la) landscape
(see above description). As at Bumble Hill,
the site occurs on a gentle upper slope (7")
but at an aspect of 164° and an elevation of
180 m ASL. Structurally, the vegetation
consists of a tall shrub layer (4-5 m, 10 %
cover) of stunted Eucalyptus haemastoma
and Banksia ericifolia var. ericifolia, with
some Leptospermum trinervium and
Angophora hispida also apparent. Below
this, a combined shrub/ground layer up to
2 m in height and 50-80 % cover occurs,
dominated by Grevillea speciosa,
Dillwynia sericea and Banksia ericifolia
var. ericifolia.

Other species present in the area included
(in decreasing order of relative abundance)
the shrubs Banksia oblongifolia, Persoonia
isophylla, Leucopogon esquamatus,
Bossiaea scolopendria, Bauera microphyl-
la, Hibbertia cistiflora subsp. cistiflora,
Hakea teretifolia, Epacris pulchella,
Pultenaea elliptica, Baeckea diosmifolia,
Hakea dactyloides, Acacia suaveolens,
Platysace linearifolia, Petrophile pulchel-
la, Persoonia lanceolata, Leptospermum
polvgalifolium subsp. polygalifolium,
Grevillea buxifolia subsp. buxifolia,
Philotheca buxifolia subsp. obovata,
Hibbertia empetrifolia subsp. uncinata; the
herbs Actinotus minor, Xanthosia tridenta-
ta, Drosera auriculata, Gompholobium
glabratum; the graminoids Leptocarpus
tenax, Lepyrodia scariosa, Ptilothrix
deusta, Xyris complanata, Patersonia
sericea, Lepidosperma filiforme,
Xanthorrhoea resinifera, Cyathochaeta
diandra, Schoenus lepidosperma subsp.
pachylepis; and the ferns/allies Selaginella
uliginosa, Schizaea bifida and Lindsaea
linearis. Allacasuarina distyla was not
observed, although Allocasuarina littoralis
was in the general area.

Vol. 119 (2) 2002

Contributions

Conservation and Management

Populations of H. procumbens at both
Bumble Hill and Strickland State Forest
represent the only known records for New
South Wales. Both populations occur on
the edge of the Somersby Plateau, an
undulating plateau of Triassic Hawkesbury
sandstone forming a northerly extension of
the Hornsby Plateau (Murphy 1993).
Neither site occurs within formal conserva-
tion reserves, although use of that part of
Strickland State Forest for timber produc-
tion is unlikely, Within New South Wales,
both the northern and southern limits of
distribution are represented by these two
Sites, covering a geographical range of
only 15 km, This area is also known for
other taxa of interest in the M. genus (e.g.
Hibbertia empetrifolia subsp. uncinata),
including some as yet undescribed species,
which require further investigation (H
Toelken, pers. comm.).

In the context of regional vegetation
communities in which this species is
found, the descriptions of habitat for the
two locations fit closely with the
Hawkesbury Coastal Banksia Woodland of
NPWS (2000), This vegetation type is spo-
radically distributed around the NSW
Central Coast on Hawkesbury sandstone
derived soils (particularly the Lambert soil
landscape), and hence there is considerable
potential for other populations of H,
procumbens to occur in the region, NPWS
(2000) have mapped 5732 ha (extant) of
this vegetation type in the Central Coast
area. Much of this already occurs within
conservation reserves managed by the
NSW National Parks and Wildlife Service,
including Brisbane Water, Dharug, Popran
and Yengo National Parks, and Parr State
Recreation Area. Consequently, potential
habitat for the species does not appear to
be immediately threatened. Several State
Forests managed by State Forests of New
South Wales also contain this habitat (e.g.
Strickland State Forest and Ourimbah State
Forest).

Although listed on Schedule 1 of the
NSW Threatened Species Conservation
Act, H. procumbens does not have an offi-
cial conservation risk code under the
Briggs and Leigh (1996) system, most
likely due to the extent of populations in
Victoria and Tasmania. Given the distance

71

Contributions

between the New South Wales and more
southern populations, further taxonomic
revision of this group will probably ulti-
mately recognise two distinct taxa.
Consequently, it is suggested that (at least
for New South Wales) this species be
given a code of 3E, indicating an endan-
gered species with a known geographical
range of greater than 100 km, but with no
known representation within formal con-
servation reserves. Should taxonomie revi-
sion determine that the New South Wales
specimens are in fact distinct from those in
the southern States, then a risk code of 2E
would be more appropriate. For New
South Wales, a regional code of 57
(Central Coast botanical subdivision)
should also be included.

The floristic composition of /7. proeum-
bens habitat on the Central Coast of New
South Wales is heavily influenced by fire,
particularly in relation to fire frequency
and intensity (Bradstock ef a/, 1997).
Management strategies for habitat known
to support H. procumbens are therefore
likely to revolve around appropriate fire
regimes, although specific research is
required to confirm this. The fire response
mechanism for other Hibbertia species in
the Sydney region predominantly involves
resprouting from basal shoots, although
high intensity fire has been reported to kill
some species (Benson and McDougall
1995). Observations made in the field sug-
gest that M. procumbens would resprout fol-
lowing fire, since specimens growing on
trail margins in Strickland State Forest were
noted to have resprouted from basal shoots
following trailbike disturbance.

Weed invasion is a concern for the
Strickland State Forest population.
Surrounding habitat is being invaded by the
exotic Pinus radiata while trail margins are
also affected by invasion of exotic Whisky
Grass Andropogon virginieus. Appropriate
management of the powerline access trail
along which several specimens occur is
paramount. Erosion, sedimentation and the
introduction of non-indigenous fill such as
basaltic road-base are currently undermin-
ing the integrity of the general habitat, and
promoting weed invasion. At Bumble Hill,
similar threats are not yet apparent,
although the site is on private land in close
proximity to a main road,

72

Future survey of heath and scrub vegeta-
tion on Hawkesbury sandstone geology
within this part of the Sydney region
should include diligent searches for H.
procumbens, particularly on and around
the Somersby Plateau. It would also be
desirable for dedicated searches to be
made within appropriate habitat in existing
conservation reserves in the region to
ensure that long term conservation of the
species is secured. Further collection of
specimens and lodgement at national
herbaria is also required to assist in taxo-
nomic revisions of this taxon.

Acknowledgments

Wyong Shire Council is acknowledged for fund-
ing the Shire-wide vegetation survey in 2000-
2001, during which the Bumble Hill population
was discovered. Assistance in the field was ably
provided by Jedda Lemmon, while confirmation
of Hibbertia procumbens specimens from
Bumble Hill and Strickland State Forest was
undertaken by stalf at the National Herbarium of
NSW. Neville Walsh (National Herbarium of
Victoria) kindly provided location details and
general information for Victorian populations,
while Hellmut Toelken (State Herbarium of
South Australia) provided comments on taxono-
my and distribution of this difficult taxon. Peter
Hind (National Herbarium of New South Wales)
provided information on NSW collections.
Travis Peake commented on the manuscript and
encouraged further searching for the Strickland
State Forest population. Steve Douglas also
commented on the manuscript and provided use-
ful information on the Strickland State Forest
population.

References

Bell SAJ (in prog.) Classification and mapping of the
vegetation of Wyong Shire. Eastcoast Flora Survey,
Unpublished report and map to Wyong Shire
Council,

Benson D and MeDougall L (1995) Ecology of Sydney
plant species. Part 3. Dicotyledon families
Cabombaceae to lupomatiaeeae. Cunninghamia 4.
217-431,

Bradstock RA, Tozer MG and Keith DA (1997) Effects
of high frequency fire on floristic composition and
abundance in a fire-prone heathland near Sydney
Australian Journal of Botany 45, 641-655.

Briggs JD and Leigh JH (1996) Rare or Threatened
Australian Plants, rev ed. (CSIRO: Ausiralia)

Douglas SM and Burcher P (in prep.) Gosford Cin
Council threatened biota profiles and assessment
guidelines. ESP Ecological Surveys and Planning
Creswick with AES Environmental Surveys Cowan.

Harden GJ (Ed) (1990-1993) Flora of New South
Wales Volumes 1-4. (UNSW Press: Kensington)

Harden GJ and Everett J (1990) Dilleniaceae. In Flora
of New South Wales Volume 1, pp 293-303. (UNSW
Press: Kensington)

Harden GJ and Murray LJ (2000) Supplement to Flora
of New South Wales Volume 1. (UNSW Press:
Kensington)

The Victorian Naturalist

Kodela PG and Tindale MD (2001) Acacia sp. B. Flora
of Australia, 11A, 223.

Murphy CL (1993) Soil Landscapes of the Gosford-
Lake Macquarie 1:100 000 Sheet Report.
Department of Conservation and Land Management.

National Parks and Wildlife Service (2000) Vegetation
survey, classification and mapping: Lower Hunter

Contributions

and Central Coast region. A project undertaken for
the Lower Hunter and Central Coast Regional
Environmental Management Strategy by CRA Unit,
Sydney Zone, NPWS.

Appendix 1. Scientific and common names of species mentioned in text. Nomenclature generally
follows Harden (1990-1993) and Harden and Murray (2000), except where recent revisions have

been published.

Common Name

Forked Comb Fern

Family Species and Authority

Class Lycopsida (Clubmosses and quillworts)

Selaginellaceae Selaginella uliginosa (Labill.) Spring -
Class Filicopsida (Ferns)

Schizaeaceae Schizaea bifida Willd.

Lindsaeaceae Lindsaea linearis Sw.

Class Coniferopsida (Conifers)

Pinaceae Pinus radiata D.Don *
Class Magnoliopsida (Flowering Plants)
Subclass Magnoliidae (Dicotyledons)

Screw Fern

Radiata Pine

Dilleniaceae Hibbertia procumbens (Labill.) DC -
Hibbertia cistiflora Wakef. Subsp. cistiflora — -

Hibbertia aspera DC

Hibbertia empetrifolia (DC) Hoogland

subsp. uncinata Toelken

Hibbertia riparia (R.Br. ex DC) Hoogland -

Euphorbiaceae Phyllanthus hirtellus F. Muell ex Muell. Arg. Thyme Spurge

Droseraceae Drosera auriculata Backh. ex Planchon -

Casuarinaceae Allocasuarina distyla (V ent.) L.Johnson =
Allocasuarina littoralis (Salisb.) L.Johnson Black She-oak

Baueraceae Bauera microphylla Sieber ex DC. -

Proteaceae Persoonia lanceolata Andrews -

Persoonia isophylla L.Johnson and P. Weston. -
Petrophile pulchella (Schrader) R.Br. -
Grevillea buxifolia (Smith ) R.Br.

subsp. buxifolia

Grey Spider Flower

Grevillea speciosa (Knight) Me Gillivray Red Spider Flower
Hakea teretifolia (Salisb.) Britten -
Hakea dactyloides (Gaertner) Cav. -

Lambertia formosa Smith
Banksia oblongifolia Cav.

Mountain Devil

Banksia ericifolia L. f. var. ericifolia

Myrtaceae Angophora hispida (Smith) Blaxell Dwarf Apple
Eucalyptus haemastoma Smith Broad-leaved Scribbly Gum

Leptospermum trinervium
(Smith) J. Thompson

Leptospermum polygalifolium Salisb.

subsp. polygalifolium
Baeckea diosmifolia Rudge

Rutaceae Philotheca buxifolia subsp. obovata
(G.Don.) Paul G. Wilson -
Fabaceae: Mimosoideae Acacia linifolia ( Vent.) Willd. Flax-leaved Wattle
Acacia suaveolens (Smith) Willd. Sweet Wattle
Acacia sp. B (Kulnura form of A. terminalis) -
Fabaceae: Faboideae — Gompholobium glabratum Sieberi ex DC. Dainty Wedge Pea

Phyllota phylicoides (Sieber ex DC.) Benth. Heath Phyllota

Pultenaea elliptica Smith

Pultenaea ferruginea Rudge var. ferruginea — -

Dillwynia sericea Cunn.

Bossiaea scolopendria (Andrews) Smith -

Bossiaea heterophylla Vent

Vol. 119 (2) 2002

Naturalist Notes

Appendix 1, (cont.)

Family Species and Authority

Common Name

Epacridaceae Epacris pulchella Cay.

Leucopogon esquamatus R.Br. -

Goodeniaceae

Dampiera stricta (Smith) R.Br. E

Scaevola ramosissima (Smith) K.Krause -

Lamiaceae
Apiaceae
AXanthosia tridentata DC

Prostanthera junonis B.J.Conn
Actinotus minor (Smith) DC

Somersby Mintbush
Lesser Flannel Flower

Platysace linearifolia (Cav.) Norman -

Loganiaceae
Stackhousiaceae

Subclass Liliidae (Monocotyledons)
Lomandraceae

Xanthorrhocaceae

Mitrasacme polymorpha R.Br. -
Stackhousia nuda Lindley

Lomandra obliqua ( Thunb.) J.F.Macbr. E
Lomandra glauca (R,.Br.) Ewart
Xanthorrhoea resinifera (Sol. ex Kite)

Pale Mat-rush

E.C.Nelson and D.J.Bedford -

Patersonia sericea R.Br.
Cryptostylis erecta R.Br.
Xyris complanata R.Br.

Iridaceae
Orchidaceae
Xyridaceae
Cyperaceae

‘Tartan Tongue Orchid

Schoenus lepidosperma (F.Muell.) K.L.Wilson

subsp. pachylepis (S.T. Blake) K.L.Wilson -
Lepidosperma filiforme Labill, -
Ptilothrix deusta (R.Br.) K.L.Wilson -
Cyathochaeta diandra (R.Br.) Nees -

Restionaceae Lepyrodia scariosa R.Br,
Leptocarpus tenax (Labill.) R.Br.
Poaceae Andropogon virginicus L.

Aristida vagans Cav,

Anisopogon avenaceus R.Br.

p Whisky Grass

Three-awn Speargrass
Oat Speargrass

* non-native species

‘a distinct form of Acacia terminalis (Salisb.) J.F. Macbr,

l la as Acacia sp. B (Kodela and Tindale 2001).

recognised in Flora of Australia Volume

Wingan Wilderness

Tucked away about half way along the
coastal strip that forms the Croajingalong
National Park in East Gippsland, and
accessible by way of the West Wingan
Road, is a very secluded section of the
Park at Wingan Inlet (Fig. 1). In recent
years the road in, about 18 km east from
Cann River and extending 34 km south
from the Princes Highway, has been
upgraded so that in normal conditions any
standard vehicle can have access. The road
is not suitable for caravans, but is quite
adequate for trailers. If in doubt, inquiries
should be made at the Parks Victoria
Office in Cann River. During holiday peri-
ods, it is necessary to book camping sites.

This particular area, with well appointed
camping sites, is a favorite destination for
fishermen, but hikers, bird watchers and
botanists will all find much of interest.
Lyrebirds and reptiles of various sorts

74

were among some of the more obvious
examples of the wildlife. I would like to
share three observations.

Near the entrance to the camping area is
à stand of the Tall Everlasting Heli-
chrysum elatum (front cover), which in
Victoria is confined to East Gippsland. It is
usually found in damp areas, and often

Fig. 1. Wingan Inlet.

The Victorian Naturalist

Fig. 2. Lilac Lily Schelhammera undulata.

along watercourses, in dry sclerophyll
forests. The dominant tree in this particular
location is Red Bloodwood Corymbia
gummifera (formerly Eucalyptus gum-
mifera) at its southern limit.

Tall Everlasting is a spectacular plant
when flowering, usually during the Spring.
Up to two metres high, the rather woolly
stems support large heads of flowers sur-
rounded by shining white bracts.

Usually tucked away in moist shady
spots beneath the understorey, but very
visible because of its distinctive coloration,
is the Lilac Lily Schelhammera undulata
(Fig. 2). This is the sole representative of
the genus in Victoria and is confined in
this State to near Orbost and to far East
Gippsland. All of the many specimens seen
around Wingan Inlet were procumbent, the
flower parts making a strong contrast with
the dark green foliage.

Of about 140 species of Srylidium found
in Australia, ten occur in Victoria. Many of
these are small herbs, most with flowering
stalks less than 30 cm long. Generally the
more spectacular members of the genus are

Naturalist Notes

Fig. 3. Tree Trigger-plant Stvlidium laricifolium.

found in Western Australia, but the Tree
Trigger-plant Stylidium laricifolium (Fig.
3), confined in Victoria to the Wingan
Inlet area, with stalks up to 150 em
crowned with a panicle of usually white
flowers, is as noteworthy as any of them,
The foliage is crowded along the stem, the
young plants in particular appearing very
much like conifer seedlings and this indi-
cates the origin of the specific name,
which is derived from the Latin ‘larix’,
meaning *larch', a European conifer.
These are but three of the many attrac-
tions of river, bush, lake and beach at
Wingan Inlet.
Ron Fletcher

28 Marjorie Ave
Belmont, Victoria 3216

For assistance with the preparation of this issue, thanks to Kate Smith (desktop
publishing), Karen Dobson (label printing), Dorothy Mahler (administrative

assistance) and Phil Bock (web page).

Vol. 119 (2) 2002

Naturalist in the Mountains

The Changing Mountains

When all else is change, mountains seem
so solid. However, we should expect
mountains to change: change was after all
what formed them. I recall one geomor-
phologist questioning a land manager's
assertion that one aim of management was
to halt erosion in the mountains — erosion,
he pointed out, is a natural process, one
that is important in building mountains in
the first place. These changes take place
over many millennia. However, we are less
prepared for changes that occur within our
own lifetimes. My thoughts turned to this
topic of change as I celebrated 25 years of
naturalising in the Snowy Mountains in
early March this year with a walk into
Blue Lake to copy that first exposure to the
mountains with Will Osborne in 1977. On
that occasion | saw a Peregrine Falcon and
Black-shouldered Kites (neither commonly
seen since at that altitude — I guess I was
just lucky) and though ticking off ravens, 1
hardly noticed the pipits. Now 1 mentally
tick off these birds, eyes open for the occa-
sional patrolling kestrel or introduced sky-
lark displaying and ears open for the hid-
den Australian Raven tucked away
amongst the hundreds of Little Ravens,
Flame Robins in season (spring) and
Pacific Black Ducks in summer and low
down near the treeline, White-browed
Scrubwrens are all searched for and ticked.
These are species that were always here
but my laboriously gained expertise now
leads me to expect them. But what of the
things that weren't there? Brumbies have
become a major feature of the alpine zone
in just a few years; certainly when I com-
pleted field studies for my PhD the pad up
to the South Ramshead was walkers only
(with the occasional fox), now it is a major
horse trail from Dead Horse Gap. Yarrow
or Milfoil has spread across the mountains
wherever disturbed ground has given it a
toehold, and pigs are now commonly
found higher than ever before.

Weather always changes from what you
remember (even if it doesn't really). One
local recalled the oft-repeated “summers
aren't as hot as they used to be. When we

76

were kids we would be swimming all the
time.’ The difference is that now we are a
bit more selective about the temperature
needed before we would go for a swim. In
the same way show was always deeper as a
kid (but then again our legs were shorter!).
Leaving aside faulty memories, there have
been documented changes in the climate
(covered in the book Snow: a Natural
History; an Uncertain Future) and some
memories... | do recall in the 1970s people
ice skating on Sponars Lake; now I haven't
seen that for a time! Other changes, possi-
bly associated with climate warming, have
been the loss of frogs; the decline of Baw
Baw Frogs in Victoria and Alpine Tree
Frogs both there and here in the Snowies.
We have just completed the annual popula-
tion monitoring of Southern and Northern
Corroboree Frogs; the Southern has
declined alarmingly and may have little
future except in captive breeding at the
Amphibian Research Centre in Victoria
(along with the one remaining Spotted
Iree Frog from north of the border). So
far, the Northern Corroboree Frog is a bit
better off.

Is this change all doom and gloom? There
are some positives, the change from the
broadscale fuel reduction burning in the
mountains and the regeneration of the
Snow Gums that used to scar the side of the
Grey Mare. Following intensive winter fox
baiting to protect the Mountain Pygmy
Possum there has been a sighting of a Spot-
tailed Quoll in the winter snow and the
finding of a quoll latrine high above the
treeline near Thredbo, reminding me that
Richard Helms in 1896 recorded traces of
quolls as being abundant in the vicinity of
Bogong Moth aestivation sites. Another
positive change has been the added interest
in the mountains. When I began my natu-
ralising there was very little support for
mountain research and very little interest,
The number of researchers involved with
the Australian Institute of Alpine Studies
and the numbers that turned up at the
Biodiversity Blitz (see last edition) are
heartening evidence of a renewed interest.

The Victorian Naturalist

It is tempting on an anniversary to reflect
on these long-term changes that have taken
place, but always of interest are the
changes on a smaller scale; these are the
ones | love to observe, particularly in this
season. Autumn is possibly the time of
greatest change — leading to that funda-
mental difference between summer and
winter: the presence of snow. It is the peri-
od when animals like the Mountain Pygmy
Possum have to put on the weight neces-
sary to get them through to spring (and
they are often into hibernation well before
the first snow comes). Other animals are
shuffling their lives about. Bush Rats deep-
en their burrows to push them into warmer
depths for winter. Despite the well known
post-breeding die-off of the male
Antechinus it is the autumn that is the most
challenging for most Antechinuses at high-
er elevations with heavier animals losing
weight through the autumn, and lighter
animals exiting the population. Insects
have to lay their eggs before the summer
runs out (this is helped in the Alpine
Thermocolour Grasshopper by daily
changes in body colour to make the best
use of the available heat energy of the sun)
while those insects that are to overwinter
must start to synthesise cryoprotectants
(antifreeze and the like) from sugars. Birds
flitting over in twittering flocks are
Yellow-faced Honeyeaters, the most con-
spicuous of the emigrant birds in the

Naturalist in the Mountains

mountains. The migration of the Bogong
Moths back to their natal sites is a less co-
ordinated and spectacular event than their
arrival, with numbers seemingly melting
away over the weeks. For most reptiles and
frogs their winter is going to be similar to
the insects, a simple case of fattening up,
finding a protected hibernaculum and sit-
ting it out, However, for the Corroboree
Frogs, their eggs are sitting awaiting the
rains that will enable eggs to hatch and the
tadpoles to move from their nests in sphag-
num moss into the ponds for the winter.

Plants too are hardening for the winter,
recovering resources from above-ground
tissue, a fact attested to by the changing
colours in the mountain heaths. Seed matu-
ration is completed and cold hardiness
increases with decreasing temperature in
much the same way as in the insects.

Spring may seem a time of major change
but it is not so absolute as autumn, The
spring thaw is incomplete. winter snow
may persist as snowdrifts into March or
April, but in autumn we are awaiting that
one day when the change becomes
absolute, a summer landscape changing
subtly into autumn suddenly becomes a
winter landscape.

Ken Green

National Parks and Wildlife Service
PO Box 2228

Jindabyne, NSW 2627

One Hundred Years Ago

AMONG THE BIRDS OF WESTERN AUSTRALIA, By ROBERT HALI

On Emus -

Emus can swim rivers if necessary, so there would be no impediment on that account, A curious
incident in the domesticated habits of this bird occurred under the notice of Mr, Price Fletcher,
who described it to me in the following terms : —"A little three-year-old child was trailing a
small tin horse and cart, when along came an Emu and swallowed all but half the string. The
remaining half dangled from its beak for half the day unitl cut sufficiently short to allow it to
swallow what had been an irritation for hours. Both child and Emu were considerably aston-
ished, Tame Emus are by no means desirable pets, for strange horses when tied to fences will get
terribly frightened by the prying bird, and eventually do damage to property. Emus are said to be
a nuisance among the flocks, as young lambs will follow any large quickly moving object and
thus get led away by Emus in broad areas. By this means lambs soon lose their “dames” and
themselves. In addition the Emu seems to take a delight in chasing both lambs and mothers.”

From The Victorian Naturalist XVI, p 165, March 1902

Vol. 119 (2) 2002 TI

Tribute

Jack Hyett oam (1915-2001)

Naturalist, teacher, author, family man and friend

When asked to write this tribute to the
late Jack Hyett, | went first to his books on
my shelves, Out fell a collection of news-
paper articles, photographs, poems and
tributes [rom various natural history maga
zines, Jack had given me the poems many
years apo written by other authors but
expressing: his love for the bush.

Enclosed with the poetry was an article
written by the late JA Leach MSc at a time
when 1 was about to launch. into a teaching
career myself in primary and secondary
schools, The last paragraph of the article
states "be happy if your pupils are discover
ing things you know nothing about, for as
Libert Bailey, the eminent nature-study

apostle, tells us truly that the best teacher of

nature-study is the one whose pupils farthest
outrun him.’

Jack was a modest man who made light
of his staggering number of achievements.
He possessed a well-modulated speaking,
voice and a slow smile which widened into
a huge prin when re-living some of his best
bush experiences,

A local newspaper (1959) wrote up the
ceremony al a primary school in Blackburn
to honour him on the publication of the
book A Bushiman's Year. The article quotes
‘a book which he wrote during his school
holidays! Jack mentioned on this day he
was rushed by excited youngsters for his
autopraph and many congratulations on the
days leading up to the ceremony - 1 can
imagine the wry chuckle as he said ‘I must
agree it has only been since they discov-
ered they would be let out of school half an
hour early so this function could be held.’

Lucky were the children whom he taught
in several primary schools, especially one:
teacher rural schools, IFT had been fortu-
nate to have had even ONE teacher like
Jack during primary school! It was not to
be, Jack's arithmetic classes were cheerful-
ly interrupted by students to point out birds
in the school ground, tortoises were
watched during lessons to observe how
they breathed, frogs, lizards and insects
brought into class by students were weleome
diversions both to students and teacher,
l'ven a rounder game was abandoned com-

78

pletely to collect and study a wattle goat
moth found by one of the girls, These and
other wonderful stories are described in 4
Bushman's Year (1959), with an introduc-
tion by the late Alan Marshall, and A
Bushman's Harvest (1961), Jack pays trib-
ute to his father ‘whose interest in the bush
and its creatures inspired mine.’ IT you are
lucky these two books can still be found in
some second-hand bookshops.

Jack with his chosen profession of teach-
ing went on to inspire so many children,
very much as the late Crosbie Morrison
had done with his radio shows and
Wildlife magazines. Jack's teaching career
never really ended, as many of us were to
benefit from his CAE courses on Bird
Study in Australia, Also his students at the
teachers’ college will remember his unique
style, | remember one evening class at a
Box Hill eollege when he enthusiastically
raced to the board and illustrated the skele-
tal structure of a gannet's body, explaining
how when they hit the water when fishing,
sometimes from a great height, their skull
structure and bone density protects them
from injury. He then proceeded to draw
other birds and how they survive in the
wild. Every adult in the room had his full
attention, As an observer of detail in the
natural world he excelled,

He was a member of the Bird Observer's
Club of Australia from 1933 and recruited
many of us to join the club during his lee-
tures. When Jack led an excursion for the
club you could be sure of large crowds and
very long convoys, For seven years he was
involved in mammal and bird surveys of
Wilsons Promontory.

He was Foundation President of the
Victorian Ornithological Research Group
(VORG), The Ringwood Field Naturalists
Club, of which he was a founding member,
nominated him for the Australian Natural
History Medallion, which he received in
1985. He was awarded the medal of The
Order of Australia (1997) for his contribu-
tion to conservation and the environment,
and Honorary Life membership in May
1999 to the Field Naturalists Club of
Victoria.

'The Victorian Naturalist

|

His achievements and awards in natural
history study are widespread and many,
and include his legacy of books written on
a wide range of natural history topics. His
talks given to many Field Naturalists and
Bird Observer Clubs are legendary. This
tribute barely scratches the surface of his
many achievements. A recent photo of
Jack appears in The Vie Nat 116 (5), 158.

Book Review

As à teacher he inspired and enthused.
His love of poetry, the classics, overseas
and interstate travel made for many fasci-
nating discussions, and he was always a
loyal and caring friend and a top naturalist.

Cecily Falkingham

27 Chippewa Avenue
Donvale, Victoria 3111

A Field Companion to Australian Fungi

by Bruce Fuhrer

Publisher: Blooming Books, 2001 revised edition, paperback, colour photographs.
ISBN 1876473401. RRP 824.95

The first edition of A Field Companion to
Australian Fungi appeared in 1985, and
has been reprinted in 1993 with some
minor changes to names inserted as an
errata sheet. Bruce Fuhrer's books are well
known to us all for their wonderful pho-
tographs. Consequently, this revised edi-
tion of the out of print and very popular
earlier edition has been anticipated keenly
for some time, The demand for books such
as this attests to the increasing awareness
of fungi and related groups by many mem-
bers within our community. This inereas-
ing awareness of fungi and cryptogams has
been fostered and encouraged by the quali-
ty of Bruce's works on the subject. These
publications have provided an introduction
to groups of organisms about which there
was formerly little readily available illus-
trated literature. This heightened interest
by an increasing number of people has
facilitated community mapping and survey
schemes for fungi, such as those conducted
during naturalist clubs" surveys and forays
and Fungimap.

The obvious major difference between this
revised edition and the previous one is that
the new edition has a soft cover and Mycena
epipterygia replaces M. sanguinolenia on
the front cover photograph. As a result of the
new sofl-cover format, the book is slightly
smaller than that of the hard cover edition,
lighter and somewhat pliable so that it rests
more comfortably in the hand.

A Field Companion to Australian Fungi:
Revised Edition maintains the same format

Vol. 119 (2) 2002

as the previous edition and commences
with a short discussion about mycological
characters, classification, ecology, names
and other aspects such as distribution and
edibility as well as a simple, visual key.
However, while there is an extension to the
"Acknowledgements! there is sadly, no
other change to the introductory text. The
arrangement of species included in the
book follows that of the major groups pre-
sented in the visual key and this arrange-
ment permits a novice to quickly find
whether a particular fungus is described.
The gilled fungi group has the greatest
number of species documented and illus-
trated with photographs and these repre-
sent approximately 32 genera. The species
range from the captivating little Mycena
austrororida to the larger and much more
robust Cortinarius radicatus. The other
groups represented are coral fungi, poly-
pores, puffballs, stinkhorns, spine fungi,
leather and shelf fungi, jelly and cup fungi.

The revised edition presents over 150
photographs of the same 138 species of
commonly occurring fungi in the previous
edition, and these exquisite photographs
remain the great strength of this book. It is
disappointing that no new photographs and
no new taxa have been included in this
revised edition. Here was a wonderful
opportunity to increase the number and
range of species featured and to illustrate
the extremes of the range of morphological
variation and colour that is found in some
of the species such as Colus hirudinosus or

79

Book Review

Dermocybe cramesina. Photographs dis-
playing the range and change of colours as
some species mature, for example, the pur-
ple tones of Cortinarius archeri that turn
brown with age would be a useful adjunct,
and reduce the likelihood that speeies will
be confused with others. The images are
supplemented with thumbnail sketches in
the text of each species, but this informa-
tion is often too brief and lacking in diag-
nostic detail to really elucidate and con-
firm identification.

The revised edition of A Field
Companion to Australian Fungi is there-
fore essentially the same as the edition that
it replaces, except that the ‘errata’ pub-
lished with the 1993 reprint, have been
incorporated in the text and the names of a
further 17 taxa in the previous edition have
also been updated or corrected. These
updates include the transfer of some of the
Cortinarius species into the genus
Dermocybe, and the use of the genus
Laccocephalum for the sclerotia-forming
polypores; for example, Polyporus mylittae
is now recognised as L. mylittae, as are P.
tumulosus and P. sclerotinius, recognised
as L. tumulosus and L. selerotinius respec-
tively. Although the new names appear in
the ‘Index’, they have not been cross-refer-
enced to the older names, and the syn-
onymy similarly does not appear within
the text for each species,

The incorporation into this new edition of
labelled illustrations to outline the princi-
ple fungal morphological characters listed
in the visual key would have enhanced the
utility of the edition. As would the inclu-
sion of terms such as glutinous, habitat,
parasite, troop and spore print in the glos-
sary to cover the use of these terms in the
text. A further suggestion would be to
update the list of suggested reading materi-
al as the present list has not been updated
to include the more recent important
mycological works, such as Bougher and
Syme (1998), Grgurinovic (1997), nor the
revised edition of Young (1994). While
these works are perhaps at the more *pro-
fessional’ end of the spectrum for some,
they are often the only remaining current
resources to use that are specifie to the
Australian mycological flora.

What is so important about books such as
this is that many people rely on them to

80

identify fungi in the field by trying to
match the fungi with the photographs. It is
a wonderful sensation to succeed. Often
these books are the sole source of readily
accessible information for the Australian
mycological flora and they perform a very
important role in educating, encouraging
and fostering interest in these neglected
groups of organisms. In the case of fungi,
there is now an increasing understanding
that fungi are a vital component of nearly
every ecosystem and there is a genuine
thirst for knowledge about them. This
demand for information needs to be satis-
fied and here was an opportunity to do so.

A Field Companion to Australian Fungi
is one of the first books that I reach for
when trying to decipher the mysteries
encountered during my rambles through
the forests, As an avid consumer of won-
derfully illustrated books such as this, |
hope that further editions will expand the
number of species featured and include
more descriptive text so that we can
expand our knowledge. Many a time of
what would otherwise have been an aim-
less ramble in the bush has been trans-
formed into a treasure hunt as 1 have
eagerly looked for fungi, The book has
opened my eyes, and we are indebted to
Bruce Fuhrer for making a new edition
available to those who do not already own
a copy of the previous edition. My com-
ments ‘for more’ are a reflection of how
beneficial | have found the previous edi-
tion and my frustration in not having
access to more of these wonderful pho-
tographs of an even greater number of
species. We rely on the author to take up
the challenge and to provide us with the
tools to increase our understanding of
these vital components of the natural
world.

References

Bougher NL and Syme K (1998) Fungi of Southern
Australia. (University of Western Australia Press;
Nedlands, WA)

Grgurinovie CA (1997) Larger Fungi of South
Australia. (The Botanie Gardens of Adelaide and
State Herbarium and The Flora and Fauna of South
Australia Handbooks Committee: Adelaide)

Young T (1994) Common Australian Fungi. 3 ed.
(University of New South Wales Press: Sydney)

Jenny Tonkin

Plant Sciences and Biodiversity

Royal Botanic Gardens, Birdwood Avenue
South Yarra, Victoria 3141

The Victorian Naturalist

Tribute

Alexander Clifford Beauglehole

26 August 1920 — 19 January 2002

Cliff was the youngest of the three sons
of Richard and Margaret Beauglehole,
dairy and potato farmers at Trewalla, near
Portland. Soon after Cliff was born the
family moved to a mixed farm at Gorae
West where Cliff attended the nearby
country school, travelling each day on
horseback down a bush track. Through nat-
ural curiosity and encouragement from his
parents he soon developed a good knowl-
edge of the natural history of his area. By
the time he left school he could scientifi-
cally name about 60 species of local
orchids and had begun sending specimens
to WH Nicholls, a leading orchid specialist
of the period. During the 1940s he discov-
ered three previously undescribed species.
Soon after leaying school he began a sys-
tematic botanical survey of the Portland
area, listing over 750 species, almost dou-
ble the number then recorded.

In 1949 Cliff married Hilda Oakley. He
was able to buy the farm at Gorae West and
subsequently their daughters Valerie and
Yvonne were born. From the outset Hilda
took a keen interest in, and often helped
with, Cliff's projects. Her assistance in run-
ning the farm was invaluable in enabling
him to give time to his explorations,

Early in 1950 Cliff began a study and col-
lection of the local native bees, in associa-
tion with the entomologist Tarlton

Rayment’, This resulted in the recognition of

30 new species, three of which were named
after the collector and five after the district.
He also worked on wasps with Tarlton
Rayment and on ants with John McAreavy.

Cliff's interest in birds led to the collec-
tion and recording of beach washed sea
birds from Discovery Bay. Between 1951
and 1963 almost 5000 carcases were col-
lected by Cliff and his helpers including
950 in one day in August 1959. The skele-
tons were prepared by natural decomposi-
tion of the flesh and were eventually
donated to museums in Australia and the
United States.

During the 1950s Cliff and Fred Davies
of Portland commenced an extensive sur-
vey of bone deposits in the many limestone

Vol. 119 (2) 2002

caves of the area. Cliff had long been aware
of the significance of these and was con-
cerned at the damage being done to the
caves through rubbish dumping and road
works. In 1963 Cliff and Fred, together
with a team of local naturalists, started the
excavation of McEachern’s Death Trap
Cave. Several tons of cave sediments were
sieved and the identifiable remains of some
2,000 animals were recovered. In 1964-5
Norman Wakefield? became involved in the
excavation. The assessment of this material
added immensely to the knowledge of the
faunal succession of the area.

Throughout these years, Cliff continued
his botanical explorations, extending his
field to include the non-vascular plants. He
covered the whole of the south-west area
of Victoria in search of mosses, liverworts,
lichens and fungi. He also studied and col-
lected the marine and freshwater algae of
his area. He had an amazing ability to rec-
ognize new or rare species and his collec-
tions are proving a treasure house for pre-
sent day taxonomists,

During the 1960s Cliff widened his hori-
zons still further and undertook two trips to
Lord Howe Island, one long trip through
the Northern Territory and West Australia
and three shorter trips in central Australia.
On each occasion he brought home large
collections of specimens, In 1967 he
embarked on a full-scale botanical survey

Cliff men uot: in the 1950s. Photo by LG
Chandler.

81

Tribute

of the Grampians. During the 1940s he had
made several trips by motor-bike to some
of the remoter areas of the Victoria Range
and believed that, in spite of a good deal of
work done by others, there was still much
to be discovered. Cliff was already corre-
sponding with Jim Willis at the National
Herbarium and had accompanied him on
field trips in the Grampians and Portland
area. Jim's encouragement in the
Grampians survey also led to the first offi-
cial recognition of Cliff's work, when he
received a grant from the Maud Gibson
Gardens Trust.

Following this first survey Cliff worked
for the National Parks Service in Western
Victoria. He then moved on to a study of
East Gippsland and eventually completed
plant surveys of the whole of Victoria.
This work was subsidised by the Land
Conservation Council of Victoria who
published his results in the form of conser-
vation recommendations. Cliff subsequent-
ly published complete checklists for the
whole State. He received some financial
support from the Western Victorian Field
Naturalists’ Clubs Association and from
individual clubs and kindred associations.
He also received a substantial grant from
the Utah Foundation (USA) toward the
alpine area surveys, and a grant from
Alcoa Australia for work on their smelter
site in Portland. He was influential in
inspiring many country naturalists to con-
tinue to monitor their environment,
enabling them to report reliably on conser-
vation issues.

Botanists throughout Australia and many
from overseas knew Cliff personally, He
willingly gave time to guiding and assist-
ing visitors doing fieldwork in his area.
His friends and associates have lasting
memories of expeditions with Cliff. His
boundless energy and enthusiasm and his
wide knowledge of the environment were
exhilarating. Lunch breaks provided an
opportunity to study marauding insects and
to watch birds. His keen eye for different
or unusual plants and his meticulous care
of his specimens were an inspiration and
example to less experienced field workers.
Above all he will be remembered for the
diversity and quality of his collections,
numbering over 90,000, which have been
lodged in herbaria all over Australia and

82

Taxa named after Cliff.

Vascular Plants

Bassia beaugleholei Ising

Caladenia beaugleholei D.Jones
Calonema beaugleholei D.L.Jones
Epilobium brunnescens subsp. beaugleholei
K.R.West & P.H.Raven

Lobelia beaugleholei D.E.Albrecht
Prasophyllum beaugleholei W.H.Nicholls
Solanum beaugleholei D.E.Symon
Stylidium beaugleholei 1.H. Willis
Utricularia beaugleholei R.J.Gassin
Villarsia unbricola var. beaugleholei H.1.Aston
Marine alga

Helminthoclada beaugleholei Womersley
Moss

Phascum beaugleholei \.G.Stone

Lichen

Heterodea beaugleholei Filson

Bees

Exoneura cliffordiella Rayment

Hylaeus cliffordiellus Rayment
Megachila cliffordii Rayment

overseas. These will continue to be an
invaluable resource and a permanent
memorial to his dedication.

Cliff participated with local naturalists in
many campaigns for environmental con-
servation. Their efforts were rewarded
when the Mount Richmond and Lower
Glenelg National Parks were gazetted. He
was actively involved in the formation of
the Western Victorian Conservation
Committee that led the battle for the
preservation of the Little Desert. He was a
foundation member of the Portland Field
Naturalists Club and of the Western
Victorian Field Naturalists Clubs
Association, a member of the National
Parks Association and of the Bird
Observers Club, In 1982 he became an
Honorary Member of the Field Naturalists
Club of Victoria in recognition of 40 years
continuous membership. In 1971 he was
awarded the Australian Natural History
Medallion’ and in 1984 the Medal of the
Order of Australia.

Cliff is survived by his wife, two daugh-
ters, seven grandchildren and two great-
great grandchildren.

! Bees of the Portland District (1953), Tarlton Rayment
The Victorian Naturalist 84, 363-83 (1967)
The Victorian Naturalist 88, 344-46 and 360-361 (1971)

Margaret Corrick
7 Glenluss Street
Balwyn, Victoria 3103

The Victorian Naturalist

Book Review

Wildflowers of Sydney and Adjoining Areas

by Alan Fairley

Publisher: Bloomings Books 2001. 268 photographs, ISBN 187647338X, RRP $32.95

The blank space on top of the cover of
this book should read *A Field Guide to
the Wildflowers of Sydney and Adjoining
Areas’. Perhaps this was a publisher deci-
sion and not the author's. This compact,
easy-to-use book is just perfect for the rain
coat, parka or rucksack pocket.

The alphabetical layout is quick and easy
to use in the field, although botanical
groupings can be difficult if one is not sure
of the family, e.g. a typical page layout is:

Acacia binervata Two-veined Hickory
Family Mimosaceae

This is followed by a coloured photo,
description, flowering period, distribution,
habitat, notes, similar species and specific
sites. In other words all you need to know
from a Field Guide.

If 1 had possessed Alan’s book on previ-
ous trips to Kuring-gai Chase, Royal
National Park and Lane Cove National
Park, my plant identification would have
been much easier with so many of the
commonly seen species all in one book.

On my first trip to the above-mentioned
places, | was just learning botany and |
remember juggling at least three books,
cross-referencing and burning the midnight
oil. For this reason I think Alan's book will
suit the beginner as well as the more expe-
rienced plant lover.

All photographs have been taken by the
author, in situ, without flashlight. Most
photos are adequate for identification with
some outstanding photos of Telopea spe-
ciosissima, the calistemons, banksias,
Philotheca, Correa and Thysanotus
tuberosus.

I was disappointed that some of the small-
er flowers were not filmed with a close-up
lens, i.e. Rulingia hermanniifolia,
Micromyrtus ciliata, Grevillea buxifolia,
Hardenbergia violacea and the baeckias.
The author mentions the twisted labellum
in Prerostylis curta but has photographed
the back and side of two specimens. The

Vol. 119 (2) 2002

same with Prerostylis concinna, where the
deeply notched labellum is mentioned but
not shown. Drosera spathulata and
Caleana major could have been sharper
and larger. Identification of eucalypt
species would have been easier with close-
up photos of fruits on all species. The
small insert photo, showing in some
instances canopy or trunk, are not helpful
in identification.

The author states that his book Wild-

flowers of Sydney aims to encourage an

appreciation of the floral beauty and vari-
ety of this area, but I was a little disap-
pointed that none of the beautiful grasses
in full bloom were included.

Attention to these few minor drawbacks
of the book in my view could have
improved an already fantastic Field Guide
to some of the most beautiful and accessi-
ble areas not far from Sydney.

A map of Sydney and adjoining areas is
provided and all the places mentioned in
the book are shaded green. This works
well with site information specified on
each page for every plant described.

The references and further reading will
be much appreciated by the reader and the
glossary is what you would expect from an
experienced naturalist, bushwalker and
author of 25 books.

This book is not just for the resident or
visitor to Sydney it also contains 122
Victorian species and 153 Queensland
species which also occur in Sydney and
adjoining areas. Only 34 are restricted in
their distribution to the Sydney region.

I would highly recommended the pur-
chase of this great, compact little book,
bulging with information to anyone plan-
ning a trip to Sydney or adjoining areas.

Cecily Falkingham
27 Chippewa Avenue
Donvale, Victoria 3111

The Field Naturalists Club of Victoria Inc.

Reg No A0033611X

Established 1880
In which is incorporated the Microscopical Society of Victoria

OBJECTIVES: To stimulate interest in natural history
and to preserve and protect Australian flora and fauna.

Membership is open to any person interested in natural history and includes
beginners as well as experienced naturalists.

Registered Office: FNCV, 1 Gardenia Street, Blackburn, Victoria 3130, Australia.
Postal Address: FNCV, Locked Bag 3, PO Blackburn, Victoria 3130, Australia.
Phone/Fax (03) 9877 9860; International Phone/Fax 61 3 9877 9860.

Patron
John Landy, MBE, The Governor of Victoria

Key Office-Bearers
President: Ms WENDY CLARK, 97 Pakenham Street, Blackburn 3130. 9877 9266
Vice Presidents: DR NOEL ScHLEIGER, | Astley Street, Montmorency 3094. 9435 8408
and DR ALAN YEN, 52-54 Brushy Park Road, Wonga Park, 3115. 9722 1665
Hon. Secretary; MRS ANNE MORTON, 10 Rupicola Court, Rowville 3178. 9790 0656
Hon. Treasurer: Ms BARBARA BURNS, 16 Montclair Court, Templestowe 3106, 9846 2608
Subseription-Secretary: FNCV, Locked Bag 3, PO Blackburn 3130. 9877 9860
Executive Editor, The Vic. Nat.: MRS MERILYN GREY, 8 Martin Road, Glen Iris 3146, 9889 6223
Editors, The Vic. Nat.: MR ALISTAIR EvANS, 28 Chandler Road, Glen Waverley 3150. 8505 4339
and Mrs ANNE MORTON, as above.
Librarian: Mrs Sneit.A HOUGHTON, FNCV, Locked Bag 3, PO Blackburn 3130, AH 5428 4097
Excursion Co-ordinator: MR DENNIS MELTZER, 8 Harcourt Avenue, Caufield 3162. 9523 1853
Book Brokerage: MR RAY Wntrk, 9 Longtown Court, Craigieburn 3064. AH 9308 3770
Newsletter Editors: DR NOEL SCHLEIGER, as above, MR KEITH MARSHALL, 8/423 Tooronga Road,
Hawthorn East 3123. 9882 3044, and MRS JOAN BROADBURY, 2 Shaun Court, Templestowe 3106, 9846 1218
Conservation Coordinator: MR Jim WALKER, 167 Balaclava Road, Caulfield 3162. 9527 5601
Group Secretaries
Botany: Ms KAREN DOBSON, 58 Rathmullen Road, Boronia 3155. BH 9877 9860
Geology: Mr Ron HAMSON, 5 Foster Street, McKinnon 3204, 9557 5215
Fauna Survey: Ms Soputt: SMALL, 107 Bondi Road, Bonbeach 3196, AH 9772 2848
Marine Research: MR MICHAEL Lyons, 18 High Street, Nunawading 3131. AH 9877 3987
Microscopical: MR RAY POWER, 36 Schotters Road, Mernda 3754. 9717 3511

MEMBERSHIP
Members receive The Victorian Naturalist and the monthly Field Nat News free. The Club organis-
es several monthly meetings (free to all) and excursions (transport costs may be charged). Field
work, including botany, mammal and invertebrate surveys, is being done at a number of locations
in Victoria, and all members are encouraged to participate.

YEARLY SUBSCRIPTION RATES — The Field Naturalists Club of Victoria Inc.

First Member
Metropolitan $44
Concessional (pensioner/student/unemployed) $33
Country (more than 50 km from GPO) $30
Junior (under 18) $16
Overseas AU$65
Additional Members
Adult $16
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Send to: FNCV, Locked Bag 3, PO Blackburn, Victoria 3130, Australia,

Printed by Brown Prior Anderson, 5 Evans Street, Burwood, Victoria 3125.

The
Victorian

Naturalist

Volume 119 (3) June 2002

Murray River Special Issue Part One

rU Published by The Field Naturalists Club of Victoria since 1884
Ga! “aw

| F.N.C.V,

Introduction — Murray River Special Issue

Terry Hillman*

Australia’s major river; a resource criti-
cal to our national rural production; a
beautiful and sometimes unique landscape;
an ecosystem under threat — the Murray is
all of these.

The Murray Darling system drains about
'/, of Australia’s surface but, despite its
size, flow volumes are small reflecting the
arid nature of most of its catchment. The
average annual (natural) discharge from
ihe system is equivalent to 16-17 hours’
flow from the Amazon. The Basin sup-
ports nearly half of Australia’s rural pro-
duction. On average, water use (almost all
irrigation) removes around % of the
Murray's water before it reaches the sea'
and dams and weirs drastically change the
pattern of flow.

Two other key aspects of the Murray's
flow are variability (*droughts and flood-
ing rains’) and unpredictability. These con-
ditions favour organisms adapted to peri-
ods of wet and dry extremes with little sea-
sonal or inter-annual reliability". Many of
the papers in this special issue of The
Victorian Naturalist deal with organisms
or whole systems (e.g. River Red Gums,
turtles, and floodplain wetlands) which
depend on rainfall or flow cues to initiate
vital functions. Again, the effects of flow
regulation are significant here,

Beauty is, of course, personal and
ascribed by the ‘beholder’ but “wetness
and dryness’ must surely contribute much
to what is beautiful in the Murray land-
scape and the response of its organisms to
these forces provides what is characteristic
and sometimes unique,

In natural resources it is a common mis-
take to confuse ‘use’ with ‘management’,
and most of what we have termed manage-
ment in the Murray to this point in time has
really been the development of more effi-
cient ways of using the resource.

' Department of Environmental Management and
Ecology, La Trobe University. PO Box 821, Wodonga,
Victoria 2689

Management implies knowledge. If we set
out to manage a natural resource we need to
know how it works — which means in this
case that we need an understanding of the
*Murray ecosystem’, its key organisms, its
flow-related processes, its biodiversity and
the factors supporting it. This special issue
provides knowledge at all these levels.
Current knowledge regarding fish, terrestri-
al vertebrates, and invertebrates (aquatic
and terrestrial) is provided. The significance
of the floodplain as a functional part of the
river is recognised through papers on Red
Gum forests, their fauna and the signifi-
cance of fallen timber, and on floodplain
wetlands. Geomorphology and connectivity
with wetlands are two important flow-relat-
ed issues discussed. Diversity at every level
from fish to microinvertebrates is ‘show-
cased’ and includes a useful key to tadpoles
and a fascinating introduction into how bio-
diversity is maintained using the example of
niche segregation by turtle species.

This special issue of The Victorian
Naturalist represents a valuable and acces-
sible compendium of knowledge about the
Murray. For those who remember the
Royal Society of Victoria symposium on
the Murray Darling held in October 1977
(1 do, sadly) it is instructive to see how far
we have developed in knowledge and
emphasis over the past 25 years.
Compendia are usually shelved for future
reference as a need arises. These papers
should be read now. We are facing impor-
tant decisions regarding the management
of the Murray — particularly flow regula-
tion — and the community (regional and
national) will need to form an opinion
regarding the trade-off between water for
production and water for supporting the
Murray ecosystem. This special issue will
provide an ecological basis on which such
an opinion can (and should) be formed.

' Reducing the output to 4-5 hours of “Amazon
flow’.

* Note that the enforced reliability of seasonal
flow, resulting from river regulation, tends to
remove the competitive advantage which such
adaptation would otherwise provide.

The

Victorian
Naturalist

Volume 119 (3) 2002
Murray River Special Issue
Part One

June

Executive Editor: Merilyn Grey
Editors: Alistair Evans and Anne Morton

This issue is dedicated to
Natalie Joanne Smith (31 May 1974 — 22 July 2002)

AGU OME gt ed IDE e e 68i HA HEAR aay nor ente 86

National Parks and Reserves on the Murray River: an Historical Perspective,
D ROI TS TOS Pee cent Ae UO ER EET ette 88

Geomorphology of the Murray River Basin, by Noel Schleiger sss B5

Floodplain Wetlands: the Jewels of the Murray River, by Rhonda Butcher
moet ee T e ee. t ed ip NIA icto f Rae e E A Da E ve LED QD) eo RS AER Cor 102

Habitat Change in River Red Gum Floodplains: Depletion of Fallen Timber
and Impacts on Biodiversity, by Ralph Mac Nally, Andrea Ballinger
ana GYegory HOPrOGQEs Lese ees ereo repr stein deenen ntar apye n eben doin Ih ttm eaten etta 107

Vertebrate Fauna of Barmah Forest, a Large Forest of River Red Gum
Eucalyptus camaldulensis on the Floodplain of the Murray River,
by Richard H Loyn, Linda F Lumsden and Keith A Ward ............. eee 114

The Distribution and Conservation Status of the Reptile Fauna of the
Murray River Region in Victoria, by Geoff Brown........... eee 133

Tadpoles of South-eastern Australia: a Guide with Keys, by Marion Anstis,
Veyicweo- Uy ANUS MAT IH E he A caseus eee Re Te ETC EET TENES UE NEST UNE Gn Me Ln 144
Rivers as Ecological Systems: The Murray-Darling Basin, edited by
WJ Young, reviewed by Gerry Quinn............... reete 147

ISSN 0042-5184

ML
Cover: Morning reflections on a Murray River billabong near Mildura. Photo by Anne

Morton.

Web address: http://www.vicnet.net.au/~fnev/vicnat.htm
email: fnev@vicnet.net.au

Murray River Special Issue

National Parks and Reserves on the Murray River:
an Historical Perspective

Daniel Catrice'

Abstract

Attempts to establish national parks and reserves along the Murray River occurred late in the history
of Victoria's conservation movement. For much of its recent history the Murray has been made to
serve industry and agriculture. The first timber reserves and State forests, which had been set aside
from 1869, confirmed a pattern of exploitation which went largely unquestioned until well into the
twentieth century. This article looks at settlement and land-use along the Murray and the efforts of
conservation groups to protect the lands bordering the river from unregulated development. It con-
cludes with a brief historical overview of the key parks and reserves along the river — Hattah-
Kulkyne, Murray-Sunset and Alpine National Parks; Barmah State Park; and Murray Kulkyne Park.

(The Victorian Naturalist 119 (3), 2002, 88-94)

Introduction

The Murray River is the main stream of
Australia's greatest and most important
river system. It rises in the Australian Alps
just below Mt Kosciuszko and enters the
sea in South Australia. For 1800 km of its
length it forms the border between New
South Wales and Victoria. This journey
takes the river through some of Victoria's
most significant and diverse natural land-
scapes, from wild forested gorges, through
riverine plains and majestic Red Gum
forests, to semi-arid desert country.

Sadly, the river has been taken for grant-
ed for most of its recent history. By the
1840s most of the river frontage had been
taken up by land-hungry squatters for
whom the river was a source of water and,
later, a convenient transport route for pro-
duce and supplies. The floodplain forests
were exploited as a timber resource and the
country beyond was partitioned into farms
and grazing runs. To promote agricultural
settlement the river was dammed and its
waters diverted for irrigation projects. The
Murray is now a regulated river. Weirs and
dams have ensured water in the Murray all
year round, but these have turned the river
into a series of connected pools.

The conservation movement turned its
attention to the Murray relatively late. The
first national park along the river, Hattah
Lakes, was reserved only in 1960, and
today there are still only a scattering of
conservation reserves. Groups like the
Field Naturalists Club of Victoria and the
Royal Australasian Ornithologists Union

Department of Natural Resources and Environment,
PO Box 41, East Melbourne, 3002
Email Daniel.Catrice@nre.vic.gov.au

88

began to show interest in the Murray from
the early 1900s, and with the assistance of
sympathetic officers in the Fisheries and
Wildlife Department and the Forests
Branch of the Lands Department, impor-
tant areas like the Kulkyne and Barmah
forests were managed with greater care.

This growing enthusiasm for nature
emerged from a sense that too much had
been lost in the years since white settle-
ment. As one writer observed: ‘Nature, as
if offended, withdraws half her beauty
from the land; the pasture gradually loses
its freshness, some of the lakes and rivers
run low, others become wholly dry. The
wild animals, the former peaceful denizens
of the soil, are not more to be found, and
the explorer, who has gazed on the district
in its first luxuriance, has seen it as it never
can be seen again’ (Henry W Haygarth
1848, cited in Dingle 1984: 138).

Over time, conservation groups became
more vocal in their demands for the estab-
lishment of national parks to preserve areas
of ‘unspoilt nature’. This article investigates
some of the human impacts of settlement
and land-use along the Murray and reviews
the various efforts to protect land and
resources in the face of the rapid, some-
times devastating changes worked upon the
landscape since European settlement,

Exploration and settlement

The first Europeans to see the Murray
River were the exploration party led by
Hamilton Hume and William Hovell. They
crossed the river near Albury in 1824,
naming it the Hume. Just six years later the
explorer Charles Sturt renamed it the
Murray in honour of Sir George Murray,

The Victorian Naturalist

then British Secretary of State for the
Colonies. Always with an eye to the future,
Sturt saw the Murray as a river highway
helping to open up vast tracts of the
Australian outback. After Sturt came Major
Thomas Mitchell who crossed the Murray
at Robinvale in 1836. Like Sturt, Mitchell
believed the river should be made to serve
a purpose. Travelling down the river
towards Swan Hill he noted ‘grassy plains
bounded by sand hills on which grew
pines’ and predicted, accurately, that the
area would be favourable for cereal crops,
grazing or irrigation (Bardwell 1990: 23).
Explorers like Hume and Hovell and
Major Mitchell travelled with an eye for
new pasture, so it is not surprising that
squatters soon followed in their footsteps.
The Murray River promised a near perma-
nent water supply, natural pasture and a

good supply of timber. Within ten years of

Mitchell's expedition, squatters had taken
up most of the river frontage.

Edward Curr, a Tasmanian merchant and
landowner, established a run at Barmah in
1841. Curr later wrote that the land was
flooded for several months of the year, but
then *abounded ... with excellent sheep
feed, in the shape of couch grass, young
reeds and so on’. It was, he said, ‘usually
as green as an emerald from November to
March when [other] pastures were with-
ered and dried’ (Curr 1965: 76).

From the outset, settlers drastically
changed the environment, Trampling by
stock compacted delicate vegetation and
soil, muddied and eroded precious water-
courses, and introduced and spread new
grasses. Cattle selected the most palatable
species, which led to a decline in the density
and diversity of native plants. They
devoured the fine fuels that carried fire,
leading to the invasion of native grasslands
by scrub. They spread exotic weeds in fur
and dung. They removed seed heads and
flowers, which disrupted plant reproduction.

After the squatters came selectors, encour-
aged by a series of Land Acts which had
been enacted in the 1860s. Selectors who
took up land were required by the govern-
ment to ‘improve’ their selections by clear-
ing vegetation. They did this with indis-
criminate zeal, spurred on by the need to
render every acre productive. Ringbarking
was the cheapest form of clearing as one

Vol. 119 (3) 2002

Part One

contributor to The Victorian Naturalist
found when he visited the country between
Kulkyne Station and Mournpall in 1914:
‘twenty years previous to our visit these
trees [Black Box Eucalyptus largiflorens|
had been ringed, and the resulting conse-
quences were acres of bare, wind-swept
earth, destitute of aught green, and a bewil-
dering interlacement of fallen trunks, and
limbs that would appall even an old-time
South Gippsland pioneer’ (The Victorian
Naturalist 31, 1914).

Settlers also made extensive use of River
Red Gum Eucalyptus camaldulensis for
huts and fences, and despite a proclama-
tion that all Murray Pine Callitris spp.
within ten miles of the river was to be left
untouched, by 1875 the local forester at
Barmah reported that all of the Murray
Pine stands at Barmah Island had been
removed (Hibbins 1991).

Land clearing upset the ecological bal-
ance. In 1858, the naturalist Gerard Krefft
explored the lower Murray and Darling
Rivers with Victorian Government
Zoologist William Blandowski. Krefft
noted that the Rabbit-rat Conilurus albipes,
Gray's jerboa Kangaroo or Burrowing
Bettong Bettongia lesueur graii and several
other species of marsupial had become rare
or extinct on the Victorian side of the
Murray (Dingle 1984). Ironically, others
complained that the kangaroo ànd possum
were multiplying at an astounding rate, due
in part to the decimation of Aboriginal
tribes who had hunted them for food.

Some of the earliest settlers looked back
in later years and lamented the changes that
they saw. Edward Curr decided that ‘fences
and tree-ringing have not improved the
scene’ nor could he enthuse over progress
when old forests along the Murray were
‘fast being converted by steam sawmills
into railway sleepers’. Fle remembered with
regret ‘the primitive scene, the Black with
his fishing canoe, the silence, the gum-
trees’ (cited in Dingle 1984; 138).

‘The Nile of Australia’

Navigation was the first commercial use
of the Murray. River boats began operating
on the Murray as early as 1853, carrying
supplies inland and returning with pro-
duce, They opened up remote areas for set-
tlement and helped to make the pastoral

89

Murray River Special Issue

and agricultural industry more profitable.
They were also able to carry goods more
efficiently than bullock dray. So great was
the promise of river transport that some
began to talk of the Murray as * The Nile of
Australia’ or “The Mississippi of New
Holland’ (Davies 1978: 34),

By the 1870s up to thirty steamers and as
many barges worked the Murray, Darling
and Murrumbidgee Rivers. The bulk of the
Riverina woolclip was thus transported
southwards through the ports of Echuca
and Swan Hill (Priestley 1984: 59). Smaller
steamers worked the ‘top end? ports of
l'ocumwal and Albury. Over the next thirty
years the railways slowly robbed the river
trade of the bulk of its traffic, By the mid-
1930s it had virtually ceased.

l'o meet the river boats" need for fuel,
vast quantities of Buloke Casuarina
luehmannii, River Red Gum and Black
Box were cul. Early steamers consumed
wood at the rate of about one ton every
eight hours; later, more powerful steamers

Ronald East of the State Rivers and Water
Supply Commission stands astride the Murtay
River near Nyah State Forest during the drought
of 1923, (Picture Collection, State Library of
Victoria)

90)

had a wood consumption of one ton per
two hours (Davies 1978: 46), Strenuous
efforts were also made to clear the river of
‘snags’, the fallen trees and limbs which
fell into the water. This assisted navigation
but had a marked effect on native fish, par-
ticularly Murray Cod Maceullochella
peeli, which used fallen trees as shelter and
breeding habitat. Fish numbers declined
dramatically during this period, from
29.064 kilograms netted in 1835 to only
4273 kilograms taken from the same areas
in 1896 (Mackay and Eastburn 1990),

Irrigation

Irrigators began tapping the waters of the
Murray in the 1870s. Headworks were
constructed to divert water from the
Murray near Gunbower Creek and an oft-
take supplied the Cohuna, Kerang and
Koondrook districts. The dry years which
began in 1877 saw other growers clutch at
the promise of irrigation. They were assist-
ed by the government which assumed con-
trol over all the natural water resources in
the State in [886 (Eastburn 1990).

The most audacious irrigation project
took shape at Mildura. In 1884 Canadian-
born brothers George and Ben Chaffey
were invited by the Victorian government
to establish an irrigation settlement on the
Murray (Davies 1978). The Chaffeys were
granted 50,000 acres of rabbit-infested
land which they cleared, subdivided and
serviced with water channels. Huge pumps
were placed in 1889 at Psyche Bend where
water was drawn from the Murray and
pumped to Kings Billabong. From there.
through a series of pumps and open chan-
nels it was distributed throughout the area,
At Psyche Bend the steam engine drove
three centrifugal pumps, each delivering
8000 gallons of water per minute.

Ultimately, irrigation along the Murray
was not the success it was envisaged by
government or farmers. Environmentally, it
was disastrous, The waste of water through
evaporation and seepage was enormous.
Worse still was the effect of clearing and
ploughing on vast areas inland from the
river, This allowed rain to run straight into
the subsoil raising the water table and
bringing saline groundwater close to the
soil surface. The introduction of irrigation
water exacerbated the problem. Irrigation-

'The Victorian Naturalist

induced salinisation was recorded at
Mildura as early as 1895 (Eastburn 1990).
By 1911 it appeared near Cohuna and
Kerang. Today it affects virtually all
regional settings along the river, including
the highlands, the riverine plain and the
mallee of the Murray-Darling Basin.

Timber reserves and state forests
The first attempts to control land and
resources along the Murray came in 1865

when the Victorian government gave itself

the power to proclaim reserves for the
‘preservation and growth of timber’
(Wright 1989: 107). By the close of the
decade 276,116 acres had been set aside as
state forests and timber reserves, including
the Barmah Forest of 19,600 acres, created
in 1869 (Wright 1989). Further reserves
were created at places like Gunbower
Island, Nyah and Walpolla, partly to halt
the damage being done to River Red Gum
forests as a result of unregulated harvest-
ing, but also to secure a valuable economic
resource.

These developments were not motivated
by ecological intent. Government action
was propelled largely by utilitarian con-
cerns. Red Gum forests along the Murray
continued to be heavily exploited for mill
logs. During the nineteenth century,
sawmillers had only to pay £15 to set up a
mill in the forest, and for £10 for cartage
and £5 for each man employed in felling
timber, he could fell the pick of the trees in
the forest (Fahey 1987). In the Barmah
Forest the Red Gum trade flourished,
prompting one observer to claim that the
timber was an ‘object of worship on the
Murray’:

The mills declare dividends because
of it; the sleeper-cutter keeps the pot
boiling in return for his daily sweat
upon it. The bullock driver draws his
Saturday night beer from it. The
axeman’s camp in the bush is kept
supplied in tucker by it. The barge-
master, the deck hand, and the skip-
per of the river tramp knows its
value well (Brady 1912, cited in
Hibbins 1991; 38).

The 1901 Royal Commission on State
Forests and Timber Reserves reported crit-
ically on the use and management of the

Vol. 119 (3) 2002

Part One

f -—

A load of Red Gum timber arrives at the Evans
Bros sawmill in Echuca, 1923. (Historic Places
Section, DNRE)

Barmah Forest. The Commission noted
that the forest had been cut over several
times, and that at the current rates of cut-
ting, the forest would not yield five years’
supply (Fahey 1987).

A similar pattern of use developed in most
of the Red Gum forests along the river. By
1932 the district forester at Swan Hill report-
ed that all the red gum at Annuello,
Mamboing and Toll Toll had been cut out
(‘Hattah-Kulkyne National Park? Resource
Collection File F/C: 3/1).

It took many decades for the State's forests
to be placed under more effective manage-
ment. Although a series of bills specifical-
ly for forest legislation were introduced
into parliament, in 1879, 1881, 1887 and
1892, none was enacted. Finally, in 1907 a
Forests Act was passed, which provided
for the establishment of a State Forests
Department under a Minister of Forests. A
Conservator of Forests was appointed,
with appropriate supporting staff and
power to declare permanent forest reserve
(Moulds 1991). Over the next decade, sev-
eral initiatives were undertaken, including
improvement works on the forest estate,
the establishment of hardwood plantations
and improved fire protection measures.

From the 1920s foresters based at Ouyen,
Swan Hill, Nyah and Mildura made
attempts to re-alforest native pine areas.
Unfortunately, until the 1960s these efforts
were largely unsuccessful. In the Barmah
Forest, thinning and coppice treatment was
carried out in the 1920s and 1930s.
Foresters also experimented with non-
indigenous species. At the Timberoo Forest
in the 1930s Red Ironbark Eucalyptus
sideroxylon and Silky Oak Grevillea robus-

91

Murray River Special Issue

ta were trialed (*Hattah-Kulkyne National
Park’ Resource Collection File).

Foresters were often the first advocates
for conservation. It was at their insistence,
during the 1930s, that settlers in the
Hattah-Kulkyne area were forbidden to
remove native pines from their selections.
In the Barmah Forest in the 1940s foresters
and scientists began to re-assess the impact
of grazing. Forester, WP Tingate drew
attention to the damage done to saplings by
cattle, “When grass was dry’ he said, *cat-
tle would eat anything green'. Tingate's
concerns were echoed by his colleague,
Stan Duncan, who observed in 1950 that
there was little hope for widespread re-
generation of the forest until grazing was
discontinued. The Forests Commission
acted on this advice and in 1950 Barmah
West, Yalca and portions of Yielima were
closed to cattle (Fahey 1987).

Water reserves

Another government concern of the
1870s and 1880s was the need to protect
water resources from the designs of squat-
ters and selectors. Thus, in 1881 all
unalienated land within one to one-and-a-
half chains of the colony's *Rivers,
Rivulets, Creeks, Channels, Aqueducts,
Lakes, Reservoirs, Swamps, Inlets, Loughs
and Straits’ was reserved (Wright 1989:
224). The order ensured the Murray River
water frontage was retained in public own-
ership and today this corridor forms the
basis of the Murray River Reserves man-
aged by Parks Victoria.

National parks

For many conservation-minded people,
the creation of State forests and Crown
land reserves was not enough. Towards the
end of the nineteenth century a growing
body of scientific opinion became increas-
ingly vocal in their demands for the protec-
tion of wildlife and the establishment of
national parks. In the broader community
too there was a growing awareness of the
natural world. Changing recreation pat-
terns brought about by extensions to the
rail network and the emergence of field
naturalist and bushwalking clubs awak-
ened people to the beauties of nature.

The Field Naturalists Club of Victoria
(FNCV) was one of the first supporters of
national parks. Established in 1880, the

92

FNCV combined the desire for open air
recreation with an interest in the natural
environment. The Club instituted walking
and camping expeditions in which the
observation of wildlife and the collection
of botanical specimens were important
activities. The FNCV was soon joined by
other societies, including the Wattle Club,
the Royal Australasian Ornithologists
Union, and the Bird Observers Club. This
small but committed conservation move-
ment, led by influential spokesmen like
Baldwin Spencer and Sir James Barrett,
began to exert pressure on the government
to protect Victoria's bushland.

In 1914, the FNCV became interested in
the Hattah Lakes system near Mildura. At its
meeting on 14 December the Club decided
to write to the Fisheries and Game
Department and request that the area be pro-
claimed a sanctuary for native game. Native
game sanctuaries were proclaimed under the
Game Act of 1890 and made it unlawful for
any person to kill any native game within a
designated area and specifically mentioned
by proclamation (Mahon 1992). Of particu-
lar concern to the FNCV was the practice of
punt-gun shooting on Lake Mournpall.
Speaking for the resolution, JG O'Donoghue
reported on a recent visit to the Lake where
he claimed that, after a single discharge of
the gun used by one individual, as many as
72 pairs of ducks had been ‘picked up’ (The
Victorian Naturalist 1915: 31). The resolu-
tion was supported by the Inspector of
Fisheries and Game, Mr F Lewis, and in
1915 an extensive area embracing parts of
Chalka Creek, Lakes Hattah, Little Hattah,
Mournpall and Brockie, together with all
land within half a mile of the creek and lake,
was proclaimed a sanctuary for native game
(The Victorian Naturalist 1915: 32).

In 1919, about 3620 acres were reserved
as the Kulkyne State Forest, and an addi-
tional 77,470 acres were dedicated in 1924,
A Kulkyne National Park was first pro-
posed in 1938 when Mr LG Chandler of
Red Cliffs outlined to the Ornithologist’s
Union and the FNCV the desirability as a
national park of the Kulkyne State Forest
and an area of Crown land to the west. A
committee of the Union investigated the
request and decided that the proposal was
premature because of the then unsatisfacto-
ry administration of national parks in

The Victorian Naturalist

Victoria. The Union recommended instead
that the area should be added to the State
forest and administered as an area for the
conservation of mallee fauna and flora. The
government agreed to the request and
42.000 acres were added to the State Forest
in 1941. At the same time, the whole
Kulkyne State Forest was declared a sanc-
tuary for native game (*Hattah-Kulkyne
National Park” Resource Collection File).

In 1956, after a long campaign by the
FNCV and the Victorian National Parks
Association (VNPA), the Victorian gov-
ernment passed Australia's first legislation
for the control of national parks. The
National Parks Act created a National
Parks Authority, one of the nation's first
statutory bodies to oversee the manage-
ment of national parks.

While the 4cr provided for the manage-
ment of national parks, it did not make
adequate provision for the selection of new
areas which were required in order to give
a broad representation of ecosystems with-
in the State. The VNPA recognised that the
State's network of national parks was far

from being a representative sample of

ecosystems. A survey by the Association
revealed that a considerable portion of the
sixty-two ‘vegetational alliances’ identi-
fied were not protected in existing reserves
(Frankenberg 1971). The Kulkyne State
Forest included important areas of the
*Saltbush, Bluebush’ alliance comprising
low shrubland and low open-shrubland. In
1959 the VNPA submitted an extensive
report to the government requesting the
reservation of the entire forest as a national
park (VNPA Newsletter, October 1959).

Consequently, in 1960 an area of approx-
imately 18,000 hectares, including Lake
Hattah was reserved as Hattah Lakes
National Park. In 1980 the surrounding
State forest which had previously been
held under grazing lease was incorporated
to form the Hattah-Kulkyne National Park
of 48,000 hectares.

Since 1970 the reservation of new national
parks and conservation reserves has been
based on recommendations by the Land
Conservation Council (LCC), now the
Environment Conservation Council. The
LCC was created in the aftermath of the Little
Desert dispute when, in the face of concerted
public opposition, the government abandoned

Vol. 119 (3) 2002

Part One

its plans to open up areas of the Little Desert
for farming. The dispute led to the reservation
of the Little Desert as a national park, but
more importantly it convinced the
Government that a more systematic system of
land-use planning was required to balance the
competing claims for uncommitted Crown
land. The outcome was the Land Conser-
vation Act 1970 which established a Land
Conservation Council to make recommenda-
tions to the Government on the balanced use
of public land in Victoria.

The Little Desert dispute was a watershed
in the history of the conservation movement
in Victoria. It changed forever the method
by which decisions were made about the
future use of Crown land. It also revealed
the extent of public support for conservation,
shaped new attitudes about the environment,
and helped the conservation movement to
refine its campaign techniques.

By 1978, the Land Conservation Council
had completed reports on fourteen investi-
gation areas, although only two of these
reports covered areas adjoining the Murray:
North East 1 (1973) and Mallee (1977),
LCC recommendations led to the reserva-
tion of 20 new parks under the National
Parks Act 1978, increasing the area reserved
under the Act from 283,000 hectares to
774,000 hectares.

Only one park along the Murray was
included under the new legislation. The
Murray Kulkyne Park, comprising 1550
hectares of Murray River lloodplain, was
proclaimed as a result of the 1977 LCC
investigation into the Mallee study area. In
1990 the Park was increased in size to 3530
hectares with the addition of the Liparoo
Block and renamed Murray Kulkyne
Regional Park.

In 1982 Barmah Forest was declared a
wetland of international importance under
the Ramsar Convention, the international
treaty to protect wetlands, and in 1987 the
LCC recommendation of 1985 to create
the Barmah State Park of 7,900 hectares
was finally implemented. The park com-
prised two sections: the first incorporating
wetlands and forests at Top Island, War
Plain and Barmah Lakes; the second cov-
ering a drier forest of River Red Gum,
Grey Box Eucalyptus microcarpa, Yellow
Box Eucalyptus melliodora and Black Box
Eucalyptus largiflorens.

93

Murray River Special Issue

Two of the largest parks along the Murray
River protect vastly different ecosystems.
After decades of argument and debate, a
series of national parks in the alpine area
were reserved by the Victorian government
in 1979, These parks, which included the
Cobberas-Tingaringy National Park along
the headwaters of the Murray, were joined
in 1989 as the Alpine National Park. Two

years later, Murray Sunset National Park of

633,000 hectares was declared in the semi-
arid desert country beyond Mildura. The
park is the second largest national park in
Victoria after the Alpine National Park and
was proclaimed, appropriately, on World
Environment Day.

'The declaration of a new national park on
the international day recognising the
importance of the environment is an indi-
cation of the changing value we place on
the land and the wildlife it sustains. The
Murray River continues to support agricul-
ture and industry, but it is also widely val-
ued for its natural and cultural values. The
river is still beset by environmental prob-
lems, including salinity, erosion, soil acidi-
ty and habitat loss, but in the national
parks and conservation reserves along the
river, nature now comes first.
Acknowledgements
I am indebted to Peter Menkhorst of the Parks
Flora Fauna Division, Department of Natural
Resources and Environment, for correctly iden-
tifying Gray's jerboa Kangaroo as an early name

for what is now the eastern subspecies of the
Burrowing Bettong Bettongia lesueur graii. |
am also grateful to Linden Gillbank of the
History and Philosophy of Science Department,
University of Melbourne, for editorial comment;

References

Bardwell $ (1990) The Major Mitchell Trail: Exploring
Australia Felix, (Department of Conservation, Forests
and Lands: Victoria)

Curr EM (1965) Recollections of Squatting in Victoria,
then culled the Port Phillip District, from 1841 to
1851. (Melbourne University Press; Melbourne)

Davies PS (1978) Man and the Murray,
a Press: Sydney)

Dingle T (1984) The Victorians: Settling. (Fairfax, Syme
& Weldon: Sydney)

Eastburn D (1990) The River Murray: History at a
Glance. (Murray-Darling Basin Commission;
Canberra)

Fahey C (1987) Barmah Forest: a History of the Barmah
Forest. (Department of Conservation, Forests and
Lands: Victoria)

Frankenberg J (1971) Nature Conservation in Victoria: à
Survey. (Victorian National Parks Association)

*Hattah-Kulkyne National Park’ Resource Collection
File F/C: 3/1, Historic Places Section, Department of
Natural Resources and Environment.

Hibbins GM (1991) Barmah Chronicles. (Lynedoch
Publications)

Mackay N and Eastburn D (eds) (1990) The Murray.
(Murray Darling Basin Commission: Canberra)

Mahon G (1992) 4m Appraisal and Review of
Sanctuaries and Wildlife Reserves Systems in Victoria,
(Department of Conservation and Environments
Victoria)

Moulds F (1991) The Dynamic Forest: A History of
Forestry and Forest Industries in Victoria, (Lynedoch:
Richmond)

Priestley S (1984) The Victorians: Making Their Mark
(Fairfax, Syme & Weldon; Sydney)

The Victorian Naturalist, various issues (full citations in
text).

Wright R (1989) The Bureauerat's Domain: Space and
the Public Interest in Victoria, 1836-84. (Oxford
University Press: Oxford)

(NSW

A THOUSAND MILES ON THE

By OW Rosenhain
Read before the Field Naturalists’ Club of Victoria, 16th April 1917

The squatter in the early days set about in a ruthless fashion ringing trees, with-
out let or hindrance—to do this work must have cost many thousands of pounds;
and now you see these dead monarchs of the forest for hundreds of miles on the
river front, and extending back as far as the eyes can reach, one mass of white,
dead timber. In places the fallen timber is so thick that cattle cannot penetrate.

After taking in sufficient fuel, a start was made for Barham Sawmills. This is a
large mill on the New South Wales side, owned by the Barham Sawmilling Co.
On account of the river being very low for many months previously, a fairly large
stock of timber had accumulated, and they estimated there were about 1,000,000
superficial feet of timber stacked there. The manager told me that timber was
becoming very scarce, and it hardly pays now to do the cutting. The nearest trees
that can be felled are about ten miles from the mill.

From The Victorian Naturalist XXXIV (4), August 1917, pp 49-58.

The Murry River in The Victorian Naturalist

RIVER MURRAY

94

The Victorian Naturalist

Part One
Geomorphology of the Murray River Basin

Noel Schleiger'

Abstract

The Murray River is an exotic stream, Since Cretaceous time it has been both a betrunked and, more
recently in the Pliocene, an engrafted river system. Since the Pliocene, the direction of the course of
the Murray has been influenced by upfaulting and jointing of the rocks over which it flows.
Geomorphological features such as the valley-in-valley structure, sharpened spurs, valley scroll
deposits, incised meanders and valleys, bahadas and prior streams are the direct consequences of
uplift, where the river rejuvenates its valley to reach its former base level in its mountain, valley or
plain tract. Features of the plain tract are sinuous meanders, because of low gradients, with levee
banks, ox-bow lakes, lagoons, anabranches, effluents and distributaries. Saline ground water entering
the Murray River has created problems for water management of the Murray. Part of this is due to
irrigation of soils developed on a salty sea bed in late Miocene and in part due to salty aquifers being
cut by the Murray bed. Aeolian features in the semi-arid areas of the Riverine Plain of the Mallee
and Wimmera involve parna deposits, deflated playas, parabolic sand dunes on the Lowan Sand for-
mation and luneties on the eastern shores of drying lakes. These are the product of a more arid cli-

mate in the Pleistocene through to the present day. (The Vietorian Naturalist 119 (3), 2002, 95-101)

The Murray River is an exotic stream
The Murray River is the major river in
Australia. It drains an area of more than
1,020,000 km? and is 2560 km long from
source to sea. When combined with the
Darling River, the tract is 5250 km, the
fourth largest in the world after the Nile,
the Amazon and the Mississippi-Missouri
system (Twidale 1968). The average

runoff per unit area is only 4% of that of

the Danube Basin in Europe, which

emphasizes the arid or semi-arid nature of

the catchment. While the average rainfall
over the whole catchment is 425 mm per
annum, only 5% of the total catchment
exceeds 750 mm per annum. Most of the
high rainfall is over the Kosciuszko and
Victorian alpine plateaus.

The Murray is thus an allogenic or exotic
stream, for it rises in high rainfall areas.
giving it sufficient flow to pass through the
semi-arid desert regions of the Victorian
and South Australian Mallee, where it has
no tributaries, to reach the sea at Goolwa
on the Coorong. Over this tract, water loss
is mainly by evaporation and partly by
downward percolation into the porous
rocks of its bed and banks. Lake Victoria
has an annual evaporation rate of 1500
mm, yet its annual rainfall can vary from
250 to 875 mm.

The Murray Gulf
The Murray River, as we know it today,
developed in the Pliocene Period 2.5 Mya

‘| Astley Street, Montmorency, Victoria 3094

Vol. 119 (3) 2002

(millions of years ago) (Twidale 1968).
When Australia began drifting from
Antarctica 65 Mya (White 1999) much of
Vietoria south of the Great Dividing Range
was successively transgressed by shallow
seas. Also at that time, the Murray Gulf,
occupying the south-west part of the
Murray-Darling Basin, was inundated by the
sea.

Hills (1975) referred to these shallow
transgressions as formed by general down-
warping or epeirogenie movements (from
the Greek epeiros = mainland or conti-
nent). Such movements can be simple
uplift or depression, tilting or broad warp-
ing which affect wide regions rather than
narrow belts (see Fig. 1).

Within the Murray Gulf, marine sediments
began from Middle Palaeocene and contin-
ued through to late Miocene when the sea
retreated in response to the epeirogenic
uplift of the bordering highlands.

^ series of parallel to sub-parallel
stranded coastal ridges successively devel-
oped as the sea retreated over a period of
falling high sea levels.

Stratigraphy of the Murray Gulf

A detailed stratigraphy of the Murray
Basin is described by Lawrence and Abele
(1988). Figs 2 and 3 summarise the struc-
ture and relative ages of the sedimentary
groups and members. The Australian depo-
sitional stages and the foraminiferal zones
which date them are indicated in the right-
hand column of Fig. 3.

95

Murray River Special Issue

T =e.

L. New South Wales

. Loxton

Mildura —-

i Adelaide =
^ * v4 SON — Murray Basin

De
ul
E EM
m
Goolwa Ser-]— M

Riverine Plain

Marine Sedimentation
Padiloisvay Horst

Great Dividing Range

PETEN

LB 3 -* y *
v.
dtp s
—— ta
Last
» RS ga
"S *"
a z e
aS "a
- bd y
3 t .
Tr%
"diea iai] NN
Manangatang «7 1 ies
. * Riverine Plain re
e Mr HN. —
.
— e TOM, oe
ù M " '
x" Kerang- >g Pee
ae

0 100

200 kms

|

Fig. 1. The Murray Gulf from Palaeocene to Miocene time. The rivers enter the Gulf via separate

mouths.

The local changes of lithology (facies
changes) across the Murray Basin from
South Australia and Victoria make it diffi-
cult to give decisive orders of age for some
units. The facies changes from west to east
are the consequence of the gradual shal-
lowing of the Gulf.

In general, the Renmark group is the old-
est, ranging in age from mid-Palaeocene to
mid-Miocene. The Ettrick Marl, and the
Geera Clay to the east and on the same
stratigraphic level, overlie the Renmark
group. The lower part of the Geera Clay is
the lateral equivalent of the Ettrick Marl.
The Duddo Limestone overlies the Ettrick
Marl and merges into the upper part of the
Geera Clay. The Winnambool Formation
overlies both the Geera Clay and the
Ettrick Marl. Finally, in late Miocene, the
Bookpurnong Beds marked the end of
marine sedimentation in the Murray Basin.

In the Pliocene, terrestrial deposition then

96

followed. Two units highlight this environ-
ment: the Torrumbarry Clay and the
Calivil Sand.

The Torrumbarry Clay

Lawrence (1975) described this member
in the Echuca North No 1 Bore (Fig. 2)
from 58 to 77 m. It consists of white
kaolinitic clay up to 30 m thick. It overlies
the Calivil Sand and is overlain by the
Quaternary Shepparton Formation. It was
deposited in lakes and on the alluvial
plains.

The Calivil Sand

Macumber (1972) described this member
in the Calivil No 2 Bore between 86 m and
105 m. It overlies unconformably the
Palaeozoie bedrock or disconformably on
the Renmark Group. The Calivil Sand is of
grey, poorly sorted quartz sand and gravel
with a white kaolinite matrix, Quite often
the member is auriferous and may contain

The Victorian Naturalist

Part One

a
2004 BookpumongBeds P
| Winnamboo! Formation

Pre- Tena B —
ll

Fig. 2. The structure of the Murray Basin, following a west to east cross section from the South
Australia border to Echuca, with eight key boreholes supplying the stratigraphy (from Lawrence 1975).

-200

ifs sagnir Fault

7,

diatoms and plant impressions, and forms
buried fan deltas up to 45 m thick. For
more detail see Bowler and Macumber
(1968) and Lawrence and Abele (1988).

Features of the Murray River

The mountain, valley and plain tracts of

the Murray River exhibit a variety of geo-
morphic features which are discussed below.

Betrunked and engrafted river systems

Fig. 1 shows the extent of the Murray
Gulf in relation to present day streams.
Prior to the incursion of the Murray Gulf,
the ancestors of the present day streams
would have entered the sea, depending on
their position then, as one river system as
at present.

Once the Murray Gulf was established,
these streams would have debouched into it
by separate mouths, e.g. Fig. 1 shows the
Murrumbidgee and Darling as a betrunked
river system.

In the Pliocene, when the sea retreated,
the Murray and Murrumbidgee joined as
one. At Wentworth, the Darling joined the
Murray to complete the system, and then
flowed across the former sea bed to be
deflected by the elevated Flinders-Lofty
block to reach the sea at Goolwa, South
Australia on the Coorong. This stream sys-
tem is an engrafted river system for the
Murray and its tributaries engrafied a new
course over the former sea bed, the slope
of which must have been to the north-west,
then westerly and finally southerly.

Twidale (1968) shows the Murray River
deflected by the Morgan fault. The Murray
was defeated at Morgan, deflecting it in a
southerly direction to Goolwa (Fig. 4a).

Vol. 119 (3) 2002

Defeated stream
The Cadell Tilt Block

The diversion of the Murray River north-
east of Echuca is an example of a defeated
stream. Harris (1938) describes ‘the great
bend of the Murray River makes to the
south when about 48 km NE of Echuca'.
His fig. 1 shows the Cadell Tilt Block
uplifted along the Cadell Fault which runs
north-north-east along the Colbinabbin
Range of Cambrian greenstones to
Deniliquin.

The now dry segment of Green Gully
which crosses the westerly-tilted uplifted-
block from Mathoura is known as a wind
gap. The Moira Marshes at the base of the
Cadell Tilt Block, with its outerop of allu-
vium, marks the ponding of the Murray

just west of its diversion point (Hills 1975;

Bowler and Macumber 1968).

The North West Bend at Morgan, SA

There is much debate among geologists
about whether the southerly deflection of
the Murray at Morgan is the result of
defeat faulting on the eastern margin of the
Lofty Ranges. Twidale (1968) states ‘the
Murray makes a sharp turn southward in
response to a series of low north to south
fault escarpments located within the
Murray plain’ between Morgan and
Blanchetown which appears to be a com-
plex but well marked fault zone.

Williams and Goode (1978) proposed
that before uplift of the Lofty and Flinders
Ranges an ancient Murray River extended
across the line of these ranges to a major
delta in the Spencer Gulf. Brown and
Stephenson (1991) have made a compre-
hensive study of the Murray Basin and

97

Murray River Special Issue

Age Epoch Series Stratigraphic units
(Mya) T.
18 |Pliocene | Upper Parilla Sand
Lower ~
z Bookpurnong Beds
2
Upper
10.5
15 Miocene
Middle
Winnambool |
Formation |
d
Lower
23.5 Ü
er
Oligocene ir
32 Lower
37.5 Upper Renmark
3 Eocene
43 Middle Group
ds Lower
53.5
Upper
56 Paleocene
Middle
62
Lower
65

Shepparton Fm

Foram Australian stages Key Foraminifera
zones

14 Kalimnan

Cheltenhamian

Torrumbarry

1 Mitchellian
Clay 3
12
11 Bairnsdalian Orbiculina universa
10 O.suturalis
9
Balcombian Praeorbiculina glomerosa

cunia

Batesfordian ^ Globigerinoides sicunus

Longfordian G. trilobus

5
3 Globoquadrina dehiscens
1
2

Janjukian
TW Globigerina euapertura
Guembolitria stowensis

Aldingan

Wangerippian

Fig, 3. Geological age-correlation chart of the stratigraphic units of the Murray Basin.

argue that this event was unlikely in the
Cainozoic era. The evidence of borelogs
suggests that the precursor to the modern
Murray emptied into the Buccleuch
Embayment at least from the Eocene.
Therefore, if Williams and Goode (1978)
are correct, the trans-Lofty tract would be
in the Cretaceous or earlier. For more
detail on this debate see Brown and
Stephenson (1991). These authors argue
that in the late Miocene a marine transgres-
sion which persisted into the early
Pliocene to the mouth of the Murray was
located near the geographic centre of the
Murray Basin. This mouth migrated west-
erly along the course of the present Murray
River as the sea began retreating from the
basin. About 2-5 Mya tectonism in the
western Murray Basin led to the formation
of Lake Bungunnia which ponded the
Murray at Morgan (Fig. 4c). About 0.7
Mya the lake waters retreated and the
Murray proceeded along a southerly down-
stream course from Morgan following a
glacial sea level drop.

Brown and Stephenson (1991) stress that
the western mouth hypothesis did not occur
for the ancient Murray at any time within

98

the Cainozoic. So whether the Murray
River at Morgan is a defeated stream is a
paradox, as it has all the features of one.

Anastomosing streams

There are streams along the Murray in its
riverine plain which form a network of
widely spaced streams, large enough and
long enough to rank a name. These streams
have diverged from the Murray to join it
further downstream, e.g. Gunbower Creek
and Barr Creek. Such streams are known
as anabranches.

Another type of diverging channel may
terminate in a shallow lake or swamp. This
is known as an effluent, e.g. Chalka Creek in
the Hattah-Kulkyne National Park termi-
nates in part at Lake Lockie one of the
Hattah Lakes. In North West Victoria near
the border with NSW and SA, Lake Victoria
terminates Rufus River, which leaves the
Murray and flows northerly. Toole Creek
actually leaves the Murray River in NSW
and joins the Wakool River in the Riverina.
This is a distributary stream.

These anastomosing streams have levee
banks built up on either side of the stream
at flood time. During flood time, erosion

The Victorian Naturalist

through a levee bank can allow the distrib-
utary to take an entirely new course as the
channels are split and shifted. This allows
a large area to be covered very thickly with
alluvium, as occurs between Kow Swamp
and Lake Charm.

Valley-in-valley structure

Between Mannum and Morgan in South
Australia, the Murray Valley shows valley-
in-valley structure, which suggests there
are two base levels due either to
epeirogenic uplift of the area, or in
response to a drop in sea level. The first
base level was a valley cut in the mature
landscape before uplift or sea level lower-
ing. The second base level is the stream
today, a valley incised in the former valley.
The first base level was cut in Pliocene
times, for Twidale (1968) reports small
outcrops of cross-bedded sandstones 6-7.5
m thick scattered over the cliff tops. There
is an oyster bed at the base of the cross-
bedded sandstone of Pliocene age. These
Pliocene sediments have not been formed
outside the confines of the wider valley,
and were laid down in a narrow estuary,
far inland up what is now the Lower
Murray Valley.

Influence of bedrock jointing

Geomorphologically, joints are important
as they form lines of weakness which can
be exploited by weathering agents, espe-
cially water and wind. The pattern and
spacing of the joint pattern (i.e. the texture
of the joint system) are imposed on the
pattern of relief. The geometry and spacing
of joints vary within the same rock and
certainly from one rock to another. The
Miocene calcarenite (sandy limestone)
forming the Murray cliffs markedly show
joint blocks exposed in the cliffs, This
joint pattern has been picked out by the
Murray in SA (Fig. 4a, b).

Sharpened spurs and valley scrolls

The Murray reaches maturity when the
amplitude of its meander belt is equal to
the width of its valley floor. The Murray
has incised a valley through the hills,
where it is still cutting laterally into its
banks to form accentuated or sharpened
spurs, then swings away to the other side
and laterally corrades hills on the other
side. In so doing it deposits valley plain

Vol. 119 (3) 2002

Part One

Ancestral Darling River

-5 7777408

Ancestral Lachlan River

Lake Hii

marsh
(€) ; d
Fig. 4. (a) The Murray River deflected southerly
by the Morgan Fault (after Firman 1966). (b)
Directional control of the Murray tract by joint-
ing in the Miocene sediments (enlargement of
the frame in a) (after O'Driscoll 1960). (c)
Palaeodrainage of Murray Basin in late Pliocene
to early Pleistocene (2.5-0.7 Mya). The Murray
system engrafted partly across the old sea floor
to enter Lake Bungunnia. After 0.7 Mya an exit
drainage line developed west of the Pinnaroo
Block to drain the lake into the Southern Ocean
and hence the modern Murray tract (after
Brown and Stephenson 1991).

alluvium between the two zones of lateral
corrasion marked by the sharpened spurs.
These zones of alluvial deposition are
known as valley scroll deposits (Hills
1975).

Alluvial fans or alluvial cones

The Murray shows valley-in-valley struc-
ture west of Wodonga. Tributaries entering
the upper terraces of the Murray drop their
sediment on approaching the Murray in the
form of a flat cone. If several tributaries
build alluvial cones side-by-side, an allu-
vial apron will form sub-parallel to the
upper edge of the terrace. This is known as
a bahada (Holmes 1969). If uplift contin-
ues or the Murray corrades its bed vertical-

99

Murray River Special Issue

ly lower, the tributary incises the flood
plain and may even build another alluvial
cone at a lower level (Hills 1975).

Parna deposits

Parna is a wind blown silt deposited in
many parts of southern Australia by rain
after dust storms, especially in the
Pleistocene when the climate was more
arid. Wide areas of the Murray Basin are
covered at the surface with a layer of clay-

loam called parna. It is like the loess of

China but of a pallid grey colour around
Melbourne. Areas of consistent loss of dust
and sand are said to be deflated. In the
northern plains are small barren swales, or
‘scalded’ areas which are believed to be
formed by the deflation of salt-affected
soils during dust storms.

Mostly sand is moved by saltation with
jumps at about 25 to 30 em above the sur-
face. Strong gale-force winds carry sand
along the ground, forming aeolian sand rip-
ples with wave lengths of 9 to 12 cm. Such
sand can blast and polish heavier pebbles
and wear flat, sloping surfaces on them.
These pebbles are known as ventifaets.

Lakes and lunettes

Hills (1940) was the first to document
lunettes. These are crescent-shaped ridges
on the eastern shores of lakes. They were
first described by Major Thomas Mitchell
from Green Lake and Dock Lake near the
Grampians in Western Victoria. They are
widespread in Australia south of latitude
32S, and there are even some in Tasmania.
They differ from sand dunes in consisting
wholly of clay-loam on the eastern shores
of lakes in semi-arid areas. The summit is
broadly rounded. The highest part is closer
to the lake shore than the outer margin
(horns). They curve parallel to the smooth
curve of the lake shore on the eastern side.
They are due to beach erosion by westerlies
when the lake level is seasonally low, In
cross section, lunettes are ty yee asym-
metrical, with the steeper slope Í facing the
lake. The bedding in lunettes is generally
parallel to the surface apart from where it is
truncated on the lake-facing side.

Lunettes are built of granular aggregates
of clay and silt on the lake floor when the
water dries up, leaving salts accumulated
on the lake bed as pellets of clayey materi-
al. Clay pellets driven by the prevailing

100

west and south-west winds mantle over the
accumulating dune. The majority of
Victorian lunettes, including many exam-
ples between Kerang and Swan Hill, are
believed to have formed between 15,000
and 17,500 ya (Bowler and Macumber
1968). Around Boort, the younger ages of
lunettes suggest that the special conditions
for lunette formation did not occur there
until later. In some lakes, younger lunettes
formed in sequence as the lake shore
shrank back, or several smaller lakes
formed within the one larger basin.

Salinity in the Murray River

Lakes consistently fed by streams, yet
inadequate to cause the lake to overflow,
accumulate all salts brought in so that their
waters become highly saline. When these
lakes dry out they become clay pans or
playas. If they are salty, yielding crystals
of halite (sodium chloride) and gypsum
(hydrated calcium sulphate), then they are
salinas, e.g. Lake Tyrell near Sea Lake on
the floor of the Murray Basin.

In the Wentworth-Mildura area of the
Murray and in South Australia between
Renmark and Morgan, groundwater from
springs seeps into the bed of the river.
Below normal river level the springs are
not visible. Some of these waters are saline.
Attempts are being made in the Sunraysia
area to intersect these saline aquifers and
stop them from entering the Murray. The
saline water is pumped into playas, where it
is evaporated and the salt is harvested. This
has minimized saline water entering the
Murray in the Mildura area.

Summary

The geomorphic features described in the
Murray Basin are of two types: tectonic
structures, which have influenced the
direction of the Murray tract; and semi-
arid features involving aeolian and fluvi-
atile processes due to the exotic nature of
the river system.

Tectonic structures

Both epeirogenic movements and local-
ized faulting have been active over the his-
tory of the Murray Basin. Epeirogenic
movements are responsible for the
betrunked and engrafted river system at the
commencement and completion respec-
tively of the Murray Gulf transgression.

The Victorian Naturalist

Faulting of uplifted blocks such as Cadell
and the Morgan faults has been effective in
diverting the course of the Murray. Joint
directions have been responsible for the
zig-zag course of the Murray in South
Australia. In its mountain and valley tract
upstream from Wodonga, valley-in-valley
structure has produced river terraces in
response to either lowering of sea level or
epeirogenic uplift of broad areas. In such a
situation, fault aprons and bahadas can
result from higher bankful discharge and
runoff. Incised streams, valley scrolls, and
sharpened spurs are common in uplifted,
rejuvenated areas of north-east Victoria
and South Australia.

Semi-arid features
Fluviatile features

The lack of tributaries joining the Murray
is a feature of the Mallee and Wimmera.
The Loddon and the Avoca Rivers fail to
reach the Murray. Due to the gentle slopes,
levees and sinuous meandering with ox-
bow lakes, lagoons and cut off meanders
are numerous. Anabranches, effluents and
distributaries are common between Echuca
and Renmark (Fig. 5).

Aeolian features

Parna from the precipitation of rain fol-
lowing dust storms, parabolic dunes on the
Quaternary Lowan Sands, as well as
lunettes on the eastern margins of lakes,
and swamps testify to the dominance of
wind in the dry summers from the
Pleistocene through to modern times.

Irrigation and movement of saline ground
water into the Murray have increased the
salinity of the Murray since European set-
tlement. The problem is complex and cost-
ly to solve. On the positive side, the
Murray Basin could be important for the
harvesting of earth resources in the 21"
century. Heavy minerals, e.g. magnetite
and ilmenite, are in evidence from
prospecting projects. As well, granite and
good quality gravel, lime, sand and clay
deposits may lay the basis for building and
engineering projects.

Acknowledgements

The author appreciated the help of Dr Ken Bell
with aspects of the manuscript. Roger Pierson of
Deakin University helped with key references.
Dorothy Mahler helped with typing and layout
of manuscript.

Vol. 119 (3) 2002

Part One

Gunbower Creek

Kow Swamp

Fig. 5. Anabranches, effluents and distributaries
in the plain tract of the Murray.

References

Bowler JM and Macumber PG (1968) The Riverine
Plain in Northern Victoria. In A Regional Guide to
Victorian Geology Excursion No 13, pp 133-144. Eds
J McAndrew and MAH Marsden.

Brown CM and Stephenson AF (1991) Geology of the
Murray Basin Southeastern Australia, BMR Bulletin
235. Bureau of Mineral Resources, Canberra,

Firman JB (1966) Stratigraphy of the Chowilla Area in
the Murray Basin: Quarterly Geological Notes.
Geological Survey of South Australia 28, 1-47.

Harris WJ (1938) The Physiography of the Echuca
District. Proceedings of the Royal Society of Victoria
51, 45-60.

Hills ES (1940) The lunette: a new form of aeolian ori-
gin. Australian Geographer 3, 15-21.

Hills ES (1975) Physiography of Victoria ~ An
Introduction to Geomorphology. (Whitcombe and
Tombs: Australia)

Holmes A (1969) Principles of Physical Geology.
(Thomas Nelson & Sons Ltd: London)

Lawrence CR and Abele C (1988) The Murray Basin.
In Geology of Victoria, Eds JG Douglas and JA
Ferguson. (Victorian Division Geological Society of
Australia Ine: Melbourne)

Lawrence CR (1975) Geology, Hydrodynamics and
Hydrochemistry of the Southern Murray Basin.
Memoir 30, Mines Department, Victoria, Australia,

Macumber PG (1972). Progress report on the
Groundwater Survey of the Avoca and Loddon
Valleys. Progress Report of the Geological Survey of
Victoria.

McAndrew J and Marsden MAH (1968) 4 Regional
Guide to Victorian Geology. Geology Department,
University of Melbourne, Melbourne.

O'Driscoll EPD (1960) The Hydrology of the Murray
Basin Province in South Australia. Bulletin of the
Geological Survey of South Australia 35.

Parkin LW (1969) Handbook of South Australian
Geology. (Geological Survey of South Australia:
Adelaide)

Stephenson AE and Brown CM (1989) The ancient
Murray River System. BMR Journal of Geology and
Geophysics 11, 387-395.

Twidale CR (1968) Geomorphology with Special
Reference to Australia. (Nelson: Melbourne)

White ME (1999) Reading the Rocks. (Kangaroo Press:
Australia)

Williams GE and Goode ADT (1978) Possible western
outlet for an ancient Murray River in South
Australia. Search 9, 442-447.

101

Murray River Special Issue

Floodplain Wetlands: the Jewels of the Murray River

Rhonda Butcher! and Michael Reid’

Abstract

Wetlands and billabongs of the Murray River floodplain are intimately connected with the main-
stream, and whilst this may imply a degree of similarity, work over the past 20 years has shown that
this is not necessarily the case. While billabongs are formed by the river, the physical and chemical
gradients of the river and floodplain wetlands are quite different and thus the biological communities
are quite different. This paper briefly describes some of the ecological differences between the main-
stream and the floodplain wetlands touching on the importance of connectivity, disturbance, and the
variability of the floodplain communities. (The Victorian Naturalist 119 (3), 2002, 102-107)

Introduction

Seasonally or intermittently flooded wet-
lands and billabongs are common in flood-
plain ecosystems and are important func-
tional components of the Murray River
(Fig. 1; Hillman 1986). Over 7000 wet-
lands covering an area greater than
200,000 ha have been mapped along the
length of the Murray River downstream of
Hume dam, with half of that area constitut-
ing floodplain wetlands (Pressey 1990).
Billabongs are probably one of the best-
known aquatic habitats in Australia, and
are a major feature of the 300 kilometre
long, confined floodplain from the Hume
dam to Tocumwal. Downstream of
Tocumwal tectonic activity, principally
uplift of an area known as the Cadell block
around 60,000 years ago (Page et al. 1996)
has contributed to the formation of three
major alluvial fans: the Barmah, Wakool
and Gunbower. Frequent flooding of these
low-lying fans has led to the development
of extensive floodplain wetland forest
(Young and Hillman 2001 ).

In 1986 Terry Hillman published a book
chapter simply entitled *Billabongs" in
which he highlighted the diversity and pro-
ductivity of billabong communities and
demonstrated their distinctiveness from
those of the mainstream Murray River. He
lamented our lack of understanding of
these systems, and whilst our understand-
ing of the ecology of billabongs and other
floodplain wetlands remains limited, it has
improved. Billabongs have a unique place
in Australian folklore, and they, along with
other floodplain wetlands, have been the

‘School of Biological Sciences, PO Box 18, Monash
University, Victoria 3800

* National Institute of Water and Atmospheric
Research, PO Box 8602, Christchurch, New Zealand

102

focus of considerable scientific interest
and research in the past 15 years.
Billabongs are floodplain wetlands that
are formed by the geomorphic action of
the stream, Most billabongs are old sec-
tions of the main channel formed by mean-
der cutoff or channel avulsion. Some may
even act as anabranches during periods of
high flow. Cut-off meanders can be quite
deep thus forming unique deep-water bill-
abongs, which can retain water for much
longer than most floodplain wetlands.
Other smaller billabongs may be formed in
floodplain depressions such as scroll
swales or when smaller tributaries dry out
to a series of pools. Superimposed upon
this variety of form is that created by the
process of sediment deposition, which
leads to a mosaic of billabongs of various
shapes and depths across the floodplain.
The process of infilling appears, in many
cases, to be very slow, meaning that indi-
vidual billabongs can be very old.
Radiocarbon dating of billabongs on the
Murray River suggests that they have been
in existence for up to 7000 years (Ogden ef
al. 2001). Even more impressive, dating of
two billabongs on the Goulburn River
indicates they were cut off from the main-
stream over 12,000 years ago — these bill-
abongs remain more than three metres
deep (Ogden et al. 2001; Reid in press).
Floodplain wetlands have been variously
categorised according to a number of sys-
tems, but mainly reflect the water regime.
They can have water for varying lengths of
time, and are generally characterised as
being shallow, less than 2 m deep. The ele-
vation of the floodplain is generally only a
few metres and the lower areas will be
prone to more frequent flooding events.

The Victorian Naturalist

Issues of connectivity

One of the features of natural floodplain
ecosystems is the degree to which water
and other materials move between the
mainstream, floodplain wetlands and the
terrestrial floodplain. Since European set-
tlement and regulation of the Murray River
there has been a physical and biotic separa-
tion of the components of the floodplain,
mainly through alteration of the flooding
regime. The connectivity of the system has
been disrupted.

Four dimensions of connectivity are
apparent in a river system as ít flows from
its headwaters to the sea. The first is the
longitudinal or upstream-downstream
dimension. The second is the lateral move-
ment of water from the river channel into
the terrestrial environment, such as occurs
in floodplain areas. The third is the subsur-
face movement of water through porous
material in the streambed both laterally
and vertically into the substrata. The final
dimension is temporal, reflecting the
change in the physical dimensions of the
various components of the river-floodplain
system through the seasons and with the
advent of floods and droughts.

Most stream biota have distinct prefer-
ences for specific conditions and will live,
or be most abundant, where they have the
greatest chance of surviving and reproduc-
ing. For example, some invertebrates are
adapted to living in the cool, highly oxy-
genated riffle areas of uplands streams.
Murray Cod prefer the warmer lowland
reaches. They have been shown to like par-
ticular snags and are quite content to wan-
der over the floodplain when there is a
flood. Leeches prefer the still waters of
floodplain wetlands (and farm dams in the
absence of natural wetlands), and River
Red Gums require certain periods of inun-
dation in order to germinate, Some biota
will occur in different areas of a floodplain
system at different stages of their life histo-
ry, a feature that highlights the importance
of connectivity across floodplain systems.

From the headwaters of a river to the
river mouth a number of physical and
chemical features change. Channel gradi-
ent, mean particle size of the substrate and
shading by riparian vegetation all decrease
as you move downstream. On the other
hand, mean flow and discharge, channel

Vol. 119 (3) 2002

Part One

Fig. 1. River Red Gum forest in flood at
Barmah. Photo by Lindy Lumsden.

width and sinuosity all inerease. In the
water column, downstream decreases
occur in dissolved oxygen concentration,
the amount of coarse particulate organic
matter and the degree of variation in water
temperature, whilst turbidity, salinity, dis-
solved organic matter, nutrient concentra-
tions and mean water temperature each
tend to increase (Boulton and Brock 1999).
The overriding influence is the directional
flow of the water.

lhe physical and chemical environment of
floodplain wetlands and billabongs is deter-
mined by the water regime: when they have
water (summer, winter, spring), when it
arrived, how long the water stayed and how
much water is present. The timing and dura-
tion of the flood pulse is a key factor in dri-
ving wetland biota life cycles (Reid and
Brooks 2000; Young 2001). Wetlands and
billabongs are relatively closed systems and
lack the longitudinal gradient seen in rivers.
Events which affect the physical and chemi-
cal environments of these systems can, and
do, occur at different times and rates (e.g.
flooding/drying) even in quite geographical-
ly close systems (Boon et al. 1990; Boulton
and Brock 1999), Water chemistry has been
shown to reflect a cyclic pattern closely
attuned to the hydrological regime. Thus,
flooding leads to dilution of wetland waters.
This is followed by gradual concentration,
which becomes extreme in periods of
drought. Associated with these changes are
biotic patterns that also affect the nutrient
cycling patterns. Considering this, billabong
and floodplain wetlands are highly variable,
rich and, at times, extreme environments
(Hillman 1986; Boon et al. 1990; Reid and
Brooks 2000).

103

Murray River Special Issue

Biological processes — primary produc-
tion and the microbial loop

When water arrives it triggers the release
of a pulse of nutrients from the sediments.
At the same time plants and algae germinate
Irom the seed bank in the sediment of the
wetland and begin to grow. The nutrients
provide food for the bacteria, which then
begin to break down the terrestrial organic
material that has been inundated. The
microbial loop is a very important compo-
nent of wetland function. It was once
thought that bacterias main role in ecosys-
tems was as decomposers of dead and dying
matter, thus making the nutrients available
to other consumers. However, research has
shown that microbes are actually a food
source in themselves and form part of the
food web, thus the term microbial loop has
become popular. Biota which feed on bacte-
ria include protozoans, such as small flagel-
lates and ciliates, and rotifers which are in
turn eaten by bigger invertebrates, and so on
(Boulton and Brock 1999).

The main groups of primary producers
include both micro and macroscopic
forms. The macroscopic primary producers
are the water plants, which include both
submerged and emergent forms that grow
in shallow areas. The distribution of aquat-
ic plants is strongly determined by water
depth and clarity because they are limited
by light availability. The microscopic pri-
mary producers include two major groups,
the periphyton, which grow attached to the
sediments and other submerged surfaces
(e.g. woody debris, plants) and the phyto-
plankton, which occur in the water col-
umn. In the Murray River and the associat-
ed deep billabongs today. the bulk of pri-
mary productivity appears to be carried out
by phytoplankton (Ben Gawne pers.
comm.; Reid ef al. in press), However,
there is evidence from the sedimentary
records of several Murray River billabongs
that prior to the arrival of Europeans aquat-
ic plants and periphvton were much more
important primary producers (Ogden 2000;
Reid er al. in press).

Habitat zones within wetlands and asso-
ciated invertebrate assemblages
Generally wetlands can have up to four
readily identified habitats or microhabi-
tats: the water surface, the open water

104

zone, the profundal zone and the littoral
zone,

The littoral or edge zone is characterised
by relatively coarse sediments and a com-
plex habitat structure. The complex struc-
ture arises because large plants can grow in
this zone as it usually equates to the
euphotic zone, which is defined as the
depth to which light penetrates in the water
column to sustain/support photosynthesis.
An abundance of woody debris often con-
tributes further to the habitat complexity of
this zone. The littoral zone is generally the
most biologically diverse region within a
wetland. For this reason, along with more
practical reasons related to access, this
zone is often the focus of biological study.

The region of the wetland that lies beyond
the euphotic zone is known as the profun-
dal zone. There is not sufficient light at this
depth to support photosynthesis so no
plants grow here. Bacteria, detrivores and
the animals that feed on them predominate
within this zone. The sediments are gener-
ally finer and there is less structural diversi-
ty than in littoral areas. The separation
between littoral and profundal zones is
highly variable, and in many shallow clear
water wetlands a true profundal zone may
not exist (Boulton and Brock 1999),

The air-water interface or the neuston of
the water surface supports a number of spe-
cialised biota, Virtually all of the biota
adapted to the neuston are predators, and
come mainly from the Hemiptera (bugs) and
the Coleoptera (beetles). Some biota are tem-
porary members of the neuston as they are
air breathers. These include the culicid mos-
quito larvae, diving beetles, and other bugs.
Because these animals are air breathers it is
believed they are relatively tolerant of poorer
water quality, especially low oxygen levels
(Boulton and Brock 1999),

The open water column is often referred
to as the pelagic or limnetic zone.
Invertebrates found in the open water zone
can be divided into the nekton, generally
macroscopic organisms that are strong,
active swimmers, and the zooplankton,
smaller organisms that are generally weak-
er swimmers or that passively drift or float
within the water column. Many zooplank-
ton such as rotifers, water fleas and cope-
pods are grazers, and can be found in large
numbers in the water column (Boulton and

The Victorian Naturalist

Brock 1999). The nekton includes the fish
and the larger invertebrates, such as many
bug and beetle species, that feed on the
zooplankton or on each other.

The micro-invertebrates of floodplain
wetlands and billabongs are particularly
well known due to the efforts of Russ Shiel
and his colleagues. A considerable body of
taxonomic and ecological work has shown
that, not only are there large numbers of
species present in these habitats, but that
many which were thought to belong to cos-
mopolitan species are actually endemic to
Australia. The most diverse billabong
assemblages are not planktonic, but are
epibenthic and epiphytic in the littoral zone
(Hillman and Shiel 1991; Green and Shiel
1992: Shiel 1995; Shiel er al. 1998).

How changes in the water regime affect
biota and the broader functioning of the
floodplain ecosystem are questions on
which considerable work has focused. A
number of studies suggest that wetland
invertebrate communities are affected by
periodicity, seasonality and duration of
flooding (Boulton and Lloyd 1992; Nielsen
et al. 1999; Quinn ef al. 2000). However,
the mechanisms driving the observed pat-
terns are likely to be complex. For exam-
ple, a recent study by Quinn ef al. (2000)
compared the changes in invertebrate com-
munities of temporary and permanent wet-
lands on the Murray (regulated) and Ovens
(unregulated) floodplains over time fol-
lowing flooding. They found that distinc-
tions between permanent and temporary
wetlands were clear and persistent for
Murray wetlands but that no distinction
could be made between temporary and per-
manent wetlands on the Ovens floodplain,
even for the samples taken just one month
after flooding. This suggests that some of
the mechanisms linking biota and flooding
regimes operate at large spatial scales.

One area of great interest is how changing
flooding regimes may affect the emergence
(hatching) of resting stages of invertebrates
and plants from floodplain sediments fol-
lowing inundation. The low and unpre-
dictable rainfall that characterises much of
the Murray-Darling basin means that aquat-
ic organisms must be able to persist through
long dry periods. Many do so by producing
desiccation-resistant eggs or seeds that can
lie dormant in dry soil or sediment for many

Vol. 119 (3) 2002

Part One

years. Studies in Australia (Boulton and
Lloyd 1992; Jenkins and Briggs 1997) and
overseas (Hairston and De Stasio 1988; De
Stasio 1989; Hairston et al. 1995) show that
eggs estimated to be up to 300 years old can
hatch under suitable conditions. The long
lived nature of the egg and seedbanks pro-
vide not only the refuges from extended
periods of drying or drought but also a
means of maintaining genetic, phenotypic,
species and community diversity (Ellner
and Hairston 1994; Hairston er al, 1996;
Leck and Brock 2000). However, despite
this apparent resilience, several experimen-
tal studies have shown that spatial variation
in the number and variety of invertebrates
emerging from flooded sediments can be
partly attributed to variation in the flooding
history of those sediments (Boulton and
Lloyd 1992; Boulton and Jenkins 1998;
Nielsen er al. 2000). Thus, changes in
flooding regime associated with river regu-
lation appear to have the potential to affect
the viability of the egg bank and hence the
abundance and diversity of emergent com-
munities.

Concluding remarks

One of the main controlling elements in
structuring aquatic communities is distur-
bance. In Australia we have high natural
disturbance with frequent floods and
droughts, which are believed to be one of
the major determinants of aquatic commu-
nity structure. Human disturbance com-
pounds environmental change and is super-
imposed upon natural variability. Virtually
all of our rivers and streams have had their
[low regime altered in some manner, usual-
ly by diversions or dams. Altered volumes
and seasonality of flows radically change
the natural settings of rivers and streams
and the delivery of water to the floodplain
and its associated wetlands. These changes
have, in turn, been implicated in the loss of
native species and an increase in invasive
exoties that are apparently better suited to
the new conditions (Stanford 1996).

The cumulative effect of these distur-
bances is the decoupling of the complex
interactive pathways that are characteristic
of the four dimensions of connectivity along
rivers in large basins (Stanford 1996).

River regulation is just one example. It
decouples the longitudinal and lateral

105

Murray River Special Issue

dimensions of rivers. In the Murray, for
example, we have whole sections of river
that have been converted from flowing
lotic environments to a series of lentic weir
pools. The weir pools act as a migration
barrier for fish and other animals, they act
as flood and nutrient sinks and they can
stimulate biophysical consistency in down-
stream sections of the river. A difficulty
associated with disturbance is identifying
causality. Particularly in lowland and mid-
land reaches of rivers, attributing alter-
ations in aquatic community structure to
any one disturbance is quite difficult.
Increased salinity, increased turbidity and
sedimentation, alterations to flow regimes
und water quality are all things that tend to
happen together, so separating out one cul-
prit can be quite a challenge at times.

Billabong and wetland environments are
extremely variable and this heterogeneity is
also reflected in the biotic community, Even
when the floodplain is inundated and would
appear on the surface to be a homogeneous
single wetland environment, environmental
patchiness still exists as different biota
respond differently to the flood event. This
variability makes the life of a natural
resource manager quite difficult. For exam-
ple if a manager is trying to assess the impact
of an environmental water allocation to a
floodplain system, the scale at which vari-
ables are measured and recorded will have a
great influence on what information can be
gained. Biota operate and respond at differ-
ent scales and rates: this spatial and temporal
variability is a critical feature of all wetland
environments and needs to be carefully con-
sidered when undertaking any ecological
assessment of a wetland (Weins 1989;
Schneider 1994; Butcher ef al. in prep).

Floodplain wetlands and billabongs form
the aquatic component of the floodplain
and have distinet differences in the physi-
cal and chemical gradients and in the bio-
logical processes and communities com-
pared to the mainstream. It is easy to think
of them as separate entities, but all three
major components (terrestrial floodplain,
aquatic floodplain habitat and the main-
stream) of floodplain river systems such as
the Murray are interdependent, They are
highly complex and variable, fascinating
environments: jewels that we should learn
to appreciate and treasure.

Acknowledgements

We would like to thank Drs Russ Shiel and
Daryl Nielsen for information and comments on
various sections of this paper,

References

Boon PL Frankenburg J, Hillman TJ, Oliver RL and
Shiel RL (1990) Billabongs. In The Murray, pp 183-
200. Rds N MacKay and D Eastburn, (Murray
Darling Basin Commission: Canberra)

Boulton AJ and Lloyd LN (1992) Flooding frequency
and invertebrate emergence from dry floodplain sedi-
ments of the River Murray, Australia. Regulared
Rivers: Research and Management 7, 137-151.

Boulton AJ and Jenkins KM (1998) Flood regimes and
invertebrate communities in floodplain wetlands, [n
Wetlands in a dryland: Understanding for manage-
ment, pp 137-146. Ed WD Williams (Environment
Australia, Biodiversity Group: Canberra)

Boulton AJ and Brock MA (1999) Australian
Freshwater Ecology: Process and Management.
(Gleneagles Publishing: Glen Osmond, SA)

Butcher RJ, Davis JA and Lake PS (in prep). Assessing
biodiversity in Australian wetlands: Trials and tribu-
lations.

De Stasio BT Jr (1989) The seed bank of a freshwater
crustacean copepodology for the plant ecologist,
Ecology 70, 1377-1389.

Ellner $ and Hairston NG Jr (1994) Role of overlap-
ping generations in maintaining genetic variation in a
lluctuating environment. The American Naturalist
143, 403-417,

Green JD and Shiel RJ (1992) Australia’s neglected
freshwater microfauna. Australian Biologist 5, 118-
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Hairston NG Jr and De Stasio BT Jr (1988) Rate of
evolution slowed by a dormant propagule pool.
Nature 336, 239-242.

Hairston NG Jr, Kearns CM and Ellner SP (1996)
Phenotyphie variation in a zooplankton egg bank.
Ecology 77, 2382-2392.

Hairston NG Jr, Van Brunt RA Jr. Kearns CM and
Engstrom DR (1995) Age and survivorship of dia-
pausing eggs in a sediment egg bank. Ecology 76,
1706-171.

Hillman TJ (1986) Billabongs. In Limnology in
Australia, pp 457-470. Eds P DeDeckker and WD
Williams. (CSIRO: Melbourne and Dr W Junk:
Dordrecht)

Hillman TJ and Shiel RJ (1991) Macro- and micro-
invertebrates in Australian billabongs. Verh.
International Verein. Limnol 24, 1581-1587.

Jenkins K and Briggs S (1997) Wetland invertebrales
and flood frequency on lakes along Teryaweynya
Creek, NSW National Parks and Wildlife Service.

Leck MA and Brock MA (2000) Ecological and evolu-
tionary trends in wetlands: Evidence from seeds and
seed banks in New South Wales, Australia and New
Jersey. USA. Plant Species Biology 15, 97-112.

Nielsen DL, Hillman TJ and Smith FJ (1999) Effects of
hydrological variation and planktivorous competition
on maeroinvertebrate community structure in experi-
mental billabongs. Freshwater Biology 42, 427-444,

Nielsen DL, Smith FJ, Hillman TJ and Shiel RJ (2000)
Impact of water regime and fish predation on zoo-
plankton resting egg production and emergence.
Journal af Plankton Research 22, 433-446,

Ogden RW (2000) Modern and historical variation in
aquatic macrophyte cover of billabongs associated
with catchment development. Regulated Rivers:
Research and Management 16, 497-512.

Ogden R, Spooner N, Reid M and Head J (2001)
Sediment dates with implications for the age of the
conversion from palaeochannel to modern fluvial
activity on the Murray River and tributaries.
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Page K, Nanson G and Price D (1996) Chronology of
Murrumbidgee River palaeochannels on the Riverine
Plain, southeastern Australia. Journal of Quaternary
Science 11, 311-326. À

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Darling Basin Commission: Canberra)

Quinn GP, Hillman TJ and Cook R (2000) The
response of macroinvertebrates to inundation in
floodplain wetlands: a possible effect of river regula-
tion? Regulated Rivers: Research and Management
16, 469-477.

Reid MA (in press). A diatom-based palaeoecological
study of two billabongs on the Goulburn River flood-
plain, south-east Australia, Proceedings of the 13th
International Diatom Symposium, Curtin University,
Perth. Australia,

Reid MA and Brooks JJ (2000) Detecting effects of
environmental water allocations in wetlands of the
Murray-Darling Basin, Australia, Regulated Rivers
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Reid MA, Fluin J. Ogden RW, Tibby J and Kershaw
AP (in press). Long-term perspectives on human
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Part One

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and Temporal Scaling. (Academic Press: San Diego)

B RJ (1995) Billabongs, Australasian Science 16,
10-13.

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biodiversity: why are there so many species?
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In Methods in Stream Ecology, pp 3-22. Eds FR
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In Rivers as ecological systems. The Murray Darling
Basin. pp 101-135. Ed WJ Young. (Murray Darling
Basin Commission: Canberra)

Young WJ (ed) (2001) Rivers as Ecological Systems
The Murray Darling Basin. (Murray Darling Basin
Commission: Canberra)

Habitat Change in River Red Gum Floodplains:
Depletion of Fallen Timber and Impacts on Biodiversity

Ralph Mac Nally', Andrea Ballinger’ and Gregory Horrocks!

Abstract

Pallen timber, or coarse woody debris, is a major habitat structural element for animals and plants
throughout forested areas worldwide, Fallen timber yolumes have been massively manipulated both
through direct means (fire-wood collection, fuel reduction) and through indirect activities (silvicultur-
al management preventing trees reaching senescence). Work in our laboratory has focused on assess-
ing the extent of fallen timber depletion in River Red Gum Eucalyptus camaldulensis Noodplain
forests of the southern Murray-Darling Basin. We estimate that contemporary fallen timber loads
average about 16% (about 19 tonne/ha) of loads prior to European settlement. We also have been
investigating relationships between fallen timber loads and biodiversity of both vertebrates and inver-
tebrates. Our observational and experimental results indicate that loads at least twice current levels
exert a positive influence on a number of ecologically critical species (c.g. Yellow-footed Antechinus

Victorian Naturalist 119 (3), 2002, 107-113)

Introduction

Ecological problems have become
increasingly evident in lowland rivers,
mostly resulting from human-induced habi-
tat changes in the streams, their riparian
zones, floodplains and in the broader catch-
ment (Dexter et al. 1986; Harding and
Winterbourn 1995; Harding ef al. 1998,
1999), Many human impacts have resulted
in simplified habitats. Simplification has
involved diminution of variability and
magnitude of important ecological process-
es, as well às changes in structural attribut-

| Australian Centre for Biodiversity Analysis, Policy
and Management School of Biological Sciences, PO
Box 18, Monash University, Victoria 3800

Vol. 119 (3) 2002

Antechinus flavipes, Brown Treecreeper Climacteris picumnus). Therefore, restoration targets of

40—50 tonne/ha seem reasonable objectives for mana

ging this critical element of habitat structure. (The

es of habitats. One example of the former is
river regulation, which reduces flow vari-
ability and may even reverse seasonal flow
patterns (Close 1990; Lake 1995; Ward and
Stanford 1995). In terms of habitat struc-
ture, one of the most striking impacts on
floodplains has been the general reduction
of structural variety, especially loss of
mature trees, ground cover and natural litter
or ‘debris’. Here, we focus on the loss of
fallen timber (also known as coarse or large
woody debris) from floodplain habitats and
evaluate effects of loss on biodiversity,
Fallen timber on floodplains is ecologi-
cally important because it: (1) yields struc-
tural habitat for fauna during both dry peri-

107

Murray River Special Issue

ods (e.g invertebrates, non-aquatic verte-
brates, Hawkins er al. 1983, Stanhope et al.
1987) and during inundation (e.g. fish,
aquatic invertebrates and microorganisms;
Bryant 1983; Thorp et a/. 1985; O'Connor
1991; Fausch and Northcote 1992); (2) may
be a nutrient source for invertebrates, flora
and fungi (Bilby 1981; Culp et al. 1996;
Edmonds and Marra 1999); and (3) traps
fine debris, in-flowing nutrients and sedi-
ment, producing complex microhabitats for
animals (Harmon ef al. 1986; Andrus et al.
1988; Aumen 1990; Naiman and Décamps
1997). Thus, fallen timber provides shelter
and nutrition for many invertebrates, rep-
tiles, small mammals and fishes.

The paper is divided into sections cover-
ing the current status of fallen timber on
floodplains of the Murray River and its
major tributaries in Victoria and New
South Wales, and how that status might
relate to pre-European fallen timber loads,
We then discuss results of some observa-
tional and experimental studies involving
the responses of terrestrial invertebrates
and vertebrates to the presence of fallen
timber in floodplain habitats.

Fallen timber—now and then

In River Red Gum Eucalyptus camaldu-
lensis forests of the Murray-Darling Basin

on public land alone, c. 115,000 tonne of

firewood and e. 122,000 tonne of timber
(including wood chips) are legally removed
annually (Crabb 1997). The total forested
area also is much contracted. It appears that
in the past, floodplain forests were more
open with fewer trees per unit area. These
forests probably had a substantially greater
number of large, senescent trees (Parkinson
and Mac Nally 2000), which produce the
greatest volumes of woody debris through
natural bough abscission and tree death
(Jacobs 1955). Centuries-long silvicultural

exploitation has led to a virtual absence of

large, old trees. Thus, recruitment of fallen
timber also is much reduced. These obser-
vations suggest that fallen timber is now
much depleted in floodplain forests of the
southern Murray-Darling Basin, but what is
the magnitude of that depletion?

To assess current fallen timber loads, we
used a combination of aerial surveys and
ground-truthing (Mac Nally et al. in press
b). Some 2442 km of the 2987 km of the

108

reaches were surveyed in this way: Murray
(Albury to Murray Bridge), Murrumbidgee
(downstream of Wagga Wagga), lower
Darling (downstream of Menindee),
Lachlan (Hillston to Murrumbidgee con-
fluence), Wakool (downstream of the
Edward River), Edward (Deniliquin to
Wakool confluence), Darling Anabranch
(downstream of Popilta), lower Goulburn
(downstream of Nagambie), Loddon
(Bridgewater to Little Murray confluence),
Campaspe (Lake Eppalock to Murray con-
fluence) and Avoca (downstream of
Charlton) rivers, Much of the residual 18%
(Gunbower Island, Barmah forest, Millewa
forest, lower Goulburn south of Nathalia,
lower Ovens floodplain around Peechelba)
was too thickly forested to survey trom the
air. The thickly forested areas were sur-
veyed using at least thirty 0.5 ha plots in
each named area, Surveys were conducted
on both public land and on private lease-
hold properties on the floodplains.

Fallen timber estimates from the aerial
survey

Fallen timber loads on more than half of
the surveyed area were estimated from aeri-
al-survey data (119,480 ha). We found that
the average fallen timber density was about
19 tonne/ha. This figure is a little less than
the complete complement of timber of
three, one-metre diameter (at breast height)
River Red Gum trees spread over each ha
(c. 21 tonne/ha). The 19 tonne/ha value is
consistent with the mean of the ground-
truthed sites, which was 22 tonne/ha. The
total amount of fallen timber on the flood-
plain outside of the forest blocks was about
2,262,900 + 100,000 tonne.

Fallen timber in the major forest blocks
Barmah had the greatest density of fallen
timber on the southern Murray-Darling
floodplain, averaging over 24 tonne/ha.
The average in the lower Goulburn area
was about half of the Barmah value. and
the other areas were between these
extremes. The estimated amount of fallen
timber at Barmah is about 741,900 tonne,
but may be as little as 565,700 tonne or as
much as 957,700 tonne given the uncer-
tainties in the estimation process. In the
five forest-block areas, comprising
101,600 ha, the mean estimate is 1,912,200

'The Victorian Naturalist

tonne of fallen timber. The range of esti-
mates was 1,483,300 tonne to 2,393,500
tonne. This wide range is due to the small
areas of each forest actually surveyed
using transects in the forest blocks:
between about 0.1% in Barmah and
Millewa (hence confidence limits of about
25% of the mean) and, at most, 0.6% on
the Ovens floodplain.

Total fallen timber load estimates

Combining results from the blocks and
aerial survey, the total fallen timber on the
southern Murray-Darling floodplain that
we surveyed was about 4,175,100 tonne,
with 9595 confidence limits of 3,645,000
to 4,753,600. Average density of fallen
timber is about 19 tonne/ha.

Pre-European levels of fallen timber on
floodplains

An exhaustive examination of plans,
forestry records and historical literature sug-
gests that the structure of River Red Gum
communities has changed considerably.
However, there are virtually no early histori-
cal data pertaining to River Red Gum
forests, and relatively little is known of the
dynamics of woody debris in these environ-
ments. In lieu of historical information, a
useful surrogate approach is identification of
sites at which modifications following
European settlement have been minimal—
such sites may yield the best picture of pre-
European fallen timber loads. Robinson
(1997) found that standing crops of live
trees, fallen timber and stags were all signifi-
cantly higher in inaccessible, old-growth
areas than in managed areas in the Millewa
State Forest, southern New South Wales.
Levels of fallen timber were about five-fold
higher in old-growth areas (125 tonne/ha),
while the values for managed areas were
comparable to those reported here (20-30
tonne/ha). There appear to be very few other
existing sites upon which to base hístorical
estimates, given the relative accessibility of
the floodplains and the demand for firewood
and timber over the past two centuries. We
think that Robinson's figure is a reasonable
one because it was not greatly different from
the maximum load (95 tonne/ha) we record-
ed in 516 half-hectare transect surveys along
the Murray River and tributaries. The figure
also is low by comparison with overseas

Vol. 119 (3) 2002

Part One

loads (Maser and Sedell 1994), so it is
unlikely to be a gross overestimate.

If the old-growth-site data of Robinson's
(1997) study are representative of pre-
impact levels, then a fallen timber density
of about 125 tonne/ha probably is reason-
able. Such a density implies a loss of
between 23.000,000 and 24,000,000 tonne
for the 221,080 ha considered here, or the
equivalent of 3.3 to 3.4 million River Red
Gums of one-metre trunk-diameter in the
form of fallen timber. This amounts to the
total timber standing on 115,000 ha of
mature, old-growth River Red Gum forest
given representative standing-timber
woodloads in contemporary, managed
forests (200-300 tonne/ha), which is about
the amount of standing timber on over half
of the area that we measured!

Loss of fallen timber: impacts on
biodiversity

Our concern about fallen timber loads in
the floodplain forests is mostly to under-
stand how broad-scale, intensive change in
a major structural element of habitats
affects the capacity of animals (and plants)
to persist, To address this question, we
have used a number of observational and
experimental methods to try to link biodi-
versity with different fallen timber loads in
three main areas of the southern Murray-
Darling Basin with extensive remnant
River Red Gum forests: Gunbower Island,
Barmah Forest and the Ovens River flood-
plain (all in northern Victoria, Australia).
The first two forests lie along the Murray
River itself, while the latter straddles a
major, essentially unregulated tributary of
the Murray River.

Terrestrial invertebrates

Due to their comparatively small physi-
cal size and limited mobility, terrestrial
invertebrates may exhibit strong responses
to localized habitat changes. Invertebrates
also represent the vast majority of biodi-
versity (Sala 2000). These features make
invertebrates important subjects for
research on the impacts of habitat-structur-
al change. However, given that most inver-
tebrate species respond at localised scales
(Hansen 2000), the impact of any localised
variability in habitat is potentially much
greater than for more mobile species, such

109

Murray River Special Issue

as birds, that can *even out’ small-scale
environmental change by moving to near-
by or more distant locations (Mac Nally in
press). Thus, the invertebrate assemblage
at any given point locality is the product of
a complex array of interacting factors, of
which fallen timber load is just one. In
addition, the sheer variety of invertebrates
makes it difficult to adequately character-
ize invertebrate assemblages (Colwell and
Coddington 1994). Through our attempts
to study invertebrates in River Red Gum
forests, we have encountered some novel
aspects of these problems.

The invertebrate fauna of Murray Basin red
gum forests reflects the geographic position-
ing of the forests. A large component of the
fauna is composed of typical dry-sclero-
phyll-forest species (e.g. Promecoderus spp.
[Coleoptera: Carabidae], Matthews 1980),
but there are also riparian-zone specialists
(e.g. Tachys spp. [Coleoptera: Carabidae],
Matthews 1980, Dolomedes sp. | Araneae:
Pisauridae]), arid-zone faunal elements (e.g.
Rhytisternus spp. [Coleoptera: Carabidae],
Matthews 1980; Neostorena sp.), as well as
widespread species (e.g. Rhytidoponera
metallica (Smith) |Hymenoptera:
Formicidae], Lycosa pseudospeciosa
Framenau and Vink [Aranae: Lycosidae]).
There are also many undescribed species for
which biogeographic affiliations cannot be
attributed at this time.

We find that the various elements of this
diverse fauna respond to habitat change dif-
ferently. For example, the removal of fallen
timber often results in forest-floor habitat
that is drier and more open, which favours
arid-zone species. Zodariid spiders (*knob-
ble" spiders; Shield 2001), which are arid-
zone specialists (Churchill 1998), increase
in abundance and diversitv in areas with
reduced fallen timber loads. In contrast,
certain species of lycosid or wolf spiders
are restricted to areas with relatively high
fallen timber loads. Thus, while the species
comprising invertebrate assemblages, and
their relative abundance, differ with
amounts of and proximity to, fallen timber,
no reduction in biodiversity per se has been
identified in areas where fallen timber had
been removed. We need to be wary of
using species richness alone as a measure
of ecological change when shifts in assem-

110

blage structure may be more informative
and indicative of system change.

Those areas of the forest that have been
most affected by removal of fallen timber
generally are the same areas that now flood
very infrequently under current flow
regimes. The confounding of flood-prone-
ness with fallen timber load makes teasing
out the particular role of fallen timber in
affecting biodiversity challenging. Focusing
on the response of obligate saproxylic
species (dead-wood specialists) is one strat-
egy for getting a clearer signal of the impact
of removal of fallen timber. This method
has been used successfully in the boreal
forests in Europe and North America, where
saproxylic species, particularly coleopter-
ans, represent a major faunal component
(Harmon er al. 1986; Jonsson and Jonsel
1999; Schiegg 2000). These highly special-
ized beetles either feed directly on rotting
timber, or on fungi growing on logs.
Despite extensive sampling in River Red
Gum forests, we trapped very few obligate
saproxylic species. The low numbers of
these species may be due to an unsuitable
combination of regular forest flooding and
the unpalatability of River Red Gum timber.
However, despite the paucity, we found
fallen timber to be an important factor influ-
encing the composition of coleopteran
assemblages in River Red Gum forests.
Thus, rather than being a critical resource to
a limited range of highly specialized taxa,
fallen timber in River Red Gum forests has
an important ecological function by provid-
ing habitat-structural complexity to a wide
range of more generalized species.

Terrestrial vertebrates

We have investigated the responses of ter-
testrial vertebrates (mammals, birds, rep-
tiles and frogs) to differences in fallen tim-
ber loads in the three major forests
(Gunbower Island, Barmah, Ovens flood-
plain; Mae Nally er al. 2001). In each for-
est, seven graded (by fallen timber loads:
1.4-60.2 tonne/ha) sites were investigated
over two years. These sites were chosen to
reflect the maximum observed range in cur-
rent loads in the forests. Our results showed
that the only native, terrestrial mammal
(Yellow-footed Antechinus Antechinus

flavipes) occupied sites in significantly

The Victorian Naturalist

higher densities when fallen timber loads
exceeded 45 tonne/ha. Ground-dwelling
birds and those using fallen timber are
more prevalent, and in richer diversities, in
the vicinity of accumulations of woody
debris. Overall, fallen timber loads did not
appear to relate significantly to avian pat-
terns apart from at the local scale (i.e. near
wood accumulations). Neither frogs nor
reptiles seemed to be influenced by fallen
timber loads. There were very few reptiles,

which may reflect broad-scale depletion of

fallen timber from these habitats; similar
impacts are evident in wood-depleted box-

ironbark forests immediately to the south of

the floodplain forests in northern Victoria
(Bennett er al. 1998; Bennett er al. 1999).
We also have conducted a reasonably
large-scale manipulation of fallen timber
loads to study causal relationships between
loads and the occurrence of birds and
mammals. About 1000 tonne of wood were
redistributed after one year’s prior moni-
toring to effect eight different experimental
treatments in 34 one-hectare plots (Mac
Nally 2001). We have established a rapid
response of a near-threatened (Garnett and
Crowley 2000), wood-dependent species
of bird, the Brown Treecreeper Climacteris
picumnus, whose densities increased sub-
stantially in all treatments with fallen tim-
ber loads exceeding 40 tonne/ha (Mac
Nally ef al. in press a). This value is about
twice the basin-wide average for flood-
plains of the southern Murray-Darling
Basin, It is important to conduct further
work to see whether the reproductive per-
formance of the birds also increases in a
corresponding fashion (Walters ef al.
1999). While analyses and studies are con-
tinuing, our results suggest that the
Yellow-footed Antechinus has a similar
preference for sites with loads exceeding
40 tonne/ha, which is consistent with our
results from the observational program.
Given the restricted breeding season and
the retention of pouched young, we should
soon be able to assess whether more wood
translates into more antechinuses.

Conclusions
Implications for restoration

There are two main avenues to approach-
ing habitat restoration. The first is to
restore a system to its pre-impact state

Vol. 119 (3) 2002

Part One

(Ward er al. 1999: Lake 2001), which we
know little about directly. If Robinson's
(1997) data are representative of pre-impact
levels, then a fallen timber load of about
125 tonne/ha may be a reasonable figure.
This load implies a restoration target of
27,635,000 tonne alone for the 221,080 ha
considered here, The magnitude of this
objective is such that it would take many
decades just to be able to grow a supply of
new timber to fell for this purpose!

Another approach is to try to reinstate
ecological function, Por example, our
objective has been to relate woodloads to
the diversity of animals. We need to recog-
nise that this restricted objective addresses
only biodiversity aims, so that ecosystem
processes such as nutrient fluxes and pri-
mary and secondary productivity are not
explicitly considered. Nevertheless, a load
averaging about 40-50 tonne/ha seems to
be the best target l'or biodiversity outcomes
given our results (Mac Nally er al. 2001; in
press a). Moreover, this figure is much
more logistically feasible than the 125
tonne/ha suggested by Robinson's (1997)
study—at least in the foreseeable future—
for restoration of fallen timber on flood-
plains, and may be achieved in relatively
few decades.

Restoring the ecological integrity of the
lowland floodplains of the Murray-Darling
basin must involve the return of the rivers
and riverine environments to something
nearer their pre-European-settlement con-
dition, One of the main components of the
structural complexity of floodplains, fallen
timber, is effectively missing over vast
areas. Restoration will be a Herculean task
for many reasons, most especially because
of the lack of replacement timber. Thus,
not only does fallen timber need to be rein-
stated, but there needs to be a massive
expansion of forested habitat on the flood-
plains to service the ecological well-being
of the floodplains. In the shorter term,
thought needs to be given to developing a
management strategy for restoring fallen
timber. For example, given a limited
amount of material, how can it be best
placed to begin the long road to recovery?
Are flood runners to be preferred given
their greater significance to both inundated
and dry-phase fauna? How specific are

111

Murray River Special Issue

animals in their use of fallen timber in
terms of its location on the floodplain? Is it
best to augment existing accumulations or
is it more effective to more widely and
thinly distribute the fallen timber? The
woody debris story is only at the begin-
nings of its rudimentary course, but our
work has established that the task is a huge
one because of the severity of the changes
that have been wrought since European
settlement.

Caution: floodplain vs in-stream fallen
timber loads

Much of the river-restoration literature
concentrates on in-stream fallen timber,
‘snags’, which are perceived to be critical
elements for fish, macroinvertebrates,
algae and ‘biofilm’. Our focus on flood-
plain fallen timber should help to place the
snag problem in perspective. What are the
relative merits of fallen timber on flood-
plains and in channels? There are surpris-
ingly few measurements of natural loads of
in-stream fallen timber in Australian
rivers, especially for lowland rivers (Marsh
et al. 1999). Marsh et al. (1999) estimated
in-stream fallen timber loads in the
Edward River to be 90 + 60 tonne/ha
(0.015 m'/m^). Given that floodplains
cover much greater areas than the associat-
ed rivers (rivers usually <100 m wide), the
severity of fallen timber depletion from
floodplains seems to be at least an order of
magnitude worse than in-stream de-snag-
ging. This is a critical point should restor-
ing snags to rivers remove fallen timber
from floodplains. This would be a severe
and certainly misguided case of robbing
Peter to pay Paul. Fallen timber manage-
ment in floodplains and in the associated
channels must be considered in a unified
way (Mac Nally and Parkinson 1999),

Acknowledgements

We thank the Murray-Darling Basin
Commission (Project R7007), the Australian
Research Council (Grant Nos F19804210,
A19927168) and the Victorian Department of
Natural Resources and Environment for funding
this work. We are grateful for the many persons
who assisted in the collection of these data and
in the logistics of setting up the experiments; the
most important contributors were: Amber
Parkinson, Chris Tzaros, Lawrie Conole,
Matthew Young, Robert Price, Keith Cherry,
Darren Ward and Nick Giles.

112

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Part One

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For assistance with the preparation of this issue, thanks to Kate Smith (desktop
publishing), Ann Williamson (label printing) and Dorothy Mahler (administrative

assistance).

Vol. 119 (3) 2002

113

Murray River Special Issue

Vertebrate Fauna of Barmah Forest, a Large Forest of River
Red Gum Eucalyptus camaldulensis on the Floodplain of the
Murray River

Richard H Loyn', Linda F Lumsden’ and Keith A Ward

Abstract

Barmah Forest is part of the largest River Red Gum forest in the world, and several studies have
examined aspects of its vertebrate fauna in recent decades. This paper summarises this work and
gives an overview of the forest fauna along with some historical information and comments on pos-
sible effects of management practices. Altogether 35 mammal, 205 bird, 20 reptile, 10 frog and 28
fish species have been recorded in or near Barmah Forest. Bats constitute a high proportion of the
mammal species. Arboreal mammals are represented by several species in generally low numbers.
The only native rodent is the aquatic Water Rat, and the only small native ground-dwelling mammal
is partly arboreal (Yellow-footed Antechinus). The most numerous ground-dwelling birds are those
that can also feed from other substrates during floods (e.g. Brown Treecreeper). Three bird species
(White-plumed Honeyeater, Striated Pardalote and Brown Treecreeper) form a high proportion of
individuals in the bird community (40%). Canopy feeding insectivores are patchily distributed and
their numbers are negatively correlated with aggressive White-plumed Honeyeaters, which even
dominate on box ridges, Hollow-nesting birds form a high proportion (34%) of individuals in the
bird community. The forest is an important habitat for several threatened species, and for large num-
bers of waterbirds, fish and frogs that breed there during floods. Deep spring foods provide the best
breeding conditions for many of these species. River regulation, grazing and logging have all con-
tributed to a range of historical changes, Mostly anecdotal evidence suggests loss of some mammal
and bird species and more recent declines in certain waterbirds, frogs and snakes. The fish fauna is

now heavily dominated by introduced species, (The Victorian Naturalist 119 (3), 2002, 114-132)

Introduction

River Red Gum Excalyptus camaldulensis
is one of the most widespread eucalypt
species in Australia, commonly growing
beside rivers and in ephemeral watercourses
and floodplains over much of the continent
(Costermans 1981; Groves 1981). It occurs
as linear strips along inland rivers with more
extensive stands of forest or woodland in
temperate areas, including the banks and
floodplain of the Murray River. The most
extensive remaining stand is the Barmah-
Millewa Forest (60,000 ha), straddling the
Murray River between Echuca, Deniliquin
and Tocumwal (Dexter 1978; Murray
Darling Basin Commission 2000).

Despite the wide occurrence of River Red
Gum forests and woodlands, little has been
published on the vertebrate fauna inhabit-
ing this distinctive forest type. This paper

describes aspects of the vertebrate fauna of

Barmah Forest. It is based mainly on multi-

! Arthur Rylah Institute, Department of Natural
Resources and Environment, 123 Brown Street
Heidelberg, Victoria 3084

Institute l'or Sustainable Irrigated Agriculture,
Department of Natural Resources and Environment,
Tatura, Victoria 3616

Author for correspondence: Richard Loyn

Email Richard. Loyn@nre.vic.gov.au

114

ple visits from 1977-80 (Chesterfield et al.
1984); intermittent observations by the
authors in subsequent years; data collected
during fauna studies in the 1990s (Bennett
et al. 1994: Brown and Bennett 1995;
Lumsden er al. 1995); results from moni-
toring projects undertaken by the Barmah-
Millewa Forum (Maunsell Melntyre 2000;
Ward ef al. 2000; Ward 2001, in prep.;
Webster 2001, in prep.); other miscella-
neous documents (e.g. Cadwallader 1977);
records in the Victorian Atlas of Wildlite
and personal communications from natural-
ists. The paper aims to give an overview
with details of selected studies only (e.g.
quantitative area searches for land birds).
The latter information has been reported
previously (Chesterfield et al, 1984), but
not analysed or presented in a widely avail-
able form. The paper focuses mainly on the
Victorian side of the river, but mentions
some known contrasts relating to the New
South Wales side.

Barmah Forest

The forest covers 29,500 ha in Victoria,
within the traditional land of the Yorta
Yorta people. It is a Ramsar listed wetland,
and sections were declared State Park in

The Victorian Naturalist

1987, with Reference Areas at Top Island
and Top End (LCC 1985). The forest is
dominated by River Red Gum over most of
this area, and periodic floods play a crucial
role in maintaining the health of these
ecosystems (Fig. 1; Bren and Gibbs 1986;
Dexter et al. 1986; Bren 1988; MDBC
2000). Stands of Grey Box E. microcarpa
and Yellow Box Æ. melliodora grow on
raised sandy ridges, and Black Box Æ.
largiflarens occupies some low-lying
ephemeral swamps, and the fringe of the
forest block. Understorey vegetation is
mostly open and dominated by grasses,
with a high proportion of introduced weeds
(Chesterfield ef al. 1984: Frood and Ward
in prep.). Scattered shrubs occur especially
on box ridges (where some species such as
Silver Banksia Banksia marginata are rep-
resented only on the New South Wales
side) and on riverbanks. Silver Wattles
Acacia dealbata and thickets of introduced
Blackberries Rubus fruticosus and Willows
Salix spp. grow mainly on riverbanks.
Various aquatic plants dominate ephemeral
swamps in low-lying areas, with extensive
beds of Moira Grass Pseudoraphis
spinescens forming on treeless floodplains
as floods recede (Chesterfield 1986: Ward
1991, 1994),

Cattle have been allowed to graze on the
Victorian side for many years, and have
impacted on swamp vegetation, with Giant
Rush Juncus ingens prospering at the
expense of Common Reed Phragmites dus-
tralis and Cumbungi Tvpha angustifolia
(Chesterfield 1986). Sheep have grazed on
the New South Wales side. Grazing has been
reduced in recent years. Logging has pro-
ceeded on both sides of the river for many
years, with railway sleepers being a major
product (Dexter 1978; Chesterfield 1986).
Limited logging continues in the State Park
(DCE 1992).

Varying proportions of the forest are inun-
dated when the river overflows its banks.
The natural season for flooding is late win-
ter and spring, following winter rain and
snow melt in the mountains. Since the
1930s, the river has been regulated by con-
struction of dams near Albury-Wodonga
(Hume Dam, 1934) and Dartmouth (1980),
and a weir at Yarrawonga (1939). The main
aim of regulation has been to store water at
peak flows and release it during summer-

Vol. 119 (3) 2002

Part One

Fig. 1. Flooded River Red Gum forest. Photo by
Lindy Lumsden.

autumn for use by irrigation farmers. Hence
there has been a reduction in the extent and
frequency of winter-spring floods and an
increase in summer-autumn flows to low-
lying areas. The dam at Dartmouth was also
intended to provide a capacity for flushing
the river to control salinity problems down-
stream. Flows of water through the forest
are further regulated by means of sluice
gates on most creeks and major flood run-
ners (distributary channels).

Methods

In the early stages of this study, all parts
of the forest were explored to gain a gener-
al picture of the fauna and its distribution.
Quantitative data on land birds were
obtained on six sites by searching areas of
4 ha (200 200 m) for an hour each in
October-November 1977 (a dry year, no
flooding) and October-November 1978 (a
wet year, with floodwater lying over about
half the forest at the time of the survey). A
canoe was used to gain access in 1978, but
the surveys were conducted on foot (wad-
ing in water up to neck-deep at one site),

115

Murray River Special Issue

The data were analysed by grouping
species into guilds, and examining effects
of sites (6) and years (2) using analysis of
variance (assuming no interaction between
sites and years). Five of the sites were in
stands of River Red Gum on the flood-
plain, and one was on a sand-ridge domi-
nated by mature Grey Box with some
Yellow Box. Four of the River Red Gum
stands were mature and three of them con-
tained many large old hollow-bearing
trees. The fourth site (Barmah Island) con-
tained mature trees but few old living vet-
erans, as intensive silvicultural treatments
had killed such trees by ringbarking and
poisoning over several decades. The fifth
Red Gum site (burnt) was regenerating
from a severe wildfire that burned ~30 ha
ten years before the study, and most of the
few old trees were dead.

Water birds were counted at various
times during this period, and attempts were
made to count colonial breeding water
birds in each of these years. Small mam-
mals were trapped at five sites in three sea-
sons, using folding aluminium traps and
small wire-cage traps (1280 trap-nights)
and harp traps and mist-nets for bats (at
four water-holes in January 1980)
(Chesterfield et al. 1984). Seats and signs
were recorded, and spotlight surveys were
conducted on foot and from vehicles.

As part of a broad scale study of verte-
brate fauna of the Northern Plains in the
early 1990s (Bennett et al. 1998) five sites
within Barmah Forest were sampled for
bats and arboreal mammals. Harp traps
were set for bats at each site for two
nights. Spotlighting was undertaken on
foot by intensively searching a 1 ha plot
for 45 minutes, on two consecutive nights,

Colonial breeding water birds were
counted from the air in most years since
the 1980s when there was a spring flood
(Ward 1993, 1996; Ward et al. 2000), with
ground truthing where possible. Earlier
records from 1955-1979 were collated by
Cowling and Lowe (1981).

The first in-depth amphibian survey for
Barmah Forest was conducted in 2000/01
(Ward 2001). Frogs and their breeding
attempts were surveyed at night on a
monthly basis between September and
February by documenting vocalising
species plus undertaking some limited

116

ground searches at approximately 20 sites
throughout the forest. Breeding attempts
were primarily recorded from tadpoles col-
lected from dip-net samples. This program
has continued into 2001/02, with monitor-
ing undertaken at various months through-
out the year (Ward 2002).

The Department of Natural Resources
and Environment has undertaken a variety
of fish surveys in and adjoining the forest
(McKinnon 1997; Stuart ef al. 2001).
Additional monitoring programs have been
established by the Barmah-Millewa Forum
to survey waterbirds and land birds (R
Webster pers. comm.) and frogs (Ward
2001, 2002).

The Atlas of Victorian Wildlife (NRE
2001) was consulted for recent and histori-
cal records of all species, examining all 5-
minute grid cells that overlapped the forest.

Results
Mammals

A total of 26 species of native mammal
and nine species of introduced mammal
has been recorded in or near the forest in
recent times (Appendix 1).

No exclusively terrestrial native small
mammals occur in Barmah Forest. The
Yellow-footed Antechinus is widespread,
although uncommon, and spends time on
the ground as well as being arboreal. This
species was trapped in low numbers
(14/1280 trap-nights on the early survey)
and several were seen during the day for-
aging from branches of River Red Gums.
The only other small mammal trapped on
the early survey was the introduced House
Mouse (5/1280 trap-nights). Even the nor-
mally common and widespread Echidna is
rarely seen in Barmah Forest,

The two native aquatic mammals (Platypus
and Water Rat) are present in the river and
creeks, though only the latter is common.

Eastern Grey Kangaroos are common and
conspicuous throughout the forest, with
concentrations often occurring in and near
extensive grass plains. White individuals
are seen occasionally. The Black Wallaby
has only recently been recorded within
Barmah Forest with the first record on the
Atlas of Victorian Wildlife from 1994. This
species has recently expanded its range into
western and northern Victoria (Menkhorst
1995). Common introduced ground-

'The Victorian Naturalist

dwelling mammals include European
Rabbits (in fluctuating numbers, e.g. more
in 1980 than 1977), Brown Hares, Red Fox,
Feral Cat (mainly on forest margins), Feral
Horse (several herds of ‘brumbies’, mainly
in central and western parts of the forest)
and Feral Pig (especially in swamps in the
west of the forest). There have been recent
records of Sambar Deer, perhaps illegally
released, but it is not known if they have
established a population.

Seven species of arboreal mammals
inhabit Barmah Forest. Common Brushtail
Possums are common among River Red
Gums while Common Ringtail Possums
occur on the box ridges. For example, dur-
ing the 1991 survey Common Ringtail
Possums were recorded at densities of 5-7
animals per ha at the two sites in box
woodlands, while none was recorded at the
three Red Gum sites. Sugar Gliders are
scarce, and mainly found near riverbanks
with Silver Wattles. This area is close to
the inland edge of their range and Sugar
Gliders appear to be uncommon compared
to elsewhere in their distribution. Squirrel
Gliders are also rare with only two records
from the forest: two animals were
observed on a Yellow Box rise on Long
Plain Track in 1982 (J Alexander pers.
comm.). and one was observed on Barmah
Island in 1978. The semi-arboreal Brush-
tailed Phascogale is also rarely recorded in
the forest with only three records on the
Atlas of Victorian Wildlife, all from the
early 1980s. More recently, one individual
was observed in 1991 on a mature Red
Gum within semi-open Red Gum forest in
the Barmah Lake day visitor area (PG
O'Connor pers. comm.). The Feathertail
Glider has been recorded only once from
the forest (in 1999: L Conole pers. comm.).
The small size and cryptic nature of this
species makes it difficult to detect, and it is
likely to occur more widely in the forest
than this one record would suggest. Koalas
have been re-introduced into the forest as
part of a state-wide translocation program.

Bats comprise the largest proportion of

the native mammal fauna of Barmah
Forest, with one species of flying-fox and
|2 species of insectivorous bats recorded
(Appendix 1). Little Red Flying-foxes are
irregular summer visitors, at times form-
ing camps of ~200 in trees near Barmah

Vol. 119 (3) 2002

Part One

Fig. 2. The Gould's Lond-eared Bat
Nyetophilus gouldi was recorded at consider-
ably higher rates in Barmah Forest than else-
where in the Northern Plains. Photo by Lindy
Lumsden.

Lake, drinking from nearby creeks (Loyn
1981) and feeding on River Red Gum
blossom. The only camp regularly used by
this species in Victoria is at the nearby
town of Numurkah, where up to 2000
individuals congregate over summer.

Barmah Forest has a high diversity of
insectivorous bats with all but one species
known from the Northern Plains occurring
there. The one exception is the Greater
Long-eared Bat Nyctophilus timoriensis
(Fig. 2), which has been recorded from a
single individual in Black Box woodland
south-west of Echuca (Lumsden 1994).
While most species occur in a range of veg-
etation types, the Southern Myotis is
restricted to riparian forests, as it forages
exclusively over water, catching aquatic
insects and small fish (Menkhorst 1995).
This species was caught on Barmah Island
during the early surveys, and has been
recorded in NSW along the Murray River at
the Barmah Bridge (Law and Anderson
1999),

Capture rates of insectivorous bats during
the 1990s surveys. were higher in Barmah
Forest than the mean for all the sites in the
Northern Plains. Some species were
recorded at markedly higher rates within
Barmah, in particular the Gould’s Long-
eared Bat, which is close to the inland limit
of its distribution in this area. The
Southern Freetail Bat, Little Forest Bat and
Gould’s Wattled Bat also had high capture
rates within Barmah. The Southern Forest
Bat, which is common throughout much of
Victoria, is rare in this region.

117

‘ial Issue

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'The Victorian Naturalist

118

Laud birds
Overview of land bird community
Altogether 199 species of native land bird
and six introduced species have been
recorded in or near Barmah Forest
(Appendix 2). Three species are numerical-
ly dominant in River Red Gum stands:
White-plumed Honeyeater, Striated
Pardalote and Brown Treecreeper, together
forming 40% of individuals in the bird
community (Table 1). Sulphur-crested
Cockatoos are common and widespread,
and make themselves conspicuous through
their incessant screeching calls. Other par-
rots, including the yellow form of Crimson
Rosella, are common and easily seen.
Three black-and-white birds that have
become common in farmland (Australian
Magpie, Magpie-lark and Willie Wagtail)
are widespread through the forest, especial-
ly in open grassy stands of River Red Gum.
Emus are conspicuous, especially in and
near grass plains and swamps, Hollow-
nesting birds form a high proportion of the

T
=

bird community, contributing means of
34% of individuals in mature stands of

River Red Gum or Grey Box and up to
54% of individuals on some sites (Table 1).

Box ridges support a surprisingly similar
fauna to the River Red Gum stands. White-
plumed Honeyeaters are less dominant but
still the most common honeyeater (Table |).
Some small insectivores are more common
than in River Red Gum stands, including
Yellow Thornbills and White-winged
Trillers. A few common farmland birds
favour box ridges, including Galahs and
Yellow-rumped Thornbills (mainly in
Yellow Box), though the latter species has
apparently increased and is now common
on treeless plains and among adjacent River
Red Gums (L Conole pers, comm.). Hooded
Robins are scarce inhabitants of box ridges,
as are Gilbert's Whistlers and Striped
Honeyeaters on the NSW side.

Many other species are localised, and
some are associated with particular habi-
tats such as box ridges or riverbanks
(Appendix 2). Open-country species such
as Richard's Pipits, Singing Bushlarks and
Brown Songlarks are found only on tree-
less flood-plains, when conditions are suit-
able (e.g. after floods). Noisy Miners are
found only in small patches of River Red
Gum on edges of treeless plains, and in

Vol. 119 (3) 2002

Part One

roadside trees outside the forest. White-
browed Scrubwrens are found mainly in
beds of rushes Juricus spp. in swamps, and
locally elsewhere (e.g. on riverbanks and
in fallen branches on logged coupes). Beds
of Giant Rush are the main habitat for
Little Grassbirds and support high densi-
ties of Superb Fairy-wrens, though the lat-
ter are common in a wide range of habitats.
Red-browed Finches are locally common
in thickets of Blackberry and other species
along the riverbank. Other species are
widespread but scarcer than in more varied
forests outside the floodplain: they include
two that forage from the bark of upper
branches ( Varied Sittella and Brown-head-
ed Honeyeater) and a group of small insec-
tivores such as Brown Thornbill and
Striated Thornbill.

Several species are more common in road-
side vegetation, farmland or forest patches
outside the forest than in the forest itself,
including Noisy Miner, Pied Butcherbird
and Grey-crowned Babbler (Appendix 2).
Bush Stone-curlews inhabit patches of rem-
nant woodland outside the forest, and are
rare within the forest. Gilbert's Whistlers
and Striped Honeyeaters are resident in low
numbers on box ridges on the NSW side
(P Maher pers. comm.) but rare on the
Victorian side. Speckled Warblers occur in a
rarely grazed patch of shrubby River Red
Gum forest near Nathalia, but apparently not
in Barmah Forest. Chestnut-rumped
Thornbills occur in a swamp dominated by
Tangled Lignum Muehlenbeckia florulenta
near Nathalia, and are rare in the forest
where lignum is scarce. Several species char-
acteristic of dry environments occur in small
numbers outside the forest ( Appendix 2).

Introduced land bird species

Four introduced bird species have been
observed within the forest: Blackbird (in
Blackberry thickets on riverbank, and near-
by gardens); European Goldfinch (occasion-
al birds, feeding mainly from introduced
Asteraceae seeds but once reported taking
nectar from box blossom; H Marshall pers.
comm.), House Sparrow (groups locally
around cattle yards) and Common Starling
(breeding in tree hollows near Barmah Lake
in spring; flocks sometimes feeding around
drying lakes in summer). These and other
species are more common in farmland and

119

Murray River Special Issue

towns outside the forest (Appendix 2), and a
fifth species (Eurasian Tree Sparrow)
occurs in small numbers at Nathalia. A sixth
species, Common Myna, has recently
reached Nathalia (2002), as part of its gen-
eral range expansion, but has not been
recorded in the forest.

Quantitative data from dry and wet years

Numbers of individuals of all species
observed on the six study sites in a dry
year and the following wet year are shown
in Table 1, by species and grouped into
guilds. Total numbers of land birds were
remarkably similar in the two years.
However, the most numerous species
(White-plumed Honeyeater) was substan-
tially more common in the dry year (1977),
and species diversity was correspondingly
higher in the wet year (1978).

Water birds were present on these sites in
1978, but not in 1977 (Table 1). Other
guilds showed little difference between the
two years, except for canopy foragers
which were markedly more common in
1978 than 1977 (Table 1). This difference
was highly significant (p — 0.003).

Total numbers of land birds were very
similar between sites (Table 1) and differ-
ences were not significant (p = 0.375).
However, substantial differences were evi-
dent for particular guilds (Table 1), includ-
ing canopy foragers (p < 0.001), aerial for-
agers (p = 0.046), bark foragers (p =
0.013), hollow nesters (p = 0.045) and nec-
larivores (p = 0.056), Nectarivores were
most numerous on Barmah Island in the
west of the forest, and scarce in the recent-
ly burnt site and the dry site at Top Island.
This guild was dominated by White-
plumed Honeyeaters. Bark foragers (main-
ly treecreepers) were most numerous at
Boals Deadwoods, which contained many
large old River Red Gums (dead and alive)
among low-lying swamp vegetation.
Hollow-nesters were most numerous at
Boals Deadwoods and Gulf, and least
numerous at Barmah Island and the burnt
site, where most old trees had been killed
by silviculture and fire respectively. This
guild included the treecreepers and a wide
range of parrots and other species, but was
dominated by Striated Pardalotes. Canopy
foragers were most numerous at the recent-
ly burnt site and at Top Island, in the drier

120

eastern part of the forest. The abundance
of canopy foraging insectivores was nega-
tively related to the abundance of White-
plumed Honeyeaters across the sites and
years (r = -0.76, p < 0.001, n= 12).

Water birds

When the forest floods, the treeless
plains become lakes and are populated by
vast flocks of water birds. During a major
flood, the treed parts of the forest provide
breeding habitat for hundreds of ducks,
mainly of three common species (Grey
Teal, Pacific Black Duck and Australian
Wood Duck), most of which nest in hollow
trees. The swamps attract a greater diversi-
ty of species, including Black Swans and a
wide range of ducks, grebes and other
birds. Flocks of Australian Pelicans may
gather on treeless plains, though breeding
has not been proved for that species.

The most spectacular breeding colonies
are those of the communally breeding
birds such as ibis and cormorants. In the
major floods of 1973-75, it was reported
that hundreds of thousands of ibis bred at
several distinct colonies in the forest
(Chesterfield er al. 1984). In subsequent
years, hundreds or thousands of pairs have
bred during spring floods, numbers vary-
ing with water depth. Beds of Giant Rush
and sometimes Common Reed are the
favoured habitat, and are typically flat-
tened by the ibis to make nesting plat-
forms. Straw-necked Ibis form dense
colonies and Australian White Ibis gener-
ally form groups of smaller sub-colonies.
Little Pied Cormorants nest colonially in
River Red Gum trees, often in association
with small numbers of Little Black
Cormorants. Great Cormorants and Darters
have nested locally in some years, select-
ing large River Red Gums near creeks,
White-necked Herons regularly nest in
small colonies, usually fewer than ten
pairs, in tall River Red Gums. Yellow-
billed Spoonbills sometimes nest singly in
tall River Red Gums, or they may nest
close to colonies of cormorants or White-
necked Herons.

Great Egrets regularly nest in large River
Red Gums on the New South Wales side
of the river during spring floods.
Previously they nested on the Victorian
side at Black Swamp until 1964, and near-

The Victorian Naturalist

by at Bullock Creek in 1966, 1968 and
1969 (Chesterfield et al. 1984). Small
numbers of Little Egrets and Intermediate
Egrets are sometimes seen with them, and
both species have bred in the forest in the
past (see Historical Changes below).
Nankeen Night Herons also periodically
nest on the NSW side of the river, with
colonies of 2000-3000 nests having been
recorded at Porters Plain during the flood-
ing of 2000/01 (Webster 2001).

Most breeding occurs in spring, and win-
ter floods rarely induce extensive breeding
in the forest. In contrast, winter rains may
induce flooding in nearby pasture, and
floodwaters sometimes attract substantial
numbers of waterbirds including some that
rarely visit the forest itself, e.g. Red-
necked Avocet. Many ducks and other
waterbirds have bred on these floodwaters.

When floods recede from the forest, open
muddy shores can attract small waders
including Palaearctic migratory species
such as Latham's Snipe and Common
Greenshank. However, there is usually too
much vegetation to provide good habitat
for most migratory species. and numbers
observed are generally low.

Reptiles

Twenty species of reptile have been
recorded in Barmah Forest incuding two
tortoises, two geckoes, six skinks, three
other lizards and seven species of snakes
(Appendix 3). Yellow-bellied Water
Skinks are conspicuous among River Red
Gums, and Boulenger's Skinks are com-
mon on box ridges (P Brown pers. comm. ).
Tortoise nests are often raided by Red
Foxes, with eggshells and footprints
remaining as evidence.

Historical records up until the 1970s listed
snakes in the Barmah Forest as being very
common (as per review by Leslie 1995).
However, numbers have greatly decreased
since this time, to the point where snakes
are now uncommon. The Red-bellied Black
Snake is the most commonly observed
snake in Barmah Forest today, though
Eastern Tiger Snakes may have once been
more common (Leslie 1995). Eastern
Brown Snakes are also occasionally seen,
along with some rarer and more cryptic
species (Appendix 3). Carpet Pythons
favour the drier Box ridges.

Vol. 119 (3) 2002

Part One

Frogs

Ten species from two families of amphib-
ians have been confirmed to reside in or
near Barmah Forest (Appendix 4). Hylidae
is represented only by Peron's Tree Frog,
while Leptodactylidae is represented by the
Southern Bullfrog, Barking Marsh Frog,
Spotted Marsh Frog, Giant Bullfrog,
Common Spadefoot Toad, Bibron's
Toadlet, Plains Froglet, Common Froglet
and Sloane's Froglet.

Common Spadefoot Toad and Bibron's
Toadlet are recent new records for the forest,
extending the known range of the latter
species. There is a single record of the criti-
cally endangered Giant Bullfrog from the
farmland area just south of Barmah Forest. A
species formerly listed as occurring in
Barmah Forest, the Warty Bell Frog Litoria
raniformis, was not found to occur in the for-
est and the single original record was found
to be erroneous (P Robertson pers. comm.).

Well vegetated, shallowly-flooded wet-
lands have been found to contain the high-
est numbers of vocalising and breeding
frogs. However, ephemeral rain-filled
depressions were also found to be signifi-
cant, and the two new species records for
the forest came from such locations. Creeks
and rivers generally support the lowest
numbers of frogs ( Ward 2001, 2002).

Fish

Twenty-eight species of fish have been
recorded in the vicinity of Barmah Forest,
with ten of these being exotic (Appendix 5).

Native fish, particularly large “commer-
cial' species such as Murray Cod and
Golden Perch, were known to be particular-
ly plentiful in the mid-1800s to the early
1900s when commercial fishing was under-
taken in the Murray River and adjoining
Barmah-Moira Lakes, However, significant
decreases in catches were recorded by the
1890s, and Golden Perch replaced Murray
Cod as the most common species (Ross
1892, cited in Leslie 1995).

Carp and Redfin were introduced to
lagoons in the Deniliquin district in
1884 and goldfish were in fountains in
Echuca in the 1890s (Leslie 1995). Even
without deliberate release into the
adjoining waterways, floodwaters would
cause an inevitable introduction into the
river system.

121

Murray River Special Issue

By 1993, a fish study in Moira Lake
failed to locate a single ‘commercial’
species of fish (Gerkhe and Brown, cited in
Leslie 1995). Instead, Carp dominated the
waterways. Although other studies have
found populations of Murray Cod, Golden
Perch and Silver Perch to persist in Barmah
Forest, some native species have declined
so much that they are now rarely encoun-
tered in the area (McKinnon 1997;
Sivakumaran and Brown 2001). Such
species include River Blackfish, Freshwater
Catfish and Macquarie Perch. Furthermore,
Carp have been found to spawn so prolifi-
cally in the shallow vegetated wetlands
within Barmah Forest that the region may be
a major point source breeding location with-
in the Murray system (Stuart er al, 2001),

Threatened Species

Of the mammal species recorded in
Barmah Forest, three are listed as threat-
ened (Appendix |). Within Victoria the
endangered Squirrel Glider is found only
in the Northern Plains and adjacent Box-
Ironbark region. It occurs in either box

woodlands with more than one species of

eucalypt, or in River Red Gum forests with
an understorey of wattles (Menkhorst er al.
1988). Areas that contain à number of tree
species appear important to provide food
resources throughout the year. Two of the
three sightings within Barmah Forest have
been in mature Yellow Box woodlands,

Ihe Brush-tailed Phascogale is listed as
vulnerable in Victoria with its core distrib-
ution through the box-ironbark forests
inland of the Divide (Menkhorst 1995),
Phere are few observations from Red Gum
forests. Further information is required on
this species to ascertain its status and dis-
tribution within Barmah Forest.

Nine land bird species listed as endan-
gered in Victoria (NRE 2000) have been
observed in or near Barmah Forest
(Appendix 2), but most appear to be
vagrants or rare visitors to the general area.
Three threatened species may have impor-
tant habitats within the forest: Barking
Owl, Superb Parrot and Grey-crowned
Babbler. The forest is one of the few
breeding habitats for Superb Parrots in the
State (Chesterfield er al, 1984; Webster
1988; Garnett and Crowley 2000), Barking
Owls have been recorded on few occa-

122

sions, and are likely to be resident in small
numbers. Grey-crowned Babblers have
declined within the forest and are now
more often seen at, roadsides outside the
forest (Chesterfield et al. 1984; Robinson
1994; Garnett and Crowley 2000). A small
group persisted on Top Island until at least
1979 (Chesterfield et al. 1984).

Several waterbird species are currently
listed as threatened (NRE 2000) on the
basis of restricted breeding colonies. This
approach is currently being revised.
Barmah Forest is sometimes one of the few
breeding colonies in the State for Great
Egrets, and has been of even greater
importance for other species of egret (see
Historical Changes, below). The forest
may sometimes be an important breeding
habitat for two threatened and poorly
known non-colonial waterbirds with cryp-
tic habits, Australasian Bitterns inhabit
beds of Giant Rush, and several may be
heard booming in the forest during floods
(Chesterfield et al. 1984). Little Bitterns
prefer beds of Cumbungi or Common
Reed, and may be found on Top Island and
Boals Deadwoods during floods, as well as
beside farm dams outside the forest. Two
or three pairs of White-bellied Sea Eagles
breed in the forest.

Two reptile species (Carpet Python and
Woodland Blind Snake), one amphibian
(Giant Bullfrog) and six fish species
(Silver Perch, Trout Cod, Murray Cod,
Golden Perch, Macquarie Perch and
Freshwater Catfish), are listed as threat-
ened in Victoria (Appendices 3-5). As dis-
cussed above, some of the fish are now
rare or absent and the Giant Bullfrog is
known only from just outside the forest.

Discussion

There is currently a very low diversity
and abundance of terrestrial mammals in
Barmah Forest. This may be a natural
occurrence, as the pattern of regular flood-
ing would result in inhospitable conditions
unless species were able to climb trees or
find high points on which to escape. In
areas where there are alternative vegetation
types on higher ground (e.g, Black Box
woodlands around the edge of the forest,
Yellow Box woodlands on sandy rises)
these may be important refuges for indi-
yiduals to shift into to escape the inunda-

The Victorian Naturalist

tion. Where clearing has extended right up
to the edge of the Red Gum, this may limit
the options for small mammals and hence
influence their survival. The low diversity
may also be a result of management
changes since European settlement. There
is evidence to suggest that other species of
terrestrial mammals once occurred in this
region (see below). Grazing and trampling
by domestic and feral animals may have
reduced the quality of habitat for these
species, and increased predation from
introduced predators may also have con-
tributed to their demise.

In most areas of Victoria, native rodents
are usually a prominent component of the
small mammal fauna (Menkhorst 1995).
However, in Barmah Forest the semi-
aquatic Water Rat is the only species of
native rodent present. It is likely that sev-
eral other species of rodents once occurred
in the Northern Plains, but are now extinct
(see Historical Changes, below).

Compared to the Grey Box woodlands of
the Northern Plains, the Red Gum forests
have higher densities of Common Brushtail
Possums, but lower densities of Common
Ringtail Possums. All the arboreal and
semi-arboreal mammals found in the region
require tree hollows in which to nest and
are more abundant in areas with higher
densities of large old trees with hollows.

The extensive area of Red Gum woodland
in Barmah Forest provides important habitat
for bats within the region. Recent studies of
the roosting and foraging requirements of
the Lesser Long-eared Bat and Gould's
Wattled Bat found that individuals fed in
remnant vegetation within the rural land-
scape, but roosted within Barmah Forest
(Lumsden et al. 2002). Both species were
highly selective of the types of trees in
which they roosted, with Gould's Wattled
Bats selecting very large live River Red
Gums (Lumsden er al. in press). In contrast
Lesser Long-eared Bats roosted predomi-
nantly in dead trees, and the females select-
ed large old ring-barked Red Gums as
maternity sites to raise their young.
Entrances were usually narrow fissures
which provided protection from predators
and competitors. Individuals regularly shift-
ed roost sites with subsequent roosts close
together. This behavioural pattern requires a
high density of suitable roost sites within a

Vol. 119 (3) 2002

Part One

small area to support a colony of bats
(Lumsden and Bennett 2000). Although the
proportion of large trees is lower in Barmah
Forest than in remnants within farmland due
to timber harvesting (Bennett er al. 1994),
the area of forest is much larger and hence
there are more suitable roost sites overall.
Both species selected areas of the forest
that had high densities of their preferred
roost trees (Lumsden er al, 2002).

As a habitat for land birds, the stands of
River Red Gum appear to be quite similar
to less extensive stands elsewhere along
the Murray River. Birds that nest in tree
hollows are substantially more common
than in other forest types (Loyn 1985),
reflecting the age of many of the veteran
trees and their tendency to form hollows
through dropping branches. Ground-feed-
ing birds respond positively to the amount
of coarse woody debris (Mae Nally er al.
2001). Birds that feed from open ground
are common if they can use alternative
substrates during floods (e.g. Brown
Treecreeper and Buff-rumped Thornbills
feeding from bark, and Superb Fairy-wrens
among aquatic plants), while obligate
ground feeders are scarce or confined to
box ridges (Chesterfield er al. 1984; Loyn
1985). This is analogous to the current sit-
uation with small native mammals, of
which the only remaining species ( Yellow-
footed Antechinus, discussed above) can
forage both from the ground and trees.
Perhaps the deduced loss of mammal
species from Barmah Forest owes more to
clearing of adjacent box woodlands, than
to changes within the forest itself.

White-plumed Honeyeaters dominate the
bird fauna of the River Red Gum stands, and
appear to exclude small insectivores through
territorial aggression. This may account for
the patchy distribution of small insectivores
such as thornbills in the forest. The typical
honeyeaters of box-ironbark forests are
Fuscous Honeyeaters Lichenostomus fuscus
and Yellow-tufied Honeyeaters £L. melanops
(Loyn 1985; Silveira et al. 1997), but they
are absent from box ridges in Barmah
Forest, perhaps because White-plumed
Honeyeaters are so prevalent. Recent data
from a large group of study sites among the
River Red Gums in the forest (Mac Nally ef
al. 2001) include White-plumed Honeyeater
as the fifth most common species (not the

123

Murray River Special Issue

first as in earlier work), and two small insec-
tivores (Weebill and Yellow-rumped
Thornbill) featured in the top four. This may
suggest a recent change in the dominance of
White-plumed Honeyeaters and supports the
notion of an inverse relationship between
numbers of small insectivores and White-
plumed Honeyeaters.

Historical changes

One of the earliest European settlers in
the Northern Plains was Edward Curr who
occupied a pastoral run near the junction of
the Murray and Goulburn Rivers in the
1840s (Curr 1883). He described the abo-
riginal women digging out rats for food
and referred to ‘kangaroo-rats’ and ‘field-
rats’. Unfortunately, as no specimens were
collected we cannot determine which
species these may have been. The Lesser
Stick-nest Rat Leporillus apicalis, a native
rodent that builds large nests of sticks, was
recorded in the Mallee region of Victoria
in 1857 (Wakefield 1966) and may also
have been in this area, It is interesting to
speculate whether ‘Rat Castle Creek” in
Barmah Forest was named after this
species. Several species of small macrop-
ods have disappeared from the Northern
Plains. The Rufous Bettong 4epyprymnus
rufescens and Bridled Nailtail Wallaby
Onychogalea fraenata were recorded from
Red Gum forests, downstream of Barmah
(near Gunbower) in the 1850s (Wakefield
1966) and may also have been present in
Barmah Forest. Both these species are now
extinct in Victoria (Menkhorst 1995).

Among land birds, both Bell Miners and
Crested Bellbirds were reported at Barmah
Forest in the 19" century (Curr 1883;
Blakers et al. 1984; Chesterfield er al. 1984)
and no longer occur in the region. Several
honeyeaters occur erratically in or near the
forest, and Blue-faced Honeyeaters may
have declined through persecution by fruit
growers. White-throated Treecreepers were
apparently rare before the 1940s and are
now common, perhaps because of increases
in stand density with regeneration after log-
ging (Chesterfield et al. 1984). Pied
Currawongs have increased over the same
period, as winter visitors mainly to the forest
edge. Long-billed Corellas increased during
the 1970s. Recent data (Mac Nally et al.
2001) and Atlas records (Appendix 2) sug-

124

gest that Yellow-rumped Thornbills have
increased since the 1970s and early 1980s
when they were scarce in the forest
(Chesterfield e¢ a/. 1984). Restless
Flycatchers may have declined and only two
were observed during recent studies (L
Conole pers. comm.). Many other changes
may have occurred, but gone unnoticed or
undocumented.

Changes in populations of water birds
may be readily noticed by visiting natural-
ists, but hard to interpret because of the
mobility of these birds, and the great varia-
tions expected from year to year with dif-
ferent flood patterns. Magpie Geese were
common in the 19" century (Chesterfield
et al. 1984) and disappeared from Victoria
until recent reintroductions. Brolgas were
also common in the 1890s (Leslie 1995).
Hundreds of Brolgas displayed and nested
on War Plain in the 1920s (Tom Galloway
in Chesterfield et a/. 1984), and the species
is now much reduced, with a pair nesting
in nearby farmland but rarely visiting the
forest. A great diversity of breeding water-
birds was recorded by Norman Favoloro in
the 1930s (in Chesterfield et al. 1984),
including Whiskered Terns. Hoary-headed
Grebes and Baillon's Crakes. Verbal
records from various sources suggest that
Intermediate Egrets were the most numer-
ous egret breeding in the forest before the
1940s (Chesterfield er a/. 1984), and they
are now the rarest. Hundreds of egrets bred
at Black Swamp during floods at the time,
whereas now breeding is more common at
Algeboia Plains on the NSW side, and
involves greatly reduced numbers, mainly
of Great Egrets. Ibis now breed in lower
numbers than they did in the peak floods
of the early 1970s. Plumed Whistling-
Ducks and Cattle Egrets colonised the
region in the early 1970s, but are found
mainly in surrounding farmland,

Azure Kingfishers became temporarily
scarce in the late 1970s, perhaps because of
the proliferation of European Carp and con-
sequent turbidity in many creeks
(Chesterfield et al. 1984). However, they
recovered quickly and were again a common
feature of the river and creeks in the 1980s.

Many factors have been implicated in the
decline of frogs, reptiles and fish. These
include commercial fishing pressures, com-
petition and predation from introduced

The Victorian Naturalist

species (e.g. fox predation on tortoise eggs
or Mosquitofish predation on frog eggs and
larvae), structural alterations to habitat
(through grazing pressure and altered flow
regimes), reduction of instream habitat (such
as through extensive de-snagging opera-
tions). and altered flooding regimes (com-
mensurate with river regulation in the mid-
19305).

Changes in habitat have occurred, espe-
cially on the swamps, through combined
effects of river regulation and grazing.
Trees are encroaching on some previously
open plains and dying back on others
(Chesterfield 1986). Beds of Giant Rush
have tended to increase at the expense of
Cumbungi or Common Reed, which are
more palatable to cattle (Chesterfield
1986).

Management

Deep spring floods clearly provide breed-
ing conditions for waterbirds, and their fre-
quency and extent have declined with river
regulation. It will be a special challenge to
provide adequate flows with increasing
demands for water use in the Murray
Darling catchment (MDBC 2000). Flood
patterns are important for all aspects of the
forest's ecology, including vegetation
(Bren and Gibbs 1986; Chesterfield 1986;
Ward 1991, 1994; Blanch et al. 1999),
health of trees (Dexter er a/. 1986), fish
(McKinnon 1997; Gehrke and Harris
2001), and waterbirds (Leslie 2001). A
specific allocation of environmental water
has been provided for the Barmah-Millewa
forest from the regulated river system, and
is primarily used to allow waterbirds (and
flora and other fauna when known to be
necessary) to complete breeding attempts
(Leslie and Ward in press).

Old hollow-bearing trees are locally con-
spicuous, especially along the river
(Newton-John 1992), although they occur
at lower densities in the forest compared to
many remnants in farmland (Bennett er al.
1994). Logging and firewood collection
have been reduced in recent years, and this
should facilitate efforts to maintain the
supply of hollows and replenish coarse
woody debris at sustainable rates. Grazing
has contributed to major changes in wet-
land vegetation (Chesterfield 1986), and
planned reductions in grazing should help

Vol. 119 (3) 2002

Part One

reverse some of those changes, with bene-
fits to certain water birds and land birds.
The ecosystem is highly complex, and a
system of regular monitoring and adaptive
management is needed to deal with the
unexpected changes that can confidently
be expected in future.

Acknowledgements

Many people contributed to the studies and
observations that formed the basis of this paper.
We are grateful to all, although there are too
many to mention individually, Several col-
leagues have worked closely with us at different
times, including Andrew Bennett, Geoff Brown,
Evan Chesterfield, Malcolm Macfarlane and
John Silins. The studies were funded by NRE,
and logistic help was provided trom the Forests
Office at Nathalia. Specific information for this
paper was supplied by Andrew Corrick and
Barbara Baxter (Atlas of Victorian Wildlife) and
Tarmo Raadik (Aquatic Fauna Database).
Lawrie Conole (Ecology Australia) provided
some recent observations and Michael Scroggie
(NRE) made useful comments on a draft. Anne
Morton invited us to write this paper. Many
thanks to all.

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Part One

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Appendix 1. Mammals of Barmah Forest, including the number of records on the Atlas of Victorian
Wildlife. Note that numbers of Atlas records may give a false impression of the abundance of
species subject to special survey effort (e.g. Koalas), while introduced species tend to be under-
recorded, + indicates personal communication. Threatened species categories: End, endangered; Vul,
vulnerable; LRnt, lower risk, near threatened (NRE 2000). I, introduced to Australia.

Common name Scientific name

Atlas records in or Threatened status
near Barmah Forest in Victoria

Short-beaked Echidna
Platypus

Yellow-footed Antechinus
Brush-tailed Phascogale
Common Brushtail Possum
Feathertail Glider

Sugar Glider

Squirrel Glider

Common Ringtail Possum
Eastern Grey Kangaroo
Black Wallaby

Koala

Little Red Flying-fox
Southern Freetail Bat
Eastern Freetail Bat
White-striped Freetail Bat
Gould's Wattled Bat
Chocolate Wattled Bat
Southern Myotis

Lesser Long-eared Bat
Gould's Long-eared Bat
Inland Broad-nosed Bat
Large Forest Bat
Southern Forest Bat

Little Forest Bat

Water Rat

House Mouse

Black Rat

Red Fox

Cat (feral)

Horse (feral)

Sambar Deer

Pig (feral)

Brown Hare

European Rabbit

Vol. 119 (3) 2002

Tachyglossus aculeatus
Ornithorhynchus anatinus
Antechinus flavipes
Phascogale tapoatafa
Trichosurus vulpecula
Acrobates pygmaeus
Petaurus breviceps
Petaurus norfolcensis
Pseudocheirus peregrinus
Macropus giganteus
Wallabia bicolor
Phascolarctos cinereus
Pteropus scapulatus
Mormopterus sp.
Mormopterus sp.
Tadarida australis
Chalinolobus gouldii
Chalinolobus morio
Myotis macropus
Nyctophilus geoffroyi
Nyctophilus gouldi
Scotorepens balstoni
Vespadelus darlingtoni
Vespadelus regulus
Vespadelus vulturnus
Hydromys chrysogater
Mus musculus

Rattus rattus

Vulpes vulpes

Felis catus

Equus caballus

Cervus unicolor

Sus scrofa

Lepus capensis
Oryctolagus cuniculus

[m

r2
T NU OU —

- wae

wey
ba OCONMOoWst

Vul

End

LRnt

127

Murray River Special Issue

Appendix 2. Birds of Barmah Forest. Note that records from imprecisely defined locations do not
generally appear as 5-minute Atlas records and are marked +: this applies mainly to records before
the 1990s. AR, Atlas records in or near Barmah Forest; BLHab, broad local habitats; FL Hab, finer
local habitats; MS, migratory status; TSVic, threatened status in Victoria. Broad and fine habitats:
W, water bird (v, among dense vegetation; m, mainly on bare mudflats; sv, in lightly vegetated
swamps; 0, in open water; t, nesting in or among trees; s, nesting colonially in swamps when flood-
ed; r, along river; f, mainly in farmland or farm dams outside forest). F, forest or woodland bird (rg,
mainly in red gum; b, mainly in box; s, most common in or near swamps or riverbanks; p, most com-
mon on treeless plains within the forest; rd, more common along roadsides or in remnant vegetation
outside the forest; g, more common in gardens outside the forest; no symbol, widespread). O, open
country species, usually found outside the forest (gp, mainly near gravel pits). Migratory or other
temporal status: S, mainly summer visitor (e.g. Sept-April); W, mainly winter visitor (March-Sept):
V, vagrant or rare visitor; X, extinct, not recorded since 19th century. Threatened or introduced sta-
tus: CrEn, critically endangered; End, endangered; Vul, vulnerable; LRnt, lower risk, near threat-
ened; DD, data deficient: I. introduced to Australia. *, reeds or cumbungi; t, rushes.

Common Name Scientific Name AR BLHabFLhab MS TSVic
Emu Dromaius novaehollandiae 44 F S

Stubble Quail Coturnix pectoralis 4 [9]

Brown Quail Coturnix ypsilophora 3 F s DD
Painted Button-quail Turnix varia 15 F

Peaceful Dove Geopelia striata 84 F TE

Diamond Dove Geopelia cuneata 2 tà TE V. Vul
Common Bronzewing Phaps chalcoptera 27 F b

Crested Pigeon Ocyphaps lophotes 42 O

Buff-banded Rail Gallirallus philippensis 4 WwW V

Spotless Crake Porzana tabuensis | W y

Australian Spotted Crake Porzana fluminea + W v

Baillon's Crake Porzana pusilla + W V S
Black-tailed Native-hen Gallinula ventralis 2 W v

Dusky Moorhen Gallinula tenebrosa 14 W r

Purple Swamphen Porphyrio porphyrio 16 W v

Eurasian Coot Fulica atra 3 W o

Bush Stone-curlew Burhinus grallarius T F rd

Great Crested Grebe Podiceps cristatus 4 W o

Australasian Grebe Tachybaptus novaehollandiae 6 WwW o

Hoary-headed Grebe Poliocephalus poliocephalus 1 W o

Great Cormorant Phalacrocorax carbo 52 W t

Little Black Cormorant Phalacrocorax sulcirostris 44 WwW t

Pied Cormorant Phalacrocorax varius 8 W o LRnt
Little Pied Cormorant Phalacrocorax melanoleucos 80 W t

Darter Anhinga melanogaster 21 W rt

Australian Pelican Pelecanus conspicillatus 37 W

Whiskered Tern Chlidonias hybridus 8 WwW SV S LRnt
Silver Gull Larus novaehollandiae 5 W o

Black-winged Stilt Himantopus himantopus t W sv

Red-necked Avocet Recurvirostra novaehollandiae + W sv/o/f

Red-kneed Dotterel Erythrogonys cinctus | W SV

Masked Lapwing Vanellus miles 39 W.O m

Banded Lapwing Vanellus tricolor | O

Red-capped Ployer Charadrius ruficapillus + W m

Black-fronted Dotterel Elseyornis melanops 15 W m

Common Greenshank Tringa nebularia ] W m S
Sharp-tailed Sandpiper Calidris acuminata i W SV S
Red-necked Stint Calidris ruficollis i W m S
Latham's Snipe Gallinago hardwickii 3 W v S

Brolga Grus rubicunda 2 W f Vul
Glossy Ibis Plegadis falcinellus 1 W SV Vul
Australian White Ibis Threskiornis molucca 85 W s

Straw-necked Ibis Threskiornis spinicollis 63 W s

Royal Spoonbill Platalea regia 24 W t Vul
Yellow-billed Spoonbill Platalea flavipes 44 W t

Little Egret Egretta garzetta 4 W t CrEn
Intermediate Egret Ardea intermedia 2] Ww t; SV CrEn

128 The Victorian Naturalist

Part One

Common Name Scientific Name AR
Great Egret Ardea alba 59
Cattle Egret Ardea ibis T
White-faced Heron Egretta novaehollandiae 90
White-necked Heron Ardea pacifica 52
Nankeen Night Heron Nycticorax caledonicus 31
Little Bittern Ixobrychus minutus 3
Australasian Bittern Botaurus poiciloptilus 5
Australian Wood Duck Chenonetta jubata 70
[Magpie Goose] Anseranus semipalmata +
Black Swan Cygnus atratus 42

Plumed Whistling-Duck
Australian Shelduck
Pacific Black Duck
Chestnut Teal

Grey Teal

Australasian Shoveler
Pink-eared Duck
Freckled Duck
Hardhead

Blue-billed Duck

Musk Duck

Spotted Harrier

Swamp Harrier

Brown Goshawk
Collared Sparrowhawk
Wedge-tailed Eagle
Little Eagle
White-bellied Sea-Eagle
Whistling Kite

Black Kite
Black-shouldered Kite
Grey Falcon

Australian Hobby
Peregrine Falcon

Black Falcon

Brown Falcon

Nankeen Kestrel
Southern Boobook
Barking Owl

Barn Owl

Musk Lorikeet
Sulphur-crested Cockatoo
Major Mitchell’s Cockatoo
Little Corella
Long-billed Corella
Galah

Cockatiel

Superb Parrot

Crimson (Yellow) Rosella
Eastern Rosella
Red-rumped Parrot

Blue Bonnet

Budgerigar

Tawny Frogmouth
Australian Owlet-nightjar
Dollarbird

Azure Kingfisher
Laughing Kookaburra
Sacred Kingfisher
Red-backed Kingfisher
Rainbow Bee-eater
White-throated Needletail

Vol. 119 (3) 2002

Dendrocygna eytoni
Tadorna tadornoides
Anas superciliosa
Anas castanea

Anas gracilis

Anas rhynchotis

Malacorhynchus membranaceus

Stictonetta naevosa
Aythya australis
Oxyura australis
Biziura lobata

Circus assimilis
Circus approximans
Accipiter fasciatus
Accipiter cirrhocephalus
Aquila audax
Hieraaetus morphnoides
Haliaeetus leucogaster
Haliastur sphenurus
Milvus migrans

Elanus axillaris

Falco hypoleucos
Falco longipennis
Falco peregrinus
Falco subniger

Falco berigora

Falco cenchroides
Ninox novaeseelandiae
Ninox connivens

Tyto alba

Glossopsitta concinna
Cacatua galerita
Cacatua leadbeateri
Cacatua sanguinea
Cacatua tenuirostris
Cacatua roseicapilla
Nymphicus hollandicus
Polytelis swainsonii

Platycercus elegans flaveolus

Platycercus eximius
Psephotus haematonotus
Northiella haematogaster
Melopsittacus undulatus
Podargus strigoides
Aegotheles cristatus
Eurystomus orientalis
Alcedo azurea

Dacelo novaeguineae
Todiramphus sanctus
Todiramphus pyrrhopygia
Merops ornatus
Hirundapus caudacutus

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129

Murray River Special Issue

Common Name Scientific Name AR BLHabFLhab MS TSVic
Fork-tailed Swift Apus pacificus | FO S
Pallid Cuckoo Cuculus pallidus 19 F S
Fan-tailed Cuckoo Cacomantis flabelliformis 16 F S
Horsfield's Bronze-Cuckoo | CArvsococeyx basalis 22 F S
Shining Bronze-Cuckoo Chrysococcyx lucidus 1] p S
Black-eared Cuckoo Chrysococeyy osculans F F S
Welcome Swallow Hirundo neoxena 132 O,W
White-backed Swallow Cheramoeca leucosternus 2 O gp

Tree Martin Hirundo nigricans 47 in rg S
Fairy Martin Hirundo ariel 6 Q,W 1 S
Grey Fantail Rhipidura fuliginosa 164 F

Rufous Fantail Rhipidura rufifrons | P SV
Willie Wagtail Rhipidura leucophrys 220 FO TE
Leaden Flycatcher Myiagra rubecula 9 F rg S
Restless Flycatcher Myiagra inquieta 58 F

Jacky Winter Microeca fascinans 168 F

Scarlet Robin Petroica multicolor 29 F

Red-capped Robin Petroica goodenovii 73 F

Flame Robin Petroica phoenicea 22 FO p W
Pink Robin Petroica rodinogaster + b WV
Hooded Robin Melanodryas cucullata 4 i b
Golden Whistler Pachyeephala pectoralis 29 F W
Rufous Whistler Pachycephala rufiventris 155 E S
Gilbert’s Whistler Pachycephala inornata 5 F b

Grey Shrike-thrush Colluricincla harmonica 283 F

Magpie-lark Grallina cyanoleuca 112 P,O rg
Crested Shrike-tit Faleunculus frontatus 9] r

[Crested Bellbird] Oreoica gutturalis t F X
Black-faced Cuckoo-shrike Coracina novaehollandiae 141 F

White-bellied Cuckoo-shrike Coracina papuensis 2 F

Ground Cuckoo-shrike Coracina maxima + oO p V
White-winged Triller Lalage sueurii 29 F S
Grey-crowned Babbler Pomatostomus temporalis 16 F rg, rd End
White-browed Babbler Pomatostomus superciliosus 3 E b
White-fronted Chat Epthianura albifrons 3 O,W p
Crimson Chat Ephthianura tricolor + [9] p SV
Southern Whiteface Aphelocephala leucopsis i [9] edge
Speckled Warbler Crhonicola sagittata * F rd
Western Gerygone Gerygone fusca 60 F S
Weebill Smicrornis brevirostris 202 F

Striated Thornbill Acanthiza lineata 53 F

Yellow Thornbill Acanthiza nana 61 F b, g
Brown Thornbill Acanthiza pusilla 7 F s
Buff-rumped Thornbill Acanthiza reguloides 169 F
Chestnut-rumped Thornbill Acanthiza uropygialis + F lignum
Yellow-rumped Thornbill Acanthiza chrysorrhoa 121 O g
White-browed Scrubwren Sericornis frontalis 32 E s

Brown Songlark Cincloramphus cruralis 7 [9] S
Rufous Songlark Cincloramphus mathewsi 35 F rg S
Little Grassbird Megalurus gramineus 7 W v
Clamorous Reed Warbler — Acrocephalus stentoreus 28 W v S
Superb Fairy-wren Malurus cyaneus 195 F 8
White-breasted Woodswallow Artamus leucorynchus 35 FW TE S
Masked Woodswallow Artamus personatus | F S
White-browed Woodswallow Artamus superciliosus 28 i s
Black-faced Woodswallow Artamus cinereus | [9]

Dusky Woodswallow Artamus cyanopterus 139 Ẹ S
Varied Sittella Daphoenositta chrysoptera 36 E

Brown Treecreeper Climacteris picumnus 362 F
White-throated Treecreeper Cormobates leucophaeus 299 F

Mistletoebird Dicaeum hirundinaceum 92 F

Spotted Pardalote Pardalotus punctatus 4l F W
Striated Pardalote Pardalotus striatus 347 F

130

The Victorian Naturalist

Common Name

Scientific Name

AR

Part One

BLHabFLhab MS TSVic

Silvereye

White-naped Honeyeater
Black-chinned Honeyeater
Brown-headed Honeyeater
Painted Honeyeater
White-plumed Honeyeater
[Bell Miner]

Noisy Miner

Red Wattlebird

Striped Honeyeater
Regent Honeyeater
Blue-faced Honeyeater
Noisy Friarbird

Little Friarbird
Richard's Pipit

Singing Bushlark
Diamond Firetail

Zebra Finch
Red-browed Finch
Olive-backed Oriole
White-winged Chough
Pied Currawong

Pied Butcherbird

Grey Butcherbird
Australian Magpie
Australian Raven

Little Raven

Common Blackbird
House Sparrow

Eurasian Tree Sparrow
European Goldfinch
Common Starling
Common Myna

Zosterops lateralis
Melithreptus lunatus
Melithreptus gularis
Melithreptus brevirostris
Grantiella picta
Lichenostomus penicillatus
Manorina melanophrys
Manorina melanocephala
Anthochaera carunculata
Plectorhyncha lanceolata
Xanthomyza phrygia
Entomyzon cyanotis
Philemon corniculatus
Philemon citreogularis
Anthus novaeseelandiae
Mirafra javanica
Stagonopleura guttata
Taeniopygia guttata
Neochmia temporalis
Oriolus sagittatus
Corcorax melanorhamphos
Strepera graculina
Cracticus nigrogularis
Cracticus torquatus
Gymnorhina tibicen
Corvus coronoides
Corvus mellori

Turdus merula

Passer domesticus
Passer montanus
Carduelis carduelis
Sturnus vulgaris
Acridotheres tristis

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Appendix 3. Reptiles of Barmah Forest. Threatened species categories: End, endangered; Vul, vul-
nerable; LRnt, lower risk, near threatened; DD, data deficient (NRE 2000).

Common name

Scientific name

Atlas records in or
near Barmah Forest

Threatened status
in Victoria

Common Long-necked
Tortoise

Murray River Tortoise

Wood Gecko

Marbled Gecko

Olive Legless Lizard

Eastern Bearded Dragon

Tree Goanna

Carnaby's Wall Skink

Large Striped Skink

Garden Skink

Yellow-bellied Water Skink

Grey's Skink

Boulenger's Skink

Peters's Blind Snake

Woodland Blind Snake

Tiger Snake

Red-bellied Black Snake

Eastern Brown Snake

Bandy Bandy

Carpet Python

Vol. 119 (3) 2002

Chelodina longicollis
Emydura macquarii
Diplodactylus vittatus
Phyllodactylus marmoratus
Delma inornata

Pogona barbata

Varanus varius
Cryptoblepharus carnabyi
Clenotus robustus
Lampropholis guichenoti
Eulamprus heatwolei
Menetia greyii

Morethia boulengeri

Ramphotyphlops bituberculatus

Ramphotyphlops proximus
Notechis scutatus
Pseudechis porphyriacus
Pseudonaja textilis
Vermicella annulata
Morelia spilota metealfei

DD

Vul

LRnt
End

131

Murray River Special Issue

Appendix 4. Amphibians of Barmah Forest. Threatened species categories: CrEn, critically endan-
gered; DD, data deficient (NRE 2000).

Common name Atlas records in or Threatened status
near Barmah Forest in Victoria

Scientific name

Peron's Tree Frog Litoria peroni 70
Southern Bullfrog Limnodynastes dumerili 59
Barking Marsh Frog Limnodynastes fletcheri 8 DD
Giant Bullfrog Limnodynastes interioris l CrEn
Spotted Marsh Frog Limnodynastes tasmaniensis 118
Common Spadefoot Toad — Neobatrachus sudelli 8
Bibron's Toadlet Pseudophryne bibroni |
Plains Froglet Crinia parinsignifera 62
Common Froglet Crinia signifera 54
Sloane’s Froglet Crinia sloanei E

Appendix 5. Fish of Barmah Forest. Threatened species categories in Victoria: CrEn, critically
endangered; End, endangered; Vul, vulnerable; DD, data deficient (NRE 2000), I, introduced to

Australia.

Common name

Scientific name

Threatened status in Victoria

Silver Perch Bidvanus bidyanus CrEn
Goldfish Carassius auratus I
Common Carp Cyprinus carpio I
River Blackfish Gadopsis marmoratus

Broadfin Galaxias Galaxias brevipinnis

Flat-headed Galaxias Galaxias rostratus DD
Mosquitofish Gambusia holbrooki I
Western Carp Gudgeon Hypseleotris klunzingeri

Carp Gudgeon (*2 spp.) Hypseleotris spp.

Trout Cod Maccullochella macquariensis CrEn
Murray Cod Maccullochella peelii peelii Vul
Golden Perch Macquaria ambigua Vul
Macquarie Perch Macquaria australasica End
Crimson-spotted Rainbowfish Melanotaenia fluviatilis DD
Oriental Weatherloach Misgurnus anguillicaudatus l
Short-headed Lamprey Mordacia mordax

Southern Pigmy Perch Nannoperca australis

Bony Bream Nematalosa erebi

Rainbow Trout Oncorhynchus mykiss I
English Perch Perca fluviatilis |
Flat-headed Gudgeon Philypnodon grandiceps

Australian Smelt Retropinna semoni

Atlantic Salmon Salmo salar |
Brown Trout Salmo trutta l
Freshwater Catfish Tandanus tandanus Vul
l'ench Tinca tinca l
Carp * Goldfish hybrid |

The Murray River in The Victorian Naturalist

Mr. JG O'Donohue said that the manager of the Kulkyne Station, in a recent
letter to him, mentioned that thousands of cormorants of several species were
nesting at the present time in the Mournpoul sanctuary. In these particular locali-
ty these birds were not hitherto known to breed so late in the season.

NATURAL HISTORY NOTES

From The Victorian Naturalist XXXII (2), 18 June 1916.

The Victorian Naturalist

Part One

The Distribution and Conservation Status of the Reptile Fauna
of the Murray River Region in Victoria

Geoff Brown'

Abstract

The lands surrounding the Murray River, from source to South Australia, support a remarkable diver-
sity of terrestrial reptiles, At least ninety-four species, representing nine families, are known from the
environs of the Murray River in Victoria, and this fauna consists of a blend of arid-adapted elements
from the north-west with temperate elements from the east. Species richness and diversity of regional
reptile assemblages tends to increase along the gradient from cool mesic (Eastern Highlands) to warm
dry regions (Mallee). Skinks are the dominant component (44%) of the overall reptile fauna in terms
of number of species. Skinks also exhibit a distinct decline in proportional species composition along
the environmental gradient from the east to the north-west, in contrast with dragons, geckoes and leg-
less lizards. reptile families that show a steady increase in their proportional representation. Twenty-
four reptile species known from the Murray River area are considered threatened in a national or state
(Victoria or New South Wales) context, and at least another 18 species are considered locally threat-
ened, The majority of these threatened species are found in the north-west of the state, and most are
classified as such because they are either at the limit of their distributional range in northern Victoria,
and consequently occur only rarely, or because they are habitat specialists and considered to be disad-
vantaged by habitat change or loss. The major determinants of reptile occurrence (and decline) are

briefly discussed. (The Victorian Naturalist 119 (3), 2002, 133-143)

The Murray River

The Murray River is one of Australia's
great rivers and, together with the Darling
River, forms the catchment for a vast por-
tion of south-eastern continental Australia.
The headwaters of the Murray River rise in
the Great Dividing Range at about 1430 m
on the New South Wales-Victoria border
and then flow westwards for 2530 km to
the Southern Ocean in South Australia
(Young and Hillman 2001).

Within Victoria the Murray River tra-
verses a range of climatic zones (or *natur-
al’ regions, after Conn 1993) over approxi-
mately 1900 km, extending from the high-
altitude Eastern Highlands at its source,
through the flat riverine plains of the
Riverina region of north-central Victoria,
to the Murrày Mallee in the semi-arid
north-west of the state, This environmental
gradient is reflected in the climate, in
increasing temperature and decreasing
rainfall along a southeast-northwest axis,
as well as in vegetation patterns and fauna
assemblages. The vegetation along the
Murray River ranges from montane forests
and alpine woodlands of the mesic high-
lands of the Great Dividing Range in the
east, through dryland and riverine wood-
lands (typically dominated by Black Box
Eucalyptus largiflorens, Grey Box E. micro-

Anhur Rylah Institute for Environmental Research,
Department of Natural Resources and Environment,
123 Brown Street, Heidelberg, Victoria 3084

Vol. 119 (3) 2002

carpa or River Red Gum £. camaldulensis)
of the plains of the Riverina, to the semi-arid
habitats on dune-fields in the Mallee,

The dominant land use along the Murray
River flood-plain is agriculture. Over the last
century widespread clearing has enabled
intensive and extensive cropping and graz-
ing across large areas, particularly in the
Riverina. Timber harvesting on public land
is carried out, mostly in the highlands and
riverine forests of the Riverina, although the
target species and the purposes for which
they are harvested vary. Fuelwood (dead and
down material) is also a relatively important
industry, for both commercial operators and
individual licensees.

While sections of the Murray River area,
notably the mallee habitats in the north-
west and the open forests of the Eastern
Highlands, are relatively well vegetated,
much of the remaining plains have suf-
fered extensive clearing, such that only a
small and highly fragmented vegetation
mosaic now exists. This mosaic includes
mallee or woodland associations and con-
sists principally of linear remnants along
streamsides and roadsides, a handful of rel-
atively large blocks that are typically river-
ine State Forests or State Parks, and a myr-
iad of small vegetation parcels, generally
located on private land. The amount of
suitable habitat within the Murray River
catchment, and its current management,

133

Murray River Special Issue

has undeniable implications for the effec-
tive conservation of reptiles.

Species composition and distribution of
reptiles along the Murray River

Ninety-four species of reptile, represent-
ing nine families, are known from the
environs of the Murray River in Victoria
(Table 1). These families include tortoises
(3 species), dragons (9), geckos (9), legless
lizards (8), skinks (41), goannas (2),
pythons (1), blind snakes (4) and elapid
snakes (17) (Atlas of Victorian Wildlife,
NRE). These species incorporate an
extremely diverse array of sizes, life-
forms, habitat requirements and distribu-
lions. Taxonomy for reptiles in this paper
follows Cogger (2000), except for Hooded
Scaly-foot Pygopus schraderi, which fol-
lows James et al. (2001) .

This reptile fauna consists of a blend of
arid-adapted elements in the north and west
with temperate elements from the south and
east. There is à strong trend of increasing.
species richness and increased diversity of
taxonomic composition of regional reptile
assemblages along the gradient from cool
mesic to warm dry regions. Skinks are the
most Conspicuous component of the overall
fauna in terms of both numbers of species
and life-forms, They comprise 44% of the
reptile species known from the area, and
exhibit a marked decline in proportional
composition of the reptile assemblage along
the environmental gradient from the Eastern
Highlands in the east (68%), to the Mallee
region in the north-west (44%) (Table 1). A
contrasting trend is apparent for dragons,
geckos and legless lizards, reptile families
that show a steady increase in their propor-
tional representation in the species assem-
blage along the same environmental gradi-
ent. The highlands in the east exhibit the
lowest species richness (27%).

Reptiles of the Murray River can be cate-
gorised into several groups, each of which
reflects a distinctive distribution. Two
major groups represent species that are rel-
atively widespread across the region, One
group, including species such as Common
Long-necked Tortoise Chelodina longicol-
lis, Large Striped Skink Crenotus robustus,
Boulenger's Skink Morethia houlengeri,
Stumpy-tailed Lizard Trachydosaurus
rugosus and Common Sealy-foot Pygopus

134

Fig. 1. The Regal Striped Skink Crenotus regius
is an agile terrestrial skink that, in Victoria, is
found in semi-arid grasslands and woodlands of
the north-west.

lepidopodus, occurs in the Mallee and
plains, but with few representative species
in the highlands. The second group, which
embraces such species as Tree Dragon
Amphibolurus muricatus, Blotched Blue-
tongued Lizard Tiliqua nigrolutea, Garden
Skink Lampropholis guichenoti and Red-
bellied Black Snake Pseudechis porphyria-
cus, occurs widely in the highlands and
plains, but not in the Mallee.

The other groups have a more limited dis-
tribution across the region. A group of
species occur only in the Mallee (e.g.
Mallee Dragon Ctenophorus fordi, Regal
Striped Skink Crenorus regius (Fig. 1),
Coral Snake Simoselaps australis,
Mueller's Skink Lerista muelleri) and at the
other end of the environmental gradient are
those species that occur primarily within the
highlands (e.g. Mountain Dragon
Tympanocryptis diemensis, White's Skink
Egernia — whitii, Delicate Skink
Lampropholis delicata, | Highland
Copperhead Austrelaps ramsayi). The
Tessellated Gecko Diplodactylus tessellatus
is an example of a true arid-zone species
(Rawlinson 1971) that is found only in the
extreme north-west. Its distribution in
Victoria is marginal to its main occurrence
in the drier interiors of the eastern and cen-
tral Australian states (Cogger 2000), and
follows the Murray River corridor (Atlas of
Victorian Wildlife). Conversely, the Garden
Skink and the Eastern Water Skink
Eulamprus quoyii are true mesic species
which also appear to use the mesic environ-
ments of the Murray River and Darling
River respectively, to extend their main dis-
tributions from eastern Australia. The

The Victorian Naturalist

Eastern Water Skink is known in Victoria
only from the Mallee around the confluence
of the Darling and Murray Rivers.

Overall, the Murray River reptile fauna
includes three aquatic species (3%), all tor-
toises, 65 species (69%) that are consid-
ered terrestrial, seven arboreal (7%)
species, eight species (9%) that are semi-
arboreal, and 11 species (12%) that are
fossorial (Table 1). The proportion of
species categorised as terrestrial (e.g. many
skinks and elapid snakes) systematically
decreases from east to west. Conversely,
the proportion of fossorial species and, to a
lesser extent, those categorised as arboreal,
increases from east to west. Fossorial
species in the Murray River area consist of
several species of legless lizards, the skink
genera Hemiergis and Lerista, blind snakes
and two species of uncommon elapid
snakes, Most of these species are restricted
to the Mallee.

Conservation status of reptiles

The conservation status of many reptiles
in rural environments of Victoria, outside
of the highlands, is alarming - while a small
number of species (e.g. Marbled Gecko
Christinus marmoratus, Boulenger's
Skink) are relatively widespread and com-
mon, most species are patchily distributed
(restricted to isolated remnant vegetation)
and occur in very low numbers (Brown and
Bennett 1995). There has been a wide-
spread decline and local extinction of rep-
tiles in the rural environment.

Overall, at least 94 reptile species have
been recorded from the Murray River area,
about one-quarter of which (25 species) are
considered officially threatened in a national
or state ( Victoria or New South Wales) con-
text (Table 1). At least another 18 species
are considered locally threatened, on the
basis of limited distribution and relatively
low numbers of records (see Table 1).

These threatened Victorian reptile species
include a locally extinct species, the Small-
scaled Snake Oxyuranus microlepidota, two
species that are ‘Critically Endangered’,
seven that are ‘Endangered’, five that are
‘Vulnerable’, eight that are ‘Lower Risk —
near threatened’ and another that is *Data
Deficient’ (Table 1). The Common Death
Adder Acanthophis antarcticus was also
considered extinct in Victoria until recently,

Vol. 119 (3) 2002

Part One

when an individual was recorded in the
north-west corner of the state (Atlas of
Victorian Wildlife, NRE). The majority
(18) of these threatened species are found in
the north-west of the state, whilst three are
known from the Eastern Highlands and
twelve from the Riverina.

Many of these species are classified as
threatened because they are at the limit of
their range in northern Victoria, and conse-
quently occur only rarely. For example,
species such as Tessellated Gecko, Beaked
Gecko Rhynchoedura ornata, Hooded
Scaly-foot Pygopus schraderi, Desert
Skink Egernia inornata, Yellow-faced
Whip Snake Demansia psammophis and
Western Brown Snake Pseudonaja
nuchalis all have broad continental distrib-
utions, yet extend only marginally into
Victoria, Conversely, the Striped Legless
Lizard Delma impar and Alpine Water
Skink Eulamprus kosciuskoi are examples
of threatened species that are typically
confined to more mesic environments of
south-eastern Australia.

Some of these threatened species, includ-
ing the Striped Legless Lizard, Hooded
Scaly-foot, Murray Striped Skink Crenotus
brachvonyx, and Carpet Python Morelia
spilota metcalfei, are classified because
they are habitat specialists and considered
to be detrimentally affected by habitat
change or loss (Robertson ef al. 1989;
Cogger et al. 1993). The Striped Legless
Lizard, for example, is in a demonstrable
state of decline because it is restricted to a
particular habitat, native grassland, which
has become exceedingly limited im area and
is fragmented and prone to further reduc-
tion (CNR 1992: Dorrough and Ash 1999).

Two species, the Hooded Sealy-foot and
the Alpine Water Skink, are considered
‘Critically Endangered’ in Victoria. The
occurrence in Victoria of the Hooded
Scaly-foot is known from three historical
(undated) museum records, two records
from the Riverina in the 1990s (Atlas of
Victorian Wildlife, NRE) and most recent-
ly from the vicinity of Terrick Terrick
National Park (P Robertson pers. comm. ).
At the opposite end of the environmental
gradient, the Alpine Water Skink is known
from a few localities, generally in alpine
bogs, in the Eastern Highlands (Atlas of
Victorian Wildlife, NRE).

135

Murray River Special Issue

Several other species that occur in the
Murray River area of Victoria have appar-
ently declined over recent years or are per-
ceived to be threatened, especially by habitat
loss or modification. Although abundant
where it occurs, the Mallee Dragon appears
to be restricted in Victoria to mallee habitat
containing Triodia (Sadlier and Pressey
1994). Triodia is an important habitat
requirement for other species, notably
Brooks’s Striped Skink Ctenotus brooksi,
Obscure Skink Morethia obscura and
Western Blue-tongued Lizard Tiliqua occip-
italis, which show only a limited or patchy
distribution in the state.

In general, the greatest threats to reptiles

(i.e. those affecting the largest number of

reptile species nationally) are habitat clear-
ance, habitat modification, overgrazing by
stock, urban development, and predation by
introduced predators (Cogger et al. 1993).
Brown and Bennett (1995) found that the
majority of reptile species in the Victorian
Riverina, the most alienated natural region
through which the Murray River flows,
occurred in very low numbers or showed a
patchy distribution. Their data, when cou-
pled with existing distribution records and
available biological and ecological data,
signal low survival status and hence seri-

ous concern for the long-term security of

the reptile assemblage of this region.
Although some reptile species are natural-
ly rare in the Riverina, some are also rare
because they are likely to have suffered
more than others from the effects of clear-
ing or disturbance. Large species, such as
Tree Goanna Varanus varius (Fig. 2) and
some of the elapid snakes (e.g. Tiger Snake
Notechis scutatus, Red-bellied Black
Snake), which have relatively large home
ranges (in the order of hectares) are likely to
be excluded from patches that are small and
isolated. These larger species have naturally
low densities, and therefore are especially
prone to reduction in suitable habitat. The
Tree Goanna, a solitary species known to
inhabit only forest and woodland communi-
ties, typically shows a strong affinity with
specific trees within its home range, which
provide hollows for shelter (Greer 1989)
and basking sites. Hence, timber harvesting
is also likely to disadvantage this species.
Some of the small reptile species, especially
skinks and legless lizards, are adversely

136

Fig. 2. Juvenile Tree Goanna Varanus varius, a
large semi-arboreal species that is found in for-
est or woodland environments across much of
Victoria.

affected by the modification of the structure
of the ground-level strata (Brown and Bennett
1995). Grazing, the commercial collection of
firewood, and the removal of litter and timber
debris by land-holders are all common prac-
tices along the Murray River, particularly in
the Riverina, and all involve the removal or
alteration of elements of the ground-layer that
many reptile species require for shelter, forag-
ing or reproduction sites.

Processes influencing the distribution
and conservation status of species

The major impact of land use along the
Murray River within Victoria has undoubt-
edly been the dramatic reduction in forest
cover. This has been particularly acute in
the Riverina, where forest cover dimin-
ished by 8495 in the period 1869-1987
(Woodgate and Black 1988). A major con-
sequence of this extensive clearing across
the Basin has been the fragmentation and
isolation of remnant vegetation, such that
an array of different-sized patches now
pepper the landscape. Many of these patch-
es are small, less than 10 ha, and isolated,
For the reptiles (and indeed, all fauna), the

The Victorian Naturalist

Part One

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Vol. 119 (3) 2002

Special Issue

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The Victorian Naturalist

138

Part One

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uoa | AQEL

———————————————————————————————————————————————————————————— M——'Á

139

Vol. 119 (3) 2002

Murray River Special Issue

immediate consequences are twofold: there
is an overall reduction in the total amount
of habitat that is available, and hence the
overall size of regional populations; and
secondly, the ability of species to persist in
the developed landscape largely depends
on their capacity to use small and isolated
remnant habitats.

Brown and Bennett (1995), using reptiles
of the Victorian Riverina as their focus,
found that the distribution and abundance
of reptiles in the rural environment is influ-
enced by a number of factors, operating at
different levels, including climate, land-
scape factors, major habitat types, micro-
habitat, grazing and timber removal. These
are explored briefly below,

Climatic gradient

The environmental gradient (reflected in
climate and vegetation) provides the over-
riding natural factor affecting reptile distri-
bution. This is most conspicuously mani-
fested in the biogeographical trend of
inereasing species diversity and abundance
from the mesic east to drier, warmer habi-
tats in the west. Reptiles are ectothermic,
relving on external heat sources to deter-
mine body temperature, and using behav-
ioural and physiological mechanisms to
regulate it. As a consequence, more reptile
species are found in arid and semi-arid
habitats than temperate ones; arid zones in
Australia support an extraordinary diversi-
ly of reptiles, in part because of their phys-
iological and ecological adaptations to a
dry, warm environment (Pianka 1986;
Bradshaw 1986; James and Shine 2000).

Landscape factors

The pattern of remnant vegetation in the
rural landscape influences the distribution
of reptiles in several ways. Most species
of reptiles depend on remnant areas of
habitat and rarely occur in cleared farm-
land. Remnant vegetation displays great
variation in its reptile fauna, and linear
remnants, particularly roadside vegetation,
typically yield the highest reptile densities
and richness. This is partly attributable to
the typically greater degree of substrate
complexity on roadsides (compared with
other remnant types), a result of the low
levels or absence of grazing and the clear-
ing of vegetation and coarse woody
debris. Roadsides are usually managed by

140

is a terrestrial lizard that is generally restricted
in Victoria to woodlands and shrublands in the
north-east.

government agencies (Brown and Bennett
1995),

The Murray-Darling Basin overall, and in
Victoria particularly, has suffered an extra-
ordinary degree of habitat loss (Woodgate
and Black 1988; MDBMC 1987; Goldney
and Bowie 1990), an unfortunate conse-
quence of which, at least for wildlife, is the
creation of a more homogeneous landscape.
Importantly, most of this clearing was
undertaken on freehold land, and freehold
land in this area now has less than 296 tree
cover (Land Information Group, NRE).
The effects on reptiles of clearing some
vegetation communities are indisputable. In
mallee vegetation, for instance, 70-95% of
the original herpetofauna has been perma-
nently lost from areas that have been
cleared (Rawlinson 1981; Cogger 1989).

Another landscape factor which has a con-
siderable impact on many species, particular-
ly those that have small home ranges or are
relatively immobile, is isolation (or degree of
connectedness) of suitable habitat. The major
impacts for reptiles arising from isolation
appear to be a reduction in species numbers, à
greater incidence and persistence of generalist
species and an increased vulnerability to habi-
tat disturbance or degradation. Recent studies
have shown that even common species that
are considered habitat generalists are affected
by habitat clearance and the subsequent isola-
tion of populations (Sarre 1995),

The fragmentation process affects reptiles
primarily by progressive reduction in the
amount of suitable habitat and the associated
increase in the degree of isolation shown by
many of these fragments. Also, fragmented
remnants in farmland are more vulnerable to
disturbance and degradation across edges.

The Victorian Naturalist

Major habitat type

Major habitat types provide another factor
affecting reptile occurrence. In the
Victorian Riverina for instance, Black Box
woodlands support more reptile species and
families and reveal greater reptile densities,
than River Red Gum forests (Brown and
Bennett 1995). Although the major habitat
types are correlated, in part, with the envi-
ronmental gradient, some importance can
be attributed to their influence as determi-
nants of reptile distribution and abundance.

River Red Gum forests are noticeably
depauperate in terrestrial reptile species;
those species typically resident are relative-
ly large, generalist species (Yellow-bellied
Water Skink Eulamprus heatwolei, Tree
Goanna, Tiger Snake, Red-bellied Black
Snake) or arboreal species (Marbled Gecko,
Carnaby's Wall Skink Cryptoblepharus
carnabyi, Tree Skink Egernia striolata,
Carpet Python) regularly recorded in semi-
aquatic and flood-prone environments, usu-
ally in low numbers (LCC 1983; Brown and
Bennett 1995). River Red Gum forest domi-
nates watercourses and low-lying areas vul-
nerable to periodic flooding across the
riverine plains. It is probably because this
forest type experiences regular inundation
that the proportion of terrestrial reptile
species is relatively low, and that most fos-
sorial species have rarely been recorded.

Microhabitats

Microhabitat characteristics are important
determinants of reptile occurrence, after
accounting for the environmental gradient.
Generally, greater reptile numbers are cor-
related with increased structural hetero-
geneity, particularly in the ground-level
and shrub strata, Open grazed grassland, or
heavily-grazed woodland with bare ground
are likely to support few, if any, reptiles.

Structural complexity near ground-level is
clearly important to many reptile species,
including a range of skinks (Downey and
Dickman 1993; Hutchinson 1993), legless
lizards (Rauhala 1993; Robertson and
Edwards 1994) and elapid snakes (Shine
1987; Schwaner 1991). For example, the
value of even modest vegetation cover for
the habitat generalist Boulenger's Skink
emerged during drought conditions in west-
ern New South Wales (Henle 1989).

Microhabitat is known to be an important
element of niche partitioning in fossorial/

Vol. 119 (3) 2002

Part One

terrestrial species, even more so than broad
habitat type. To illustrate, the Spotted
Burrowing Skink Lerísta punctatovittata is
an arid-zone resident and a habitat general-
ist for which abundance has been correlated
with litter build-up and other ground-level
substrates, such as timber and discarded
corrugated iron, that afford shelter (Henle
1989). Other fossorial/terrestrial reptile
species have also shown strong connections
with various components of the terrestrial
microhabitats, such as bark that has been
shed naturally or piled following forestry
activities (Bynoe’s Gecko Heteronotia
binoei; Bustard 1968a), well-developed lit-
ter (Three-toed Skink Hemiergis decresien-
sis; Robertson 1981), litter depth and pro-
portions of bare ground and grass sub-
strates (Coventry’s Skink Niveoscineus
coventry’; Brown and Nelson 1993).

A number of arboreal or semi-arboreal
reptile species occur along the Murray
River, and all require, to varying degrees,
elevated microhabitats as foraging, basking
or shelter sites, These species include
geckos, dragons, skinks and goannas, and
while some of them (e.g. some dragons,
Sand Goanna Varanus gouldii) are flexibly
arboreal, other species seem to be depen-
dent on elevated microhabitats. For exam-
ple, numbers of the strictly arboreal gecko
Tree Dtella Gehyra variegata in western
New South Wales are strongly correlated
with the size, shape and density of Black
Box (Henle 1990).

Modification of the microhabitat is clear-
ly one of the most important processes gov-
erning reptile occurrence and abundance,
and can arise in myriad different ways (e.g.
fire, introduced animals, salinity changes,
recreational impacts, altered water regimes,
pollutants and biocides). However, the two
activities that arguably have the most influ-
ence on habitat structure in rural environ-
ments along the Murray River are grazing
and timber removal (both standing timber
and coarse woody debris).

Grazing

Though few studies exist which examine
the effects of different grazing regimes on
reptile occurrence and density, the limited
evidence suggests that excessive grazing
has a significant impact on many terrestrial
reptiles, both in Australia (where much of
the evidence is inferential) (Recher and

141

Murray River Special Issue

Lim 1990; Cogger ef al. 1993; Scougall et
al, 1993; Smith er al. 1996; Brown 2001)
and overseas (Busack and Bury 1974;
Jones 1981; Brooks 1999). Grazing affects
the microhabitat principally through the
simplification of the understorey and
fround-layer, seen in lowered structural
and floristic diversity, altered floristic
composition and soil disturbance. Species
which live underground (e.g. Aprasia spp.,
Ramphotyphlops spp., Coral Snake, Bandy
Bandy Vermicella annulata) or forage in
the sub-soil and litter layers (e.g.
Hemiergis spp., Lerista spp.) are likely to
be most affected by impacts such as soil
compaction and soil erosion.

Timber removal

Timber extraction clearly disadvantages
those arboreal species that depend on elevat-
ed sheltering, basking and foraging sites (e.g.
many geckos, Tree Skink, Tree Goanna)
simply through the reduction of suitable
habitat, The Tree Goanna, for example, uses
tree hollows for shelter, exposed trunks and
branches for basking, and includes eggs or
nestlings of tree-nesting birds in its diet
(Greer 1989; Green and King 1993). While
limber extraction adversely affects a number
of reptile species, this practice is known to
benefit some species. For example, stumps
resulting from the harvesting of Callitris
eventually provide suitable microhabitats for
Bynoe's Gecko and Tree Dtella, relatively
widespread gecko species that are able to
exploit the apertures that form between the
bark and the stump (Bustard 1968b).

Many terrestrial species which rely on
fallen timber for cover, activity and breed-
ing sites are also disadvantaged by fire-
wood removal. Eleven species were found
to exploit Jogs in the Riverina (Brown and
Bennett 1995); many species closely asso-
ciated with them either for shelter or activ-
ity sites (e.g. Carnaby’s Wall Skink,
Garden Skink). The value of logs for a
number of terrestrial and/or arboreal reptile
species is well known (Webb 1985; Greer
1989). The decline of the Carpet Python in
the semi-arid region of Victoria and New
South Wales has been attributed to a num-
ber of different processes, amongst them
the loss of hollow-bearing trees due to tim-
ber extraction, and firewood collection
(Sadlier 1994; Shine and Fitzgerald 1996),

142

Conclusions

In summary, habitats along the Murray
River are diverse and this is reflected in
the wonderful variety of reptiles that are
known from the river’s environs. Whilst
the conservation status for many of these
species is stable in Victoria, a substantial
number of species are considered threat-
ened. The threats to reptiles generally vary
according to land management practices
along the river’s course. What remains
constant, however, is that the responsibility
for ensuring the survival of reptiles, indeed
all native fauna, is ours.

Acknowledgements

This paper benefited from the scrutiny of Rick
Shine (University of Sydney), Nick Clemann,
Paul Close and Phoebe Macak (all Arthur Rylah
Institute for Environmental Research) and the
provision of information by Peter Robertson
(Wildlife Profiles Pty Ltd), the Atlas of
Victorian Wildlife and Land Information Group
(both NRE). Much of what is presented here
was stimulated by discussions and collaborative
research with ecologists, particularly Dr Andrew
Bennett (Deakin University).

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143

Murray River Special Issue

Tadpoles of South-eastern Australia: a Guide with Keys

by Marion Anstis

Publisher: Reed New Holland, Frenchs Forest 2002, 281 pp, ISBN 1876334630.
RRP $59.95

In the context of the Murray River, or
indeed of any and all aquatic habitats in
south-eastern Australia, this is the book I
wish I'd written! But I didn't; and even if I
had I wouldn't have been able to do it half
as well as Marion Anstis has, Australian
herpetology has been extraordinarily
enriched by contributions from people with-
out primary training in the field: such shin-
ing examples as Mike Tyler, Eric Worrell,
George and John Cann, John Coventry and
Charles Tanner come readily to mind. And
now Marion Anstis has joined that distin-
guished company, with a flourish! Studying
frog life histories has been a spare-time
occupation for Marion, a full-time music
teacher, for the years of field and laboratory
work, photography, drawing and writing
that went into the book. What if she had
been in a position to pursue her passion for
tadpoles full-time? — I dare say we would
now be in a position to savour her books on
the tadpoles of Australia, the southern hemi-
sphere or the world...

The subtitle of Marion's book — 4 Guide
with Keys — sums up its outstanding
strength: it responds comprehensively to
the naturalist's need for assistance in iden-
tifving tadpoles. If you want to know more
about how tadpoles work, their anatomy
and physiology, their behaviour, ecology
and evolution, then you can turn to Roy
McDiarmid and Ron Altig’s encyclopaedic
Tadpoles: The Biology of Anuran Larvae
(University of Chicago Press, 1999), But if
your interest is in your local tadpoles, the
sorts of places to find them in, how they
develop, and most particularly the sorts of
frogs they'll turn into, then Marion's book
is the very one you’ve been waiting for,

Tadpoles, it must be acknowledged, are
difficult to identify; undoubtedly that’s
why it’s taken so long for us to come to
terms with them. For the taxonomist the
saving grace of tadpoles is the mouth
structure, of which more in a moment. For
the rest, they are seriously deficient in

144

measurable or countable things such as
appendages, scales, fin rays or body seg-
ments; you have to rely on much less
quantifiable characters, including body
shape and form, fin depth and eye position.
Very occasionally a tadpole has a bizarre,
the-only-one-of-its-kind characteristic: the
Glandular Frog Litoria subglandulosa of
northern New South Wales provides one
such example. For the most part, however,
tadpoles are tadpoles are tadpoles. But
their mouths are truly a marvel, the more
so because not a vestige of the larval
mouth structure remains in the frog, which
has pretty regular jaws and teeth much
more resembling yours or mine. Most
commonly in tadpoles the jaws have a
beak-like configuration, rather like that of
a squid or octopus; they're made of keratin
(horn) and are jet-black, with serrated
edges. Surrounding them, typically, is a
fleshy, folded disc, roundish or oval in out-
line, fringed by a single row or multiple
rows of short papillae. The area between
the jaws and the papillary fringe bears a
number of rows of tiny, keratinous teeth,
giving the appearance of a series of minia-
ture black combs. The size and shape of
the jaws, the disposition of the papillae
and, most particularly, the number of rows
of teeth and their arrangement, show con-
sistent variations among species, meaning
that the details of mouth structure are usu-
ally the most useful attribute of tadpoles
for the identifier to home in on. And I
guarantee that your first glimpse of a tad-
pole's mouth structure will absolutely
delight you with its neatness, its complexi-
ty, its symmetry. The downside? — it’s too
small to really appreciate without a hand-
lens or — vastly better — a stereoscopic
microscope. Even then you need to gently
manipulate the mouthparts if you want to
be sure of seeing the fine details properly.
(The lower photograph on p 141 of the book
gives you a good idea of how much — or
how little! — of the mouth structure you can

The Victorian Naturalist

discern just by inspection. And the drawing
below reveals what it’s really like.)

What all this means, in the end, is that
mere descriptions of tadpoles, no matter
how good they are, are not going to unrav-
el the mysteries of their identity. You need
a key, and you need copious, excellent
illustrations. 1 don't think you could ask
any tadpole book to fulfil those prescrip-
tions with greater inclusiveness, elegance
and accuracy than Marion’s has achieved,
More than that, she has squarely confront-
ed the problem that tadpoles change so
much throughout their development that
what will suffice to identify them at one
stage will be of no use whatever at another.
The magical mouth morphology, for
instance, is not fully expressed in early lar-
vae, so it isn't of much help. But you may
well be put on the right track by, say, the
presence or absence of external gills in à
very early stage, structures which vanish
altogether as the tadpole grows.

So something over one-sixth of the book
is devoted to two very substantial and
detailed keys, one to tadpoles and the other
to eges and embryos. To prevent the tad-
pole key from becoming too unwieldy, tad-
poles are first classified into 17 general
body types and their mouths (or *oral
discs’) into 15. These are individually illus-
trated with line drawings; so when you
encounter in the key ‘Body type 10, oral
disc type 8° (for instance), you can find out
at once what you should expect to see. But
there are also plentiful illustrations of other
features of tadpoles: the position of the
eyes, the shape of the body and tail, the
locations of the spiracle (opening of the gill
chamber) and the anus, the various pigmen-
tation patterns, and so on, Similarly, for the
egg and embryo key, there are colour pho-

tographs of egg-masses and drawings of

egg and embryonic features.

Preceding the keys there is a general
introductory section covering the classifi-
cation of frogs, methods of collecting and
caring for tadpoles (with reminders about
legal and ethical constraints), issues relat-
ing to the conservation of frogs and tad-
poles, and — perhaps most useful of all — an
illustrated account of the stages of tadpole
development, from Stage | (newly fertil-
ized egg) to Stage 46 (newly metamor-
phosed froglet). The staging system is a

Vol. 119 (3) 2002

Part One

standard one developed by American
authors, but Marion has adapted it to better
fit the local scene, and included extra
drawings which illustrate some of the vari-
ation among Australian species, If you're
describing tadpoles or taking measure-
ments of their dimensions, it's crucial to
note what stage they're at, and on pages
13-19 you'll find the means to do so.

The bulk of the book (190 pages) consists
of species accounts; no less than 84 of the
89 species and subspecies of frog known to
occur in New South Wales, Victoria and
Tasmania are included. Since more than
half of them were barely known or com-
pletely unknown before Marion's work, this
is indeed a remarkable accomplishment.
What is just as meritorious is that there are
colour photographs of living tadpoles of all
but three forms. Each account is headed
with a colour photograph of an adult frog
and a distribution map of the species; then
follow descriptions of breeding sites, eggs,
embryonic and larval development, tadpole
behaviour, metamorphosis and similar
species. The tadpole photographs usually
include several views and sometimes sever-
al stages (the Holy Cross Toad Notaden
bennetti and the Red-crowned Toadlet
Pseudophryne australis are good examples;
even though the caption of the P. australis
photo is awry!). And even if you don't like
tadpoles much you'll have to admit that
some of them are simply gorgeous to look
at: the Dwarf Tree Frog Litoria fallax on p
117, for example, or Haswell's Froglet
Paracrinia haswelli on p 234. As well as
the photos there are excellent drawings, first
of a tadpole in left lateral view (to show the
position of the spiracle, which is an impor-
iant aid to identification, but rarely shows
up in photos) and, second, of the oral disc.
Although I don't believe tadpoles will ever
be easy to identify, | do think that if you
take full advantage of all the explanatory
and illustrative material that accompanies
Marion's keys, and if you make yourself
generally familiar with the species accounts,
you'll have a better chance than people any-
where else in the world of establishing what
sort of frog the particular tadpole in your
sights is going to turn into.

Errors and inconsistencies are few and
minor as far as | can see. Tasmania's Moss
Froglet is listed as Brvobatrachus nimbus;

145

Murray River Special Issue

it should (according to the latest opinion)
be included in Crinia. Here and there a
spot on a distribution map has gone astray:
for instance the Victorian Smooth Froglet
Geocrinia victoriana has acquired an out-
lier population near Jerilderie, NSW, and
an even more remote one somewhere in
the vicinity of Sydney! On the other hand,
Banjo Frogs Limnodynastes dumerilii, cor-
rectly said to occur on Flinders Island,
Tasmania, don't feature there on the map,
and their occurrence on King Island,
Tasmania, is overlooked altogether.
Although there is a Glossary on p 8, you'll
search it in vain for such technical terms as
adhesive organ, pronephric bulge, and sto-
modaeal pit. And finally — in the interests
of user-friendliness — | would have liked
the page-headings to be more informative.
Quite frequently, in the species accounts,

you'll find yourself looking at a page of

text and illustrations with the species name
inconspicuous or absent: pages 134, 136
and 138-139 are a few among many exam-
ples. It means that if you're flipping
through the book and glimpse a tadpole or
mouth like the one you're trying to identi-
fy, you then have to turn back a page to
find the name. Use of species names as
page-headings would have avoided this
minor inconvenience.

In sum, then, if you want to take advantage
of the fact that — at last! — being able to iden-
lify south-eastern Australian tadpoles is a
realistic prospect, this is the book you sim-
ply must have. Even if you're not a tadpole
enthusiast, you may buy the book (as one
friend has done) not for its usefulness, but
just because it’s such an outstanding exam-

ple of how attractive modern field guides
can be. In either case, warmest congratula-
tions to Marion Anstis on a truly invaluable
piece of work. Additional commendations to
the Australian Biological Resources Study,
to Rio Tinto and to the World Wide Fund
for Nature, all of whom supported Marion’s
research and the production of the book.
Well done, Reed New Holland, too, for
achieving a quality of production that
matches the quality of the research, and for
holding the price at a reasonable level.

And my favourite tadpole? — the one fea-
tured on pp 130-132, the Heath Frog
Litoria littlejahni. Why? Well, for one
thing my respected mentor and colleague
Murray Littlejohn and | were the first to
study the life history and describe the tad-
pole of this species. That it was named for
Murray, the person who has done more
than anyone else to enrich our knowledge
of Victorian frogs, is particularly pleasing.
If it’s permissible to dedicate a book review
then I dedicate this one to him, on the occa-
sion of a milestone birthday that he
achieves on the very day that I’m writing
this. But finally, the Heath Frog tadpole is
just a delightful animal — big, velvety-
black, always active (Marion describes it as
‘agile’), easy to feed and care for, and near-
ly always successfully reaching metamor-
phosis in suitable captive conditions. If
only every home could have one!

Angus Martin

Senior Associate in Zoology
Department of Zoology
University of Melbourne
Victoria 3010

Email biotica(dbigpond.com

he tadpole of Peron's Tree Frog Literia peronii, a common and abundant species in eastern and
northern Victoria, including the Murray Valley. Photos by Marion Anstis.

146

'The Victorian Naturalist

Part One

Rivers as Ecological Systems: The Murray-Darling Basin

Edited by WJ Young

Publisher: Murray-Darling Basin Commission, 2001. GPO Box 409, Canberra, ACT 2061
Paperback, 336 pp. colour illustrations, 860. ISBN 1876830034

This ambitious book aims to summarise
our knowledge of the ecology of the river
systems of the Murray-Darling Basin,
especially the effects of flow and flow
change in these ecosystems. This is a laud-
able and timely objective as the availabili-
ty and quality of water for irrigation, con-
sumption and the environment is one of the
major resource management issues cur-
rently facing Australia and the world.
Overall, the book achieves this aim, pro-
viding a comprehensive summary of scien-
tific and other literature related to the
Murray-Darling Basin up to about 1999, ]
use the word summary rather than review
deliberately here because review implies
some critical evaluation and this book
tends to report other people's results
uncritically. The book seems to have been
edited very consistently; the chapters are
cross-referenced with no obvious changes
in style between them, although the level
of detail does vary between chapters. The
contributing authors generally represent
scientists with considerable expertise on
their particular topic within the Basin, The
book is also a remarkable collection of
photographs of physical and biological
characteristics of the Murray-Darling
Basin. These photographs would make a
very valuable resource for educators, both
secondary and tertiary, if they were avail-
able in electronic form. I urge the Murray
Darling Basin Commission to consider
releasing a CD based on the book.

The range of topics covered is impressive
and appropriate. The Introduction claims
that ‘It presents, for the first time, a compre-
hensive conceptual model for the major
rivers of the basin ..." (page vii), This model
is really the first part of the book
(Understanding river systems) and while the
material required for a conceptual under-
standing is included, there is not really any
model, even informal, presented. The second
part of the book deals with specifics of land-
scapes, flows and biota. Some of these chap-

Vol. 119 (3) 2002

ters are the best summaries of relevant infor-
mation for the Murray-Darling Basin avail-
able; in particular, | found the section on
large plants very useful.

Like any book of this type, especially one
with numerous authors across different
chapters, there are some things that don't
work so well, I have already mentioned the
lack of critical evaluation of the literature.
Similarly, there is usually no distinction
made between information based on pub-
lished data versus information based on
observation versus information based on
pure speculation. It is not that any of the
information is obviously incorrect but that it
is presented with no indication of the uncer-
tainty associated with it. The chapters also
vary quite a bit in the amount of detail and
quantitative information provided. For
example, chapter 4 (Landscapes, climates
and flow regimes) is detailed with flow
duration curves and hydrographs provided.
In contrast, chapter 2 (River flow, process-
es, habitats and river life) is very descrip-
tive, even though details such as cross-sec-
tions illustrating changes in river morpholo-
gy would have been really helpful. There
are also a few minor annoyances with the
way the book is structured. There are no
summaries of main points at the end of each
chapter so main messages in each chapter
tend to be lost. It is also an awkward book
to cite because authors are associated with
both chapters (e.g. Chapter 1) and sections
within chapters (e.g. Chapters 5 and 6).

In the end, the main difficulty this book
will face is finding its target audience. It is
clearly not a ‘coffee table’ book designed
for the general publie, neither is it a text-
book suited for a university course in river
ecology, although I would imagine both
these groups would find parts of the book
very informative. It seems designed for
professionals in a range of fields who are
associated with the science or management
of our river systems. Specialist ecologists
or hydrologists will find useful information

147

throughout, and agricultural scientists,
economists, engineers, etc. will find the

book a valuable summary of the effects of

flows on river ecology. My final comment,
which is not meant to devalue the book in
any way, is that | hope that parts of this
book are quickly out of date. This will
mean that research on the effects of flow
on riverine ecosystems, and the Murray-
Darling Basin in particular, is continuing
and our knowledge base is improving.

Despite the confident tone in some sec-
tions of this book, our limited understand-
ing of the links between flow and river
ecology is still a major constraint on suc-
cessful management of one of Australia's
most important resources, the rivers of the
Murray Darling Basin.

Gerry Quinn

CRC for Freshwater Ecology
School of Biological Sciences
Monash University, Victoria 3800

The Field Naturalists Club of Victoria Inc.

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Established 1880
In which is incorporated the Microscopical Society of Victoria

OBJECTIVES: To stimulate interest in natural history
and to preserve and protect Australian flora and fauna.

Membership is open to any person interested in natural history and includes
beginners as well as experienced naturalists.
Registered Office: FNCV, 1 Gardenia Street, Blackburn, Victoria 3130, Australia.

Postal Addre

: FNCV, Locked Bag 3, PO Blackburn, Victoria 3130, Australia.

Phone/Fax (03) 9877 9860; International Phone/Fax 61 3 9877 9860,

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and Mrs ANNE Morton, as above.

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and DR NOEL SCHLEIGER, as above,

Conservation Coordinator: MR Jim WALKER, 167 Balaclava Road, Caulfield 3162, 9527 5601
Group Secretaries

Botany: MS KAREN DonBSON, 58 Rathmullen Road, Boronia 3155. BH 9877 9860

Geology: MR Rog HAMSON, 5 Foster Street, McKinnon 3204. 9557 5215

Fauna Survey: Ms Sopit SMALL, 107 Bondi Road, Bonbeach 3196, AH 9772 2848

Marine Research: MR MicttAkt. LYONS, 18 High Street, Nunawading 3131, AH 9877 3987

Microscopical: MR RAY PowrR, 36 Schotters Road, Mernda 3754. 9717 3511

MEMBERSHIP
Members receive The Victorian Naturalist and the monthly Field Nat News free. The Club organis-
es several monthly meetings (free to all) and excursions (transport costs may be charged). Field
work, including botany, mammal and invertebrate surveys, is being done at a number of locations
in Victoria, and all members are encouraged to participate,

Printed by Brown Prior Anderson, 5 Evans Street, Burwood, Victoria 3125,

The
Victorian

Naturalist

Volume 119 (4) August 2002

Murray River Special Issue Part Two

Published by 'The Field Naturalists Club of Victoria since 1884

The Murray River in The Victorian Naturalist
NOTES OF A SHORT COLLECTING EXCURSION TO THE UPPER MURRAY

By Messrs C French and D Best
Read before the Field Naturalists" Club of Victoria, Nov 17 1884

In crossing a paddock, close to the railway line, we found quantities of a small
but pretty composite, the name of which, with those of the other plants collected,
have been identified and named by the kindness of Baron von Mueller. Some of
the plants here collected, as also portion of those from Mulwala, he found to be of
interest, and a complete list of all our specimens appears as an addendum to this
paper. Turning over some logs, we were fortunate in finding a few specimens of a
very handsome beetle, Carenum sp. which were the first of the kind taken by us.

We had not proceeded far before we came across a couple of the large Iguana
(Hydrosaurus varius) common to the district, one being beautifully marked with
yellow bands; but feeling sure of getting some on the day of our intended depar-
ture we did not attempt to capture it, which we now regret, as we did not again see
a similar one, although we did many of the plain coloured, and of these we
brought down two; the former although only a variety is much rarer than the latter.

We drove towards the Murray pine scrubs ... a halt was here made, and a cock-
roach hunt suggested and improvised. This, we can assure our friends, was no
ordinary cockroach hunt, for these bulky and somewhat repulsive looking fel-
lows, burrow to a depth of nearly 2ft. into the sand, and forcibly reminded us of a
wombat hunt on a small scale. We were fortunate enough, after digging for our
bare lives, to find some of these Titanic cockroaches, which were speedily bot-
tled up for our collections.

Proceeding along we came across our first Quandong tree, Santalum acumina-
tum, which to our great delight had a few ripe fruit on the top branches.

The Quandong here seems to be fast dying out, the cause being generally attrib-
uted to the clearing off for the improvement of the runs for sheep.

| We] afterwards did a little botanising in the surrounding pine scrubs, collecting
a few orchids, amongst them being Prerostylis rufa, var. Mitchelli, this being
considered somewhat of a rarity here.

Prom The Victorian Naturalist 1 (13), January 1885, pp 135-139.

The Murray River in The Victorian Naturalist
WANDERINGS ON THE MURRAY FLOOD-PLAIN

By JG O’ Donohue
Read before the Field Naturalists’ Club of Victoria, 8th February 1915

Whilst ranging through the Mallee we realized, at a very early stage in our pere-
grinations, that caution had to be exercised in the choice of a site whence to view
a foraging bird, At every few yards the extensive and crater-like nests of the
Myrmecia nigricipes were to be encountered, These ants appeared to be always
spoiling for fight, and if one of us inadvertently approached within half a dozen
yards of a citadel, or, for that matter, allowed his shadow to fall upon it, out they
would pour in myriads, and, with gaping mandibles, skirmish over a large area in
quest of the offender.

From The Victorian Naturalist XXXI (2), 10 June 1915, pp 25-35.

The

Victorian
Naturalist

Volume 119 (4) 2002
Murray River Special Issue
Part Two

August

Executive Editor: Merilyn Grey
Editors: Alistair Evans and Anne Morton

Fish of the Murray River, by John Koehn .............. eret 152
Niche Segregation between Three Species of Freshwater Turtle in a Large
Billabong During Flood, by Catherine E Meathrel, Phillip J Suter and
Noda cA OTs eeu edet RERUM es We Ee ee er TRES HS AIRSS EM dg eras 032 160

Invertebrates of the River Red Gum Forests of the Murray River,
by Andrea Ballinger and Alan L Yen ........... eese 174

A Preliminary Survey of the Arboreal Invertebrate Fauna of Two
River Red Gum Trees Eucalyptus camaldulensis near the Murray River,
by Alan L Yen, Simon Hinkley, Peter Lillywhite, John Wainer and
HEAT en ode ena BA Hg Pues creep iR ua PLES LED eder TER Tenere PARIT 180

Aquatic Macroinvertebrates of the Murray River, by Phillip J Suter and
EGRE PLWH TALE Utd UIN ASR: NAD a 186

The Swamp Yabby (Cherax sp.) of the Murray River Catchment,
by Geoffrey N Edney, Dale G McNeil and Susan H Lawler ............. ees 200

Murray River Microfauna, by Russell J Shiel.................. eee 205

ISSN 0042-5184

Cover: Sweeping bend of the Murray River near Mildura. Photo by Anne Morton.

Web address: http://www.vicnet.net.au/~fnev/vicnat.htm
email: fnev@vicnet.net.au

Murray River Special Issue
Fish of the Murray River

John Koehn'

Abstract

Fish are an integral component of the Murray River, contributing to its biodiversity, ecology and cul-
tural heritage, as well as providing commercial and recreational fishing opportunities, Fish are an
important way of connecting the community to the river system, The number of species in the
Murray River is low by world standards; species range from the large, well known Murray Cod to
smaller, lesser known species, such as Australian Smelt. Seven fish species are considered to be
nationally threatened, with several other species threatened on a regional basis. Many of the threats
to the fish species relate to the use of water and its associated infrastructure. Changes to flows, barri-
ers to fish passage, cold water pollution, snag removal and habitat alterations, commercial and recre-
ational utilisation, and interactions with introduced fish species have all contributed to the decline in
fish populations. Many of these threats are currently being addressed or under consideration for
changed management regimes to reduce their impact. Issues such as the provision of more water for
improved environmental flows will pose ongoing challenges. The restoration of riverine ecological
processes is a key way by which fish populations may be restored, and this needs to be undertaken

with the support of the community. (The Victorian Naturalist 119 (4), 2002, 152-159)

Introduction

Although they largely remain hidden
under the water surface, fish are an integral
part of the fauna of any river system, This
is no more so than in the Murray River,
where the legendary ‘mystique’ of large
species such as Murray Cod Maecullo-
chella peelii peelii transforms their impor-
tance from being merely fish species to
becoming important components of our
folk lore and cultural heritage. Fish pro-
vide a major way in which the community
can be connected to the river and its fauna
(Sinclair 2001). Whilst Murray Cod are
well known because of their size and status
as a species of commercial and recreation-
al significance, there are many other
species which are also important but lesser
known. Fish are an important component

of the biodiversity, ecology and culture of

the Murray River.

The native fish and their status

The number of fish species in the Murray
River is relatively low by world standards,
totalling only about 30 native species
(Table 1), several of which are restricted to
the lower river zones and associated with
marine or estuarine reaches. Whilst this
number of species may be expected of a

! Cooperative Research Centre for Freshwater Ecology,
Arthur Rylah Institute for Environmental Research,
Department of Natural Resources and Environment,
123 Brown Street, Heidelberg, Victoria 3084

Email John.Koehn(Zinre.vic.gov.au

152

river with a relatively low overall dis-
charge, it can be compared to the more
than 1300 fish species described for the
Amazon Basin (Cadwallader and
Lawrence 1990) and highlights the impor-
tance of the need for conservation of indi-
vidual species. The majority of the species
is widespread along the river, although
some have distributions more suited to
either the upper or lower zones. For exam-
ple, Two-spined Blackfish Gadopsis
bispinosus occurs in the higher reaches,
whilst species such as the Bony Herring
Nematolosa erebi are only found in the
mid and lower river zones. A further group
of seven diadromous species, which
require access to marine/estuarine waters
to complete part of their life cycles, are
found in the lower zones of the river.
These species include the Pouched and
Short-headed Lampreys; Geotria australis
and Mordacia mordax, Short-finned Eel
Anguilla australis, Common Galaxias
Galaxias maculatus, Tupong Pseuda-
phritis urvilii, Estuary Perch Macquaria
colonorum and Blue-spot Goby Pseudo-
gobius olorum. Other essentially marine
species such as Mulloway Argvrosomus
hololepidotus would also have entered the
lower, estuarine river reaches more fre-
quently in the past.

The exact number of fish species present
is not really known, with the taxonomy of
several groups undergoing revisions,
including the hardyheads (Atherinidae),

The Victorian Naturalist

Part Two

Table 1. Species list (From Pierce 1988 and Cadwallader and Lawrence 1990) and conservation sta-
tus for freshwater fish of the Murray River. EPBC, Environment Protection and Biodiversity
Conservation Act 1999, Protection Act; ASFB, Australian Society for Fish Biology 2001 listing; CE,
critically endangered; E, endangered; V, Vulnerable; Epop, endangered population in New South
Wales; FFG, listed under the Flora and Fauna Guarantee Act, Victoria. P, New South Wales
Protected species (i.e. no take); (P), Protected from commercial take; UC, under consideration.

Common name Scientific name

Listing

Vic NSW

Native freshwater species

River Blackfish Gadopsis marmoratus DD

Two-spined Black fish Gadopsis bispinosus

Broad-finned Galaxias Galaxias brevipinnis

Flat-headed Galaxias Galaxias rostratus V DD

Mountain Galaxias Galaxias olidus DD

Murray Cod Maccullochella peelii peelii UC V, FFG

Trout Cod Maccullochella macquariensis E CE CE, FFG EP

Golden Perch Macquaria ambigua V

Macquarie Perch Macquaria australasica E E ERE AE’

Silver Perch Bidyanus bidyanus V CE PEG VP

Southern Pygmy Perch Nannoperca australis V

Australian Smelt Retropinna semoni

Freshwater Catfish Tandanus tandanus V V.FFG (P)

Bony Herring Nematalosa erebi

Southern Purple Spotted Mogurnda adspersa E CE, FFG Epop
Gudgeon

Western Carp Gudgeon Hypseleotris klunzingeri

Midgeley’s Carp Gudgeon Hypseleotris spp.

Flat-head Gudgeon Philypnodon grandiceps

Dwarf Flat-head Gudgeon — Philypnodon sp. FFG

Crimson Spotted Melanotaenia fluviatilis DD, FFG
Rainbowfish fluviatilis

Murray Hardyhead Craterocephalus V E,;EPG Œ

stercusmuscarum fluviatilis

Agassiz’s Chanda Perch Ambassis agassizi

Native diadromous species
Short-headed Lamprey
Pouched Lamprey
Short-finned Eel

Common Galaxia

Mordacia mordax
Geotria australis
Anguilla australis
Galaxias maculatus

Ex, FFG Epop

Tupong
Estuary Perch
Blue-spot Goby

Pseudaphritis urvillii
Macquaria colonorum
Pseudogobius olorum

Introduced species
Brown Trout
Rainbow Trout

Salmo trutta

Oncorhynchus mykiss

Carp Cyprinus carpio

Tench Tinca tinca

Goldfish Carassius auratus

Redfin (English Perch) Perca fluviatilis

Gambusia Gambusia holbrooki
Weatherloach Misgurnus anguillicaudatus

galaxiids (Galaxiidae) (T Raadik pers.
comm.) and gudgeons (Eleotridae), which
include hybrid species (Bertozzi et al.
2000). New genetic techniques are also
being used to investigate several other
species and these taxonomic revisions
could result in the description of new
species. Eight introduced species are also

Vol. 119 (4) 2002

present in the Murray River. One addition-
al native species, the Broad-finned
Galaxias Galaxias brevipinnis, is now pre-
sent in the upper and mid reaches where it
has been introduced from east coast rivers
through flows from the Snowy Mountains
Scheme (Waters et al. 2002). This normal-
ly diadromous coastal species appears to

153

Murray River Special Issue

be completing its life cycle landlocked in
Ireshwater, as occurs in the highland lakes
in Tasmania (Fulton 1990).

Fish species range in size from the
Murray Cod, which is Australia’s largest
freshwater lish (recorded up to 113 kg), to
smaller species such as the Australian
Smelt Retropinna semoni which weighs
only a few grams. Photographs, illustra-
ions and detailed information on these fish
species can be found in the following texts:
Cadwallader and Backhouse (1983),
Merrick and Schmida (1984), Koehn and
O'Connor (1990a), McDowall (1996) and
Allen et al. (2002).

It has been recognised that most native
fish species in the Murray River have suf-
fered major declines (c.g. Cadwallader
1981; Cadwallader and Gooley 1984), The
causes of such declines have been dis-
cussed by many authors (e.g. Cadwallader
1978; Koehn and O'Connor 1990b;
Kearney et al. 1999). There is concern
about the long-term future of many
species, with seven already being consid-
ered nationally threatened (Table 1),
including the eritically endangered ‘Trout
Cod Maceullochella macquariensis (Fig.
1) (Australian Society for Fish Biology
2001). The natural range of this species is
now restricted to about 120 km of the

Murray River, immediately downstream of

Lake Mulwala, Whilst this species has
been stocked from hatcheries into the
upper Murray River and other sites, these
fish have not yet formed self-sustaining
populations. Many other species have
reduced or fragmented ranges and are list-
ed as threatened in either NSW or Victoria
(Table 1). Concern has been expressed at
the status of many of these species in
South Australia (Pierce 1988).

l'he capture of Trout Cod and Silver
Perch is now prohibited in both New South
Wales and Victoria, Of particular commu-
nity concern is the decline of many ‘fag-
ship’ species such as Murray Cod, Trout
Cod, Silver Perch Bidvanus bidvanus and
Catfish Tandanus tandanus. These species
are readily identified by the public, and
their loss indicates that all is not well with
the health of the river. The fish community
in the lower Murray River has been listed
us endangered in New South Wales (New
South Wales Fisheries 2002) and under the

154

Vig. L. Trout cod Maceullochella macquariensis

is now critically endangered. The only remain-
ing natural recruiting population occurs in the
Murray River, downstream of Lake Mulwala.

Flora and Fauna Guarantee Act in Victoria
(WWw.nre.vic.gov.au).

Commercial fisheries for species such as
Murray Cod, Golden Perch Maequaria
ambigua, Silver Perch and Catfish have all
been greatly reduced or ceased (Reid ef al.
1997). Recreational angling remains popu-
lar and many native species such as
Murray Cod and Golden Perch are keenly
sought. Lake Mulwala provides one of
Australia's premier Murray Cod fisheries
but eod from it appear to use both the
Murray and Ovens rivers upstream for
spawning (Koehn 1996). This highlights
the importance of tributary streams to the
fish of the Murray River system.

Fish habitats, needs and threats

The state of fish habitats and threats to
them vary along the river. Recent scientific
assessments of flow and environmental
impacts along the Murray River (Thoms ef
al, 2000; Jensen er al, 2000) highlight how
threats ehange through the reaches (Table 2).
For example, cold water pollution is an issue
downstream of Lake Hume and Dartmouth
Dam (on the Mitta Mitta River) and barriers
to fish passage more an issue in the lower
reaches. Many of the threats are related to
the human use of water and associated infra-
structure and the issue of environmental
flows was à reoccurring one. In essence, fish
need instream structural habitat and access
to it, suitable water conditions in terms of
quantity and quality and a functioning
ecosystem to provide food resources.

The native fish present in the Murray
River could be described generally as a
‘warmwater’ species assemblage. Water

The Victorian Naturalist

Part Two

Table 2. Major threatening processes for fish in the Murray River summarised from Thoms er al.

(2000) and Jensen ef al. (2000).
Reach

Major threatening processes relating to fish

Dartmouth Dam to Lake Hume

Constant flows, unseasonal high flows, reduced flooding,

unseasonal water temperatures, reductions in snags

Lake Hume to Tocumwal

Constant flows, unseasonal high flows, reduced flooding,

unseasonal water temperatures, reductions in snags

Tocumwal to Torrumbarry

Unseasonal high flows, reduced flooding, constant flows,

reductions in snags

Torrumbarry to Wentworth

Reduced flows variation and flooding, reductions in snags,

constant flows, barriers to fish passage, increased turbidity

Wentworth to Wellington

Reduced flooding, barriers to fish passage, reductions in snags,

possible fish health impacts from Lake Victoria.

Lower lakes and Coorong

temperature is important for the function-
ing of fish populations, particularly for
reproduction, larval survival and optimal
growth. The release of cold water from
low level outlets in impoundments poses a
major problem to warmwater fish, restrict-
ing the success of spawning and may have
detrimental effects on metabolic function
and growth rates (Koehn 2001). This is
highlighted in the upper zones of the
Murray River where high volume, cold
water irrigation releases occur from the
low-level outlets of Dartmouth Dam and
Hume Dam. Three species of warmwater
native fish (Trout Cod, Macquarie Perch
Macquaria autralasica and Murray Cod)
have disappeared from the reaches of the
Mitta Mitta River downstream of the
Dartmouth Dam. It is likely that recruit-
ment of these species has not been possible
due to cold water releases during the
spawning season of this species (Koehn er
al. 1995). Low water temperatures from
Lake Hume is likely to continue to restrict
Murray Cod recruitment in the river down-
stream, The Murray River also receives
water of unknown temperatures from the
Snowy Mountains Scheme (Jacobs 1990).
Cold water does however favour intro-
duced species such as Brown and Rainbow
Trout Salmo trutta and Oncorhynchus
mykiss that can prey on smaller native
species.

Reproductive strategies vary among
species. Some species such as Golden
Perch and Silver Perch produce up to
500,000 eggs which are laid in the water
column and left without care by the par-
ents. In contrast, species such as the River
Blackfish Gadopsis marmoratus lay only a
few hundred adhesive eggs on a wood sub-

Vol. 119 (4) 2002

Reduced freshwater flows, barriers to fish passage

strate that are dependent on parental care.
It has often been contended that many fish
species have their recruitment aided by
flooding, being able to exploit the high
food availability on floodplains in times of
floods, Whilst there is little evidence for
use of the floodplain (Humphries et al.
1999), nutrients from the floodplain can be
released during flooding and support an
increase in production of algae, aquatic
plants, zooplankton and other invertebrates
that provide rich food sources for juvenile
fish (Gehrke 1991). Reduced flooding,
together with diminished flow levels could
significantly decrease juvenile habitat
availability and food production areas. A
further complication is the clearance and
replacement of native vegetation with
flood intolerant exotic crops and pastures.
Under such conditions, floodwaters rapidly
become de-oxygenated as microbial com-
munities decompose the flooded vegeta-
tion. This may adversely affect the survival
of certain fish species, particularly when
movement to other areas is restricted
(Gehrke 1991; McKinnon and Shephead
1995).

Fish can be mobile, and many have a
need to move widely throughout the river
system. Species such as Golden Perch are
mobile (Koehn and Nicol 1998), with
some individuals migrating over 1000 km
(Reynolds 1983). It has recently been dis-
covered that high numbers of juvenile fish
of species, such as Silver Perch, also move
upstream (Mallen-Cooper et al. 1996), pre-
sumably to recolonise. Species, such as
Murray Cod, which were previously
thought to be sedentary have been shown
to undertake pre-spawning migrations
(Koehn 1997), Whilst these species may be

155

Murray River Special Issue

able to survive and reproduce even if such
movements are not able to take place, their
ability to recolonise and their ultimate sur-
vival and distribution over the longer term
may be detrimentally affected. Lake
Hume, Lake Mulwala and Torrumbarry
Weir all pose barriers as do the many locks
and weirs in the river downstream of
Euston.

Many other species, especially those in
the lower zones of the river, have parts of
their life cycle that must be completed in
saline water (normally the sea) and hence
have to migrate to complete their life
cycles. Barriers at the barrages (Murray
mouth) prevent movements to and from
the sea as well as preventing the occasional
entry of many marine species. Some
migrations are understood to occur in a
particular season, for a particular reason,
e.g. for spawning, and may often be affect-
ed by changes to flows. However, our
understanding is not complete for all
movements, Hence fish passage should be
available to all species throughout the year.
Aggregations of fish migrating upstream
often occur immediately downstream of
barriers (weirs and dams) making these
fish very susceptible to capture by anglers.

Increases in water levels, both large and
small, can stimulate the movement of fish
(Mallen-Cooper e al, 1996). Reductions in
flooding may restrict such movements, as
might constant flow levels. The limiting of
cues and the barriers to movement may
affect spawning success and the distribu-
tion of species. Sudden reductions in flow
levels can also lead to the stranding of lish.

Another form of fish movement is the
drift of larvae (Koehn and Nicol 1998;
Humphries and Lake 2000; Humphries et
al. in press), This has the purpose of
recolonisation and distribution of off-
spring, and can be affected by altered flow
rates and impounded waters. High irriga-
tion flows during early summer may mean
that larvae are carried greater distances
than. would have occurred naturally.
Impounded waters can trap larvae and pre-
vent their distribution downstream. Such
effects can alter the structure of fish popu-
lations. Larvae can also be transferred into
irrigation channels where they are unlikely
to survive (Koehn and Nicol 1998).

Fish numbers are often related to the

156

amount of habitat available. Of course
there must also be access to this habitat,
which can be blocked by barriers. Snags or
large woody debris are the major form of
structural habitat in lowland rivers and are
widely used by many species (Koehn
1993; Koehn and Nicol 1998). The use of
this habitat by species such as Murray Cod
has long been recognised (Koehn 1997).
Snag removal has been widespread
throughout the river, with snag numbers
now considerably less than those that
occurred naturally. Snags are used as home
sites for territory formation, predator
avoidance and prey detection. They offer
protection from high water velocities, and
are sources of food and spawning sites for
many species. Snag removal has caused a
major loss of fish habitat.

Many other floodplain habitats, such as
billabongs and wetlands, have also been
degraded, removed or made inaccessible.
Habitats in the form of pools and scour
holes can also be lost through infilling by
sedimentation, Removal of snags and bank
erosion can lead to a more uniform channel
without a diversity of habitats. Variations
in depth and velocities are also important to
provide the suitable habitat for all species
throughout their life cycle, and the presence
of snags promotes such habitat diversity.

Eight introduced fish species are also pre-
sent in the Murray River. Of these, Carp
Cyprinus carpio, Redfin Perca fluviatilis
and Gambusia Gambusia holbrooki are the
most widespread. Carp receive the most
public attention and are often blamed for
many of the ills of the river. Recent reviews
(e.g. Koehn er al. 2000) indicate that they
are typical invasive species, which are
tough and well adapted to making the most
of already degraded riverine environments.
With minimal predation pressure from
reduced populations of native species such
as Murray Cod and Golden Perch, Carp
populations have expanded rapidly. Now in
large numbers they contribute to water tur-
bidity, uproot aquatic plants and utilise
large amounts of habitat space. It is esti-
mated they make up 90% of fish biomass in
many reaches of the Murray River (Harris
and Gehrke 1997).

The introduced salmonid species (mainly
Brown and Rainbow Trout) are restricted
mainly to the upper river zones (above

The Victorian Naturalist

Lake Mulwala) and are keenly sought by
some anglers, Together with Redfin, these
species are formidable piscivores, which
can impose predation pressure on smaller
native fish species. Redfin populations
appear to be reduced in many areas com-
pared with the past (Cadwallader 1977),
but this species persists throughout the
river, particularly in still waters. Gambusia
have been implicated in the harassment of
fish through fin nipping and potential pre-
dation of eggs (Cadwallader and
Backhouse 1983), but their impact on
native species is not known.

Goldfish Carassius auratus are common,
often in high numbers in the lower river
reaches. Oriental Weatherloach Misgurnus
anguillicaudatus are an aquarium species
now present in the Murray River from
Lake Mulwala to Barmah. Their distribu-
tion has increased downstream from the
Ovens River. The impacts of this hardy
species, which can survive in damp mud, is
not known. Other introduced species such
as Roach Rutilus rutilus, Atlantic Salmon
Salmo salar and Brook Trout Salvelinus
fontinalis have been recorded in the
Murray-Darling Basin (Cadwallader and
Lawrence 1990). Whilst not yet present in
the Murray River, the coastal species
Spotted Galaxias Galaxias truttaceus has
also recently been recorded in the
Campaspe River (P Humphries pers.
comm.), presumably after a translocation.

Many of the fish species of the Murray
River are or have been sought after for their
commercial or recreational significance.
Commercial fishing has undoubtedly taken
a toll on numbers of some fish species in
the past (Rowland 1989). Today however,
most catch is from recreational take and
this is not well quantified. Commercial
native fin fishing now only continues in
South Australia and this is likely to be
reduced or phased out in the near future.
The reduction of species other than those
sought by fishers indicates that this was not
the only cause for the decline of Murray
River fish species.

Conservation efforts and the future

A Fish Management Plan for the Murray
River was initiated in 1991 (Lawrence
1991) outlining management action to be
taken to improve fish populations. This

Vol. 119 (4) 2002

Part Two

plan is to be superseded by a Native Fish
Strategy (Murray Darling Basin
Commission 2002) which will address key
components for the restoration of native
fishes across the Murray-Darling Basin,
including the Murray River, over a 50 year
time frame. Recently, the river has been
assessed in relation to environmental flows
and other threats, with many options for
actions being canvassed (Thoms er al.
2000; Jensen et al. 2000). Several of the
options discussed, together with other
management actions, are already planned
or under way. Some of these actions can be
initiated readily whilst others must be
structured over the longer term. The provi-
sion of better environmental flows was one
recommendation which was seen to have
the most benefit for the whole river
(Roberts et al. 2001) and negotiations to
improve environmental water allocation
and its delivery are being undertaken
through many different forums.

Options are now being considered for the
remediation of cold water releases from
Lake Hume. A fish lift has been installed
at Lake Mulwala to assist with fish passage
past that structure and a fully operational
fishway has been installed at Torrumbarry
weir (Mallen-Cooper et al. 1995). There is
anecdotal evidence of increased numbers
of Silver Perch upstream of Torrumbarry
following the installation of that fishway.
Modifications to existing. poorly function-
ing fishways and the construction of new
fishways are intended for all structures
downstream of Torrumbarry including the
Barrages at the river mouth (J Barrett pers.
comm. ).

A considerable amount of research has
been undertaken in the Murray River to
improve our ecological knowledge of fish
in this ecosystem. The general movements
and habitat use of Murray Cod, Trout Cod
and Golden Perch have been investigated
(Koehn and Nicol 1998). Particular
emphasis has been placed on investigating
the downstream movements of fish, espe-
cially Golden Perch, with respect to inter-
ference to this type of movement caused
by weirs (J O'Connor unpubl. data). Study
of the downstream movement of fish lar-
vae has also been undertaken. A resnag-
ging project (adding large woody debris)
has been undertaken downstream of Lake

157

Murray River Special Issue

Mulwala to provide additional habitat for
Trout Cod and other species. This project
has determined the structure and pattern of
snags and is currently monitoring recoloni-
sation by fish and macroinvertebrates.
Study of movements and recruitment of
carp around Barmah has highlighted this
area as a likely key source of recruitment
for the river (Stuart ef al. 2001). This new
information has been included in a new
carp management plan for the area.

National recovery plans have been pre-
pared for Trout Cod (Brown et al. 1998),
Silver Perch (Clunie and Koehn 20002)
and Freshwater Catfish (Clunie and Koehn
2000b). Whilst the latter two plans are yet
to be implemented, the ecology of Trout
Cod has been investigated and monitoring
of the Murray River population undertak-
en. Recent data indicate that this popula-
tion is at least stable and may be expand-
ing downstream. The provisions of fish
passage at Lake Mulwala may allow this
species to colonise the lake and expand its
population upstream into the Ovens and
Murray Rivers. Ideally, this would link the
population to the fish currently being
stocked in the Ovens River as part of the
national recovery plan.

The restoration of ecological processes,
such as migration, recruitment and organie
production and cycling, must be recog-
nised as key components for restoring the
Murray River. With these processes in
place, the fish populations then have a
chance to re-establish and inerease. This
ecosystem approach to management must
include the tributaries and catchments of
the river. Ultimately, the management of
the river must include and be supported by
community ownership.

As a result of increased knowledge and
recent and intended management actions,
there are some positive signs for fish in the
Murray River. The provision of fish pas-
sage throughout the river will allow the
entry of some marine species, allow
species with marine life stages to
recolonise the river successfully and allow
migratory fish within the river to move and
recolonise freely. This should improve the
distribution of species such as Tupong, and
Silver Perch. More natural water tempera-
tures downstream of Lake Hume can only
be a positive for the recruitment of warm

158

water species such as Murray Cod and
Golden Perch. The provision of fish pas-
sage at Lake Mulwala may allow the Trout
Cod and Silver Perch populations down-
stream to establish in the lake and rivers
upstream. Additional water to meet envi-
ronmental needs is still required to provide
adequate environmental flow conditions to
restore some of the key flow components,
which maintain ecosystem processes. This
will remain one of the biggest challenges
for the restoration of fish populations in
the Murray River.

Acknowledgements

The author wishes to thank Simon Nicol and
Paul Humphries for comments on the manu-
script.

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Ann Williamson (label printing) and Dorothy Mahler (administrative assistance).

Vol. 119 (4) 2002

159

Murray River Special Issue

Niche Segregation between Three Species of Freshwater Turtle
in a Large Billabong During Flood

Catherine E Meathrel', Phillip J Suter' and Nada M Radford'

Abstract

During the summer of 1996/97 habitat and dietary overlap of three species of freshwater turtle,
Chelodina longicollis, C. expansa and Emydura macquarii in Lake Moodemere, a large billabong in
north-east Victoria, were studied to assess their level of niche segregation. Lake Moodemere was
inundated by flood waters and was connected to the Murray River for the first four weeks of the
study. Five separate habitat types of varying complexity were identified within the billabong, rang-
ing from open water to dense reed beds. All three species of turtle preferred the more complex habi-
tats in which to forage, but very few turtles contained food. During times of flood, all turtles
appeared to become opportunistic and consumed any food items available. Prey refugia were unim-

portant during times of flood. (The Victoriam Naturalist 119 (4), 2002, 160-173)

Introduction

The biodiversity of billabongs is greater
than that of the parent river with which
they are associated, and this is thought to
be promoted by high productivity and
environmental variability (Hillman 1986).
The high levels of productivity of lower-
order organisms in billabongs provide a
plentiful food supply for higher-order con-
sumers such as waterfowl, carnivorous fish
and freshwater turtles (Carpenter ef al.
1985). Turtles are an important element in
the dynamics of Australian freshwaters.
They occupy a complex trophic role, feed-
ing on a large variety of plants, animals
and microbes and via overland migrations
can enter temporary water bodies that are
inaccessible to other aquatic vertebrates
(e.g. fish) (Chessman 1978),

Three species of turtle (family Chelidae)
occur on the floodplain of the Murray
River — the Eastern Snake-necked Turtle
Chelodina longicollis (Shaw), the Broad-
shelled River Turtle Chelodina expansa
(Gray) and the Murray Turtle Emydura
macquarii (Gray). Table 1 summarises the
major differences between these three
sympatric species (information taken from
Goode 1967; Goode and Russell 1968;
Legler 1978; Chessman 1983, 1984, 1988;
Georges 1984; Georges ef al. 1986;
Kennett and Georges 1990; Legler and
Georges 1993; Cogger 1996).

Most research into the ecology of C.

' Department of Environmental Management and
Ecology, La Trobe University, Wodonga, Victoria
3690

Author for correspondence: Catherine Meathrel

Email c.meathrel@aw.latrobe.edu.au

160

longicollis, C. expansa and E. macquarii
has been species-specific and comparative
between rivers and billabongs (Chessman
1988; Kennett and Georges 1990). Limited
previous research on sympatric chelids has
suggested some degree of habitat and
dietary overlap in the prey consumed,
especially between the long-necked
species (Chessman 1988; Legler and
Georges 1993). There remains a paucity of
quantitative research detailing the turtles*
ecology.

Analyses of community dynamics
depend on measurements of how organ-
isms utilise their environment (Krebs
1989). The choice of niche by freshwater
turtles is likely to be influenced by varying
degrees of intra- and interspecific competi-
tion, The measurements of niche parame-
ters of one species or population may be
compared to that of another species or pop-
ulation. Food is one of the most important
dimensions of the niche and the analysis of
dietary preference of an organism is
strongly related to the niche specifications
of that organism (Krebs 1989). The three
species of Murray River turtles all
occurred in a large, permanent billabong
on the Murray floodplain, Lake
Moodemere, north-eastern Victoria. To
assess competition between the three
species of turtles two interactive niche
dimensions were analysed (habitat and
diet) over a four-month period from
October 1996 to January 1997. Lake
Moodemere was flooded at the start of the
research. The billabong was connected to
the Murray River (usually separated by

The Victorian Naturalist

Part Two

Table 1. Comparison of the habits and size of the three turtle species found in the River Murray.

Species Chelodina expansa

Chelodina longicollis

Emydura macquarii

Range
central and coastal
Queensland

Feeding strategy obligate carnivore on
motile prey, especially
Hemiptera, fish and
decapod crustaceans

Preferred aquatic
habitat

Egg-laying season

permanent

late autumn

Nest substrate heavy soils

Murray-Darling system, Coastal and inland
waterways from
Queensland to South
Australia

opportunistic carnivore
including carrion, decapod
crustaceans and benthic
invertebrates

shallow, ephemeral
billabongs

early summer

heavy soils

South Australia through
Murray-Darling system
to inland and northern
NSW

omnivore, scavenger,
especially on periphyton

large, permanent

early summer

light sands

Position within top bottom middle
the water column

Distance found moderate high low
from parent river

Average adult 43 25 3l
carapace length (cm)

Neck morphology long long short

about 150 m) for the first month of the
study and, as river levels subsided, the bill-
abong became isolated, Turtles were cap-
tured in a number of different habitat types
of varying complexity in order to test the
null hypothesis that there would be no spa-
tial overlap between the species. The null
hypothesis that there would be no dietary
overlap between species in any habitat
type was tested by assessment of the type
and abundance of prey available to each
chelid species and this was compared with
the prey actually consumed.

Methods
Study site

Lake Moodemere is a crescent shaped,
ox-bow billabong situated 5 km west of
Rutherglen in north-eastern Victoria (36°
03° S, 146° 23° E) that receives an annual
rainfall of between 500-600 mm (Land
Conservation Council 1983). Lake
Moodemere was chosen for this study
because large numbers of each species of
turtle had been caught during the previous
summer by researchers from La Trobe
University.

On average, the billabong is 600 m wide
and 2 km in length with a surface area of
approximately 0.86 km’. Because a 150 m
channel to the Murray River was dug for
water regulation, the billabong never dries

Vol. 119 (4) 2002

out (Land Conservation Council 1983).
Lake Moodemere is therefore classified as a
perennial billabong (mean annual depth of
1.2 m) although surface area and depth vary
seasonally and annually. At peak flooding
in October 1996, the billabong measured
1.8 by 5.1 km, had reached 10 kn? in area
and was, on average, 3.8 m deep.

The billabong is surrounded by a commu-
nity of closed grasslands and tall open forest
dominated by River Red Gums Eucalyptus
camaldulensis. Common species surround-
ing the billabong and its periphery are Giant
Rush Juncus ingens, bulrush Schoenoplectus
sp., Common Reed Phragmites australis,
spike-rush Eleocharis sp., milfoils
Myriophyllum spp., Floating Pond Weed
Potamogeton tricarinatus, flat-sedge Carex
sp. and Water Ribbons Triglochin procerum
(Land Conservation Council 1983).

Classification of habitat types

In total, five distinct habitat types were
determined for investigation (Table 2).
These habitats were chosen on the basis of
turtle prey refugia. For analysis, habitats
were ranked by complexity. Habitat com-
plexity was assessed on the basis of sub-
jective estimations of the surface area
available to turtle prey. It was assumed
that the greater the surface area available,

161

Murray River Special Issue

Table 2. Classification of habitat complexity within Lake Moodemere, Victoria. * Reed beds con-
tained areas of 1-5m' of open water within the reed bed. ** No areas of open water present.

Habitat Habitat Relative width of Relative abundance Reed bed Complexity
type stems or branches of stems or branches width (m) ranking

| Open water Nil Nil 0 I

2 Woody debris Broad Low 0 a

3 Reed bed Narrow Low 2-3 3

4 Reed bed Narrow Medium 18 -= 20* 4

5 Reed bed Narrow High 20.—25** 5

the higher the complexity of the habitat,
and the greater the number of prey items
available to turtles. Estimations of surface
area were calculated by the approximate
width of reed stems or snags, multiplied by
the relative abundance of the given refuge
(reeds or snags). Water depth and tempera-
ture were measured in each habitat type
during each sampling trip.

Assessment of prey in the habitat types

Samples of potential prey were collected
from each habitat type during each trip.
Invertebrates and small fish were collected
from the water column, substrate and
among refuge-type reed beds or snags to
ensure a reasonable sample through the
habitats. All reference samples were col-
lected using a 200 jtm mesh dip net with a
mouth dimension of 0.29 * 0.25 m. All
sweeps were for 20 seconds using a vigor-
ous and continuous action. Invertebrates
within the water column were collected by
walking forward, so as not to collect
organisms stirred up from the substrate,
with the dip net circling up and down, back
and forth. Invertebrates within the sub-
strate were collected by walking backward
and kicking up the substrate while circling
the dip net 10 em above the bottom. Reed
beds were sampled for invertebrates and
fish by vigorously thrusting the dip net
amongst the reeds using an up, down and
around motion. Invertebrates and fish asso-
ciated with submerged branches were sam-
pled by agitating the surface of the branch-
es with the leading edge of the dip net.
Specimens were preserved in 70% alcohol
and later identified to order or family and
counted. Only presence and absence of
taxa were used for comparison with those
found in the gut of turtles as the sweep net
sampling method did not resemble turtles?
hunting actions.

162

Collection and processing of turtles

Lake Moodemere was sampled continu-
ously for turtles on two consecutive days
each month from October to December
and twice in January. Each field trip was
conducted continuously between 10:00 hr
and 13:15 hr the following day. This
allowed each net in each habitat to be in
the water for a total of 24 hours. All turtles
were caught under the authority of the
Victorian Department of Natural
Resources and Environment (permit num-
ber RP-96-147) and the La Trobe
University Animal Experimentation Ethics
Committee (permit number LSC 9624).

Habitat preference for each species of
turtle was measured by the abundance of
each species captured in the five habitat
types, Within each habitat in Lake
Moodemere, two standard design fyke nets
were placed randomly. Only areas that
included the habitat type for at least 10 m
either side of the net were sampled.
Previous researchers have caught turtles
using baited nets (Georges 1982; Georges
et al. 1986; Bjorndal 1991). However, bait-
ed nets were not used in this study because
it was thought that baits would attract tur-
tles and cause an artificial habitat shift.

Nets in all five habitat types were
checked and cleared every four hours.
Once captured, turtles were returned to the
shore in water-filled bins. There, all turtles
were etched on the edge of a scute for
recognition of having been captured. This
ensured that no one turtle was sampled
more than once for its gut contents. The
first ten turtles above 15 em carapace
length for each species per trip were sam-
pled for dietary analysis using stomach
flushing (Legler 1977). This procedure
took place as soon as possible, lasted
approximately two minutes per turtle and
always occurred within one hour of collec-

The Victorian Naturalist

tion. Stomach contents were preserved in
70% alcohol. Once their stomach contents
had been sampled, turtles were held until
the next visit to their site of capture (i.e.
maximum holding time of 4 hr). Turtles
were not released immediately, as this
would have required excessive disturbance
in the vicinity of other nets.

In the laboratory, gut samples from tur-
tles were examined under a stereo-micro-
scope. Sand was assumed to have been
accidentally ingested by turtles and was
not included in the analysis of food items.
Turtles that yielded no digesta were
assumed to contain no material in the gut
and were excluded from the analysis (fol-
lowing Georges 1982). Percentage compo-

sition by number, mass and frequency of

occurrence were used to evaluate the rela-
tive importance of different foods eaten by
turtles. The numerical method involved
counting the number of items belonging to
each taxonomic group. Aquatic insects and
small crustaceans usually remained intact
and were easy to count. Items that did not
occur as discrete units (such as filamentous
algae and carrion) could not be counted
and were omitted from the numerical
analysis. The diet of a turtle was assessed
by combining the percent numeric, mass
and frequency of food items in the stomach

of a turtle and calculating an Index of

Relative Importance (IRI) (Pinkas 1971) as
follows:

Numerical percentage
. occurrence of FC, in I,

— er 49 voa ae? dr qr DD 0
total occurrence of all FC for all J 100

Mass percentage
. mass of FC, in I,

- total mass of all FC for all l 100

Frequency of occurrence percentage
_ total proportional occurrence of FC, — 100
total proportional occurrence for all FC
where FC, and I, are food category and
individual number ‘n’ respectively. The
IRI gives the relative occurrence of the
items in the diet for the numerical or mass
value of a food type.

In total, 45 different categories were
counted from stomach samples (Table 3).
For analysis, these 45 categories were
pooled into 12 (for turtles) and 11 (for ref-

Vol. 119 (4) 2002

Part Two

Table 3. Stomach content items found in
Chelodina expansa, C. longicollis and E. mac-
quarii sampled in Lake Moodemere, Victoria. a,
adult; |, larval stage.

Aquatic plants
Aquatic and terrestrial plant debris
Aquatic invertebrates
Aschelminthes: Nematoda
Annelida
Hirudinea
Oligochaeta
Arachnida
Arthropoda: Crustacea
Cladocera
Copepoda
Decapoda
Atyidae: Paratya australiensis
Palaemonidae
Parastacidae: Cherax destructor
Arthropoda: Insecta
Coleoptera (a)
Dytiscidae (a)
Hydrophilidae (a)
Coleoptera (1)
Diptera (a)
Diptera (1)
Ephemeroptera (1)
Hemiptera (a)
Corixidae
Notonectidae
Mesoveliidae: Mesovelia sp.
Odonata (1)
Plecoptera (1)
Trichoptera (1)
Mollusca
Bivalvia
Gastropoda
Terrestrial invertebrates
Arachnida
Insecta
Lepidoptera (1)
Vertebrates; Chordata
Teleostomi: Cyprinidae
Cyprinus carpio and Carassius auratus
Eleotridae sp.
Mucous and undetermined

erence collections) functional groups.
These functional groups were based on
general location and size of food items in
relation to assumptions of how a turtle
would distinguish them. Pooled groups
only took into account organisms that were
relatively similar in size and generally
remained in the same area (as per Williams
1980 and Ingram et al. 1997).

Aquatic plants were separated from the
rest of the plant debris found because these
items were fleshy and of bite-sized pieces of
stems and shoots. All other plant material,
including terrestrial River Red Gum leaves,

163

Murray River Special Issue

Table 4. Functional grouping of organisms found in the stomachs of turtles and reference collec-

tions, Lake Moodemere, Victoria.

Functional group Invertebrates

Hirudinae
Trichoptera larvae
Plecoptera nymphs
Odonata nymphs

Benthic organisms

Justification

All either crawl, sprawl or climb across
(or just a centimetre above in the case
of freshwater worms) the benthos

Ephemeroptera nymphs

Oligochaeta

Terrestrial insects
Diptera adults

Surface film organisms

Mesoveliidae adults

Littoral/pelagic/surface
adult insects

Hemiptera

Littoral/pelagic/surface
larval inseets

Coleoptera

Littoral/pelagic/surface
and benthic adult insects

Coleoptera

Cladocera
Copepoda

Pelagic zooplankton

Pelagic — active movement
Littoral and mobile

Littoral and sedentary Mollusea

Fish larvae and adults

Decapod crustaceans

All occur at or near the surface of the
water

Adults move throughout the water column
Larvae move throughout the water column
Adults move throughout the water column
and on the benthos

Zooplankton throughout the water column

Fish throughout the water column
Occur on plants and substrate in the littoral zone

Occur on plants and substrate in the littoral zone

reed stems and small grass-like plant mater-
ial, was termed plant debris.

Aquatic invertebrates were grouped into
functional groups as described in Table 4.
Benthic invertebrates included:
Hirudinea, Oligochaeta, Ephemeroptera
nymphs, Odonata nymphs, Plecoptera
nymphs, Trichoptera larvae, because all
either crawl, sprawl or climb across the
benthos (Williams 1980; Ingram ef al,
1997). Invertebrates associated with the

surface included Diptera adults, adults of

the family Mesoveliidae (Hemiptera) and
all terrestrial invertebrates that had fallen
onto the water. Except for Mesoveliidae,
all other Hemiptera adults were swim-
mers and moved about the water column
in the littoral, pelagic and surface zones.
For this reason Hemiptera adults were
kept as a separate functional group.
Diptera larvae were kept separate as a
group as not all occur near the surface;
some can be found associated with the
benthos. Coleoptera adults remained as a
separate group because, although they
were associated with the littoral and
pelagic zones like Hemiptera, they were
also associated with the benthos.
Coleoptera larvae were retained as a sep-

164

arate group for this reason as well but
they were not grouped with the adults as
they occupy different niches. Cladocera
and Copepoda were grouped as zooplank-
ton. The decapod crustaceans, Atyidae,
Palaemonidae (shrimps and prawns) and
yabbies Cherax destuctor were all
grouped together. The Mollusca
(Bivalvia and Gastropoda) was classed as
a distinct group. All fish adults and fish
larvae were grouped because both were
active within the water column.

Analyses of the dietary data were per-
formed using hierarchical clustering and
ordination programs in the numerical
analysis package PATN (Belbin 1992),
Similarity of diet was assessed using the
Bray-Curtis dissimilarity measure clus-
tered using Unweighted Pair Group
ArithMetic Averaging (UPGMA ).
Hierarchical clustering was used because
with a small to moderate data set (0 — 100),
most relationships could be determined
from a single dendogram (Belbin 1992),
Resultant groups were confirmed by exam-
ining their positions in ordination space
using Semi-Strong Hybrid multidimen-
sional scaling (SSH). This grouped turtles
or habitats into three dimensions.

The Victorian Naturalist

The dietary indices of niche breadth and
dietary overlap were calculated. Niche
breadth was calculated using Levins’ mea-
sure, B = 1/2 p, where p = proportion of a
food item | to # in a species (Levins 1968,
cited in Krebs 1989), to assess the uniformi-
ty of distribution of dietary food items
among the turtle species. Levins’ measure
was minimal when a turtle species con-
tained only one food type (minimum niche
breadth, maximum specialisation). The
range of B was from 1 to n, where 7 was the
total number of food types. To standardise
niche breadth to generate By, B was
expressed on a scale from 0 to 1.0, and was
calculated by dividing B by the total num-
ber of food types (i.e. B4 =(B- 1)/(n- 1)).

Dietary overlap was calculated using
Horn’s index (R, Horn 1966) because it
minimised biases associated with human
observations of resource categories that
animals may not distinguish (Krebs 1989)
and was calculated as:

Ro r1 (x (Pj + Pik) log.pj; y Pr) -È Pi log.p;, *
X pj, log,pj) / 21082

Data were analysed using either PATN or
the SPSS statistical package (SPSS Inc.
1993) and all tests followed Sokal and
Rohlf (1995). All means are presented + 1
standard error unless otherwise stated.

Results

Lake Moodemere changed in depth sig-
nificantly (one-way ANOVA, Fs jo =
60.27, p < 0.001; Fig. 1) over the summer
of 1996/97. The water level during
October was at its highest when floods

covered the region. An extra 2.8 m of

51

-— " riv

E

=

E

gai

5

$

e?| T

E] n

= | ?
1 : i2

ae.
04 uu = E - "
Sap Oct Now Dec marly Jan — (aie Jan
Sampling trip

Fig. 1. Mean (+ | se) water depth at Lake
Moodemere, Victoria during the 1996/97 turtle
breeding season.

Vol. 119 (4) 2002

Part Two

water covered the billabong and joined it
to the Murray River. In November the lake
became isolated from the river and the
water level was approximately 0.5 m
above December and January levels. From
December to late January the water levels
decreased a further 0.2 m, Temperature
increased gradually from October (17.2 +
0.3*C) to late January (27.4 + 0.5°C) (F; j4
= 326.57, p= 0.0001).

There was a significant difference in the
number of freshwater turtles caught at
intervals during the 1996/97 breeding sea-
son (x? = 91.252, df= 3, p < 0.001). Only
one C. longicollis was caught during
October and three of this species in
November. Most turtles were caught in
early January (n = 46). During late
January, turtle numbers decreased slightly
(n = 31) but still exceeded the numbers
caught in October and November.

The number of turtles caught in each
habitat was dependent on habitat complex-
ity (x^ = 42.021, df= 1, p < 0.001; Table
5), Turtles were caught most often in habi-
tat five (highest complexity) and the least
often in habitat one (lowest complexity).
The total number of turtles caught per
habitat was positively related to complexi-
ty (linear regression: y = 0.675x + 0,275;
p= 0.01). The coefficient of determination,
however, was low. Only 15% of the varia-
tion in turtle numbers caught could be
explained by habitat complexity.

Prey availability and habitat complexity
The UPGMA classification enabled
recognition of three habitat groups with a
dissimilarity of 0.87 and showed a seasonal
separation (Fig. 2). Group | consisted of all
habitats during the flood and open water
(lowest complexity) samples plus habitat
five late in the season (highest complexity).
Within the main cluster of this group was a
tighter cluster of the open water habitats
and habitat four and five of the November
sampling. This indicated a strong similarity
between the low complexity habitats and
flooded habitats. This group was largely
formed due to the greater amount of zoo-
plankton present. Group two contained
only habitat four, collected in December.
This sample split was largely due to the
high abundance of benthic invertebrates
and the presence of aquatic Acarina and

165

Murray River Special Issue

Mollusca. Group three contained the late
season samples from habitats of greater
complexity. A greater amount of fish and
Decapoda at this time of sampling were
responsible for this grouping.

These groupings were confirmed by SSH
ordination (Fig. 3). The stress for the three-
dimensional plot was 0.067 and was
accepted as a very good fit (Belbin 1992).
For ease of presentation, the figure shows
the three-dimensional configuration of ref-
erence samples as projected into the first
two dimensions (first two axes). Group one
clustered closely and indicated a strong
association while group 3 was more broad-
ly spaced.

Dietary preference of turtles

Stomach contents were flushed from a
total of 15 C. expansa, 17 C. longicollis
and 24 K. macquarii in Lake Moodemere
between October 1996 and January 1997.
The major component of food found in all
species was plant debris representing >80%
for all species (Fig. 4). The next most abun-
dant food category in the diet of turtles was

fish. Fish constituted approximately 20% of

C. expansa’s diet and a minimal percentage
in the other two species. Emyvdura mac-
quarii was the only species to contain a
minimal amount of aquatic plants. The
remaining, food categories were trivial to
the overall proportion of dietary items con-
sumed by each species of turtle.

All three groups contained the larger
lood categories such as Decapoda and fish.

0., 01l 0.2093 0.52

| |
Mab tat 3, Nov
Habitats, bale Jan
Habirat2,Nov
llabitat4, Nov
Habitats, Noy |
Habiratl,Late Jan | |
Habitat, Nov |
Habitat h, barby dan | |
Habitat, per
Habitata, Barly Jan
Habitatbe, Late Jan
Habitatz,Early dan
Habitat4d, arly dan |
Habirats, Barly Jan | |
MHabitat3,Late Jan I |
Habitat4, bate Jan

| |

0.0141 )

bray-(

Table 5. l'requencies of freshwater turtle
species caught in Lake Moodemere, Victoria
1996/97 turtle breeding season. Habitat com-
plexity is ranked from 1, lowest to 5, highest.

Habitat
Species 15.2. Be MS 3Edfal
Chelodina (0 ae? 3 5 —:5 IO
expansa
C. longicollis 0 5 i 20) fe 425
Emydura eee 6 22 48
macquarii
Total 2- 4. 15.21. 40.92

There was a significant difference in the
amount of fish, Hemiptera and Coleoptera
larvae between species (one-way ANOVA,
E554 = 5.50, p € 0.01, F554 = 3.89, p <
0.05, Fz s; = 4.76, p = 0.05). Chelodina
expansa consumed more fish than both C.
longicollis and E. macquarii; C. longicollis
consumed more Hemiptera and Coleoptera
than E. maequarii (p < 0.05, Scheffé's
post-hoc tests).

The UPGMA analysis produced three
groups at a level of 0.35 dissimilarity (Fig.
5). This low level of dissimilarity and each
species occurring in each group indicated a
high level of similarity of diet. The ordina-
tion plot for the analysis of turtles versus
[ood categories eaten supported the group-
ings of the dendogram once food types
responsible for the groupings were known
(Fig. 6). The stress for the three-dimen-
sional plot was 0.123 and accepted as a
good fit (Belbin 1992).

2500

+0348 1.2

"urtis Dissimilarity

Fig. 2. Dendogram based on turtle food found within habitats over the 1996/97 season, from a Bray-

Curtis association.

166

The Victorian Naturalist

* group *

oe | wunup 2
pa
ri á group
*
DA +
N n2| *
wn rs
x Us
iE 5 FA üs 1 5
n w
ELI
UR
n T -
mI
axis 1

Fig. 3. SSH ordination plot in two-dimensions
based on turtle food found within habitats from
a Bray-Curtis association. Stress — 0.067.

Group one turtles clustered broadly, indi-
cating a broad association between the tur-
tles. The broad clustering on the ordination
plot supported the high level of dissimilarity
shown in the dendogram within this first
group than the other two. This first group
was responsible for a greater amount of
food in the categories — plant debris, zoo-
plankton and surface invertebrates. The tur-
tles within this group had been eating more,
and the variation of food eaten was reflected
by the higher level of dissimilarity between
turtles.

Groups two and three were tightly clus-
tered on the ordination plot and indicated a
close association between stomach con-
tents of these turtles. Turtles within these
two groups contained very little in their
stomachs. They separated out on the den-
dogram because each contained slightly
different food categories. Group two
included aquatic plants and benthic inver-
tebrates, and group three included very low
amounts of fish, plants, aquatic plants,
zooplankton and benthic invertebrates. The
majority of turtles were in group three and
indicated that most turtles did not contain
very much, if any, food in their stomachs.

On a scale of 0 to 1, the Levins’ value for
niche breadth was low for all species.
Chelodina longicollis had the most spe-
cialised diet with a niche breath of 0,0039.
Emydura macquarii had a niche breadth of
0.0085, whereas C. expansa had the broad-
est (albeit still low) breadth of 0.0424.

The dietary overlap between C. expansa
and C. longicollis (R, = 0.9329) was high;
so too, was the overlap between C.
expansa and E. macquarii (R, = 0.9244).
This indicated that a number of similar

Vol. 119 (4) 2002

Part Two

food types were shared. C. /ongicollis and
E. macquarii had the least overlap between
their dietary niches (R, = 0.1939).

Discussion

The number of turtles captured in this
study was known to be lower than actual
turtle numbers in the billabong, but were
assumed to be a reflection of actual turtle
abundance. The number of turtles caught
could have been affected by a number of
factors: being washed away by the flood,
decreased net effectiveness with higher
volumes of water, low temperatures and/or
absence due to migration.

Turtles are ectotherms and consequently
have a body temperature close to that of
their environment (Begon ef al. 1990).
During lower temperatures in October, tur-
tles may have been less active throughout
the water column due to their reduced
metabolism (Parmenter 1981, Zimmerman
and Tracy 1989),

During the flood, the five habitats studied
within Lake Moodemere were completely
submerged. No reeds or snags were visible
when the water depth reached 4.1 m and
the submerged refugia no longer provided
invertebrates with a substrate close to the
water's surface. Williams (1980) has
shown that many aquatic invertebrates
move along or amongst the protective veg-
etation to the water's surface to breathe.
Increased water levels and flow rates meant
that these animals were probably swept
away, or were more vulnerable to preda-
tion. However, given the large dilution
effect of the flood, turtles were most likely
unable to find prey in the deeper waters.
Hence, turtles may have moved to the shal-
lower habitats, leaving the main body of
the billabong (the area of netting) in search
of prey.

In November, the number of turtles
caught remained low yet the water temper-
ature within the billabong had increased
and the water level had returned almost to
normal, as indicated by water marks on the
tree lines and reed beds. The bottom of the
main body of the billabong was cooler and
turtles may have sought warmer water in
which to forage where the ectothermic
invertebrates may have been more abun-
dant (Chapman et al. 1996). When the
water level receded, some turtles were

167

Murray River Special Issue

100 5
= 80 | — =
[^]
E
S |
SR OD
o | heather
A [m Aquatic Plants |
[e] ElFish |
EE 3 Plant Debris |
He
[s]
S
& 2 -
0^ =
C. expansa C. longicollis E. macquarii
n=15 n=17 n=24

Fig. 4. Cumulative proportional IRI values of diet items in the three turtle species (Chelodina
expansa, Chelodina longicollis and Emydura macquarii) from Lake Moodemere, Victoria during the

1996/97 breeding season,

stranded in smaller pools of water and
were observed to cross land to return to the
billabong. Crossing back to the permanent
billabong was gradual as the smaller pools
dried up and/or became devoid of food
resources before larger pools.

The number of turtles caught in Lake
Moodemere was maximal during January.
This was probably the result of three fac-
tors: increased water temperature,
increased immigration and decreased water
level. The warm water in the main section
of the billabong may have increased the
activity levels of turtles, making them
more apt to encounter a net during move-
ment. Also, the immigration of turtles
(Graham et al. 1996) from ephemeral bill-
abongs (that were becoming too warm or
too dry) into the main billabong would
have increased turtle abundance within
Lake Moodemere and hence, the probabili-
ty of capture. Finally, an increase in the
number of turtles moving around in the
water column, coupled with the decreased
water levels, would have increased the
catchability of turtles.

Shallower areas in which turtles were
thought to have occurred in the cooler
October and November months were, in
January, probably unfavourable habitats
for turtles due to very warm and hypoxic
water (Chapman er al. 1996). Turtles that

168

had been using the shallower outer region
of the billabong earlier in the season may
have moved back into the main water body
of the billabong in January. The turtles
could now perhaps sustain a higher meta-
bolic rate. Collections taken during this
study indicated that invertebrates and fish
were most abundant among the reeds and
snags by this time. Consequently, the need
for turtles to seek out other areas in which
to forage may have become unnecessary.

No turtles were recaptured over the dura-
tion of this study, either from their original
habitat of capture, or any other habitat.
This suggested that turtles disperse widely
throughout the billabong, and showed no
habitat preference. Given the fluctuations
in water level and temperature of Lake
Moodemere during the 1996/97 turtle
breeding season, it seems reasonable to
suggest that preferences may not have
been detected simply on the basis of the
billabong being so dynamic. In years of
greater billabong stability (i.e. non-flood
years), habitat partitioning by turtles may
be evident given enough netting over a
variety of habitats.

After flooding, none of the turtle species
appeared to have a preference for any partic-
ular habitat type. This suggested that all
three species of turtle shared the same habi-
lats within Lake Moodemere, and that the

'The Victorian Naturalist

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Part Two

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macqu Indiv55 [
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expan Indiv08
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longi Indiv20 |
longi Indiv30 - |
.expan Indiv35 | |
.expan Indivi9 "uS
expan Indiv25 I

.expan Indiv32 A

macqu Indiv06
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macqu Indiv48| |
.macqu Indiv29 | |
longi Indiv34 | |
macqu Indiv27_
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longi Indiv03
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macqu Indiv22
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longi Indiv45 T
expan Indiv42 -
longi Indiv51 |
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oo60BHNOSOO090000N80058O555o000o00to0httboionoo0nmio

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expan Indiv47 I
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expan Indiv56 |
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QOO00o0t mA

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0.0036 0.3189 0.6341

|
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0.9494 1:2647 Do

Bray-Curtis Dissimilarity

Fig. 5. Dendogram of turtles versus food eaten based on a Bray-Curtis association. C. /ongi —

Chelodina longicollis, C. expan — C. expansa and E. macqu — Emydura macquarii.

niche requirements of each individual
species were met within these shared habi-
tats.

Turtles were caught more frequently
within the medium to high reed bed areas
with higher complexities, compared to
open water areas of a billabong. This sug-
gested that there were favourable condi-
tions for turtles in areas associated with
higher habitat complexities and a greater
amount of prey. Unfortunately, the ordina-

Vol. 119 (4) 2002

tion of turtles by habitat type (Fig. 6) did
not support this assumption, probably due
to the limited number of dietary samples
taken from each of the five habitats.

The reference collections of prey types
were not distinguishable by habitat groups
but rather by seasonal groups (Fig. 3), and
suggested that although the same food cate-
gories were generally found within each
habitat, their abundances varied temporally.

Plant debris was the dominant food item

169

Murray River Special Issue

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*

A A Group 3
[72]

IE:

+ =
*
x
SSH Axis 1

Fig. 6. SSH ordination plot in two-dimensions based on turtle diet from a Bray-Curtis association.

Stress — 0.123.

in the stomachs of all species of turtle sam-
pled from Lake Moodemere, even surpris-
ingly, the obligate carnivore Chelodina
expansa. Plant debris included reed stems
and small particles such as seeds from ter-
restrial and aquatic plants. This material is
primarily cellulose and, being difficult to
digest, yields little nutritional value,
Georges (1982) studied the diets of fresh-
water turtles in eastern Australia and con-
sidered this plant material to be unimpor-
tant and eliminated the food category from
further analysis. Yet the ubiquitous pres-
ence of plant debris in the diet of turtles
from Lake Moodemere suggested that
some value must be gained (Bjorndal
1989). Vogt and Guzman Guzman (1988)
studied the dietary preferences of freshwa-
ter turtles in Mexico. There, turtles
retained a large proportion of plant debris
in their diet (more than any other food
type), especially when co-existing with
other species of turtle. This same study
also found that the absence of one species
of turtle in a water body influenced the diet
of the remaining species, with the domi-
nant food source shifting from plant debris
to insects. It seems sensible to suggest the
presence of plant material in the diet of
turtles may mirror a balance between

170

quantity and quality. Turtles may sate
themselves on plants when food of higher
nutritional value (invertebrates) is less
abundant. Conversely, plant debris may be
passively ingested while turtles forage for
benthic or littoral invertebrates (Legler
1978). The presence of plant debris in the
obligate carnivore, C, expansa, may sim-
ply have resulted from passive ingestion.

Metabolic and digestive rates in reptiles
vary in a mass-specific and thermal depen-
dent manner such that gut passage time
will affect the interpretation of stomach
contents (Coulson and Coulson 1986).
Smaller prey, such as zooplankton or
insects, and softer bodied prey, such as
shrimps Paratya australiensis, digest faster
than larger or more indigestible prey. This
implies that, if turtles are feeding on small-
er, more easily digestible prey, they will be
under-represented in the sample (Bjorndal
and Bolten 1992). Paratya australiensis
filter-feed on reeds and/or snags (Pringle e/
al. 1993) and the higher proportion of
plant debris from reed remains may be
linked to turtles passively ingesting reeds
while feeding on shrimps.

Fish rated highly on the index of relative
importance (IRI) scale, presumably
because of their mass. Fish matter, though

'The Victorian Naturalist

highly digestible, was more abundant than
smaller prey in the diet of the obligate car-
nivore, C. expansa. Research by Legler
(1978) demonstrated that, when all species
co-oecur in highly-productive, ephemeral
billabongs, resources are partitioned. Fish
and shrimp are consumed primarily by the
carnivorous species, with the omnivores
concentrating on invertebrates and aquatic
plants. When the turtles migrate back into
more permanent billabongs, such as Lake
Moodemere, resource levels are relatively
lower and prey items are no longer parti-
tioned (Chessman 1978).

After fish, the next most common dietary
item sampled from Lake Moodemere tur-
tles was aquatic plants. Interestingly, Æ.
macquarii is the only omnivorous turtle in
this billabong, and the only species
observed directly to eat aquatic plants. This

is consistent with the findings on the diet of

this species by Chessman (1986).

After aquatic plants, the next most com-
monly occurring, item in the diet of turtles
from Lake Moodemere was aquatic
inseets. Adults of both Coleoptera and
Hemiptera trap air beneath their hard,
shell-like forewings (elytra) and in
between microscopic, hydrofuge hairs on
the body. The entrapped bubble of air may
last several hours to days at low tempera-

tures (Ingram et al. 1997). At the start of

this study, the lower water temperatures
would have meant that the activity of these
diving beetles and bugs would have been
less, owing to both their ectothermic exis-
tence and that a given amount of entrapped
air would have lasted longer. As the water
temperature increased towards summer,
trips to the surface for oxygen replenish-
ment by these insects would have
increased in frequency, thereby increasing
the time vulnerable to predation by turtles.
Species of Coleoptera and Hemiptera
known to stay under water permanently
(Elmidae, Hydraenidae, Naucoridae)
(Ingram e al. 1997) were not found in
stomachs sampled during this study,
Corixidae, Notonectidae and Nepidae are
totally aquatic families of Hemiptera, liv-
ing within the water column (Williams
1980). These families come to the surface
to renew air supplies, with the Corixidae
and Notonectidae being very active.
Mesoveliidae are associated with the sur-

Vol. 119 (4) 2002

Part Two

face film or on floating aquatie plants
(Williams 1980). Of the Hemiptera found
in stomach samples of turtles from Lake
Moodemere, all were found in the water
column or near the surface.

Overall, then, the diet samples taken
from turtles in Lake Moodemere suggested
that resources were shared over all habi-
tats, The high proportion of plant debris in
their diets indicated that most turtles were
essentially omnivorous, eating solter-bod-
ied animals and passively ingesting plants,
However, caution must be exercised when
suggesting the method of prey selection
based on so few samples. Also, sampling
was conducted over a 24 hour period on
five oceasions. There is no information in
the literature regarding how frequently tur-
tles feed in the wild. Clearly, samples from
daily foragers would differ from those of
weekly foragers. Hence, the samples
obtained in this study may not truly indi-
cate the degree of resource partitioning
between these species.

All of the turtle species occupying, Lake
Moodemere showed an extremely narrow
niche breadth and a high degree of dietary
overlap, which suggested that turtles were
competing for a limited resource, in this
case food, The relationship between com-
petition and niche overlap is complex
(Krebs 1989). For instance, zero niche
overlap does not mean that interspecific
competition is absent. Conversely, niche
overlap does not always imply competi-
tion. In many cases niche overlap is only
used as a deseriptive measure of communi-
ty organisation (Krebs 1989). However,
high overlap occurs in both diet and habitat
during periods of resource scarcity if ani-
mals do not disperse widely,

On a niche breadth scale of 0 to 1, all tur-
tle species in this study would he classified
as dietary specialists. Recall, however, that
C. expansa is an obligate carnivore, €.
longicollis is an opportunistic carnivore and
E. maequarii is an generalist omnivore in
ephemeral billabongs or when all species
do not co-occur within a given habital
(Chessman 1983, 1984, 1986), Assuming
that sufficient samples were obtained dur-
ing the study to show the degree of dietary
specialisation of each turtle species, if
would seem that there was a low diversity
of prey species available to the turtles, This

171

Murray River Special Issue

may have resulted from the post-flooding
dilution effects making only certain prey
taxa vulnerable to predation.

Given the high degree of dietary overlap
found between the turtles in Lake
Moodemere, a particular type of food may
have been a limiting resource. Dietary
overlap was highest between the two carni-
vores (Chelodina expunsa and C. longicol-
lix), as expected. It was also high between
the obligate carnivore (C, expansa) and the
omnivore (E. maequarii). Interestingly,
dietary overlap was lowest between the
opportunistic carnivore (C. /ongicollis) and
the omnivore. Relative to other billabongs,
Lake Moodemere is lacking in macro-
phytes (pers. obs.). Therefore, it was possi-
ble that the omnivore shifted toward a
more carnivorous diet, partitioning the ani-
mal prey between all three turtle species.

When food is both diverse and abundant
within the billabong, in this case after the
flood, interspecific competition for food
resources may be lessened. Too few turtles
were sampled during and immediately
after the flooding event to discern if inter-
specific competition was reduced at that
time, When water levels dropped and the
temperature within the billabong increased,
a substantial number of turtles were cap-
tured (n — 54). The high degree of dietary
overlap, suggesting competition for food,
may have been an artefact of competition
with other predators at the billabong,

Other major consumers of zooplankton
and maeroinvertebrates in billabongs
include several species of water birds and
native and introduced fish (Hansson et al.
1978; Bunn and Boon 1993). Lake
Moodemere is considered to have an over
abundance of Red-fin Perca fTuviatus (G
Closs pers, comm.); many small members
of this species were caught in this study.
he perch population is considered density-
dependent because there were many stunt-
ed individuals. As for many density-depen-
dent fish, their large numbers and reduced
size result from high levels of both inter-
and intraspecific competition (Liebold
1996). Studies on the diets of fish in bill-
abongs (Harris 1985; Barlow er al. 1986;
Rowland 1992 in Ingram er al. 1997) have
shown that fish commonly eat Coleoptera,
Diptera, Hemiptera, Odonata and
Trichoptera, all of which are eaten by tur-

172

tles. Competition from Red-fin and occa-
sional Golden Perch Macquaria ambigua
may decrease the amount of food available
to turtles, forcing the turtles to seek non-
preferred foods. This may have bee
reflected in the amount of plant debris.

The majority of turtles caught in Lake
Moodemere contained very little in their
stomachs, reflecting the limit in food avail-
ability. Turtles that had empty stomachs
clustered more tightly on the ordination
plots and dendograms simply because they
were empty. The turtles with more food
items in their stomachs tended to eat a
wider variety of food types, rather than
specialise, although their niche breadths
were still quite low. This explains why
there was no separation of turtle species on
the dendograms, because only nine indi-
viduals contained a substantial amount of
food. Given the limited sample size, it was
decided to include the rarer food types in
the diet of turtles. Exclusion of rarer taxa
may have reduced ‘noise’ around patterns
seen in the dendogram and ordination plots
(Faith and Norris 1989), but the end result
would, most likely, not have changed.

The spread of all three species throughout
the dendograms groups indicated that all
three species of turtle were eating essential-
ly the same food, from the same habitats,
Turtles did not appear to favour any one
particular habitat type or prey item,

The habitat and prey selectivity displayed
by turtles in this study may not reflect the
present conditions, but those learned prev i-
ously, For instance, the three species of
turtles may have shared resources in this
study, not as a reflection of the current
post-flood conditions when resources were
presumably abundant, but as a reflection of
other seasons. In these perhaps more ‘typi-
cal years, water levels are lower, habitat
diversity is lower and food resources may
be limited. As such, the turtles may over-
lap in time, space and the utilisation of all
resources in this closed system.
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173

Murray River Special Issue

Invertebrates of the River Red Gum Forests

of the

Murray River

Andrea Ballinger! and Alan L Yen’

Abstract

The Murray River is surrounded by River Red Gum forests for much of its length. These forests are
habitat toa rich and diverse invertebrate fauna, both in the dominant River Red Gum trees and in the
surrounding ground layer, A brief introduction is provided to some of the important ecological roles
played by invertebrates in these forests, The responses of some of these invertebrates to flooding is
discussed, (He Victorian Naturalist 19 (4), 2002, 174-179)

Introduction

River Red Gum Zuealyprus eamaldulen
sis forests are a dominant feature of the
Murray River floodplains. Yen er al
(2002) recorded 6,666 invertebrates from
over 450 species when the canopy of two
individual trees was sprayed with inseeti-
cide, What do all these invertebrates do?
While inventory of the invertebrate fauna
is an essential first step in understanding,
what they do and how they interact with
ench other, there is still much to be done

before we have a clear understanding of

the structure and function of the inverte
brate Fauna; Yet, with the limited informa
tion available and with research currently

in progress, we ean still outline some of

the important roles that invertebrates play
in these Forests,

In this paper we provide examples of

invertebrate fauna typical of River Red
Guin forest and present a guild approach as
a means ol coping with the sheer diversity
of arthropods, We then use flooding as an
example of how the guild approach can be
used to understand the impact of ecolopi-
cal processes on invertebrate assemblages,

Invertebrates of the River Red Gum tree

The canopy invertebrate fauna ol Æ
camaldulensis is dominated by herbivorous
insects and their associated parasitoids and
predators, There are, however, other types
of invertebrates present, such as pollinators
and scavengers, The invertebrates on River
Red Gum ean be grouped into three broad
types; (1) species that feed on River Red
Gums; (2) species that feed on other inver-
(ebrates; and (3) the remainder, The follow-
ing information is based on Moore (1966),
Morgan (1986), Stone and Bacon (1994),

School of Biological Setences, Monash: University,

Clayton, Victori SOQ
52-54 Brushy Park Road, Wonga Park, Vietora 5115

174

Collett (2001), Yen (unpubl. data), and
Ballinger (unpubl, data).

Species that feed on River Red Gums

Foliage (leaf) ehewers are insects that bite
off whole chunks of leaf. There is a range of
species, primarily amongst the beetles (such
as the chrysomelid Paropsis atomaria, the
scarab Anoplognathus montanus, the wec-
vils Gonipterus spp.), as well as tree crick-
ets, Black masses of the pergid sawfly
Perga dorsalis are often seen moving on the
trees in search of suitable foliage.

Other chewing-type insects feed differ-
ently on the leaves, Some insects, predom-
inantly the caterpillars of several species of
moth, only feed on one surface of the leal;
their feeding exposes the leaf veins to give
the appearance of a skeleton. One of the
most common caterpillars to feed in this
way is the Gumleaf Skeletoniser, Uraba
lugens. A variation of this feeding behav-
iour involves moth larvae that bind two
leaves together with silk and skeletonise
the leaves on the inner surfaces,

Leaf miners feed between the upper and
lower epidermis of leaves, The adult female
oviposits into the leaf, and the larvae feed
as they move through the tissue, The feed-
ing produces an often species-characteristic
pattern on the leaf) Most leaf miners are
flies or moths, Examples include the
agromyzid fly Japanagromyza eucalvpti
and the ineurvariid moth Perthida sp.

Gall-inducing insects also lay eggs into
the plant tissue, However, unlike leaf min-
ers, ovipositing gall-inducing insects
release chemicals that induce the plant to
produce extra tissue, which the developing
larvae then feed on, The shape of the gall
is usually characteristic of the species,
Gall-inducing insects generally belong to

The Victorian Naturalist

the flies, wasps and true bugs. Examples are
the fly Fergusonina sp. (Fergusoninidae)
and the coecoid 4piomorpha frenchi
(Eriococcidae).

The Hemiptera (or true bugs) use stylet-
like mouthparts to feed, Some are sap-
suckers on leaves, such as the white lace
lerp, Cardiaspina albitextura while others,
such as the scale insect, Eriococcus cori-
aceus, feed on twigs. Leafhoppers
(Cicadellidae) are especially common on
River Red Gums, and include the wafer-
thin nymphs of Stenocotis spp.

High numbers of invertebrates are found

when the River Red Gum flowers. Adults of

many species depend upon nectar and
pollen, although their immature stages have
completely different feeding habits:
buprestid beetle larvae are wood borers, ich-
neumonid wasp larvae are predators, and
calliphorid fly larvae are carrion feeders.
Other visitors include thrips (Phlaeothrips
spp.). flower wasps, and ants. One of the
visitors that arrived after European settle-
ment is the Honey Bee Apis mellifera.

The bark of River Red Gums is shed, but
not completely detached from the tree, pro-
viding ideal habitat for subcortical fauna
(Baehr 1990). Bark probably functions pri-
marily as a protective environment, rather
than acting as a direct source of food.
Many foliage active invertebrates (herbi-
vores, predators, parasitoids) may shelter
beneath the bark. The main insects that
feed on bark are bark beetles and ambrosia
beetles that leave carvings, scribbles or
galleries on the wood beneath the bark
(New 1988).

Insects that feed on the wood of the liv-
ing tree are not as commonly seen because
of their habitat. They are generally the
immature stages of longicorn (in branches
and trunk) and buprestid beetles (in twigs
and branches) or cossid and hepialid
moths. They bore through the timber, leav-
ing galleries that can be used by fungal and
detritus feeders. The larvae of longicorn
and buprestid beetles can be parasitised by
ichneumonid and braconid wasps or
preyed on by clerid and cucujid beetles.
Root feeders include the larvae of scarab
beetles, such as Heteronyx and
Anoplognathus, and of the giant ghost
moth Trictena atripalpis. The ends of the
empty brown pupal cases of the latter are

Vol. 119 (4) 2002

Part Two

often found protruding from the ground
beneath the canopy.

Species that feed on other invertebrates

These can be divided into two groups:
predators and parasitoids. The main insect
predators in the canopy include: cantharid
beetles, mantids and neuropterans (which
use mandibles to chew their prey), asilid
flies and reduviid bugs (which use stylet-
like mouthparts). Spiders are abundant
under bark, particularly Clubiona spp.
(Clubionidae), and various jumping spiders
(Salticidae), White-tailed spiders,
Lampona cylindrata (Lamponidae) are
common, and are thought to prey on the
numerous sac spiders (Brunet 1994),
Family groups of the social huntsman,
Delena sp. (Sparassidae) are found under
larger sheets of bark, There are certain
groups of carabid beetle that rely on the
subcortical habitat provided by the bark of
River Red Gums (Baehr 1990), such as
Sphallomorpha sp., which is believed to
prey on ants moving up and down the
trunk (Matthews 1980), Other non-insect
predators include: pseudoscorpions, the
scorpion Lyehas marmoreus and cen-
tipedes. Many predators are highly mobile
and can move freely within and between
trees.

The Gumleaf Skeletoniser, is a good
example of how much diversity can be
associated with an individual species,
Austin and Allen (1989) recorded 22 para-
sitoid species supported by the immature
stages of the Gumleaf Skeletoniser. The
eggs of the Gumleaf Skeletoniser may be
parasitised by a small wasp, Trichogramma
sp. There may be egg predators such as the
predatory mite Microsmaris goannae
(Erythraeidae). There are larval parasitoids
such as ichneumonid and chalcid wasps
and tachinid flies (Campbell 1962). To
complete the complex, there could also be
pupal parasitoids such as the chalcid wasp
Brachymeria rubripes.

Other invertebrates

The remainder of the invertebrates
include species that scavenge upon detri-
tus, fungal feeders (such as booklice), for-
agers (e.g. ants such as Crematogaster sp.
seeking the honeydew exudations of sap-
sucking insects or nectar), and accidental

175

Murray River Special Issue

visitors (species that use the tree for shel-
ter, such as adult stages of aquatic insects),
Some invertebrates also use microhabitat
features, such as bird nests and tree hol-
lows, Bird nests provide shelter for beetles,
fies, thrips and moths. There are ectopara-
sites such as ticks and lice associated with
the vertebrates,

River Red Gums can have parasitic mistle-
toe plants on them. One of the interesting
associations is the feeding on mistletoe by
larvae of the lyeaenid butterfly, Ogyris
olane, These caterpillars are attended by ant
species from genera such as /ridomprmex,
Camponotus, Crematogaster, Monomorium,
Podomyrma, Ochetellus, Anonvehomyrma
and lroggattella (Eastwood and Fraser
1999), Many of these ant species have
pround-layer nests, so their movement
between the nest and the canopy is an
important link between these two strata,

Invertebrates of the ground and soil lay

lhe diversity seen in the arboreal fauna
extends to [orest-Hoor invertebrate assem-
blages, Although the ground layer is not as
rich in herbivorous species as the canopy,
there are many species ol more mobile her-
bivores, such as grasshoppers, as well as
the subterranean root feeders mentioned
previously, The pround layer has many
microhabitat features that provide habitat
for invertebrates, These include coarse

woody debris, logs, and rocks, Some of

these ean be microcosms of their own, with
a diverse tauna, but very different com-
pared to the canopy fauna, There are more
detritivores, scavengers, seed removers and
predators active on the ground.

Fallen leaves, twigs and bark form a layer
of organie litter on the forest floor, Although
the litter laver in River Red Gum forest is
not as deep as in wetter Forests (Glazebrook
and Robertson 1999), the decaying organic
matter is still an important resource to many
invertebrates, Many small beetles, such as
silvanids, feed on the moulds growing on the
litter (Matthews 1992),

Moths in families such as the oechophorids
have specialised to feed on dead leaves, thus
forming an important first step in the recy-
cling process, There are numerous other
detritivores, such as cockroaches and Ties,
as Well as a very rich micro-invertebrate
Fauna dominated by mites and springtails.

176

Many species of ants nest underground or
in logs, so their foraging, activity is greater
within this stratum. Ants are a major scav-
enping and predatory group, but some
species are also important seed removers,

he role of ants in seed dispersal is likely
to be relatively limited in River Red Gum
forest because these forests are largely
devoid of the shrubby understorey plants,
such as Acacia spp., which typically have
seeds adapted to dispersal by ants, Instead,
the main seeds available would be those of
River Red Gums and grasses. The green-
head ant, Rhytidoponera metallica, one of
the most commonly trapped ants in River
Red Gum forest, is recognized as an
important seed remover in other dry scle-
rophyll forests (Drake 1981, Hughes and
Westoby 1992), Seeds of shrubs adapted to
dispersal by ants tend to have hard coat-
ings that are not penetrated by the
mandibles of A. metallica (Rodgerson
1998), In contrast, Drake (1981) identified
Eucalyptus spp. seeds with relatively weak
coats as being vulnerable to being eaten by
R, metallica. Thus, the role of ants in River
Red Gum forest may be one of seed eaters
rather than seed dispersers,

Given the abundance of logs in River Red
Gum forests, wood feeding invertebrates are
not as common as might be expected. The
reasons for this are discussed in Mac Nally
er al, (2002), In areas that do not flood regu-
larly, termites (Coptotermes sp.) are very
abundant in fallen timber, Saproxylic beetles
found in River Red Gum forest include
Avion spp. (Bostrichidae), which bore into
the sapwood (CSIRO 1991), Like bark, logs
provide shelter, rather than a direct food
souree, to numerous species. Many ants,
including, — Zridomyrmex spp. and
Monomorium spp., nest in rotting logs.

lhe subterranean stratum is home to
many detritivores, predators, and immature
stages of canopy species. The soil species
perform several important functions,
including burrowing (which assists soil
aeration and water percolation) and mixing
of decomposing organic matter, The
important groups of soil animals include
earthworms, spiders, beetle larvae and
moth larvae, Living in à burrow potentially
confers an advantage in a semi-arid envi-
ronment, because temperature and humidi-
ty tend to be moderated in the burrow,

The Victorian Naturalist

However, it is likely that flooding prevents
burrowing taxa obtaining great diversity in
River Red Gum forest. The burrowing
wolf spider Lycosa leuckartii is found in
relatively open areas that flood infrequent-
ly. The beetle Clivina quadratifrons
(Carabidae) is captured in pitfall traps in
damp areas. Its powerful forelegs are used
for burrowing in sand (Matthews 1980).

Guilds

Mac Nally et al. (2002) and Yen er al.
(2002) emphasize the sheer diversity of
invertebrates found in River Red Gum for-
est. The composition of invertebrate
assemblages is not static, but varies both
spatially and temporally. Yen ef al. (2002)
found that there was considerable seasonal
turnover of species on two individual trees
located in close proximity to each other.
While acknowledging the limitations of
pitfall trapping, Mac Nally et al. (2002)
found the composition of ground-dwelling
invertebrate assemblages to vary between
seasons and years, as well as in response to
forest-floor characteristics. It is most likely
that there is considerable species turnover
associated with time (day/night, seasonal),
habitat condition (tree age, tree flowering,
soil moisture, organic litter loads on forest
floor), in space (both on highly-localized
and broader scales e.g. between individual
trees, and between trees in different geo-
graphical locations), environmental change
(floods, drought) and anthropogenic fac-
tors (clearing, grazing, fragmentation,
inappropriate fire).

Yen et al. (2002) identified the canopy
invertebrates trapped, to species using the
usual taxonomic characters that separate
them at the class, order, family and species
levels, However, with such a large number
of invertebrate species and with changes in
the composition of the fauna, it is difficult
to describe what each invertebrate species
is doing in River Red Gum forests. This
can be overcome to a certain extent by
assessing the ecological functioning of the
invertebrates. As many closely related
species fulfill similar ecological roles,
invertebrates can be classified according to
guilds as defined by Root (1967): ‘groups
of species that exploit the same class of
environmental resources in a similar way.’
Moran and Southwood (1982) developed a

Vol. 119 (4) 2002

Part Two

guild classification for arboreal insects in
the United Kingdom. The major guilds that
they defined are (1) phytophages (plant
feeders), divided into (1a) chewers and (1b)
sap-suckers; (2) epiphytic fauna; (3) scav-
enging fauna (excluding ants, but including
nectarivores); (4) predators, divided into
(4a) insect predators and (4b) other preda-
tors (excluding ants); (5) parasitoids; (6)
ants; and (7) tourists. This idea has been
extended to arboreal invertebrates in Borneo
(Stork 1987) and Australia (Yen 1989;
Abbott et al. 1992; Peters ef al. 2001),
although Abbott ef al. (1992) and Peters et
al. (2001) modified the guild definitions to
suit their particular studies, The use of
guilds permits examination of how groups
of animals, not necessarily taxonomically
related, utilise a resource. For example, a
possum that chews the leaves of a particular
species of eucalypt may be placed in the
same guild as a leaf chewing species of bee-
tle. A particular species can change guilds
in different life cycle stages. For example,
the larvae of buprestid beetles feed on wood
in the trunk (guild 1a), while the adults feed
on flowers of the tree (guild 3).

Invertebrate response to inundation

Under natural conditions, River Red Gium
forests are characterized by frequent flood-
ing, with extensive areas of forest inundated
for prolonged periods. Those invertebrates
usually found on the forest floor must either
move to higher ground or ascend the trunks
and/or canopy of River Red Gums. Thus,
ground-dwelling guilds in River Red Gum
forest are expected to display relative niche
plasticity and high mobility compared to
their counterparts in less variable habitats.
Regulation of the Murray River has sub-
stantially reduced the frequency and extent
of flooding. Flood mitigation potentially
alters the structure of ground-dwelling
invertebrate guilds, allowing species not
adapted to inundation to invade. In addition,
many species regarded as canopy inhabi-
tants spend some part of their life cycle on
the forest floor, and hence are affected by
alterations in flood regime, The next section
of this paper examines how changes created
by flood reduction ramify throughout all the
guilds, demonstrating that various guilds are
profoundly interconnected elements of a
single ecosystem,

177

Murray River Special Issue

In spring/summer 2000 the Barmah
Forest experienced major flooding for the
first time in several years. Immediately fol-
lowing the recession of floodwaters, we
sampled ground-dwelling, invertebrates al
different localities throughout the forest

that had been subjeet to inundation of

varying extents. The fauna on sites that
had been extensively flooded was very dif-
ferent from the fauna of sites that experi-
enced moderate or minor/no flooding.
Predatory carabids (ground beetles) com-
prised only 4% of the beetle fauna trapped
on sites which experienced minor or no
flooding, compared with 63% on sites
which experienced extensive flooding.
Similarly, numbers of wolf spiders
(Lycosidae) increased dramatically in
areas that had recently been inundated for
prolonged periods, Wolf spiders and cara-
bid beetles comprise a guild of ground-
dwelling, generalist predators. The domi-
nance of this guild immediately following
emersion is likely to reflect an inerease in
both live prey and carrion availability
resulting from aquatic taxa becoming
stranded by receding floodwaters, Many
carabids are adapted to hunt in the riparian
zone, l'or example, Cutadromus lacor-
dairei, a common species on llood-prone
sites, has been recorded preying upon frogs
(Littlejohn and Wainer 1978). Several ol
the dominant woll-spider species trapped
on sites soon after lloodwaters receded
also are associated with the edge of water-
ways, such as Artoria howquaensis
(Framenau submitted).

In order to exploit this short-term inerease
in prey availability, wolf spiders and cara-
bid beetles must be able to colonize areas
rapidly as floodwaters recede. All carabids
captured in areas that experienced exten-
sive flooding were fully winged, and hence
potentially capable of quickly dispersing
into new areas by flight. However, vertical
migration, onto tree trunks, or into the
canopy, also represents an important flood-
survival strategy for wolf spiders and cara-
bids, We trapped invertebrates on tree
trunks in both flooded and dry areas of for-

est, During periods when the forest floor
was inundated, searching under sheets of
bark on trees standing in floodwater

revealed high numbers of invertebrates. In
addition to the usual subcortical fauna, such

178

as pancake spiders (Hemicloea spp.) and
sac spiders (C/ubiona spp.), specimens of a
number of species commonly trapped on
the ground in newly emersed areas were
collected, including small wolf spiders (e.g.
Artoria howquaensis) and carabids (e.g.
Catadromus lacordairei), Thus, many
species that are typically thought of as
ground-dwelling may utilize arboreal habi-
tat when necessary. Since much terrestrial
prey must also climb to avoid rising flood-
waters, tree trunks potentially provide rich
pickings to the generalist-predator guild
during the inundation phase. Other general-
ist predators, such as marbled scorpions,
house centipedes, earth centipedes and
scolopendrid centipedes are commonly
trapped on both the forest floor and on tree
trunks in River Red Gum forest.

Ants, as a guild, potentially demonstrate
the greatest plasticity in terms of habitat
usage in response to flooding. The ground-
foraging ant fauna of River Red Gum for-
est is not particularly diverse, with only
about seven species being commonly pit-
fall trapped in Barmah forest, including:
Iridomyrmex sp., Rhytidoponera metallica,
Camponotus sp. and Paratrechina sp.
Although these species usually nest on the
ground, or in rotting logs, when the forest
floor flooded all common species were
found to shift the nest, complete with
reproductives, larvae and pupae to under
bark on tree trunks. Again, vertical migra-
tion enabled rapid recolonisation of areas
following the recession of floodwaters,
Although ant activity was temporarily
reduced by about half in areas that had
recently experienced extensive flooding,
ants were still pitfall trapped in high num-
bers on such sites,

Flooding also impacts on other guilds.
Inundation produces changes in canopy
structure, with increased leaf longevity and
size (Stone and Bacon 1995), This poten-
tially impacts on phytophagous guilds. In
addition, increased soil moisture resulting
from flooding is known to control popula-
tions of the Gumleaf Skeletoniser Uraba
lugens (Campbell 1962). Thus, the effects
of flooding are manifold and extend to all
areas of River Red Gum forest and to the
invertebrates that live there. A guild
approach to invertebrates lets us consider
the ecological processes in River Red Gum

The Victorian Naturalist

forest without being overwhelmed by the
great biodiversity.

Acknowledgements

The Murray-Darling Basin Commission provid-
ed funding for research on forest-floor inverte-
brates in River Red Gum forest. Thanks to
Volker Framenau (The University of
Melbourne) and Martin Baehr (Munich
Museum) for kindly identifying the wolf spiders
and carabid beetles collected. Ralph Mac Nally
and PS Lake provided doctoral supervision to
Andrea Ballinger.

References

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(1992) Arthropod fauna of jarrah (Eucalyptus mar-
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tion. Australian Journal of Ecology 17, 263-274.

Austin AD and Allen GR (1989) Parasitoids of Uraba
lugens Walker (Lepidoptera: Noctuidae) in South
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Braconidae. Transactions of the Royal Society of
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Baehr M (1990) The carabid community living under
the bark of Australian eucalypts. In The Role of
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Hampshire)

Brunet B (1994) The Silken Web: A Natural History of
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Campbell KG (1962) The biology of Rosella [ugens
(Walk,), the Gum Leaf Skeletoniser moth, with par-
ticular reference to the Eucalyptus camaldulensis
Dehn. (River Red Gum) forests of the Murray valley
region. Proceedings of the Linnean Society of New
South Wales 87, 316-338.

Collett N (2001) Biology and control of psyllids, and
the possible causes for defoliation of Eucalyprus
vamldulensis Dehnh, (River Red Gum) in south-east-
ern Australia. Australian Forestry 64, 88-95,

CSIRO Division of Entomology (1991) The Insects of
Australia: a textbook for students and research work-
ers, 2 ed, Volume IL. (Melbourne University Press:
Melbourne)

Drake WE (1981) Ant-seed interaction in dry sclero-
phyll forest on North Stradbroke Island, Queensland,
Australia: Australian Journal of Botany 29(3), 293-
310.

Eastwood R and Fraser AM (1999) Associations
between lycaenid butterflies and ants in Australia.
Australian Journal of Ecology 24, 503-537.

Framenau VW (submitted) Revision of the wolf spider
genus Artoria Thorell with the description of six new
species from floodplain habitats in Victoria, South-
eastern Australia (Araneae, Lycosidae).

Glazebrook HS and Robertson Al (1999) The effect of
flooding and flood timing on leaf liter breakdown
rates and nutrient dynamies in a River Red Gum

Part Two

(Eucalyptus camaldulensis) forest. Australian
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Hughes L and Westoby M (1992) Fate of seeds adapted
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Littlejohn MJ and Wainer JW (1978) Carabid beetle
preying on frogs, The Victorian Naturalist 95, 251-
352.

Mae Nally R, Ballinger A and Horrocks G (2002)
Habitat change in River Red Gum floodplains: fallen
timber and biodiversity. The Victorian Naturalist 119,
102-108.

Matthews EG (1980) 4 Guide to the Genera of Beetles
of South Australia; Part 1 Archostemata and
Adephaga. (South Australian Museum: Adelaide)

Matthews EG (1992) 4 Guide to the Genera of Beetles
of South Australia: Part 6 Polyphaga, Lymexyloidea,
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Adelaide)

New TR (1988) Associations between insects and
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Peters PJ, Read J and Sanson GD (2001) Variation in
the guild composition of herbivorous insect assem-
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Rodgerson I, (1998) Mechanical defense in seeds
adapted for ant dispersal. Ecology 79, 1669-1677.

Root, RB (1967) The niche exploitation pattern af the
blue-gray gnateateher. Ecological Monograplis 37,
317-350.

Stone C and Bacon PE (1994) Insect herbivory in a
River Red Gum (Eucalyprus camaldulensis Dehnh.)
forest in southern New South Wales. Journal
Australian Entomological Society 33. 51-56.

Stone € and Bacon PE (1995) Leaf dynamics and
insect herbivory in a Eucalyptus camaldulensis forest
under moisture stress, Australian Journal of Ecology
20, 473-481.

Stork NE (1987) Guild structure of arthropods fram
Bornean rain forest trees. Ecological Entomology 12,
69-80,

Yen AL (1989) Overstorey invertebrates in the Big
Desert, Victoria. In Mediterranean Landscapes in
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(CSIRO: East Melbourne)

Yen AL, Hinkley S, Lillywhite P, Wainer J and Walker
K (2002) A preliminary survey of the arboreal inver-
lebrate fauna of two River Red Gum trees Eucalyptus
camaldulensis near the Murray River. The Victorian
Naturalist 119, 180-185-

THE STORY OF THE MURRAY RIVER
By AS Kenyon, Melbourne

Those interested in the Murray River story should read The Nile of Australia, by

David John Gordon; Paving the Way, by Simpson Newlands; Half-crown Bob,
by Price Warung, and Knocking About, by August Pierce.

From The Victorian Naturalist LIX (1), 8 May 1942, pp 16-19.

Vol. 119 (4) 2002

179

Murray River Special Issue

A Preliminary Survey of the Arboreal Invertebrate Fauna of
Two River Red Gum Trees Eucalyptus camaldulensis
near the Murray River

Alan L Yen", Simon Hinkley', Peter Lillywhite', John Wainer"
and Ken Walker'

Abstract

In February and October 1999, the canopy and bark invertebrates from two individual River Red
Gum Eucalyptus camaldulensis trees in the Moira State Forest NSW were collected. A total of 458
morphospecies was collected from the canopy and 69 morphospecies from beneath the bark. In view
of the wide distribution of River Red Gum across Australia, this preliminary survey suggests that
there is a high level of invertebrate biodiversity associated with this tree species. (The Victorian

Naturalist 119 (4), 2002, 180-185)

Introduction

Many eucalypt species have a diverse
invertebrate fauna, primarily insects, asso-
ciated with them (Majer eż al. 1997). There
is considerable information available on a
few of these insect species, particularly
those that have attained pest status, but
there have been relatively few studies on
the total fauna. One of the common land-
scape components associated with the
Murray River is the River Red Gum
Eucalyptus camaldulensis Dehnh. A pilot
study, in an attempt to document the inver-
tebrate fauna of a single River Red Gum
tree, is reported here. There is considerable
seasonal variation in the composition of
the invertebrate fauna, so the project
involved assessing individual trees at dif-
ferent times of the year.

River Red Gum is a widespread species,
probably the most widely naturally distrib-
uted species of eucalypt aeross Australia
(Chippendale 1988). Invertebrate studies
on River Red Gum have involved studies
of pest species such as the Gum Leaf
Skeletoniser Moth Uraba lugens
(Campbell 1962) and psyllid bugs (Collett
2001) to estimates of the amount of insect
herbivory on foliage (Stone and Bacon
1994), There are no published studies on
the composition of the arboreal inverte-
brate fauna associated with this species.

' Museum Victoria, GPO Box 666E, Melbourne,
Victoria 3001

52-54 Brushy Park Road, Wonga Park, Victoria 3115
3/17 Fisher Street, East Malvern, Victoria 3145

180

Methods

Field work was conducted in the Moira
State Forest (off Coolamon Road), Riverina
Region, New South Wales (35*52'46" S
144*55'36" E). Sampling was undertaken
on two occasions: 8-10 February 1999 and
6-7 October 1999, [n February, a River Red
Gum was selected, sampled and then felled
to take foliage and wood samples. In
October, the invertebrate fauna of a nearby
tree was sampled. This tree was not felled
for foliage and wood samples.

Arboreal invertebrates

Arboreal invertebrates were collected by
spraying the canopy with synthetic
pyrethroid Reslin® (manufactured by
Wellcome) diluted 1:40 with water. The
insecticide was applied with an engine dri-
ven spraying machine (Stihl& SG17
Electronic backpack sprayer) that dispels
droplets of insecticide. The operator was
elevated to a height of up to 15 m with a
cherry picker. Spraying was conducted
early in the morning (about 0700) in order
to utilise rising warm air currents. Spraying
the whole tree took about one hour.

Pyrethroid insecticides work as knock-
down insecticides; insects succumb to the
insecticide after contact. The active life of
the insecticide is measured in terms of a
few seconds after exposure to sunlight.

The sprayed invertebrates fell from the
canopy, and were collected by placing a
continuous layer of plastic sheeting
beneath the canopy. The invertebrates
were lett to fall for an hour after cessation
of spraying, and then the invertebrates on

The Victorian Naturalist

the sheets were collected by forceps or
paint brushes. This continued for 4 hours
after spraying, after which there were no
more invertebrates falling on to the sheets.

The first tree was sprayed in fine weather
on 9 February 1999, A tree approximately
23 m in height, single trunk, with a good
head of foliage was used. One of the con-
ditions for the permit was that the tree was
not very old (less than 1.5 m dbh), did not

have tree holes or pipes, and was not of

Aboriginal cultural significance. The tree
had only one mistletoe plant. The tree was
estimated to be about 80 years old by New
South Wales Forest Commission staff.

A similar sized tree was sprayed on 7
October 1999 using the same regime. This
tree was located approximately 100 m
from the February tree.

Bark invertebrates

Invertebrates dwelling beneath the bark
were collected by direct searching before
spraying. The bark was peeled off from
ground level to a height accessible with a
step ladder (about 3-4 m). Invertebrates were
collected with forceps or paint brushes.

Invertebrates from other microhabitats

The leaf litter and woody debris beneath
the canopy of the tree was raked over and
invertebrates were collected by forceps
before spraying. Two samples of litter,
each from ground area of approximately |
* | m, were brought back to the laboratory
for extraction in Tullgren Funnels (heat
funnels). Samples were left in the Tullgren
Funnels for 5 days. The soil directly
beneath the litter searched was raked and
dug up to a depth of 10 em for inverte-
brates. Invertebrates were collected using
forceps. Only a small number of inverte-
brates were collected in February using
this technique, so it was not repeated in
October. Several samples of timber (trunk
and branches) were taken back to the labo-
ratory to see if any invertebrates emerged.
The timber was kept for 12 months.
Leaves were also kept, but little emerged
from these samples.

Identification

All specimens were sorted initially to a
higher taxonomic level - generally to the
ordinal level. However, millipedes and
centipedes were identified only to Class,

Vol. 119 (4) 2002

Part Two

and Hymenoptera was divided into ants
and non-ants (mainly wasps). Specimens
were then identified to the morphospecies
level. In some groups (Coleoptera,
Neuroptera, Diptera and Lepidoptera),
immature stages were treated as separate
from adults.

Specimen sorting and storage

All material was labeled and stored either
in 70% ethanol or pinned. Specimens are
to be initially used as part of a Museum
Victoria exhibition on River Red Gum bio-
diversity, after which it will be deposited
into the Museum collection,

Results and discussion
Total invertebrate fauna

A total of 8456 invertebrates, identified
into 547 morphospecies from 24 orders,
was collected from the two trees with all
the collecting techniques (Yen 2000).
Results presented in detail here are only the
canopy and bark samples because the other
microenvironments were probably under-
sampled. The identification of specimens to
morphospecies may result in the double
counting of certain species if (1) immature
and adult stages of the same species are
considered different morphospecies, or (2)
if there is marked sexual dimorphism
which has not been considered.

The potential effects of distinguishing
immature and adults as different morphos-
pecies can be estimated when the number of
morphospecies collected from the canopy is
considered (Table 1). The spraying resulted
in 458 morphospecies, of which 31 were
larval morphospecies. Consequently,
31/458 (6.8%) of morphospecies, were lar-
vae, When the number of individuals is con-
sidered, 316/6666 (4.7%) were larvae, For
the bark fauna, the percentages are 11,60%
(8/69) for morphospecies or 0.7% (8/1175)
for individuals (Table 1).

Canopy invertebrate fauna

In February 1999, a total of 297 morphos-
pecies, represented by 4092 individuals from
19 orders, was collected from the canopy
spraying, and 260 morphospecies (2574 indi-
viduals) were collected in October. A total of
458 morphospecies (6666 individuals) was
obtained from the two trees (Table 1),

The most dominant invertebrate groups
in terms of number of morphospecies were

181

Murray River Special Issue

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The Victorian Naturalist

182

Part Two

Table 2. Number of species of invertebrates collected only in February, only in October and in both

February and October (total).

Order Canopy Bark
February October Total February October Total

Araneae 12 16 12 2 11 4
Scorpionida 0 0 0 0 | 0
Pseudoscorpionida 0 1 l 0 0 |
Acarina 3 l 0 l 5 |
Chilopoda 0 0 0 0 l 0
Diplopoda 0 0 0 0 2 0
Collembola l l 0 0 3 0
Thysanura 0 l 0 0 0 0
Odonata 3 0 0 0 0 0
Blattodea 2 l 0 0 0 0
Mantodea 1 0 0 0 0 0
Orthoptera 4 2 | 0 2 0
Phasmatodea | 0 0 0 0 0
Psocoptera 0 2 0 0 l 0
Hemiptera 59 19 9 2 8 0
Thysanoptera 2 | 2 l 0 0
Neuroptera l l | 0 0 0
Neuroptera larvae 0 | 0 0 0 0
Coleoptera 29 37 16 | D 0
Coleoptera larvae 2 6 2 0 5 0
Strepsiptera | 0 0 0 0 0
Diptera 29 21 25 0 0 0
Diptera larvae l 0 0 0 0 0
Lepidoptera 0 0 1 0 0 0
Lepidoptera larvae 7 9 3 0 3 0
Hymenoptera - ants i 3 S 2 4 2
Hymenoptera - other 42 38 12 0 | 0
Total 207 161 90 9 52 8

bugs (Hemiptera), flies (Diptera), wasps
(Hymenoptera), beetles (Coleoptera), spi-
ders (Araneae), and ants (Hymenoptera) in
February (Table 1), and in October the
descending order of dominance was beetles,
flies, wasps, spiders and bugs (Table 1).

In terms of numbers of individuals, the
most dominant groups were ants, beetles,
bugs, flies, thrips (Thysanoptera) and mites
(Acarina) in February (Table 1), while bee-
tles, flies, bugs and wasps dominated the
October sample (Table 1). The reduction in
numbers of ants between February and
October was most marked.

Bark invertebrate fauna

In February 1999, a total of 17 morphos-
pecies, represented by 64 individuals from
14 orders, was collected from beneath the
bark, and 60 morphospecies (1111 individ-

uals) were collected in October. A total of

69 morphospecies (1175 individuals) was
obtained from the two trees (Table 1).

At the morphospecies level, the dominant
groups were spiders and ants in February,

Vol. 119 (4) 2002

and spiders, bugs, mites, beetles, and ants
in October (Table 1).

When the number of individuals under
bark is considered, the dominant groups in
February were ants, spiders and mites (Table
1). In October, the ants still dominated the
bark fauna, followed by millipedes
(Diplopoda) and pseudoscorpions
(Pseudoscorpionida) (Table 1). It is interest-
ing to note that only seven invertebrate
orders were collected under bark in February
compared to 14 orders in October.

Differences between February and October

Of the 458 morphospecies collected
from the canopy, 207 were collected
only in February, 161 only in October,
and 90 (19.7% of morphospecies) were
collected in both February and October.
Of the 69 morphospecies collected from
beneath bark, nine were collected only
in February, 52 only in October, and
eight (11.6% of morphospecies) were
collected in both February and October
(Table 2).

183

Murray River Special Issue

Table 3. Number of species only on canopy or
on bark, or on both canopy and bark (C^ B).

Order Canopy Bark C+B
Araneae 33 10 7
Scorpionida 0 1 0
Pseudoscorpionida 1 0 |
Acarina | 4 3
Chilopoda 0 l 0
Diplopoda 0 2 0
Collembola | 2 |
Thysanura l 0 0
Odonata 3 0 0
Blattodea 3 0 0
Mantodea | 0 0
Orthoptera 6 1 l
Phasmatodea l 0 0
Psocoptera 2 I 0
Hemiptera 81 4 6
Thysanoptera 4 0 |
Neuroptera 3 0 0
Neuroptera larvae I 0 0
Coleoptera 80 5 2
Coleoptera larvae 9 4 |
Strepsiptera 1 0 0
Diptera 85 0 0
Diptera larvae | 0 0
Lepidoptera | 0 0
Lepidoptera larvae 18 l 0
Hymenoptera - ants 7 0 8
Hymenoptera - other 92 1 0

Total 436 37 31

The total numbers of individuals collected
from the canopy in February was 4092
compared to 2574 in October (Table 1). The
increase was due to more spiders (154 in
February: 75 in October), mites (230:19),
bugs (646:474) and significantly, ants
(1201:115). In contrast, there were more
beetles (625:765) and non-ant Hymenoptera
(132:274) in October. For the fauna beneath
bark, the number of individuals in October
was 1111 compared to 64 in February.
Practically all invertebrate orders had more
individuals beneath bark in October, and the
major components were spiders, scorpions,
pseudoscorpions, millipedes and ants.

These seasonal differences may be attrib-
utable to seasonal activity patterns in the
different invertebrate groups. Some may
simply have been more active at one time of
the year, while others may have been utilis-
ing different microhabitats or had different
patterns of seasonal development. Some
species may be inhabiting the soil or litter
layers during one season and move into the

184

bark and canopy in another; this could
occur for groups such as scorpions and ants.

Links between canopy and bark

A total of 436 morphospecies were col-
lected only from spraying the canopy, 37
morphospecies collected only from
beneath bark, while 31 morphospecies
were found in both canopy and bark sam-
ples (Table 3). However, the canopy spray-
ing could have sampled bark invertebrates
from further up the tree.

River Red Gum invertebrate diversity

Stone and Bacon (1994) studied insect
herbivores in River Red Gum forests in the
nearby Gulpa Island State Forest (NSW).
They found 49 species of phytophagous
insects in the canopy, and the dominant
orders were the Phasmatodea, Hemiptera,
Thysanoptera, Coleoptera, Diptera,
Lepidoptera and Hymenoptera, In our
study, Phasmatodea and Lepidoptera
occurred in very low numbers.

Majer er al. (2000) used canopy knock-
down on four species of eucalypts:
Eucalyptus creba and E. moluccana in east-
ern Australia and E. marginata and E, calo-
phylla in western Australia. They found 27
orders of invertebrates; 967 morphospecies
in eastern Australia and 687 morphospecies
in western Australia. Their method differed
in that their sampling regime involved
spraying 10 trees of each species and collec-
tions were made with 10 * 0.5 m° funnel-
shaped nets under each tree. The procedure
was repeated seasonally for four seasons.
The larger number of morphospecies col-
lected by Majer er al. (2000) may be attrib-
utable to the larger number of eucalypt
species in their study, more intensive sea-
sonal sampling, and geographical differ-
ences in the fauna.

Eucalyptus camaldulensis is the most
widely naturally-occurring species of euca-
lypt in Australia. Our study involved two
individual trees in one locality, and even
allowing for over-estimation of number of
morphospecies by distinguishing between
immature stages and adults, some 458 mor-
phospecies were collected from the canopy
and 69 morphospecies from beneath the
bark. We could expect this number to
increase substantially if the study was
repeated to allow for species turnover asso-

The Victorian Naturalist

ciated with seasonal changes, changes in the
state of the tree (e.g. flowering), differences
between individual trees in the same area,
differences between geographical locations,
differences between extreme environments
(temperate versus arid), and allowing for the
fact that the fauna dwelling within the tim-
ber is under-represented.

Since European settlement, River Red
Gum woodlands have undergone a range of
environmental changes such as timber har-
vesting, fragmentation, and altered hydrol-
ogy regimes. If the effects of these and
other changes on the invertebrate fauna of
River Red Gums is to be understood, basic
information on the composition of the
invertebrates associated with River Red
Gums over a broad range is required.

Acknowledgements

The authors wish to thank the following
Museum staff and volunteers for assisting with
field work: John Coventry, Ross Field, Alan
Henderson, Rhys Jones, Carolyn Rasmussen,
Chris Rowley, Dean Smith and Rodney Start.
The New South Wales Forest Commission pro-
vided logistic support in terms of staff assis-
tance, chain saw work and use of the cherry

Part Two

picker, and we wish to thank Geoff Heagney,
Nick Saunders, David Leslie, Lisa Whiting,
Wayne O'Brien and Paul Childs for their help.
The project was undertaken under the conditions
of the State Forests of New South Wales Special
Purposes Permit No 05352 issued to AL Yen.

References

Campbell KG (1962) The biology of Roselia lugens
(Walk), the gum-leaf skeletonizer moth, with partic-
ular reference to the Eucalyptus camaldulensis
Dehnh. (River Red Gum) forests of the Murray val-
ley region, Proceedings of the Linnean Society of
New South Wales 87, 316-338.

Chippendale GM (1988) Eucalyptus, Angophora
(Myrtaceae), Flora of Australia 19. (Australian
Government Publishing Service; Canberra)

Collett N (2001) Biology and control of psyllids, and
the possible causes for defoliation of Eucalyptus
camaldulensis Dehnh. (River Red Gum) in south-
eastern Australia. Australian Forestry 64, 88-95.

Majer JD, Recher HF and Ganesh S (2000) Diversity pat-
terns of eucalypt canopy arthropods in eastern and
western Australia, Ecological Entomology 25, 295-306.

Majer JD, Recher HF, Wellington AB, Woinarski JCZ
and Yen AL (1997) Invertebrates of eucalypt forma-
tions, In Eucalypt Ecology: Individuals to
Ecosystems, pp 278-302. Eds J Williams and JCZ
Woinarski. (Cambridge University Press; Cambridge)

Stone C and Bacon PE (1994) Insect herbivory in a
River Red Gum (Eucalyptus camaldulensis Dehnh.)
forest in southern New South Wales. Journal of the
Australian Entomological Society 33, 51-56.

Yen AL (2000) Biodiversity in River Red Gum. The
Bush Telegraph May-July 2000, 7.

The Murray River in The Victorian Naturalist
NOTES ON A COLLECTING TRIP TO THE MURRAY AND LODDON RIVERS

By C French, Jun.
Read before Field Naturalists’ Club of Victoria, 11 January 1897

4th November. — Drove to Salt Lake and caught a couple of Lizards—a
Monitor and a lace Lizard, or Iguana; on the latter were noticed a great many ticks
similar to the ones found on Opposums and Native Bears. Bee-eaters were seen in
numbers, but they were only just commencing to tunnel, and none had eggs.
Orange-fronted Ephthianuras were seen in fairly numerous quantities, but though
some hours were spent searching round the edges of this lake amongst the
Salicornia bushes, only one nest was taken... Much to our gratification a nest of
Chestnut-eared Finch was found underneath and attached to a Brown Hawk's nest.

10th November — crossed into New South Wales to work some swampy coun-
try ... birds were numerous, amongst which the following were conspicuous:
Painted and New Holland Snipe, Dottrel (of four species), Rails (of two species),
Bee-eaters, Ibis, Black-tailed Tribonyx, &c.

Altogether upwards of 100 different species of birds were noticed. Plants to the
number of about seventy were collected, in flower, which was a very fair collec-
tion, considering the dry season. Beetles were fairly numerous, about eighty
species being collected. Appended are lists of the specimens obtained on our
excursions.

From The Victorian Naturalist XM (10), January 1897, pp 127-134.

Vol. 119 (4) 2002 185

Murray River Special Issue

Aquatic Macroinvertebrates of the Murray River

Phillip J Suter' and John H Hawking

This article is dedicated to the memory of Prof WD (Bill) Williams (1936-2002)
whose early books on the aquatic invertebrates of Australian inland waters stimu-
lated interest and research on the ecology of these important groups.

Abstract

The aquatie macroinvertebrate fauna of the Murray River is presented based on surveys and scientif-
ic literature. Insects dominate the aquatic macroinvertebrate fauna but the non-insect groups such as
the crustaceans, molluscs, worms and other primitive classes are also well represented, The distribu-
tion of the fauna is discussed in relation to the four physical and biological zones of the river; the
Headwater, Riverine, Mallee and Lower Murray tracts. The biology and ecology of many groups is
documented and, where possible, the impacts of human influences on the invertebrates of the Murray
River are given. (The Vietorian Naturalist 119 (4), 2002, 186-200)

Introduction

The aquatic invertebrates are to a certain
extent the forgotten fauna of the Murray
River. With the exception of the Murray
River crayfish and of course the nuisance
insects such as mosquitoes the aquatic
invertebrates come a distant second to the
more obvious aquatic icons such as
Murray Cod, Golden Perch, Macquarie
Perch and other native fish. However, the
aquatic invertebrate fauna are an important
component of a healthy functioning river

and should not be viewed as just a part of

the food supply for fish. The aquatic
macroinvertebrates are an essential compo-
nent of a functioning aqualic ecosystem.
They are primarily decomposers with her-
bivores and predators making up the func-
tioning feeding groups (Bolton and Brock
1999). Their feeding activities mobilise
carbon and energy from plants through to
the higher aquatic orders. In addition the
aquatic macroinvertebrates have been used
successfully as indicators of water quality
and river condition (Norris ef al. 2001).

The aquatic macroinvertebrate fauna of

the Murray River as a whole were not
examined systematically until 1980 when
the then River Murray Commission com-
menced their biological monitoring pro-
gramme which is still ongoing today. Over

Department of Environmental Management and
Ecology, La Trobe University Albury/Wodonga
Campus, Cooperative Research Centre for Freshwater
Ecology, PO Box 821 Wodonga, Victoria 3689
' Murray Darling Freshwater Research Centre,
Cooperative Research Centre for Freshwater Ecology,
PO Box 921 Albury, NSW 2640

186

430 macroinvertebrates taxa were recorded
from the Murray River between Jingellic
and Lake Alexandrina during 1980-1985
(Bennison ez al. 1989). Insects dominated
the aquatic invertebrates with crustaceans,
molluses and worms also well represented.
Sources of information for this paper
include (Bennison er al. 1989; Boulton and
Lloyd 1991; Suter er al. 1993) and person-
al observations by the authors, Throughout
the paper reference will be made to the
highly modified flow regimes caused by
the consumptive use of water from the
river. This has undoubtedly influenced the
macroinvertebrate composition and distrib-
ution (Geddes 1990; Walker 1990), but
detailed discussion of these effects is
beyond the scope of this paper. An excel-
lent presentation of the hydrological and
geomorphological structural changes to the
river can be found in Thoms er al. (2000).
The Murray River physically and biolog-
ically can be divided into four major zones
or tracts (Bennison ef al. 1989; Thoms et
al, 2000) (Fig. 1);
ethe Headwaters tract reaches from the
Murray's source to Albury. The gradient of
the river is steep and the streams are domi-
nated by cobble, boulder and sand sub-
strates with little organic material. This
zone includes the alpine areas down to the
foothills (Fig. 2);
* ihe Riverine tract reaches from Albury to
the junction of the Edwards River. The
gradient is low, the substrate is dominated
by sand and silt with moderate levels of
organie material and the river flows across

'The Victorian Naturalist

South Australia :
i Darling River

|
Murray River \ l

\
i
1 |
|
ren af
+
i
\

Mallee
-— Tract N. ow
Q Murray |
TAM Tract |

! Victoria \

Part Two

New South Wales

Lachlan River

Murrumbidgee River

Riverine Tract
5 |

A Murray River
|
p |
I A.
| ‘a
| Headwater~
Tract E

Fig. 1. The four physical and biological tracts of the Murray River.

a broad floodplain and is multi-channelled
with extensive floodplain wetlands;

ethe Mallee tract reaches from the
Edwards junction to the Darling junction.
The low gradient continues, and the sub-
strate is sand and silt with moderate levels
of organic material. Throughout this tract
the river flows along a single narrow
watercourse; and

ethe Lower Murray tract reaches down-
stream of the Darling junction to the
Murray mouth (Fig. 3). The river gradient
is the lowest in this tract, the substrate is
dominated by sand and silt with moderate
levels of organic material, and the river is
now highly regulated with locks and weirs
which have altered the characteristics of
the river from lotic to lentic.

€ 2

Fig. 2. Headwater tract near Jingellic.

Vol. 119 (4) 2002

This paper provides an overview of the
aquatic macroinvertebrate fauna of the
Murray River, including both the main
channel and the floodplain wetlands that
together constitute the ecological compo-
nent and integrity of the river, and relates
their distribution to these zones.

Cnidaria

The Cnidaria is represented in the
Murray River by at least three genera, two
hydrozoans (Hydra and Cordylophora)
and the freshwater jellyfish Craspedacusta
sowerbyi. The solitary hydroid Hydra is
found in the Riverine, Mallee and Lower
Murray Tracts but the colonial hydroid
Cordylophora appears restricted to the
Lower Murray Tract (Bennison ef al.

Fig. 3. Lower Murray tract in South Australia.

187

Murray River Special Issue

1989). Both these genera occur on logs and
other such stable substrates. Their distribu-
tion may be greater than the artificial sub-
strate data suggests but other sampling
methods used in rivers may limit the ade-
quacy of collections.

The freshwater jellyfish Craspedacusta
sowerbyi has been recorded in the Hume
reservoir but it is rarely seen because it is
small (25 mm diameter) and almost trans-
parent. This limited distribution in the
Murray system may be a product of sam-
pling effort. Little is known about this
species although Williams (1980) recorded
that it is usually found floating near the
surface of large freshwater lakes and reser-
voirs.

Turbellaria (Flatworms)

Flatworms are recorded throughout the
river from the headwaters to Lake
Alexandrina. In the headwaters Spathula
sp., a very dark animal, is commonly found
on rocks. Ball (1977) recorded that all
species of Spathula are found in cool, well
oxygenated mountain water. Cura pingus is
a lowland species and has been recorded
from above Lake Hume to the end of the
Murray River. The taxonomy of this group
is limited and formal identification requires
serial sectioning of properly preserved ani-
mals (St Clair et al. 1999), It is highly like-
ly that there are more species than these
two in the Murray, particularly in the
Headwater tract (St Clair et al. 1999).

Temnocephalidea

The temnocephalans are small oval-
shaped animals with 2-6 apical tentacles.
The temnocephalans are ectosymbionts on
freshwater crayfish. Some species live on
the carapace of these large crustaceans and
others in the gill chamber where they feed
on small insect larvae, rotifers and nema-
todes, and algae (Williams 1980). An indi-
vidual crayfish can often have multiple
species on its body. A study by Cannon
and Sewell (1994) suggested that each
crayfish species had a distinct temnocepha-
lan fauna associated with it. The taxonomy
of this group is currently being revised. At
least nine species have been recorded from
the Murray River with Temnocephala
caeca in the Headwater tract and Temnoce-
phala chaeropsis mainly trom the Mallee
and Lower River tracts.

188

Fig. 4. Tubificid worm Branchiura sowerbyi.

Nematomorpha (Horsehair worms)

Adult horsehair worms, genus Gordius,
have been recorded in pools in the
Headwater and Riverine tracts of the
Murray River down to Corowa. The larvae
and juveniles are parasitic in terrestrial
invertebrates (insects) where they encyst
after being eaten and later break out of the
host as an adult when the host is near water
(Williams 1980).

Annelida (Worms and leeches)

The oligochaetes (segmented worms) are
common throughout the Riverine, Mallee
and Lower Murray tracts. Worms do occur
in the Headwater tract, but numbers are
generally low. Bennison er al. (1989) only
identified the worms as non-tubifids and
tubificids. The non-tubificid worms were
abundant in the Riverine and Mallee tracts
particularly in the depositional zones
where organic material accumulates. The
tubificid worms were low in abundance
along the length of the river suggesting
very low levels of organic pollution. One
of the most noticeable tubificid worms is
Branchiura sowerbyi, a distinctive hairy-
gilled worm recognised by the posterior
half having conspicuous gills (Fig. 4). It is
common but not abundant in all tracts
downstream of Albury occurring in deposi-
tional zones where there is a build up of
fine organic material.

The leeches were represented by two
families, Glossiphoniidae and Richard-
sonianidae. The Glossiphoniidae are small
translucent leeches that prey on inverte-
brates particularly freshwater gastropods
(Williams 1980) and are low in abundance
in the Headwater, Mallee and Lower
Murray tracts. The larger Richard-
sonianidae were rare in the river and were
only recorded in the Lower Murray tract.

'The Victorian Naturalist

Part Two

Fig. 5. Murray River Mussel A/athyria jacksoni.

Mollusca (Snails and mussels)

The molluscs are well represented in the
Murray River with 5-10% of the communi-
ty composition in the main channel of the
Headwater tract, but they decline to less
than 5% in the Riverine, Mallee and Lower
Murray tracts (Bennison ef al. 1989). The
distribution of many species may have
changed since regulation of the Murray
began in the early 1900s (Sheldon and
Walker 1993),

Bivalvia (Mussels, pea-clams)

The small bivalves Sphaerium sp.,
Pisidium sp. and Corbiculina australis
occur in the main channel of the river with
Sphaerium most abundant above Lake
Hume, Pisidium in the headwater streams
and C. australis along the length of the
river from the headwaters to Lake
Alexandrina.

Two large bivalve molluscs are associat-
ed with the Murray River. The common
riverine mussel Alathyria jacksoni (Vig. 5)
occurs in the main stream channel in areas
with flow, and the floodplain mussel
Velesunio ambiguus is found in the slow or
still waters of backwaters and billabongs.
In the Lower Murray tract the hydrology
has been highly modified by the presence
of locks and weirs resulting in
stationary/slow flowing reaches rather than
a natural flowing river. Walker (1990) sug-
gested that this change has provided the
opportunity for Velesunio ambiguus (the
floodplain species) to invade the main

channel of the river at the expense of

Alathyria jacksoni (the riverine species).
Velesunio ambiguus lives in the calm
waters behind the weirs and A. jacksoni in
the turbulent waters below the weirs
(Walker 1990).

A population of a third species of mussel
A. condola exists below Lake Mulwala.

Vol. 119 (4) 2002

Fig. 6. Gastropod snail.

The mussels have a larval stage (glochidia)
which is parasitic on fish (Walker 1990)
and it is most likely that A. condola has
colonised the Murray on fish migrating
from the Murrumbidgee River via the
Mulwala Canal.

Gastropoda (Snails)

About 18 species of gastropod molluses
or snails (Fig. 6) have been recorded from
the lower Murray River and were abundant
in the first half of the 20" century (Sheldon
and Walker 1993). Recent work along the
lower reaches suggests that many species
have very low abundances or no longer
occur (Bennison ef al. 1989; Boulton and
Lloyd 1991; Sheldon and Walker 1993).
One such species, Notopala sublineata
hanleyi, is a live bearing snail that was
common from the lower Murray but now
appears extinct in the river. However, an
extant population was reported by Sheldon
and Walker (1993) taking up residence in
the Kingston and Loveday irrigation
pipelines near Barmera, South Australia.
Sheldon and Walker (1993: 298) suggest
that ‘this is possibly the last surviving pop-
ulation of N. hanleyi’. In addition a num-
ber of other species of snails have a greater
abundance in pipelines than in the lower
river, including, Ferrissia petterdi, Gabbia
australis, Glyptophysa aliciae, Thiara
balonnensis and *Angobia angasi’. All
these species are detritivores feeding on
organic and microbial accumulations and
the inside of the pipelines provides ideal
habitat compared with the highly regulated
lower Murray River (Sheldon and Walker
1993). The pipelines provide the variabili-
ty of wetting and drying that once occurred
in the lower Murray, but does not occur
now due to regulation. This refuge in the
irrigation pipelines is also a nuisance for
irrigators as the snails can block sprinklers

189

Murray River Special Issue

and drip irrigation lines. The sculptured
snail Thiara balonnensis still occurs in the
river from Lake Mulwala to South
Australia but is now rare in the lower
reaches of the river except in the irrigation
pipelines. The freshwater limpet Ferrissia
petterdi is still common throughout the
Murray and its flood plain being found in
fast flowing waters as well as in billabongs
and backwaters.

There are seven species of the family
Planorbidae represented along the Murray,
with riverine and floodplain species.
Glyptophysa gibbosa, Isidorella hainesii,
Glyptophysa aliciae, Bayardella cosmeta
are found mainly in the river, whereas
Gyraulus meridionalis, G. tasmanicus and
Helicorbis australiensis are more common
in the backwaters and billabongs. The
riverine species have helicoil shells (nor-
mal coiled snail) and the floodplain species
have flat, ammonite-like shells. These
snails all graze on aquatic plants.
Glyptophysa gibbosa, Isidorella hainesii,
Gyraulus meridionalis, Helicorbis aus-
traliensis, Glyptophysa aliciae and
Bayardella cosmeta, occur along the
length of the Murray whereas G. fasmani-
cus only occurs in streams of the upper
Headwater tract. Regulation has been
implicated in the loss of aquatic plants in
the Murray River (Thoms et al. 2000) par-
ticularly where there is raised turbidity
reducing light availability (Lower Murray
tract) and river bank erosion (Riverine,
Mallee and Lower Murray tracts). This
reduction of aquatic plants in the main
channel may also have affected the distrib-
ution of the planorbid snails. In floodplain
wetlands where the aquatic plants are still
abundant the planorbid snails still can be
found.

A number of introduced gastropod snails
have successfully colonised the Murray
River. Physa acuta, a snail introduced
from Europe or North America, was
known from only the lower Murray prior
to 1970 (Smith and Kershaw 1979).
However, this species has spread the
length of the Murray and now competes
with the planorbid snails and in many
cases has replaced the native species.
Bennison ef al. (1989) recorded the planor-
bid snails /sidorella sp. and Physastra gib-
bosa downstream of the Hume weir until

190

Fig. 7. The water mite Peza ops

1980 after which it was replaced by Physa
acuta. Potamopyrgus niger was introduced
from New Zealand and occurs in the lower
Murray tract. It is possible that this species
will also become widespread throughout
the Murray and its tributaries.

Arachnida (Water mites)

Water mites are a common fauna of the
Murray River and its floodplain wetlands
but are not well documented because they
are difficult to identify. Reliable identifica-
tion of water mites of the Headwater tract
has shown that there are representatives of
three major groups: Hydracarina,
Halacaroidea and Oribatida. The last two
groups are few in number with only single
species of each group being recorded,
whereas the Hydracarina are abundant and
diverse. An unknown species of Orabatida
is commonly collected from gravel beds in
the river, Also commonly collected is the
halacaroidean Peza ops, a terrestrial-like
mite, whose gnathosoma has a long,
curved, slender rostrum, with the pedipalps
inserted above the rostrum (Fig. 7). The
common Hydracarina are Coaustralio-
bates, Austroliobates, Killimobates,
Flabellifrontipoda. Hydroma and
Unionicola, Watermites are also common
in floodplain wetlands with over 20 mor-
pho-species recorded from wetlands along
the Murray in South Australia (Suter ef al.
1993).

Water mites have also been found para-
sitising the freshwater mussels, with six
species being associated with Velesunio
ambiguus and 3 of those with Alathyria

The Victorian Naturalist

z + " *
Fig. 8. The Common Yabby Cherax destructor
jacksoni. Five species of the genus
Unionicola have only been recorded from
the Murray in South Australia. The sixth,
Unionicolopsis opimipalpis, has been
found on V. ambiguus collected from the
Murray River near Albury (Viets 1980).

Crustacea (Shrimp, yabbies and crabs)

The erustaceans are a major component
of the aquatie invertebrate fauna of the
Murray River, and are the dominant inver-
tebrate group in the lowland tracts of the
river. In the Headwater, Riverine and
Mallee tracts the Crustacea represent
between 2-5% of the aquatic invertebrate
community, but in the Lower Murray tract
the proportion of crustaceans increases to
10-15% of the macroinvertebrate commu-
nity (Bennison et al. 1989).

Amphipoda (Sideswimmers, scuds)

The amphipods, side swimmers or scuds,
are small fast swimming crustaceans that
have a very distinct distribution in the
Murray River. Generally they are rarely
found in the main channel upstream of the
Darling River junction. Amphipods
become a major component of the lower
river fauna in South Australia with

Austrochiltonia spp. and two species of

gammarid amphipods reaching high abun-
dances downstream of Morgan.

Decapoda (Yabbies, crayfish, shrimps and
prawns)

Shrimps and prawns are represented
along the Murray River by three species,
Paratya australiensis, Macrobrachium
australiense and Caridina mecullochi. AN
three species occur in both the main chan-
nel and floodplain wetlands. Paratya
becomes more abundant in the main chan-
nel in the lower sections of the river sys-
tem and this could be a response to altered

Vol. 119 (4) 2002

Part Two

Ihe Murray Crayfish Euustacus

Fig. 9.
armatus.

flow regime due to the presence of locks
and weirs, In contrast, Macrobrachium is
more prevalent in the foothills of the
Headwater tract. Caridina is rare at all
sites along the river.

The Murray River has two species of
freshwater crayfish, the Common Yabby
Cherax destructor (Fig. 8) and the Murray
River Crayfish Euastacus armatus (Fig. 9).
Both species are omnivores feeding on
detritus, algal biofilm and carrion. The
yabby is the smaller of the two species
(7250 g in weight; Geddes 1990) and is
common in billabongs, backwaters and
other wetlands on the floodplain.
Following floods, numbers can increase
rapidly and large numbers are caught from
wetlands along the floodplain. Cherax
destructor now occurs in the main channel
of the river in the Lower River tract where
the locks and weirs have changed the flow
regime from a river to a series of long nar-
row lakes. The Murray River crayfish is
the second largest freshwater crayfish in
the world (reaching up to 3 kg in weight;
Geddes 1990) second only to the giant
freshwater crayfish As/acopsis gouldi from
Tasmania. It lives in the main channel of
the river and tributaries where flow rates
are higher. Geddes (1990) noted that C.
destructor is adapted to slow flowing,
warm water and is active in the warmer
months (September to May) whereas, Æ.
armatus is adapted to cool waters and
strong. flowing rivers and is active in the
cooler months (May to October) when
oxygen concentrations are high. The
occurrence of the two species in the
Murray River follows a similar pattern to
the mussels with the riverine species (X.
armatus) being replaced by the floodplain
species (C. desiructor) in those sections of

191

Murray River Special Issue

the river which are controlled by the locks
and weirs, Geddes (1990) suspects that
natural populations of E. armatus are now
extinct in South Australia. The distribution
of the Murray River crayfish is also influ-
enced by fishing pressure and habitat mod-
ification. Snags in the Murray River pro-
vide large stable substrates for macroinver-
tebrates and both number of species and
abundance is higher on snags than on the
associated river bed (Lloyd et al. 1991).
The removal of over 13 million snags from
the river (Thoms ef al. 2000) not only has
affected the fish populations but also the
macroinvertebrates, and most probably the
Murray River crayfish.

The small spider crab Amarinus lacustris
is commonly found in streams with elevat-
ed salinities. This species is only found in
the lower Murray downstream of Murray
Bridge where it lives in the roots of the
willow trees that dominate the riparian
zone.

Syncarida

The syncarids are primitive crustaceans
that lack a carapace over their thorax and
although extinct syncarids were marine
inhabitants all living forms are now
restricted to fresh waters (Williams 1980).
A single specimen of a syncarid crustacean
Koonunga sp. has been recorded from the
Murray River near Albury. Syncarids of
this genus are usually associated with deep
hyporheic gravels, and so are rarely found
in rivers unless washed out during high
flows.

Isopoda

Two species of isopod occur in the
Murray River, Heterias pusilla and
Tachaea picta. H. pusilla is a small isopod
which lives in fine organic deposits in the
Riverine, Mallee and Lower Murray tracts.
They are occasionally recorded from the
upper catchment, but they generally are
rare. Tachaea picta has a more restricted
distribution occurring only in the Lower
Murray tract downstream of the Darling
junction, but rarely downstream of
Morgan. The distribution of Tachaea is
usually restricted to the Darling River
where it is parasitic on shrimps and
prawns, During Darling River flood,
Tachaea picta is washed into South

192

Australia and becomes common in the
lower Murray River.

Insecta

The insects are the dominant macroinver-
tebrate fauna in the Murray River repre-
senting over 70% of the communities in
the main channel and wetlands in all tracts.

Ephemeroptera (Mayflies)

The mayflies are a primitive order of
insect with an aquatic nymphal stage and
two terrestrial winged stages, the subimago
and imago or spinner. They are diverse in
the Murray River with over 20 species
being recorded along its length but they
represent less than 5% of the aquatic com-
munity. The number of species present
declines along the river’s length with
approximately 15 species in the Headwater
tract, less than ten species at Albury and
downstream in the Riverine tract and only
four species downstream of the Darling
River junction. This reduction in taxa can
also be seen at the generic level with
Coloburiscidae (Coloburiscoides) Onisci-
gastridae (Tasmanophlebia) Baetidae
(Edmundsiops) Caenidae (/rpacaenis and
Tasmanocoenis) and the Leptophlebiidae
(Atalophlebia, Nousia, Koorrnonga,
Ulmerophlebia, Neboissophlebia and
Austrophlebioides) common in the
Headwater tract. In the Riverine tract
downstream of Albury only Atalophlebia,
Neboissophlebia, Edmundsiops, Irpacaenis
and Tasmanocoenis are found. Atalo-
phlebia, Cloeon and Tasmanocoenis are
the only genera that occur in the Lower
Murray tract. The reduction of taxa may
reflect a natural gradient from upland to
lowland river conditions as Coloburis-
coides, Edmundsiops, Nousia, Koor-
rnonga, Austrophlebioides and Neboisso-
phlebia all occur in cool, highly oxygenat-
ed flowing streams that are usually associ-
ated with upland reaches. The genera
Atalophlebia, Cloeon and Tasmanocoenis
are more commonly associated with warm,
slow flowing habitats.

Pardo et al. (1998) recorded a decline in
mayfly diversity in the Mitta Mitta River
downstream of Dartmouth dam (11
species) compared with an unregulated trib-
utary (17 species). There were fewer lep-
tophlebiid mayflies in the regulated com-

The Victorian Naturalist

pared with the unregulated river. They
argued that the pattern of flow and cold
water releases in summer in the regulated
Mitta Mitta Rivers adversely affected the
mayfly community. The relative abundance
of nymphs of Coloburiscoides was not dra-
matically affected by the regulated flows
(21% in unregulated and 27% in regulated),
but Edmundsiops hickmani declined from
53% to 13% (Pardo et al. 1998). These two
species are restricted to the fast flowing
mountain streams where they are found on
rocks or on logs in the fast flowing water.
Coloburiscoides wedge their bodies
between rocks using the spines on their
gills as anchors (Dean and Suter 1996) and
use the long hairs on their legs and mouth-
parts to trap particles of organic material
flowing by in the fast flowing water.

The two genera of the Baetidae recorded
in the Murray River, Edmundsiops and
Cloeon, occupy the lotic and lentic habitats
respectively. Edmundsiops is restricted to
the upper catchment, whereas Cloeon is
associated with the lower tracts in station-
ary waters and wetlands particularly in
beds of aquatic plants.

The nymphs of the Oniscigastridae
(Tasmanophlebia) occur in standing waters
and slow flowing reaches of rivers and
streams from the mountains to near sea
level. They are normally found on sandy
substrates (Dean and Suter 1996), but they
do not occur downstream of Albury in the
Murray.

The caenid mayflies, the smallest
mayflies in Australia, are found along the
river's length and also in the floodplain
habitats, These mayflies are detritivores
and live in silt and sand particularly on
logs and in leaves on the bottom. They are
rarely seen as they emerge from the river
to fly as dusk falls and they only fly for a
short period, maybe two to three hours,
during the night. Swarms may continue all
night but emergence ceases as the sun
starts to rise. Dawn swarms continue for an
hour or two after sunrise. The caenids are
poor fliers and dispersal is also restricted.
Five species are commonly found along
the Murray River. /rpacaenis deani and an
undescribed species Tasmanocoenis sp. B
is found in the Headwater tract down to
Albury; Tasmanocoenis sp. B, Tasmano-
coenis tillyardi, Tasmanocoenis tonnoiri

Vol. 119 (4) 2002

Part Two

and Tasmanocoenis arcuata occur from
Albury downstream to Echuca; and
Tasmanocoenis tillyardi and Tasmano-
coenis arcudta are common in the
Riverine, Mallee and Lower Murray tracts.
The two headwater species /. deani and
Tasmanocoenis sp. B appear to be influ-
enced by inter-basin transfers of water
from the Snowy River catchment.
Irpacaenis deani has higher abundances
above the transfer point at Khancoban and
Tasmanocoenis sp. B increases down-
stream of the transfer point (McInerney
2000). In years when transfer volumes are
high Tasmanocoenis sp. B abundances
increase and years when transfer volumes
are low favours /rpacaenis deani.

Odonata (Damselflies and dragonflies)

The dragonflies and damselflies have
aquatic larval stages and terrestrial adults,
Sixteen species of damselflies and 28
species of dragonflies have been recorded
from the Murray River and its floodplain.
They represent less than 5% of the
macroinvertebrate community in the main
channel of the river (Bennison ef al. 1989),
but up to 10% in the wetlands. Many of the
species are obligate stream-dwellers,
restricted to the flowing part of the river
channel. However, downstream from
Echuca the river flow has been reduced by
the construction of many locks which form
large impoundments of still-waters which
have allowed colonisation by standing
water species.

In the montane section of the Headwater
tract the damselflies (Zygoptera) are
restricted to three species, Austro-
argiolestes calcaris, A. icteromelas and
Synlestes weyersii. The fauna changes after
Khancoban to a valley fauna, with
Nososticta solida and Rhadinosticta sim-
plex the dominant damselflies down the
length of the river with Pseudagrion aure-
ofrons from Lake Mulwala to South
Australia. The floodplain wetlands contain
standing water species Xanthagrion ery-
throneurum, Austrolestes analis, A. leda,
Ischnura heterosticta and I. aurora.

The dominant species of dragonflies
(Anisoptera) in the upper montane reaches
of the Headwater tract are dustroaeschna

flavomaculata, and Synthemis eustalacta

the larvae of which survive the winter

193

Murray River Special Issue

snow conditions. Below the snow line the
dragonfly fauna is characterised by the
gomphids Austrogomphus guerini,
Hemigomphus gouldii, the aeshnids,
Austroaeshna atrata, A. inermis, A. pul-
chra and the synthemistid, Eusynthemis
brevistyla. The fauna changes after
Khancoban to a valley
Austroaeschna unicornis, Austrogomphus
ochraceus, Apocordulia macrops,
Cordulephya pygmaea and Eusynthemis
virgula. Austroaeschna unicornis ranges
from the Khancoban valley reach through
to South Australia, whereas the other
species are only common to above Lake
Mulwala, except for 4. macrops which has
been recently collected from the Barmah
Forest, near Mathoura (Hawking unpubl,
data). The gomphid A. ochraceus is
replaced by its congener 4. australiae from
Lake Mulwala to the lower reaches in
South Australia. Memicordulia tau is very
abundant in the lakes and locks but it also
found along the total length of the river.
The opportunistic species, Hemianax
papuensis (Fig. 10), Orthetrum cale-
donicum, Diplacodes bipunctata and D.
haematodes, can be found in the slow
flowing and backwater areas of the lower
Murray now occurring in both the wet-
lands and main channel.

Plecoptera (Stoneflies)

Stoneflies are cold water adapted and the
nymphs of most stoneflies occur through-
out winter and spring, and are rarely found
during summer. McInerney (2000) record-
ed reduced abundance of stoneflies in
spring to summer as they emerged to terres-
trial adults. Stoneflies are generally found
in fast flowing cool water streams and
rarely oceur in wetlands on the floodplain.

av.

Fig. 10. Larva of the dragonfly Hemianax
papuensis,

194

fauna of

Therefore they are almost all restricted to
the Headwater tract where their relative
abundance is less than 5% of the macroin-
vertebrate community. Stoneflies do occur
in the Riverine tract in areas of fast flows
and usually a constricted river channel (e.g.
the Barmah Choke). The Murray River
fauna has representatives from four fami-
lies, with the Gripoterygidae being domi-
nant with at least 15 species in the genera
Dinotoperla, Riekoperla, Newmanoperla,
Illiesoperla and Leptoperla. MclInerney
(2000) recorded abundant Dinotoperla ser-
ricauda and Riekoperla rugosa down-
stream of the Khancoban pondage due to
the reduced temperature of the water from
the Snowy inter basin transfer.
Downstream of Albury stoneflies are rare
with Dinotoperla serricauda being record-
ed in the Mallee tract where the velocity of
water is high. In the lower river tract only
one species has been found. Dinotoperla
evansi was recorded in the Murray River
near Murray Bridge. This warm water
species also lives in farm dams (Suter and
Bishop 1990) and may well have colonised
the lower Murray from this source,

The other families represented are the
Austroperlidae, Notonemouridae and
Eustheniidae. The Eustheniidae are preda-
tory stoneflies and are represented by three
genera, Cosmioperla, Thaumatoperla and
Eusthenia. These large insects are only
found high in the headwaters of the upland
tributaries of the Murray River. Also only
in the headwater tributaries are the
Notonemouridae (Austrocerca tasmanica)
and the Austroperlidae (Austropentura vic-
toriae, and Acruroperla atra). Acruroperla
atra is found associated with accumula-
tions of dead leaves or leaf packs in pools
in the upper catchment.

Megaloptera (Dobsonflies)

The Corydalidae are large (up to 50 mm)
predatory aquatic insects with a short lived
terrestrial adult. They are represented in
the Murray River by a single species
Archichauliodes (Riekochauliodes) gut-
tiferus (Walker) which is restricted to the
Headwater tract where the river is fast
flowing and the substrate is dominated by
rocks and cobbles overlying sand.
Archichauliodes larvae can also be found
under rocks in dry stream beds in upper
catehment intermittent streams.

The Victorian Naturalist

Fig. 11. A backswimmer (Notonectidae).

Hemiptera (True bugs)

Aquatic bugs are characteristic of still
waters and backwaters, and so are often
associated with wetlands rather than rivers.
In the Murray over 30 species are recorded
from along the bank edge of the main river
usually in areas where flows are slow but
their relative abundance is less than 5%.
The aquatic bugs are nearly all predators
and consume gaseous oxygen from the air
rather than dissolved oxygen from within
the water. They lay their eggs in or on
plants or attached to a firm substrate
(Lansbury and Lake 2002) and the females
of one genus of giant water bugs
(Belostomatidae: Diplonychus) lay their
eggs on the back of the males (Hawking
and Smith 1997). The aquatic bugs occupy
two distinct habitats, being either fully
aquatic or surface dwellers.

The fully aquatic bugs are dominated by
the water boatmen (Corixidae), with
Micronecta, Sigara, and Agraptocorixa the
most common genera, and the backswim-
mers (Notonectidae; Fig. 11) represented in
the Murray River by Anisops and Enithares.
Micronecta annae is common along the
whole length of the river and in floodplain
wetlands. M. gracilis and M. robusta are
also common in the Riverine, Mallee and
Lower Murray tracts. In the main channel of
the Murray, Sigara sublaevifrons and
Agraptocorixa hirtifrons appear restricted to
the Headwater tract, but it is widespread in
wetlands from Albury to South Australia
(Suter et al, 1993). Sigara truncatipala is
restricted to the Lower Murray tract where-
as Agraptocorixa eurynome was throughout
the river's length.

Four other families, Nepidae, Naucoridae
and Belostomatidae (Fig. 12) and Pleidae

Vol. 119 (4) 2002

Part Two

Fig. 12. The giant water bug Diplonychus eques
(Belostomatidae).

are also fully aquatic and occur in the
Murray, but usually are rare and mainly
associated with floodplain wetlands. The
Nepidae are represented by the water scor-
pion (Laecotrephes tristis) and the needle
bug (Ranatra dispar). These are large
species (up to 50 mm) with a long respira-
tory tube extending from their abdomen
which enables them to hunt below the
water surface and breath air through the
tube which they extend beyond the surface
tension of the water. The giant water bug
Diplonychus eques (Belostomatidae) and
the creeping water bug Maucoris congrex
(Naucoridae) are also quite large (>15 mm
long) and occur in macrophytes particular-
ly in floodplain wetlands. As the name
suggests the pygmy backswimmers
(Pleidae) are quite small (<2.5 mm) which
makes this group rarely seen, but they do
occur in wetlands along the Murray.

The surface dwelling bugs include the
water striders Tenagogerris, Rheumato-
metra, and Limnogonus (Gerridae), the
small water striders Microvelia (Veliidae),
the water — treaders Mesovelia
(Mesoveliidae) and the water measurers
Hydrometra (Hydrometridae). These taxa
occur in both the main channel and flood-
plain wetlands of the Murray River.
Tenagogerris euphrosyne and Rheumato-
metra philarete occur in the Headwater
tract and Limnogonus sp in the Riverine,
Mallee and Lower river tracts. Microvelia
oceanica occurs in the Riverine tract,
Microvelia peramoena along the full
length of the river including floodplain
wetlands and Mesovelia hungerfordi is
found in all tracts of the Murray both in the
river and its wetlands.

195

Murray River Special Issue

Neuroptera (Lacewings)

There are two families of lacewings that
oecur in the Murray River, the Osmylidae
and the Sisyridae. The osmylids
(Kempynus sp.) are often found on large
rocks and boulders in the splash zone in
small upland streams, and have not been
recorded downstream of Jingellic. The
sisyrids or sponge flies (Sisyra sp.) occur
in the Murray but are rarely found as they
live in sponges on logs or rocks. Sponge-
flies lay their eggs on vegetation overhang-
ing a stream and the emergent larvae drop
to the water and swim to a freshwater
sponge. The larvae use their long straight
jaws to probe the sponge tissue and feed
on the contents (New 1991). They have
been collected from logs in the Barmah-
Millewa forest but may be more wide-
spread, wherever sponges occur.

Coleoptera (Beetles)

Bennison ef al. (1989) recorded the bee-
tles as the second most species rich group of
aquatic macroinvertebrates in the Murray
River. Over 90 species were recorded with
greatest richness in the Headwater and
Riverine tracts. The beetles represented 15-
2095 of the composition of the macroinver-
tebrate community in the Headwater tract,
less than 1594 in the Riverine tract, less than
394 in the Mallee tract and less than 1096 in
the Lower Murray tract. Both the riffle bee-
tles (Elmidae) and tiger beetles (Dytiscidae)
dominated the Headwater tract but many
species of the tiger beetles were also com-
mon in the Riverine, Mallee and Lower
Murray tracts, particularly in the floodplain
wetlands.

The riffle beetles can be found feeding
on algae attached to logs and rocks in fast
flowing streams in the upper catchment
where they are both abundant and species
rich. The headwater streams have at least
10 species within the genera Kingolus,
Simsonia, Notriolus and Austrolimnius.
Downstream of Albury the numbers of
species and genera declines with only
Kingolus, Austrolimnius and Coxelmis
(total of five species) in the Riverine tract
and only a single species of Kingolus and
Coxelmis in the Mallee and Lower Murray
tracts. In the lower gradient stretches of the
Murray the elmids are found associated
with logs and snags.

196

The predatory diving/tiger beetles (Fig.
13) are very active hunters that use
gaseous oxygen to breathe and therefore
must return often to the water surface to
replenish their oxygen supply. They are
usually associated with pool or pond habi-
tats, Tiger beetles range from small (<5
mm), medium (5-15 mm) to large (15-34
mm). The small species in the Murray
River include Antiporus with three species
in the headwaters (4. blakei, A. femoralis
and A. gilberti), but only a single species,
A. gilberti, downstream of Albury.
Liodessus gemellus, L. amabalis and
Sternopriscus multimaculatus occur in the
Lower Murray tract, whilst Limbodessus
compactus and Allodessus bistrigatus are
found along the full river's length. The
medium sized tiger beetles include
Rhantus suturalis found in the Headwater
and Riverine tracts only, and the wide-
spread Eretes australis and Megaporus sp
that are found in wetlands in all tracts and
occasionally in the main channel. The
large predatory diving beetles Cybister sp
has only been recorded from the Mallee
tract.

Diptera (True flies)

The true flies are the most species rich of
the aquatic insects in the Murray River
with 158 different species recorded by
(Bennison ef al, 1989). It is likely that the
number of species of dipterans is higher
than this due to the improvements in the
taxonomic literature since the study. The
flies represented between 20 and 30% of
the total species composition in the
Headwater tract, but increased their domi-
nance in the Riverine and Mallee tracts
with 40 to 50% of the total species rich-
ness. In the lower Murray tract the impor-

Fig, 13. The tiger beetle Sternopriseus
(Dytiscidae).

The Victorian Naturalist

tance of the dipterans declined to between
30 and 40%.

Adults of the biting midges (Ceratopo-
gonidae), blackflies (Simuliidae) and mos-
quitoes (Culicidae) are the main nuisance
insects associated with the river. The lar-
vae of the ceratopogonids are found along
the length of the Murray, but there are
more species in the Riverine and Mallee
tracts than in the Headwater or lower
Murray tracts. These animals are more fre-
quently found in the drying mud of wet-
lands and in the lower Murray in saline
seepages along the cliffs near irrigation
areas where they can cause significant nui-
sance value (e.g. Loxton; Suter unpub-
lished data). The blackflies are usuallv
associated with flowing water and firm sta-
ble substrates like rocks and logs. There
are few species in the Murray all upstream
of Swan Hill, with Austrosimulium furio-
sum in the Headwater tract, Austro-
simulium montanum, A. bancrofti, Simu-
lium ornatipes and Simulium nicholsoni in
the Riverine tract. Mosquitoes are rare in
the main channel of the Murray River, but
they do breed on the floodplain particular-
ly following floods when water becomes
trapped in isolated pools and ponds.

The non-biting midges (Chironomidae)
are the most diverse of the dipterans, and
are found in all habitats along the length of
the river. There are predatory midges
(Tanypodinae), organic feeders
(Chironominae, blood worms), filter feed-
ers (Chironominae, Tanytarsini) and wood
borers (Chironominae). The wood borer
Stenochironomus watsoni is found in all
iracts of the river associated with logs and
submerged bark [rom riparian trees. Many
of the filter feeders (Kiefferulus martini,
Rheotanytarsus spp. and Tanytarsus spp.)
also are associated with logs and snags
where they build tubes from which they
can catch organic material flowing past.
Rheotanytarsus spp., which are only found
in the Headwater tract, construct a silk net
attached to its tube to trap its food. The
Chironominae dominate the Riverine,
Mallee and Lower Murray tracts.

An interesting observation was made by
Pettigrove (1989) that the larvae of
Procladius paludicola showed abnormali-
ties of the mouthparts in the Mallee region
of the Murray River. Similar abnormalities

Vol. 119 (4) 2002

Part Two

have been shown to be associated with
exposure to chemicals such as heavy met-
als and pesticides (Madden er al. 1995;
Madden ef al. 1992: Warwick 1990). In the
lower Murray tract P. paludicola is rare in
the main channel of the river, but is the
major predacious midge in saline evapora-
tion basins and other wetlands on the
floodplain (Suter et al. 1993; Suter et al.
1995). This species is also abundant in rice
fields and with Chironomus tepperi can
attain very high numbers and become a
nuisance insect. Chironomus tepperi is an
early coloniser of wetlands following
floods and can attain very high numbers
providing a primary food supply for native
waterfowl (Maher and Carpenter 1984).
However, in rice fields these large num-
bers can cause damage to the rice crop
(Stevens 1995) thereby escalating these
beneficial insects to an economic nuisance,
The rapid colonisation and population
growth of C. tepperi in floodplain wetlands
and rice fields appears associated with the
high levels of organie material present in
the sediment (Suter er a/. 1995).

Trichoptera (Caddisflies/Casemoths)

The caddisflies have a terrestrial stage that
resembles a moth, but the larvae are aquatic
and construct a diverse array of cases in
which they live. In the main channel of the
Murray the caddisflies represent approxi-
mately 1596 of the aquatic community in the
Headwater tract and between 5-15% in the
Riverine, Mallee and Lower Murray tracts
(Bennison ef al. 1989). Although they are
found in a wide range of habitats the greatest
number of species occur in cool flowing
streams and as such have a very distinct
upper catchment distribution in the aquatic
macroinvertebrate community of the Murray
River. At least 12 families (Atriplectididae,
Calamoceratidae, Calocidae/Helicophidae,
Conoesucidae, Ecnomidae, Hydrobiosidae,
Hydropsychidae, Hydroptilidae, Leptocer-
idae, Limnephilidae, Odontoceridae,
Philorheithridae) with 730 species of caddis-
flies are recorded from the Headwater tract
whereas the Riverine, Mallee and Lower
Murray tracts have only 3 families present
(Ecnomidae, Hydroptilidae and
Leptoceridae) with fewer than 15 species.

The caddisflies are sensitive to environ-
mental change, particularly in the high

197

Murray River Special Issue

quality waters of the upper catchment.
They have a diverse range of habitat use
and diet and so alteration to water quality,
hydrology or addition of pollutants affects
their community structure. McInerney
(2000) found that there was a reduction of
the community structure of the caddisflies
as a response to the inter-basin transfer of
water from the Snowy Scheme. The preda-
cious caddisflies Taschorema evansi and
Ecnomina sp. show a reduction in abun-
dance downstream of the inter-basin water
transfer while the leptocerid caddisflies
Triplectides ciuskus and Triaenodes sp.
became more abundant downstream of the
inflows.

The diet of larval caddisflies can be
divided into three main types based on
method of collection of food (Neboiss
1991). There are the net spinners,
(Hydropsychidae) that construct silken
retreats with a silk net at the entrance used
to capture organic particles carried past in
the current. The Hydropsychidae (Fig. 14)
are almost restricted to the Headwater tract
where three species of Cheumatopsyche
and one species of Diplectrona occur.
However, Cheumatopsyche also occurs
downstream of Hume and Yarrawonga
weirs where there is fast flowing water.
The case makers (Calamoceratidae,
Calocidae, Helicophidae, Conoesucidae,
Hydroptilidae, Leptoceridae, Limno-
philidae, Odontoceridae and Philorheith-
ridae) shred and chew on leaves, or graze
on algae. The Leptoceridae is the most
species rich family in the Murray. In the
headwaters at least 12 species are found
mainly in the genera Notalina (5 spp.)
Triplectides (4 spp.), Oecetis (2 spp.) and
Triaenodes (1 sp.). Triplectides australis
and Triplectides australicus occur along
the length of the Murray with T. australis
mainly in the main channel and T. australi-
cus in the floodplain wetlands. Lectrides
varians, an endemic species to South
Australia, occurs in the lower Murray near
Murray Bridge.

The third group include the free living
Ecnomidae and Hydrobiosidae. Species in
these families are predators. The ecnomids
occur throughout the river and its flood-
plain, but at the species level there are dis-
tinct changes from headwaters to Lake
Alexandrina. In the headwaters three

198

"E i» & - Pip has" na " 4
Fig. 14. Larva of the Hydropsychidae (Tri-
choptera).

species occur with Eenomus pansus, E.
continentalis, and Ecnomina F sp. AV9
(Cartwright 1997) common in the rivers
and streams, but downstream of Albury Æ.
pansus is only found in the Riverine and
Mallee tracts. In the Albury area E. pansus
occurs in the main channel of the river and
E. turgidus and E. cygnitus are found in
wetlands on the floodplain. In South
Australia E. pansus and E, russellius are
both in the main channel and E. turgidus
occurs with E. pansus in the wetlands.

Lepidoptera (Aquatic moths)

Moths of the family Pyralidae, subfamily
Nymphulinae (Fig. 15), are the only lepi-
dopteran group to have aquatic larvae or
aquatic caterpillars (Hawking 2001), The
taxonomy of this group is still in its infan-
cy although recent work has enabled dif-
ferent species to be recognised (Hawking
2001). Five species of pyralid are found
along the Murray River. One species
inhabits rocks in the fast flow regions of
the Headwater tract. The larvae construct
silk retreats on rocks and feed on the
attached algae on the rock surface. In con-
trast the other four species are found on
macrophytes. Three species occur in the
ribbonweed beds (Vallisnera) in the fast
flow of the upper Murray from Khancoban
to Yarrawonga. They construct a case of
Vallisnera strips attached to the plant leaf
blades. One species is restricted to the
pondweed Potamogeton in the slow flow-
ing areas. The larvae construct a case by
cutting a section of leaf and folding it back
onto the leaf. This species is common in
the Potamogeion beds of lower Murray
River from Echuca to Woods Point.

'The Victorian Naturalist

Fig. 15. Pyralid larva in its case (Nymphulinae).

Conclusion

This paper has attempted to give an
overview of the aquatic macroinvertebrate
fauna of the Murray River without
attempting to give an assessment of the
condition of the river. Norris et al. (2001)
recently concluded that the macroinverte-
brate communities along the Murray River
from Dartmouth Dam to Lake Alexandrina
were in poor condition. They also high-
lighted the absence of pristine or minimal-
ly modified sites in lowland rivers in the
Murray Darling Basin and recommended
caution in the interpretation of the results.
The current distribution of the macroinver-
tebrates has certainly been modified hy the
influence of regulation, particularly in the
Lower Murray tract where the river now is
a series of narrow lakes and some species
have benefited at the expense of others
(e.g. the yabby replacing the Murray cray-
fish), and introduced species have success-
fully invaded the river (e.g. Physa acuta).
However, it is not possible to determine
the extent of modification of the inverte-
brate fauna because there are only limited
data prior to construction of the locks and
weirs. In addition, the other lowland rivers
in the Murray Darling Basin with which
the Murray communities could be com-
pared are also affected by regulation. This
lack of a reference condition limits the
ability to assess the condition of the
Murray but we are now in the position to
monitor the effects of management deci-
sions (e.g. environmental flows) on the
aquatic macroinvertebrate communities of
the Murray River.

Vol. 119 (4) 2002

Part Two

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Murray. Australian Water Research Advisory Council
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Madden CP, Austin AD and Suter PJ (1995) Pollution
monitoring using chironomid larvae: What is a deformi-
ty? In Chironomids: From genes to ecosystems, pp 89
94. Ed P Cranston, (CSIRO Publications: East
Melbourne)

Madden CP, Suter PJ Nicholson BC and Austin AD
(1992) Deformities in chironomid larvae as indicators
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Maher MT and Carpenter SM (1984) Benthic studies of
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Murray Rivers on the Ephemeroptera, Plecoptera and
Irichoptera. (Unpublished Honours thesis, La Trobe
University, Wodonga)

Neboiss A (1991) Trichoptera. In The inseets of Australia,
pp 787-816. (Melbourne University Press: Melbourne)

New TR (1991) Neuroptera, In The Insects of Australia,
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Prosser | and Young B (2001) Snapshot of the Murray-
Darling Basin River Condition, (Murray Darling Basin
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dam operation on mayfTy assemblage structure and lile his-
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Pettiprove V (1989) Larval mouthpart deformities in
Procladius paludicola Skuse (Diptera: Chironomidae)
from the Murray and Darling Rivers, Australia
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St Clair R, Doep | and Winsor L (1999) A survey of
Spatula tryssa Wall and other freshwater flatworms in
the Victorian Alps with an evaluation of the conserva-
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Sheldon F and Walker KE (1993) Pipelines as a refuge for
freshwater snails, Regulated Rivers Research &
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Smith, BJ and Kershaw RC (1979) Field Guide to the
Non-marine Molluses of South-castern Australia
(Australian University Press: Canberra)

Stevens MM (1995) Biology and control of. Chironomus
tepperi Skuse, a pest of rice in New South Wales, In
Chironomids: Prom genes to ecosystems, pp 235-239,
Ed P Cranston, (CSIRO Publishers: East Melbourne)

Suter PJ and Bishop JE (1990) Stoneflies (Plecoptera) ol
South Australia, In Mayv/lies and Stoneflies: Life
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(Kluwer Academie Publishers: Dordrecht)

Suter PJ, Goonan PM, Beer JA and Thompson TH (1993)
| biological and physico-chemical monitoring study of
wetlands from the River Murray flood plain in South

Australia. (Australian Centre for Water Quality
Research: Adelaide)

Suter PJ, Goonan PM, Beer JA and Thompson TB (1995)
The response of chironomid populations to flooding
and drying in flood plain wetlands of the lower River
Murray in South Australia, In Chironomids: From
genes to ecosystems, pp 185-195. Ed P Cranston,
(CSIRO Publishers: Fast Melbourne)

Thoms M, Suter P, Roberts J, Koehn J, Jones G, Hillman
T and Close A (2000) Report of the River Murray
Scientific Panel on Environmental Flows, River Murray

Dartmouth to Wellington and the Lower Darling
River. Murray-Darling Basin Commission, Canberra,

Viets KO (1980) New Unionicolidae (Acari,
Hydrochnellae) from Australia. Transactions of the
Royal Society of South Australia 104, 27-40.

Walker KF (1990) Mussels, In The Murray, pp 309-307.
Eds N Mackay and D Eastburn. (Murray Darling Basin
Commission: Canberra)

Warwick WF (1990) The use of morphological deformi»
fies in chironomid larvae For biological effects monitor-

Institute, Saskatoon,

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invertebrates of Australian inland waters. (Macmillan
Company of Australia Pty Ltd: South Melbourne)

The Swamp Yabby (Cherax sp.) of the
Murray River Catchment

1

Geoffrey N Edney', Dale Gi McNeil and Susan H Lawler

Abstraet

The Swamp Yabby has been recently collected from Barmah State Forest, the Ovens River, Victoria
and Deniliquin, NSW. Here we present the species in general terms and provide figures for identili-
cation, The Swamp Yabby differs from other Cherax species both in morphology and ecology. This
animal resides primarily in burrows rather than the river. Burrowing Cherax were previously

unknown in Victoria. (The Fierarian Naturalist 119 (4), 2002, 200-204)

Introduction

lhe Swamp Yabby has been known to
locals, fishermen, rangers and scientists for
some time but remains undeseribed,
despite specific promotional efforts by the
Victorian Department of Natural
Resources and Environment who even
releused a range of material identifying the
Swamp Yabby as a conservation priority,
Ihe species is called Cherax sp. C in the
preliminary key by Horwitz (1995), A full
description cannot be achieved however,
primarily due to the lack of specimens col-
lected, and importantly, the absence of a
female specimen, This prevents any taxo-

' Edney Ecological Services, 61 Wilson Street,
Wodonga, Vietoria 3690

` Department of Environmental Management and
Leology, La Trobe University, PO Box 821, Wodonga,
Victoria 3689, and The Cooperative Research Centre
for Freshwater Ecology, Albury, New South Wales
Email s.Tawlergboaw.latrobe.edu.au

200

nomic description, leaving the species
without conservation status.

We have collected Swamp Yabbies
(Cherax sp.) trom three locations, the
Ovens River floodplain in Victoria and the
Murray and Edward/Gulpa floodplains in
New South Wales. Specimens were col-
lected from the Ovens near Wangaratta
and the Edward/Gulpa near Deniliquin
through excavation of burrows. Depending
on the water table, crayfish were located in
burrows up to one metre in depth, whilst
some large burrows were excavated to a
depth of over a metre without locating a
specimen. During a very minor flood on
the Murray floodplain at Barmah, Swamp
Yabbies were collected using submerged,
baited hoop nets,

Other investigators have reported collect-
ing Swamp Yabbies from two more

The Victorian Naturalist

Fig. 1. The collecting site in the Barmah State
Forest. a) The small stream in which the Cherax
sp. were captured. b) The burrow entrance of
Cherax sp. Note the cattle hoofprints trampling
the burrow entranee. Matchbox indicates size.

Victorian tributaries of the Murray; at Lake

Nagambie, on the Goulburn River, south of

Shepparton (Royal 2002) and Lake
Nillahcootie, near Mansfield on the Broken
River (Brian Woodbridge pers. comm.).
This suggests that their range covers not
only the floodplains of the Murray and its
associated anabranches, but also the three

most significant Murray tributaries of

north-eastern Victoria.

Description of collections

Eleven specimens were examined (Table
1). Three were caught in open water in the
Barmah State Forest, Victoria, and seven
were dug from burrows near the Ovens
River, Victoria, and the Murray River at
Deniliquin, New South Wales. An addi-
tional juvenile specimen was kindly pro-
vided by Brian Woodbridge of the Murray
River Aquarium in Echuca, Victoria, who
successfully bred them in captivity.

The Barmah State Forest site was a low
dry floodplain with a small, slow-flowing
creek at its centre. Cattle were grazing in
the area, These crayfish were found in

Vol. 119 (4) 2002

Part Two

Fig. 2. The collecting site near the Ovens River.
At this site Cherax sp. was collected in a Red
Gum floodplain forest.

water that was 10-20 cm deep and they
were caught late at night. Nearby burrows
were clearly visible (Fig. 1).

The burrows at the Ovens River were on
elevated land between billabongs. This land
would be underwater only during flood
events and is usually about two to three
metres above the water level. It is currently
used for cattle grazing. The chimneys
(entrances to Swamp Yabby burrows) at the
Ovens River were 30-40 m from the closest
billabong (Fig. 2).

Identification

The Swamp Yabby has a unique mor-
phology and is easy to recognise.
Anecdotal evidence suggests a community
awareness of a ‘large black yabby with
huge claws'. An unknown ranger at
Barmah State Forest called the animals
‘Moonclaws’, in response to its large, cres-
cent shaped claws.

The species is easily recognised by its
claws, which are very broad and have a
paddle-like protrusion (Fig. 3). It is also
often larger than Cherax destructor col-
lected from the same site. Three Swamp
Yabbies from Barmah showed occipital
carapace lengths (OCL) of 70, 70 and 75
mm, while we caught over fifty C. destruc-
tor, the largest of which was 60 mm OCL.
We expect that larger specimens of both
species occur.

Sympatric crayfish species

The Swamp Yabby does not occur in iso-
lation. It coexists with the Common Yabby
C. destructor throughout its entire range,
and with the Murray River Crayfish
Euastacus armatus throughout most of its

201

Murray River Special Issue

Table 1, Swamp Yabbies examined by the authors. Sizes are given in mm OCL (occipital carapace
length: from the eye to where the carapace joins the tail). ' at 2 years of age

Site Sex Number Size range Collected by
collected (*raised by)
Murray River at Barmah State Forest, Victoria male 3 70-75 GNE
Ovens River, Wangaratta, Victoria male 2 50-60 GNE & DGM
Edward/Gulpa Junction, Deniliquin, NSW male 5 50-70 DGM
Murray River Aquarium, Echuca, Victoria male | 34! BW*

range. It is sympatric with the burrowing
species Engaeus lyelli and Engaeus cymus
(Horwitz 1990), and with the genus
Geocharax (Tyler et al. 1983),

Behaviour, inter- and intra-specific
competition

From our observations, it seems that the
Swamp Yabby is more aggressive than C.
destructor and is likely to outcompete it in
the wild. When divided by a barrier within
the same aquarium, the Swamp Yabbies
made extreme exertions to breech the bar-
rier. When no barrier was provided, C
destructor were immediately attacked and
swifily killed unless they left the water via
an emergent stick or rock.

Swamp yabbies also exhibit strong
intraspecific aggression. Four Swamp
Yabbies were placed in aquaria (160 x 40
cm), and isolated in four small burrows
separated by dry earth barriers. After the
first night the smallest individual had been
cut in half and only one individual survived
the first week. This individual subsequently
survived in the aquarium for over a year.
Swamp yabbies appear to have quite a soft
carapace compared with C, destructor.
Although this may be an adaptation to bur-
rowing, it is surely a disadvantage during
intraspecific competition. There was no
evidence of cannibalism after death from
these competitive bouts.

The degree of habitat partitioning
between Swamp Yabbies and C. destructor
in the wild is uncertain. Cherax destructor
were never found in Swamp Yabby bur-
rows although they did exist in shallow
simple burrows in the same dry billabong
as the Swamp Yabby. Only single speci-
mens of Swamp Yabbies were found with-
in each burrow, suggesting a non-colonial
burrowing system.

Crayfish burrows
Australian freshwater crayfish burrows

202

(a)

PN

= c^

(b)

"ig. 3. a) The Swamp Yabby (Cherax sp.). b)
To identify a Swamp Yabby, the width of the
claw (B) is more than 50% of the length (A),
and a paddle (C) is present on the claw.

have been classified by Horwitz and
Richardson (1986). Three types are recog-
nised: burrows opening below the water
surface in a permanent water body, bur-
rows connected to the water table but not to
a stream or lake and burrows independent
of the water table. Ellen Clark anticipated
this scheme with her terms aquatic, semi-
aquatic and terrestrial (Clark 1936).

The genus Cherax is known to construct
burrows which are always connected either
to a permanent water body or to the water
table (Horwitz and Richardson 1986).
There are three burrowing Cherax known
from Queensland: C. punctatus, C. robus-
tus and C. cartalacoolah (Short 1993). All
of them are smaller than the Swamp Yabby
and burrow in sandy substrates. The bur-
rows of Cherax plebejus, from southwest

The Victorian Naturalist

Western Australia, descend vertically to a
small chamber where one often finds a
male and a female (Horwitz and Knott
1983),

Burrowing by the Swamp Yabby

The Swamp Yabby constructs a burrow
that is always connected to a permanent
watercourse or to the water table (Type 2
as described by Horwitz and Richardson
1986). Some entrances were freshly sealed
with mud indicating the presence of water
within the burrow system. The Swamp
Yabby appears to remain in its burrow for
much of the time. The burrow covers more
than one cubic metre and could be much
larger underground. A surface chimney
leads to a tunnel which descends for a
short distance and then will branch or join
a chamber.

The burrow has several chambers which
are about the size of a football. Some are
used for storing water. There are often
multiple tunnels connecting chambers and
several surface holes. Burrows of the
Swamp Yabby can be distinguished from
those of C. destructor and Engaeus by the
sheer size of the entrance hole: usually 5-8
cm in diameter vs 1-2 cm for Engaeus and
generally <4 em for C. destructor.

To date we have only collected males,
and other workers have confirmed an
extremely skewed sex ratio (Tarmo Raadik
and Brian Woodbridge pers. comm.). The
reasons for this are unknown. It is possible
that there is some sexual habitat segrega-
tion, however current collecting techniques
may somehow be biased towards finding
males. Females may live in extremely deep
burrows and remain non-emergent during
small floods.

Associated fauna

Burrowing crayfish are known to support
a large and often unique fauna termed the
‘pholeteros’. Their burrows are refuges for
other animals during drought. We have
observed an Engaeus lyelli and a large
number of the frogs Limnodynastes tas-
maniensis in burrows during our excava-
tions. The burrows of other crayfish often
support faunal assemblages (Horwitz and
Knott 1981; Suter and Richardson 1977).
In addition, the crayfish themselves sup-
port a diverse ectofauna, which are animals
that live on their shells, tails and gills.

Vol. 119 (4) 2002

Part Two

^A oF T
Fig. 4. A Swamp Yabby surrounded by
ectosymbiotic worms Temnosewellia minor.
Photo by Karli Hawking.

Temnocephalans (Fig. 4) are ecto-symbi-
otic worms that lay eggs under the tailfins
and live on the carapace of crayfish
(Cannon 1991; Cannon and Sewell 1994,
1995: Sewell and Cannon 1995, 1998),
They filter feed and are attached by a large
caudal sucker. Large numbers of temno-
cephalans were observed on all Swamp
Yabbies collected. Some of these were sent
to Lester Cannon of the Queensland
Museum, who identified them as
Temnosewellia minor, a species known to
be common on Cherax in the Murray River
catchment. Temnocephalans are turbellari-
an flatworms known to be symbiotic on
freshwater crayfish throughout Australia
(Cannon 1991), The worms are nearly as
variable as their hosts, and may be species
specific in many cases (Cannon and Sewell
1994).

Impacts and conservation

All of our sites were grazed by cattle.
The soil showed extensive disturbance
from trampling and grazing. There is anec-
dotal evidence that cattle impact heavily
on Swamp Yabbies and many Swamp
Yabby burrows were found within hoof-
prints and with flattened chimneys.
Crayfish chimneys were scarce in the open
and more common around fallen logs and
roots. These habitats provide protection
from soil disturbance such as during floods
or trampling, and may provide a food
source, attracting invertebrates during
decomposition.

In 1936 farmers were complaining about
losing cattle and horses in *crab-hole coun-
try’ near Benalla, Victoria (Clark 1936),
and anecdotal evidence suggests that these
complexes were always ploughed in for
safety. Although the species responsible

203

Murray River Special Issue

has never been identified, similar collapses
have been reported due to extensive bur-
rowing by Engaeus (Clark 1936). The
Benalla area, however, is now known to be
central to the Swamp Yabby distribution
and it is possible that these ‘crab’ hole
areas may have been the result of large
aggregations of Swamp Yabby burrows.
Furthermore, the Murray, Edward/Gulpa,
Broken, and Goulbourn Rivers are all
highly regulated. High winter and spring
flows, that would have traditionally Mood-
ed Swamp Yabby habitats no longer occur.
Ihe effect of these impacts on Swamp
Yabbies and their habitat warrants further
investigation. The Ovens River, which
remains mostly unregulated, provides a
good opportunity to observe Swamp
Yabby habitat in à more natural flooding
regime. The destruction of such large bur-

row complexes, and the deterioration of

natural flow conditions suggest that past
land practices may have had a large impact
on Swamp Yabby habitat. *Crab-hole
country’ is virtually gone now from the
Benalla area. It is now difficult to ascertain
how these land practices may have impact-
ed upon Swamp Yabby populations in the
past and how these may have affected its
distribution and ecology,

Disturbance has been shown to affect
crayfish populations elsewhere in Australia
(Growns 1995), and it is possible that
many former populations have disappeared
due to human disturbance. It should also
be noted that the large complex of chim-
neys which first drew our attention to the
presence of the Swamp Yabby on the
Ovens River was graded to make a
drainage channel during roadmaking activ-
ities in 1998. Although some burrows still
persist there, it is clear that current land
use also has an impact upon the Swamp
Yabby and its habitat.

Acknowledgements

We'd like to thank Brian Woodbridge of Murray
River Aquarium, Echuca, for the juvenile
Swamp Yabby and his observations, l'armo
Raadik of the Department of Natural Resources
and Environment for his observations, Lester
Cannon for identifying the temnocephalan
worms, and Karli Hawking for photographing
them. Thanks to De Adam Campbell for assist-
ing with specimen collection and extremely
brave assistance during burrow excavation Also

204

thanks to an anonymous reviewer for their com-
ments. Extra big thanks to the Campbell and
Williamson families for their hospitality, access
to their properties, and the use of their shovels.
These crayfish were collected under research
permits from the Victorian Department of
Natural Resources (RP:-562) and New South
Wales Fisheries (F96/187HP).

References

Cannon LRG (1991) Temnocephalan symbionts of the
freshwater cray fish Cherax quadricarinatus from
northern Australia, Hydrobiologia 227, 341-347.

Cannon LRG and Sewell KB (1994) Symbionts and
biodiversity. Memoirs of the Queensland Museum 36,
33-40.

Cannon LRG and Sewell KB (1995) Craspedellinae
haer, 193] (Platyhelminthes; Temnocephalida)
ectosymbionts from the branchial chamber of
Australian crayfish (Crustacea: Parastacidae).
Memoirs of the Queensland Museum 38, 397-418.

Clark FE (1936) The freshwater and land erayfishes of
Australia. Memoirs of the National Museum of
Victoria 10, 5-58.

Growns LO (1995) dsracopsis gouldi Clark in streams
of the Gow Range, Northern Tasmania: the effects of
catchment disturbance. Papers and Proceedings of
the Roval Society of Tasmania 129, 1-6.

Horwitz P (1990) A taxonomic revision of species in
the freshwater crayfish genus Engueus Erichson
(Decapoda: Parastacidae), Jnyertebrate Taxonomy 4,
327-614.

Horwitz P (1995) 4 preliminary key to the spectes of
Decapoda (Crustacea, Malacostraca) found in
Australian inland waters. Identification Guide No 5
(Co-operative Research Centre for Freshwater
Ecology: Albury)

Horwitz P and Knott B (1981) The faunal assemblage
in freshwater crayfish burrows in sedgeland and for-
est at Lightning Plains, Western Tasmania. Papers
und Proceedings of the Royal Society of Tasmania
125, 29-32.

Horwitz P and Knott B (1983) The burrowing habit of
the koonac Cherax plebejus (Decapoda:
Parastacidae). Western Australian Naturalist 15, 113-
117

Horwitz PHJ and Richardson AMM (1986) An ecolog-
ical classification of the burrows of Australian
Freshwater Crayfish, Australian Journal of Marine
and Freshwater Research 37, 237-242.

Royal D (2002) www.eray fishworld.com

Sewell KB and Cannon LRG (1995) A scanning elec-
tron microscope study of Craspadella sp. trom the
branchial chamber of redelaw crayfish, Cherax
quadricarinatus, from Queensland, Australia.
Hydrobiologia 305, 151-158.

Sewell KB and Cannon LRG (1998) New temnoceplia-
lans from the branchial chamber oF Australian
Euastacus and Cherax hosts, Proceedings of the
Linnean Society of New South Wales 119, 21-36.

Short J (1993) Cherax cartalacoolah, a new species of
freshwater crayfish (Decapoda, Parastacidae) trom
Northeast Australia. Memoirs of the Queensland
Museum 33, 55-59,

Suter PJ and Richardson AMM (1977) The biology of
two species of Angaeus (Decapoda: Parastacidae) in
Tasmania H1. Habitat, food, associated fauna and dis-
tribution. Australian Journal af Marine and
Freshwater Research 28, 95-103.

Tyler MI, Twidale CR, Ling IK and Holmes JW
(1983) Natural History of the Southeast. (Royal
Society of South Australia Inc)

The Victorian Naturalist

Part Two

Murray River Microfauna
Russell J Shiel'

Abstract

The microfauna of the Murray and tributaries is a speciose, albeit neglected, component of the river-
ine biota. The community derives from diverse in-stream ,and, seasonally, off-stream sources. There

are regional differences in species composition -

tributaries may contribute different taxa to the

mainstream river. A mixed assemblage of protists, rotifers and microcrustaceans persists into the
lower Murray, with longitudinal changes in species composition during long travel times to the river
mouth. Effects of river regulation, land use and salinisation on microfaunal biodiversity are sum-
marised. (The Victorian Naturalist 119 (4), 2002, 205-211)

Introduction

In marked contrast to the recent surge of

research activity on macroinvertebrate tax-
onomy and ecology in the Murray-Darling
Basin, spurred by river health initiatives
and pressure for biodiversity information,
the microfauna of Murray-Darling waters
remains largely neglected. Microfauna reg-
ulate bacterial and algal abundance, and
occupy important links in food chains for
higher order consumers such as macroin-
vertebrates, fish and birds. In most
Australian freshwater ecosystems, but par-
ticularly in the Murray-Darling system,
these connections have been under-esti-
mated or ignored (Green and Shiel 1992),

What little information exists on the
microfaunal component of the Murray
River and tributaries derives largely from a
few local studies. Species composition and
successional events in littoral microcrus-
tacea from a year-long study of a Goulburn
River billabong were documented by Shiel
(1976). The zooplankton of Lake Hume
was reported by Walker and Hillman
(1977), that of several other impoundments
by Powling (1980). A basin-wide survey,
published in part by Shiel er al. (1982),
demonstrated the disparate nature of the
microbiota of the west-flowing Murray
River and the south-flowing Darling River.
The former contained a cool-temperate
microcrustacean-dominated lacustrine
(limnoplankton) assemblage, attributed to
the serially-impounded nature of the
Murray River, the latter a true riverine
(potamoplankton) community dominated
by rotifers, many of which are warm-
stenotherms or tropical in affinity.

The role of turbidity in structuring zoo-

Department of Environmental Biology, The
University of Adelaide, South Australia 5005

Vol. 119 (4) 2002

plankton communities in Lake
Alexandrina, at the Murray mouth in South
Australia, was examined by Geddes
(1984). Boon ef al, (1990) reviewed infor-
mation on upper Murray billabong ecolo-
gy, including microbiota, for the Murray-
Darling Basin Commission volume *7/e
Murray’; information on riverine zoo-
plankton was reviewed for the same vol-
ume (Shiel 1990). Other relevant and more
recent studies include: invertebrate emer-
gence from flooded sediments (Boulton
and Lloyd 1992), rapid responses to flood
events by billabong rotifer assemblages
(Tan and Shiel 1993), Lakes Hume and
Dartmouth zooplankton in the context of
biomanipulation (Matveev and Matveev
1997), heterogeneity of habitat and micro-
faunal biodiversity across ephemeral wet-
lands (Shiel er al. 1998), fish predation on
zooplankton (Nielsen ef al. 2000a, b) and
the role of propagules in sediments
(Langley et al. 2001; Nielsen er al. 2002;
Shiel et al. 2001; Skinner er al. 2001).

Composition of Murray River microfauna

Three broad groups of microinvertebrates
comprise the bulk of pelagic and littoral
communities in Murray-Darling waters,
viz. protists (Protista), rotifers (Rotifera)
and a suite of microcrustacea dominated
by copepods (Crustacea: Copepoda),
cladocerans (Crustacea? Branchiopoda:
Anomopoda/Ctenopoda) and (occasional-
ly) ostracods (Crustacea: Ostracoda). Other
groups occurring in microfaunal samples
include larvae or smail adults of various
insects, particularly Diptera and
Hemiptera, and water mites (most often
Hydracarina or Oribatida). These latter
groups may be abundant seasonally and be
important in structuring microfaunal

205

Murray River Special Issue

assemblages through food web interac-
tions, however, for the purposes of this
review, only the major groups are treated
further.

Protista: Protists (or protozoa, unicellular
heterotrophs) include flagellates, ciliates
and amoebae, both naked and testate. They
are commonly the numerically dominant
animals in any sample taken from Murray
River standing (lentic) or slow-flowing
(lotic) waters. Protists are under-represent-
ed in rapidly-flowing lotic waters; they are
relatively fragile, and do not survive.

Protists, remarkably, have received only
cursory mention in Murray River ecologi-
cal studies, if they are mentioned at all.
'The lack of study of Australian ciliates, for
example, was regarded as ‘regrettable and
somewhat astonishing’ when an endemic
Murray River loricate ciliate was described
(Foissner and O' Donoghue 1990). The age
of the continent and the potential for isola-
tion to drive speciation events were seen as
significant in predicting a diverse indige-
nous protist community.

^ suggestion that Australian protist
assemblages may differ from those of the
northern hemisphere was made by
Laybourn-Parry et al, (1997), who collect-
ed S/entor, a mixotrophic (photosynthetic)
ciliate, from two Murray-Darling reser-
voirs (Tuggeranong, ACT and Hume,
NSW). Stentor blooms colouring the water
black have long been reported from
Murray River waters (e.g. as 'cf.
Climacostomum in Walker and Hillman
1977), where densities >50,000 I^ have
been recorded (cited in Laybourn-Parry ef
al. 1997). Stentor is uncommon in northern
hemisphere waters, where it is subjected to
heavy predation by, inter alia, cyclopoid
copepods. The latter are rare in Murray
River reservoirs, which tend to be domi-
nated by herbivorous calanoid copepods
(see below), hence predation pressure on
Stentor appears to be low. Stentor also
reaches bloom populations seasonally in
some Murray River billabongs (cf. Fig. 1,
Shiel 1990).

Further evidence for distinctive protist

assemblages comes from a recent study of

testate amoebae in the upper Murray catch-
ment (Meisterfeld and Tan 1998). They
reported a rich testate community in the

206

à —— E d

Fig. 1. S/entor cf. amethystinus in Ryan's #1
billabong, Murray River floodplain near
Bonegilla, Victoria. Photo by Russell Shiel.

environs of Lake Catani, Mt Buffalo, with
89 taxa, 34 of them new to Australia, record-
ed from eight Sphagnum/sediment samples.
Six of the testates were noted as ‘strictly
Gondwanan' in distribution. Several others
were undescribed and possibly Australian
endemics. New records of naked amoebae
(Heliozoa) also were reported from ponds in
the Mt Buffalo region (Mikrjukov and
Croome 1998), Given that almost half of the
amoebae reported in these two small surveys
were new to the Australian fauna, it would
seem that a diverse and unrecorded protist
fauna awaits discovery!

The only group of protists which has
been studied intensively in Murray River
waters is the freshwater Acanthamoeba
Naegleria, particularly in the lower
Murray in South Australia, from which
Adelaide and other regional centres draw
drinking water, As the causative organism
of amoebic meningitis, Naegleria, and
other pathogenic protists such as Giardia,
are continuously monitored by the
Australian Water Quality Centre at
Bolivar, South Australia.

Given the paucity of studies on protists in
Murray-Darling waters, little can be said of
their potential for bioindication or their
biodiversity in any of the Basin's waters,
which is remarkable given the level of
research effort applied to protists else-
where (see, for example, Foissner and
Berger 1996).

The Victorian Naturalist

a s - a aa =
Fig. 2. Collecting sediment cores from Ryan's
33, Bonegilla. Photo by Russell Shiel.

Rotifera: Rotifers, the smallest metazoans,
with most «200 um in size, are commonly
the most abundant microfauna after pro-
tists. More than 600 of the 720 rotifer
species now known from Australia have
been recorded from Murray-Darling waters
(Shiel unpubl. data). The greatest rotifer
biodiversity is found in ephemeral waters,
where 7100 co-occurring species have
been collected in single net tows (Shiel er
al. 1998). Billabongs also support rich
rotifer assemblages by virtue of the vege-
tated, partitioned habitat; 2350 microfau-
nal species, including 7200 rotifer spp.,
have been recorded from an ephemeral
pool over more than 20 years of sampling
(Langley et al. 2001; Shiel er al. 2001).
More commonly, a collection will contain
5-50 rotifer spp., depending on structural
complexity of the habitat — densely vege-
tated and therefore partitioned billabongs
have more species than does the open
water of reservoirs and rivers. Floating and
submerged vegetation provides some pro-
tection from visual predators. Rotifers
occupy all feeding niches in such parti-
tioned habitats — detritivores, bacteri-
ovores, herbivores, carnivores and para-
sites, Suites of each of these feeding
groups may co-occur. For example, several
co-occurring species of Brachionus,
Filinia, Keratella or Trichocerca may
overcome the problem of competition for
resources by taking different size food
items. Experimental evidence suggests that
even bacteria may be partitioned in a simi-
lar manner, by size, morphology or chem-
istry (Boon and Shiel 1990). Rotifers are
able to persist in ephemeral habitats by
production of resistant resting eggs, which

Vol. 119 (4) 2002

Part Two

4500,
F = 66.09 (p < 0.001) 12
"
5 1000}
[7]
D
p 14
$
500
1:25 12
0
pH6.28 pH632 pH675 pH641

Fig. 3. Resting egg counts in sediments from
three flood frequencies: annual, biennial and
every c. 25 years (from Shiel eż al. 2001).

remain in dry sediment until rewetting and
appropriate cues to hatching. All resting
eggs do not respond to the same cues,
hence there is a succession of hatching
events, with concomitant compositional
changes the longer a previously dry habitat
is flooded (Tan and Shiel 1993). Fig. 2
shows collection of dry sediment from the
previously-wetted margin of Ryan’s #3, a
shallow ephemeral pool on the floodplain
of the Murray River adjacent to Ryan's #1,
a permanent billabong. Resting stages
counted from 1 mm slices of 1 em diame-
ter cores taken from these sites showed
significant differences in density (Fig. 3),
up to 1200 cm’, with highest densities at
the margin of Ryan’s #3, the biennially
flooded (ephemeral) site (Shiel er al.
2001).

Microcrustacea: Of the three main groups
of microcrustaceans abundant in Murray
River waters, copepods tend to predomi-
nate, with cladocerans either perennial in
low numbers or markedly seasonal, when
they may reach high densities. Ostracods,
except in salinized waters, tend to be rare
incursions from the littoral, their preferred
habitat. Most abundant copepods in reser-
voirs and billabongs, and also often in weir
pools and downstream reaches of the
Murray and tributaries, are calanoids of the
southern hemisphere family Centro-
pagidae, commonly Boeckella and
Calamoecia species, with Hemiboeckella
species in ephemeral waters, and (rarely)
the usually-coastal Gladioferens in down-
stream sites such as Lake Cullulleraine.
Two species of the northern affinity
Diaptomidae occur across northern

207

Murray River Special Issue

Australia, and may extend into the north of

the Murray-Darling, Basin (ef. Bayly
1966),

Of about 50 centropapid species known
from Australia, the most common in
Murray River waters, and across southern
Australia generally, is Boeckella triarticu
lata, which occurs across a wide range ol
habitat types (Bayly 1992), In fish-free
ephemeral habitats the large (4 mm) 8
major may be a seasonal predator on other
plankters (Green ef ul. 1999), with B
pseudochelae and. Hemiboeckella searli
also common in ephemeral waters,
Billabong species include Bo /Tuvialis, B
minuta and Bo symmetrica, which may co
occur with one or more Calamoecia
species, commonly C, ampulla or C
lucasi, Two or three co-occurring
calanoids in a habitat is usual, four or five
less so, and six is rare, Notably, salinized
waters of mid-Murray regions around
Keranp/Swan Hill have halophile
ealanoids, Calamoecia salina or C, elitel
lata, suggesting a much longer evolution
ary history of salinization than that
induced by human activities in the region
in the last 200 years,

Cyclopoid copepods are less well known
in Murray River waters, They are not com-

mon or abundant in the open water of

reservoirs or rivers, but ean be seasonally
speeiose and abundant in billabongs and
ephemeral waters, They fill both herbivo-
rous and carnivorous niches, prey includ-
ing rotifers, other copepods, eladocerans
and occasionally, small macroinverte-
brates, ep, mussel glochidia (juveniles).
Most species are from the Family
Cyelopidae, and about 100 spp, are known
from the continent, Common genera are
lustralocvelops (a large predator), Zuev-
clops, Mesocyelops, and Microevelops
he latter two genera are the most speciose
based on present taxonomie information
(ep, Holynska 2000), As for calanoids,
two or more species co-occurring at a site
is common, often markedly different in
sive, thereby reducing competition,

A third group of copepods, the harpacti-
cords, are benthie in habit, rarely collected
in open water, and poorly studied in
Australasia, Contliocamptus species appear
to be the most common in Murray River
billabonys and ephemeral waters, associat

208

ed with sediments and vegetation (cf.
Hamond 1987),

Plankton collections are usually dominat-
ed by juveniles of the resident copepods,
both nauplii and copepodite stages, and
this presents a major taxonomie impedi-
ment to biodiversity studies, Not only does
a researcher have to determine the species
present, but also may need to discriminate
the life stages, This is not an easy task
when three or four species co«occur, and
samples may include several copepodite
und sub-adult instars, with one sex or the
other required for species determination.
Juveniles also tend to predominate in
downstream river reaches; adults apparent-
ly are able lo avoid outflows from weir
pools or reservoirs,

Cladocerans (water Neas’) may make up
a significant part of microfiunal communi-
ty diversity in shallow vegetated habitats,
but less so in open water of reservoirs and
rivers, where their size renders them sus-
ceptible to predation by macroinverte-
brates and fish, More than half of the e,
200 spp. of eladocerans now known from
the continent occur in Murray-Darling
waters, The family Chydoridae is the most
diverse, and in excess of 100 species have
been recorded from Australia, with more
than half of them endemie (Shiel and
Diekson 1995), More than 20 species of
chydorids co-occurred in a Goulburn bill-
abong (Shiel 1976) where they filled detri-
livore and herbivore niches; Many chy-
dorids are adapted to scraping biofilms
from surfaces, hence are littoral or epiphyt-
ie in habit, Only species of Chvedorus tend
to be collected in open water, and then
associated with filamentous algal blooms
Where they collect food by seraping along
filaments,

Only a few other eladoceran families
have truly planktonic representatives in
Murray River waters: Bosminag (Bosmin-
idac) is common in reservoir and river
plankton, as is Diaphanosoma (Sididae),
and seasonally, Moina (Moinidae),
Daphnia (Daphnidae), one of the larger
cladocerans, tends to be seasonal in bill-
abongs and ephemeral habitats, or perenni-
al in low numbers in reservoirs, where it is
probably subjected to heavy predation
pressure, Smaller daphnids, such as
Ceriodaphnia, are more often collected in

The Victorian Naturalist

plankton tows. Other daphnid genera —
Simocephalus, Scapholeberis — are bill-
abong/shallow water dwellers. One daph-
nid genus, Daphniopsis, is a halophile,
occurring in salinized waters throughout

the Basin. Species of other families of

cladocerans — Macrothricidae, Ilyocryp-
tidae and larger sidids (e.g. Latonopsis) are
usually found in shallow, vegetated habi-
tats, particularly regularly inundated
ephemeral pools.

Notably absent from Australia, possibly
evolving after the breakup of Gondwana,
are the northern hemisphere predatory
cladoceran families Cercopagidae,
Leptodoridae and Polyphemidae, which
are important “structurers’ of plankton
communities in northern hemisphere
waters (Shiel and Dickson 1995; cf. Rivier
1998). Species of Podonidae are reported
from coastal/marine waters around
Australia but not from inland Australia
(Smirnov and Timms 1983).

Ostracods, which include detritivores,
herbivores and predators, occasionally are
abundant in collections from billabongs
and ephemeral pools. In some billabongs,
five or six species of ostracods is not
unusual — large species (e.g. Australo-
cypris, Mytilocypris) may co-occur with
smaller (e.g. Cypretta, Limnocythere,
Newnhamia). Ostracods rarely appear in
open water collections.

Sources of microfauna in the mainstem
rivers

In Murray River waters there is no exper-
imental evidence to demonstrate the contri-
bution of headwater reservoir limnoplank-
ton to downstream rivers, nor of contribu-
tions from floodplain lentic waters. From
circumstantial evidence in the studies cited
earlier, and from studies of river microfau-
na elsewhere (e.g. Baranyi et al. 2002), the
persistent microfaunal community in
downstream reaches reflects disparate con-
tributions from upstream impoundments,
floodplain waters which may at times have
a connection to the river, regions of slow-
flow such as backwaters or braided chan-
nels, waste stabilization ponds from river-
side communities which may discharge
into Murray tributaries, in fact any stand-
ing water which connects to the river at
any time.

Vol. 119 (4) 2002

Part Two

Geographical differences in source
waters, regionalism in the microfauna and
regional rainfall events will influence what
particular assemblage is being inoculated
into tributaries. Downstream weirs and
locks on the Murray River provide low- or
no-flow conditions and a longer retention
time, which permits reconstitution of a
microcrustacean assemblage. This is
referred to in European studies as ‘age’ of
the water, with rotifers dominating in
waters of low ‘age’ (e.g. short retention-
time storages such as Lake Mulwala) and
microcrustacean assemblages appearing in
waters of greater ‘age’ (e.g. long retention-
time storages such as Lakes Dartmouth
and Hume) (cf. Baranyi er al. 2002).

What this phenomenon reflects is the life
cycle of the respective microfauna. At
ambient temperatures in Murray River trib-
utaries, rotifers are reproducing in days,
microcrustacea in weeks. Rotilers are able
to get through their life cycles in the short
retention-time storages, microcrustacea are
not. The latter require stable conditions for
a longer period to reach adult reproductive
stages and are unable to complete life
cycles in turbulent or rapid through-flow
storages. This life cycle disparity is the
basis for the differences noted earlier
(Shiel er al, 1982) between Murray and
Darling microfauna, The microfauna of the
Murray traverse a series of lentic or at least
slow-flowing weir pools; those of the
Darling below the headwater rivers a lotic
system, at least until Menindee Lakes.
Unlike the Murray system, which has an
extensive network of lentic waters — a het-
erogeneous array of billabongs (Hillman
1986), intermittently flooded Barmah-
Millewa Forest waters, the Macquarie
Marshes, irrigation returns from tributary
systems — the Darling is deeply incised (up
to 10 m) into its floodplain, and lacks these
sources of lentic microfauna, It remains a
lotic, largely unimpeded system, albeit one
of low flow,

Notably, a rotifer-dominated river micro-
fauna (Brachionus, Keratella and
Synchaeta dominants among 22 rotifer
species) was collected from the Murray
River near Morgan early in 2002 (MC
Geddes pers. comm.), over 20 years after a
very similar autumn river plankton was
reported from the same reaches of the

209

Murray River Special Issue

lower Murray (Shiel et al. 1982). Despite
widely publicised alterations to the flow
regime, abstractions, declining water quali-
ty, the spread of carp and other deleterious
effects of human interference with Murray
waters, the microfauna appears remarkably
little changed. Twenty years may be
insignificant in the time frame of the
Basin's ecological evolution.

Biogeography/endemism

Microfaunal biogeography is subject to
two interpretations — that of the *cosmopoli-
tanists', for whom everything is every-
where, and the ‘regionalists’, for whom it is
not. Many of the microfauna species found
throughout Murray River waters are the
same species found in rivers and lakes
everywhere; they are indeed cosmopolitan.
But some are not. If they occurred world-
wide, surely they would have been recorded
in some 300 vears of microfaunal research
in the northern hemisphere,

As noted earlier, the protists remain enig-
matic. A small block of rotifers are more
restricted, Gondwanan or Australasian on
present distribution information. About 13-
15% of rotifers are Australian endemics, or
possibly more correctly ‘Australasian’;
some Brachionus species previously
thought to be indigenous have been report-
ed from Thailand (Sanoamuang er al.
1995). Most of the apparently indigenous
rotifers are known from only one or a few
habitats, primarily billabongs or small
ephemeral waters, and appear to be
restricted to them. The implication that
they are thus endangered by loss of habitat
is clear, but loss of species remains undoc-
umented for the continent.

Cladocerans presently stand at c. 48%
endemism, but this figure is likely to
increase when taxa carrying ‘northern
hemisphere’ names are examined more
closely using modern methods (Shiel and
Dickson 1995). The work of Bayly (cited
in Bayly 1992) indicates that the level of
endemism in the Australian calanoids
approaches 90%, Cyclopoids are less
intensively studied, but evidence to date
(e.g. Holynska 2000) suggests that greater
endemism will be revealed as the
Australian ‘cosmopolitan’ species shed
their ‘European’ names,

210

The future?

In the Murray-Darling Basin, as in salin-
izing wetlands in the southwest of Western
Australia and in other degraded wetlands
on continental Australia and Tasmania,
species diversity is inversely related to
salinity. The diverse microbiota of fresh
waters is replaced by a halophile or halo-
biont assemblage tolerant of the new con-
ditions. The new taxa may even be more
abundant than the assemblage they
replaced, but they are invariably less spe-
ciose. For example, the 100-species rotifer
assemblage of upper Murray billabongs is
replaced by one or two species of
Brachionus/Hexarthra around Kerang, and
the multispecies assemblage of Boeckella/
Calamoecia/Hemiboeckella of upper
Murray billabongs is replaced by
Calamoecia salina or C. clitellata in the
salinized downstream lakes. In the human
time frame this decline in biodiversity is
well documented for macrofauna . What is
not documented is the loss of species at the
microfaunal level due to loss of habitat.
With greater than 70% of billabongs now
gone from some floodplain reaches (e.g.
Goulburn River downstream of Eildon),
how many species previously found only
in those billabongs are now lost?

Persistence of propagules in sediments,
despite the loss of wetlands, may be used
to counter the suggestion that microfaunal
Species are being lost at an increasingly
rapid rate. But as mentioned earlier, resting
stages may have specific cues for emer-
gence, and habitat changes such as salin-
ization may remove these cues. Decreased
flooding frequency may exceed the life
expectancy of a propagule — resting stages
have a finite life, be it decades or perhaps
centuries, Further, propagules evolved to
resist desiccation, not trampling by cattle,
or tractor and plough. The introduction of
exotic planktivorous fish (e.g. Gambusia),
either deliberately or accidentally, must
also have had a profound impact on micro-
faunal populations.

In summary, it is evident that microfau-
nal heterogeneity persists in the more pris-
tine (upper) Murray-Darling floodplain
waters, and it is the floodplain which is the
source of the biodiversity, not the reser-
voirs, which are relatively recent in an
evolutionary time-frame, nor the rivers,

The Victorian Naturalist

which transport microfauna and propag-
ules. With loss of the more pristine wet-
lands due to agriculture, salinization, or
other causes, the microfauna is lost, as ulti-
mately, if not replenished, is the bank of
cysts, resting eggs and ephippia in the sed-
iments.

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Boulton AJ and Lloyd LN (1992) Mean flood recur-
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Part Two

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Shiel RJ (1976) Associations of Entomosiraca with
weedbed habitats in a billabong of the Goulburn
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284. Eds N Mackay and D Eastburn. (Murray-
Darling Basin Commission: Canberra)

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from Australia. Transactions of the Royal Society of
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Shiel RJ, Green JD and Nielsen DL (1998) Floodplain
biodiversity: Why are there so many species?
IHydrobiologia 387, 39-46.

Shiel RJ, Green JD and Tan L-W (2001) Microfaunal
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Shiel RJ, Walker KF and Williams WD (1982)
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Skinner R, Sheldon F and Walker KF (2001)
Propagules in dry wetland sediments as indicators of
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211

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The
Victorian

Naturalist

Volume 119 (5) a October 2002

aN

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Treasures from the Kingdom of Fungi

by Taylor F Lockwood

Publisher: Taylor Lockwood, 2001. 128 pp, 250 colour photographs. RRP $55
Available from Fungimap, Royal Botanic Gardens Melbourne, phone 03 9252 2374

The aptly titled Treasures from the
Kingdom of Fungi is the long-awaited hard-
bound collection of Taylor Lockwood's
amazing photographs. Taylor has been pre-
senting these in slide shows around the world
for the past 15 years. This 128 page book
contains approximately 250 colour pho-
tographs of fungi, illustrating 157 taxa from
19 countries (including 25 Australian taxa).
Text is limited to the preface, foreword and a
brief introduction; most of the book consists
of page after page of delightful photographs
including many mushrooms but also a good
selection of puffballs, stinkhorns, coral and
bracket fungi.

The images are grouped together in chap-
ters that divide the book aesthetically
rather than scientifically. Chapters include
‘The Span of the Rainbow’ showing fungi
of every colour of the rainbow, ‘Foods of
the Gods’ illustrating some of the edible
fungi, and a ‘Lovers’ Lane’ of naturally-
occurring, artistically-twinned examples.
Each photograph is captioned merely with
identification of the fungus (although some
20% are ‘unidentified’, not even to genus),
and the general location where the photo-
graph was taken (usually the country name
only, but states are also listed for the USA
examples), Taylor’s enthusiasm for the
topic is clearly evident with the layout and
backgrounds on each carefully designed
page showing great attention to detail.

Having spent many days in the field in
search of perfect fungi to photograph, and
having referred to many other books of
field photography, I can personally attest
to the difficulty of finding a good quality
specimen and the skill required to capture
it adequately for posterity. The broad
range of superb field examples chosen as
the subject matter in this book are evi-
dence of Taylor's efforts *on the ground',
and his technical and creative talents as
photographer are clearly demonstrated in
the spectacular images that result.

One drawback of this book is the number
of photographs labelled as ‘unidentified’.
Taylor does not collect specimens to allow

identification at a later date, and some of the
photographs are artistic rather than scientific,
and so lack all the diagnostic features
required for identification. However, scien-
tific presentation is not the aim of this book.
Taylor's goal was to inspire interest in and
appreciation of fungi, which I feel he has
well and truly accomplished with this col-
lection. Treasures from the Kingdom of
Fungi is not a field guide or scientific text
(which should have few unidentified
photographs and preferably include a scale),
but a brilliant ‘coffee table book’ on fungi.

To produce a book without some typo-
graphical or grammatical errors would be
an almost impossible task. There were a
small number of general mistakes in this
book as well as a few names spelt incor-
rectly; Cyptotrama aspratum (spelt as
asprata), Mycena leaiana (incorrectly spelt
leiana) and Tremella fuciformis (incorrect-
ly spelt fusiformis), also in the index
Boletus is spelt Bolteus, and Cladonia
comes before Cladina. However, these cer-
tainly do not detract from the overall high
standard of presentation.

The retail price of $55 for this book may
seem a little high, but it is well worth con-
sidering for the bounty of beautiful, high
quality images. Taylor's aim ‘to inspire
others into deeper appreciation of this spe-
cial part of the natural world’ has been
admirably achieved by superbly illustrat-
ing the diversity of this often weird and
wonderful kingdom. Although this is not a
scientific text, I still feel it should be pre-
sent on the book shelf of every mycologist
and naturalist as an inspiration and delight
if nothing else.

Simon H Lewis
Royal Botanic Gardens Melbourne
Birdwood Avenue, South Yarra, Victoria 3141

Twelve Fungimap Target species àre included (most
showing diagnostic features); Amanita phalloides (var.
alba) p. 45; Ascocoryne sarcoides p. 106; Boletellus
obscurecoccineus p. 51; Coprinus comatus p. 118;
Craterellus cornucopioides p. 121; Cyptotrama aspratum
pp. 37, 53: Lepista nuda p. 118; Mycena interrupta pp.
31, 42: Mycena leaiana p. 86; Podoserpula pusio p. 28;
Pseudohydnum gelatinosum pp. 20, 79; Tremella fuci-

formis p. 106,

The
Victorian

Naturalist
Volume 119 (5) 2002 E October

F.N.C V,

Executive Editor: Merilyn Grey
Editors: Alistair Evans and Anne Morton

Research Report — Seasonal and Altitudinal Differences in the Abundance and
Species Richness of Some Macroinvertebrates in
Kosciuszko National Park, Australia, by Michelle Stock

Ora Cathermedanina PIREN voe retra oriens reete dans 216
Contributions Observations on the Feeding Behaviour of Uchidanurinae
(Collembola: Neanuridae) in Australia, by Penelope
Greenslade, Sean Moore and Roger Farrow iesirea 221
Butterfly Management Uses Pollard Walk,
by Bryan T Haywood and Christopher J Wilson... ess 224

First Record of the Asian Freshwater Leech Barbronia weberi
(Blanchard, 1897) (Euhirudinea: Erpobdellidae) in Australia,
by Fredric R Govedich, Bonnie A Bain and Ronald W Davies .....227
Dragonfly Nymphs with Dipteran *Hitch-hikers': an Example
of Phoresy Found in Dandenong Creek,

DIA DORT ESSI QUU, crs Pet neh erre aderant per UIDES sete eR 229
Acacia obtusifolia — Introduction and Spread in Native Bush,
a AL AET Deu uns E t Ms tom M. C c Cm Ol

The Discovery of Leadbeater's Possum Gymnobelideus
leadbeateri along the Woori Yallock Creek, Yellingbo,

hari ato or E SRM A Pre MES EC Be ae TA AN A 233
Ooline Cadellia pentastylis F.Muell.: a Survivor,
DIR OR ECL CHIE Med, A Arden e eed Meee AN eee, EO
Foraminifera from Lake Corangamite, Victoria, by Ken N Bell......237
Naturalist inthe Winter Keeps Us Warm, by Ken Green... $238
Mountains Snow in Spring, by Ken Green ..........
Tribute William Nigel Balcombe Quick, by Michael F Bi wb and
USPC NIE CAE UTRAM... MO. een Ee MC UE 242
Book Reviews Treasures from the Kingdom of Fungi, by Taylor F Lockwood,
SAU ECT bay Tat phe c (o 8 ID ECT Ye ele rei M e TUO cp EN. EN 214

Butterflies of Australia: their Identification, Biology and
Distribution, by Michael F Braby, reviewed by Andrew Atkins ....244
Dragonflies of the World, by Jill Silshy, reviewed by

OIOV ETE T Te pco 7 ORCC SS QI ia t3 REDDIT eee 245
Possums: The Brushtails, Ringtails and Greater Glider,
by Anne Kerle, reviewed by Ken Bell .................. eere 246

ISSN 0042-5184
Cover: Dragonfly Aeshna sp. — a representative of the invertebrates covered in this issue, e.g.
dragonfly nymphs (p. 229) and dragonflies of the world (p. 245). Photo by Roger Gaymer.

Web address: http://www.vicnet.net.au/~fnev/vicnat.htm
email: fnev@vicnet.net.au

Research Report

Seasonal and Altitudinal Differences in the Abundance and
Species Richness of Some Flying Macroinvertebrates in
Kosciuszko National Park, Australia

Michelle Stock! and Catherine Marina Pickering

Abstract

Seasonal and altitudinal effects on invertebrate abundance and the diversity of flying macroinverte-
brates, including potential pollinators, were examined at an alpine, subalpine and montane site in
Kosciuszko National Park, Australia. Sampling using traps consisting of sticky white balls was con-
ducted at four times during the growing/flowering season (between December and March). Only
insects greater than 4 mm (macroinvertebrates) were identified. Of the total of 9011 invertebrates
collected the most diverse and abundant macroinvertebrate taxa were Diptera. The next most abun-
dant and diverse were the Hemiptera and Hymenoptera, with few Lepidoptera and Orthoptera. There
were both seasonal and altitudinal effects, with the abundance and diversity of macroinvertebrates
and Diptera (74 mm) highest between December and February, corresponding to the main flowering
periods. Abundance at all sites declined in March as temperatures declined. The abundance and
diversity of macroinvertebrates in the alpine site was higher than expected compared to the montane
site during the first three sampling periods. This differs from studies overseas that have found a
decline in macroinvertebrate diversity and abundance with increasing altitude. (The Victorian Naturalist

119 (5), 2002, 216-221)

Introduction

Invertebrate diversity, abundance and
activity have been found to decline with
increasing altitude in mountain regions
(Primack 1978; Arroyo er al. 1982, 1985;
Inouye and Pyke 1988; Kearns 1992;
Primack and Inouye 1993; Green 1997;
Bingham 1998). This is thought to be due
to the increasing severity of the environ-
ment, with lower temperatures, less bio-
mass of vegetation, longer periods of snow
cover, increased risk of frosts etc. at higher
altitudes (Mani 1968; Arroyo ef al. 1985;
Inouye and Pyke 1988; Green and Osborne
1994). For example alpine areas are char-
acterized by atmospheric cold, high atmos-
pheric aridity, snow and ice, high ultra-
violet radiation, large diurnal fluctuations
in temperature, and rapid desiccation of
delicate and soft-bodied organisms. These
factors constrain the survival, growth and
reproduction of invertebrates limiting their
abundance and distribution in high altitude
environments (Mani 1968; Arroyo et al.
1982, 1985; Green and Osborne 1994;
Green 1997; Korner 1999).

Invertebrate abundance, activity and
diversity can also vary seasonally in sub-
alpine and alpine areas, including in the
Australian Alps (Inouye and Pyke 1988;
! School of Environmental and Applied Sciences,
Griffith University, Gold Coast Campus, PMB 30

Gold Coast Mail Centre, Queensland 9726. Email
c.pickering@mailbox.gu.edu au

216

Green 1997). For example, very few
Diptera and other flying insects overwinter
in alpine areas with most either there acci-
dentally (lifted by air currents), or migrat-
ing into the alpine for the summer, but
returning to the lower areas to breed during
winter (Mani 1968; Green and Osborne
1994), During the snow free period insect
activity can vary with weather conditions,
particularly temperature, and wind, This
can result in seasonal and diurnal variation
in the abundance and diversity of pollina-
tors in alpine sites (Arroyo er al. 1985;
Inouye and Pyke 1988; Bergman er al.
1996; Green 1997),

The most common type of insects in the
alpine region of Kosciuszko National Park
are Orthoptera (crickets and grasshoppers),
Coleoptera (beetles), Hemiptera (bugs),
Lepidoptera (butterflies and moths),
Diptera (flies) and Collembola (springtails)
(Inouye and Pyke 1988; Green and
Osborne 1994; Green 1997). The main
potential pollinators of the Kosciuszko
alpine flora are thought to be the Diptera
with 60 species visiting flowers (Inouye
and Pyke 1988). Hymenoptera (33 species),
along with several species of Lepidoptera
and Coleoptera, were also found on flowers
(Inouye and Pyke 1998). Unlike other
alpine areas, there is a lack of social bees
and other types of specialist pollinators in
the Australian Alps and in New Zealand

The Victorian Naturalist

alpine areas matching an apparent lack of

specialist pollination syndromes (Primack
1978; Inouye and Pyke 1988; Pickering
1997). The decline in the abundance and
diversity of insect pollinators in subalpine
and alpine environments is thought to have
influenced pollination syndromes, timing
of flowering and duration of stigmatic
receptivity of alpine plants (Arroyo et al.
1982, 1985; Inouye and Pyke 1988; Kearns
1992: Totland 1993; Bergman ef al. 1996;
Bingham 1998; Pickering 1997; Korner
1999),

The aim of this study was to examine if

altitude and seasonality affected the diversity
and abundance of some flying macroinverte-
brates in Kosciuszko National Park. [t would
be expected that the abundance of inverte-
brates including pollinators varies within a
flowering season, with the highest abun-
dance in the warmest months (January-
February in Australia). Peak flowering in the
alpine region of Kosciuszko National Park
also occurs in the warmest months, although
some species flower early in the season,
soon after snow-melt or later in the season,
often in March (Costin et al. 2000; pers.
obs.). At lower altitudes insect numbers
could peak earlier in the season during
spring, when most plants are flowering.
Numbers could also remain higher for
longer, as temperatures are higher on aver-
age, and do not fall as fast late in the grow-
ing season (March-April; Green and
Osborne 1994), It would also be expected
that at higher altitudes macroinvertebrate
activity/abundance would be lower than at
subalpine or montane sites at any given time.

Methods
Study site
Macroinvertebrate diversity and abundance
was sampled using passive traps consisting
of white Styrofoam balls (11 cm diameter)

coated in Tangle Trap® (modification of

method by Prokopy 1968) attached to 2
mm diameter wire, with the balls posi-
tioned approximately 5 cm above the vege-
tation. Traps were placed at three altitudes
in Kosciuszko National Park, New South
Wales. The lowest altitude site was in the
Eucalyptus woodland (montane vegetation
zone) 1400 m ast near Sawpit Creek, The
next site was in subalpine heath near
Prussian Creek below Smiggin Holes

Vol. 119 (5) 2002

Research Report

(1640 m ASL) and the final site was in tall
alpine herbfield in the alpine region above
Charlotte Pass Village on Mt Gutherie
(1860 m ASL).

Sampling occurred four times during the
flowering season. Initial sampling was on
18-20 December 1998 when all sites had
been snow free for several weeks and tem-
peratures were starting to increase, Sites
were sampled again on 16-18 January 1999
at the peak of the alpine flowering season.
The third sampling period was 1-3 February
1999 when flowering was declining at all
sites. The final sampling was 27-29 March
1999 when flowering was finished and cli-
matic conditions were less favorable for
insect activity.

At each site one passive trap was ran-
domly placed along each of five 30 m tran-
sects approximately 50 m apart. The traps
were left for two days. Invertebrates
greater than 4 mm in length were collected
and identified to morphospecies in the
field laboratory. It was not feasible to iden-
tify invertebrates less than 4 mm in the
field laboratory.

Statistical analysis

Effects of altitude and season on species
richness and abundance were examined by
two-way analysis of variance using SPSS
10.00 (Coakes and Steed 2001), However,
due to the relative lack of spatial dispersal
of the traps at each altitude, the altitudinal
effect must be interpreted with caution. To
satisfy the assumptions of the analysis the
total number of invertebrates, total number
of Diptera, and number of invertebrates >4
mm data were transformed using a natural
log (x + 1) transformation, Tukeys post hoc
test was used to compare means at each
time and site.

Results
Diversity and abundance

A total of 9019 invertebrates was collect-
ed over the four sampling periods at the
three sites, For macroinvertebrates (2490
>4 mm), the most common were Diptera
(flies 69%; Table 1). The other groups of
macroinvertebrates in decreasing abundance
were Hemiptera (bugs ~11.5%),
Hymenoptera (bees and wasps ~11%),
Lepidoptera (moths ~4%), Orthoptera

Á

(grasshoppers ~4%), Blattodea (cockroach-

217

Research Report

es — less than 1%), Arachnida (spiders — less
than 1%) and Coleoptera (beetles — less
than 1%). Approximately 76% of the inver-
tebrates caught were smaller than 4 mm
(Table 1) and were not identified.

Diptera were the most diverse macroin-
vertebrates as well as the most abundant
(Table 2). There were 31 morphotaxa rep-
resenting at least 10 families. Hymenoptera
were represented by three bee morphotaxa
and four wasp morphotaxa. Within the
Lepidoptera there were five morphotaxa,
with at least one representative of the
Oecophoridae, Geometridae and Noctuidae
families. There were four Orthoptera mor-
photaxa, one type of Blattodea, one type of
Coleoptera, and two types of Hemiptera.

Effect of altitude and time

The total number of invertebrates varied
seasonally at each of the three altitudes
with a clear decline in invertebrate num-
bers in March compared to earlier in the
season (Fig. 1). There were significantly
more invertebrates collected in December
in the alpine site than at any other time at
any altitude (Fig. 1; two-way ANOVA,
effect of time by altitude interaction, df —
6, F — 19.819, p « 0.001). The peak num-
ber of invertebrates was highest in January
for both the subalpine and montane sites.

The data for just the small invertebrates
(less than 4 mm) showed the same pattern as
for total invertebrates. The largest number of
invertebrates was found on traps in the alpine
site in December, and there was a decrease in
the number of invertebrates caught at the
three altitudes in March (Table 3; two-way

Table 1. Number of invertebrates belonging to
different orders trapped at the three altitudes
(montane, subalpine and alpine sites) in
Kosciuszko National Park during the 1998/1999
growing/flowering season.

Invertebrate Total % all %

order number invert. >4mm
Micro- and meso-

invertebrates 6931 76.85
Diptera 1432 15.88 68.58
Hemiptera 240 2.66 11.49
Hymenoptera 226 2.5] 10,82
Lepidoptera 87 0.96 4.17
Orthoptera 84 0.93 4.02
Blattodea 10 0.11 0.48
Arachnidae 8 0.09 0.38
Coleoptera l 0.01
Total 9019

218

ANOVA, effect of time by altitude interac-
tion, df= 6, F = 12.733, p < 0.001).

The data for Diptera only also showed
the same pattern, There was a significant
interaction between altitudes and times
(Table 4; two-way ANOVA, effect of time
by altitude interaction, df= 6, F = 6.378, p
< 0.001). The number of Diptera also
declined dramatically in March at all alti-
tudes (Table 4).

There was considerable variation in
diversity of macroinvertebrates at the dif-
ferent altitudes and times (Fig. 2; two-way
ANOVA, effect of time by altitude interac-
tion, df = 6, F = 3.835, p = 0.003). All
three altitudes showed a relatively constant
number of morphospecies between
December and February, with a clear
decrease in March (Fig. 2).

Discussion

The abundance and diversity of inverte-
brates was lower at the end of the grow-
ing/flowering season in March than in
December and January at all three alti-

Table 2. Diversity of insects trapped at the three
altitudes during the 1998/1999 flowering season
at Kosciuszko National Park.

Order
Diptera

Family No. morphotaxa

Total 31
Muscidae
Tachinidae
Calliphoridae
Syrphidae
Tabanidae
Asilidae
Tipulidae
Simulidae
Tephritidae
Culicidae
Unknown

Total
Anthophoridae
Ichneumonidae
Unknown
Total
Oecophoridae
Geometridae
Noctuidae
Unknown
Total
Acrididae
Pyrgomorphidae
Total

‘Total

Cleridae

Total
Cicadidae
Unknown

= — n= co ae eal ee ge a ON OS ee eos gn aloo

Hymenoptera

Lepidoptera

Orthoptera

Blattodea
Coleoptera

Hemiptera

The Victorian Naturalist

800

200 —
= ur = x

0 IE
December January February March

Mean number of invertebrates per trap
b
e
©

Alpine

800

200
== i

Mean number of invertebrates per trap

E vw
0 ==
December January February March
Subalpine
&
$ 600
=
s
2
2 400
B 200: p
A dr
s
E!
E]
z

December January February March

Montane

Fig. 1. Mean and standard error of the total
number of macroinvertebrates per trap at four
times during the flowering season at three alti-
tudes in Kosciuszko National Park.

tudes. This pattern is consistent with other
studies in alpine and subalpine sites in
Kosciuszko National Park that used differ-
ent sampling methodologies (Inouye and
Pyke 1988; Green 1997), as well as with
studies of alpine and subalpine environ-
ments elsewhere in the world (Arroyo ez
al. 1982, 1985; Primack and Inouye 1993;
Bergman et al. 1996; Bingham 1998). The
reduction in invertebrate numbers in
March is likely to be due to factors such as
lower temperatures at the end of the flow-
ering season. Factors such as temperature,
wind speed and light levels have all been
found to affect insect activity, including in
sites close to those used in this study
(Primack 1978; Arroyo et al. 1985; Inouye
and Pyke 1988; Primack and Inouye 1993;
Bergman er al. 1996). These changes in
invertebrate numbers are likely to affect

Vol. 119 (5) 2002

Research Report

E
5
È
g 8
Bg
S d
5
e EN
5
E o»
0
December January February March
Alpine
14
a
Eam f
& 10 pee
Bs " DR
5
$ 6
ij
Ss 4
=
2 EL
0
December January February March

Subalpine

e T.

Mean diversity per trap

December January February March

Montane

Fig. 2. Mean and standard error of the diversity
of macroinvertebrates per trap at four times dur-
ing the flowering season at three altitudes in
Kosciuszko National Park.

the potential for insect pollination for those
few alpine plants that flower late in the
season (Arroyo et al. 1985; Inouye and
Pyke 1988; Primack and Inouye 1993;
Bergman eft al. 1996). The bulk of the
flowering in the Kosciuszko alpine
occurred in December and January, when
macroinvertebrate diversity was high. By
March, however, when only a few species
were flowering (e.g. Mueller's Snow-gen-
tian Chionogentias muelleriana), most
plants had finished releasing their seeds
and insect diversity had decreased to only
one morphospecies (a hymenopteran).

The high abundance and diversity of
macro-Diptera compared to other orders of
macroinvertebrates found in this study are
consistent with observations in nearby
sites. Diptera were the most diverse (31
species, at least 10 families) and abundant

219

Research Report

E AO E L o Å o o aM A
Table 3. Mean (and standard error) of the number of <4 mm invertebrates trapped at each sampling
period at alpine, subalpine and montane sites.

Sampling date Alpine Subalpine Montane Total

20 December 1998 469 + 109 11248 110 218 2302-57
18 January 1999 73414 120 + 24 166+ 46 120+ 19
3 February 1999 314 18 66 + 24 121 +43 824-18
29 March 1999 2341 1528 TO SIED 34-210
Total 168 + 54 78 + 13 117417 120+ 19

TES Ss eh a Se TN aT LES Saas wen ae EES Te,
Table 4. Mean (and standard error) of the number of Diptera trapped at each sampling period at
alpine, subalpine and montane sites.

Sampling date Alpine Subalpine Montane Total
20 December 1998 75 +27 143 34 - 10 41-11
18 January 1999 2943 31+7 2347 2843
3 February 1999 27-6 194-5 245 23-3
29 March 1999 0 1441 942 aem
Total 33 10 16-3 2244 2443

taxa (69% of insects greater than 4 mm) in
the study presented here. Inouye and Pyke
(1988) also found that Diptera were the
most abundant and diverse flower visitor,
with a total of 60 species of Diptera,
belonging to 18 families visiting flowers
from a range of species. The most abun-
dant flower visitors were Diptera, account-
ing for 62.3% of flower visitors, followed
by Hymenoptera (30.8% flower visitors),
then Lepidoptera (11.2%) and Coleoptera
(1.9%) (Inouye and Pyke 1988).

Several studies have shown that species
richness decreases with increasing altitude
(Mani 1968; Arroyo er al, 1985; Primack
and Inouye 1993; Green 1997; Bingham
1998), From this, it was expected that there
would be a greater diversity of taxa in the
montane site, than the alpine and subalpine
sites. This was not the case. Overall there
were forty-three morphospecies of macroin-
vertebrates in the alpine, thirty-three in the
subalpine and forty in the montane site. In
addition, during peak flowering in the
alpine, there were either more or similar
numbers and diversity of macroinvertebrates
than in the subalpine and montane sites.
Therefore, it could be that there are quite
high numbers of macroinvertebrates, partic-
ularly Diptera, available to act as pollinators
at the time of peak flowering in the alpine.
The results obtained in this study should be
viewed with some caution, as they could be
due to site specific factors, and hence may
not be representative of a general trend with
altitude. Also, the diversity data only applies

220

lo insects greater than 4 mm and hence does
not reflect total invertebrate diversity.

The relative dominance of Diptera
amongst alpine insect fauna, particularly
among potential pollinator suites, has been
found in a range of alpine regions (Inouye
and Pyke 1988; Kearns 1992: Totland
1993). It is thought, in part, to be due to
Diptera numbers not declining, or not
declining as fast with increasing altitude as
the abundance and diversity of other insect
groups, such as bees (Primack 1978; Arroyo
et al. 1982, 1985; Kearns 1993). For exam-
ple, the diversity of Diptera was higher at
high and mid elevations than at low eleva-
tions in study across an altitudinal gradient
in the Rocky Mountains, USA (Kearns
1992). The abundance of Diptera did not
decline with increasing altitude, unlike other
groups of insects (Kearns 1993), resulting in
the increasing importance of Diptera as pol-
linators at higher altitude. Similar patterns
have been found in some other alpine
regions (Primack and Inouye 1993; Totland
1993) with Diptera abundance and diversity
found to be fairly constant or declining more
slowly across altitudinal gradient. The effect
of this is that at higher elevations, which
often have fewer bees and other specialized
pollinators, Diptera were most common
(Arroyo ef al. 1982; Primack and Inouye
1993). In the study presented here, Diptera
were also found to be the most diverse and
abundant insects in the alpine zone, adding
support to their prominence as potential pol-
linators in such environments.

The Victorian Naturalist

Acknowledgements

Thanks to New South Wales National Parks and
Wildlife Service for access to the site, and to
Janice Harrington, Karen Rudkin and Kylie
Catteral for their assistance in the field.

References

Arroyo MTK, Primack R and Armesto J (1982)
Community studies in pollination ecology in the
high-temperate Andes of central Chile, I: Pollination
mechanism and altitudinal variation. American
Journal of Botany 69, 82-97.

Arroyo MTRK. Armeston JJ and Primack RB (1985)
Community studies in pollination ecology in the high
temperate Andes of central Chile. II: Effects of tem-
perature on visitation rates and pollination possibili-
ties, Plant Systematics and Evolution 149, 187-203.

Bergman P, Molau U and Homgren B (1996)
Micrometeorological impacts on insect activity and plant
reproductive success in an alpine environment, Swedish
Lapland. Arctic and Alpine Research 28, 196-202.

Bingham RA (1998) Efficient pollination of alpine
plants. Nature 391, 238-239.

Coakes SJ and Steed LG (2001) SPSS Analysis Without
Anguish, (Wiley: New York)

Costin AB, Gray M. Totterdell CJ and Wimbush DJ
(2000) Kosciusko Alpine Flora. (CSIRO: Collingwood)

Green K (1997) Inter-annual, seasonal and altitudinal
differences in invertebrate activity in the Snowy
Mountains. The Victorian Naturalist 114, 222-229.

Green K and Osborne W (1994) Wildlife af the

Contributions

Australian Snow Country, (Reed Books; Chatswood)

Inouye DW and Pyke GH (1988) Pollination biology in
the Snowy Mountains of Australia: Comparisons
with montane Colorado, USA. Australian Journal of
Ecology 13, 191-210.

Kearns CA (1992) Anthophilous fly distribution across
an elevation gradient. American Midland Naturalist
127, 172-182.

Korner C (1999) Alpine Plant Life: Functional Plant
Ecology of High Mountain Ecosystems. (Springer:
Berlin)

Mani MS (1968) Ecalogy and Biogeography of High
Altitude Insects. National Library of Australia, Series
Entomologica 4.

Pickering CM (1997) Reproductive strategies and con-
straints of alpine plants as illustrated by five species
of Australian alpine Ranunculus. Opera Botanica
132, 101-108.

Primack RB (1978) Variability in New Zealand mon-
tane and alpine pollination assemblages. New
Zealand Journal of Botany 1, 66-73.

Primack RB and Inouye DW (1993) Factors affecting
pollinator visitation rates: a biogeographic compari-
son. Current Sciences 65, 257-262,

Prokopy RJ (1968) Sticky spheres for estimating apple
maggot adult abundance, Journal of Economic
Entomology 61, 1082-1085,

Totland O (1993) Pollination in alpine Norway: flower-
ing phenology, insect visitors, and visitation rates in
two plant communities. Canadian Journal of Botany
71, 1072-1079.

Observations on the Feeding Behaviour of Uchidanurinae
(Collembola: Neanuridae) in Australia

Penelope Greenslade!', Sean Moore’ and Roger Farrow’

Abstract

Two observations of feeding on the plasmodial stage of a slime mould (Mycetozoa) in Eucalyptus
forest in Victoria and Tasmania by species of the log-inhabiting Uchidanurinae are documented. It is
suggested that feeding on Protozoa may be much more common in members of the Neanuridae than
is currently considered, The Uchidanurinae are of conservation significance as they tend to be inhab-
itants of old growth vegetation, highly diverse in south-eastern Australia and Tasmania and very sus-

2, 221-223)

ceptible to human impacts. (The Victorian Naturalist 119 (5), 2002, 22

Introduction
The Uchidanurinae are among the largest
and most spectacular Collembola.

Individuals may be up to 10 mm long and are
dark blue, grey or black with a flattened, seg-
mented body furnished with lobes and digita-
tions often conspicuously tipped with orange
or yellow. The animals live in or under old
decaying logs in old growth forests or in
heathland undisturbed by fire for long peri-
ods, and unlike most other Collembola, lack
a jumping organ. Although only three
Australian species have been described so

! Division of Botany and Zoology, Australian National
University, Canberra, ACT 0200

1 7 Girdwood Road, Boronia, Victoria 3155

' Tilembeya Consulting, 777 Urila Road, Queanbeyan,
NSW 2620)

Vol. 119 (5) 2002

far, collections from south-east Australia and
Tasmania indicate that about thirty species
can be distinguished on gross morphology
and it is likely that a much larger number of
cryptic species exist (Greenslade 1991,
1993). The subfamily is also found in New
Zealand, New Caledonia and parts of south-
east Asia and is considered to be Gondwanan
in origin (Greenslade 1991).

Based on their mouthparts, Collembola
can be divided into two groups. The first
group possesses a mandible with a toothed
molar plate while the second group has a
simpler mandible without a toothed molar
plate. Observations made on easily cultured
species from families belonging to the first

221

Contributions

group suggest that Collembola have rela-
tively unspecialised feeding habitats and
consume fungi, bacteria and other microor-
ganisms, with some preferences for certain
fungal species, possibly based on nutrition-
al value (Hopkins 1997). However, by defi-
nition, easily cultured species are likely to
show relatively generalised feeding prefer-
ences so the unspecific nature of feeding
habits may not be as widespread as these
observations indicate (Greenslade 1993).
Families lacking the molar plate, such as
the Neanuridae to which the Uchidanurinae
belong, show a greater morphological varia-
tion in mouth parts, suggesting that a range
of specialised feeding habits may be a char-
acteristie of this group. It is known that sev-
eral species of Neanurinae, a closely related
subfamily, have been kept alive for long
periods as long as temperatures are kept low
and moisture regimes high, but feeding and
reproductive behaviour were never observed
in culture (L Deharveng pers. comm.). The
Uchidanurinae are typical of the Neanurinae
in that individuals have been kept at a tem-
perature of 4°C for over five months, but
feeding on microorganisms such as bacteria,
yeasts or fungi was never observed nor were
any eggs seen (P Greenslade pers. obs.). In
addition, studies of the alimentary canals of
Neanurinae have failed to find any identifi-
able contents (L Deharveng pers. comm. ).

Observations

Two independent observations in the
field have been made of Uchidanurinae
feeding in natural undisturbed environ-
ments. In both cases a number of individu-
als were observed during the day feeding,
on the plasmodial stage of a yellow slime
mould, probably Fuligo septica. The first
observation was made by R. Farrow in tall
wet sclerophyll forest in the Florentine
Valley, southwest Tasmania, in February
1992. Farrow photographed two individu-
als apparently feeding on a conspicuous
yellow slime mould growing on the lower
side of a well-rotted log, possibly of a
Eucalyptus species, in deep shade about
midmorning. The second set of observa-
tions were made by S Moore on both the
29 July, 2001 and the 5 August, 2001, at
the same site in semi-open eucalypt forest
in Kinglake National Park, northeast of
Melbourne, Victoria. These animals were

222

feeding on the slime mould growing on the
outside of a rotting log, possibly Acacia
sp., in eucalypt forest with an understorey
of bracken Preridium esculentum, Again,
the observations were made during the
day. On the first occasion, two individuals
were noted at 2 pm feeding in overcast,
calm and wet conditions at a temperature
of approximately 12°C. On the second
occasion, ten individuals were observed at
10 am in overcast, calm and damp condi-
tions at a temperature of approximately
8°C, This site was extremely humid, being
adjacent to a rainforest gully. Photographic
records of both observations were made
and specimens were collected, The two
species were different and belonged to two
different available genera in the
Uchidanurinae. The Tasmanian species
belonged to Acanthanura sp, cf. dendyi
(Lubbock 1899) and the Victorian species
to Womersleymeria sp. bicornis group
(Womersley 1940), but cannot be identi-
fied further until a revision of the subfami-
ly in Australia can be carried out.

Specimens of both slime mould and
Collembola from the Kinglake National
Park were retained for examination in cul-
ture but proved difficult to observe under a
normal microscope light.

Discussion

Slime moulds are found on shed bark,
well-rotted logs and in leaf litter, mainly in
moist environments. They are most easily
cultured from cut bark and survive desic-
cation and rehydration (Stephenson and
Stempen 1994). All collections of
Uchidanurinae from forests have been
made from similar habitats, that is either
within or under logs or in leaf litter with an
abundant bark content. Individuals are usu-
ally collected from just under bark on logs
or between bark and the thick layer of
moss growing on it.

Slime moulds were, in the past, consid-
ered to be closely related to fungi, as indi-
cated by the scientific name, Myxomycetes.
However they are now considered to have a
closer relationship and ancient ancestry with
Protozoa, and are more correctly referred to
as Mycetozoa (Wheeler 1984). Although
soil Protozoa have sometimes been record-
ed as food of Collembola there have been
no direct data to support the suggestion. In

The Victorian Naturalist

fact, reviews of feeding in Collembola usu-
ally fail to mention Protozoa (Christiansen
1964) or mention them but give no support-
ing reference (Rusek 1998). Andren and
Schnurer (1985) assume Protozoa constitute
a considerable proportion of the diet of
Folsomia fimetaria (Isotomidae), as only a
small percentage of the animals’ growth
could be accounted for by fungi, but direct
evidence was lacking. Although Ing (1967)
recorded adventitious Collembola probably
feeding on slime moulds in culture, the only
published observation of an identified
collembolan species feeding on the plas-
modial stage of Mycetozoa is that of
Chassain (1973). He observed feeding of
Neanura muscorum Templeton (Neanuri-
dae: Neanurinae) on two species of slime
moulds, Dictydiaethalium plumbeum Rost
and Trichia varia (Pers.). Chassain (1973)
also noted that when the slime mould trans-
forms to the sporulation phase, some
Collembola become trapped and die. A
number of other organisms, Coleoptera,
Diptera and Acari (Blackwell 1984;
Wheeler 1984; Lawrence and Milner 1996)
have been reported as forming an associa-
tion with slime moulds (Frederick 1990) but
rarely with the plasmodial stage since most
species feed on mature sporocarps.

The Uchidanurinae are nearly all heavily
pigmented and have eight ocelli on each
side of the head. This is suggestive of diur-
nal activity in these organisms as nocturnal
species normally have reduced ocelli and
reduced pigment of the body. However,
foraging during the day of cryptic, humid
log-inhabiting species might not be expect-
ed because these animals would be vulner-
able to the lower humidities found outside
the log habitat. It is significant that the
diurnal activity recorded here was at a time
when light intensity in these forests was
low and at sites where humidities were
particularly high. Observations have also
been made of a different species foraging
during the day on the surface of a rotting
log in rainforest at Cooloola National Park,
Queensland (P Greenslade pers. obs.).

We suggest that as all members of the sub-
family Uchidanurinae have very similar
mouthparts, they may be specialist feeders
on slime moulds. This would explain not
only the observations recorded here, but also
the difficulty of culturing these animals and

Vol. 119 (5) 2002

Contributions

also explain their preferred habitat under the
bark of rotting logs. Moreover, the literature
indicates that the feeding habitats of the
family Neanuridae are still unknown
because identifiable gut contents are not nor-
mally observed in mounted specimens, Our
observations, together with those of
Chassain (1973), indicate that it is possible
that other members of the family Neanuridae
may also show a preference for or feed
exclusively on slime moulds and perhaps
even more commonly on other Protozoa.

Acknowledgements

The authors would like to thank the anonymous
referee for the suggestions of improvements to
the manuscript.

References

Andren O and Schnurer J (1985) Barley straw decom-
position with varied levels of microbial grazing by
Folsomia fimetaria (L) (Collembola, Isotomidae),
Oecologia 68. 7-62.

Blackwell M (1984) Myxomycetes and their arthropod
associates. In Fungus-Insect relationships:
Perspectives ia Ecology and Evolution, pp, 67-90,
Eds Q Wheeler and M Blackwell. (Columbia
University Press: New York)

Chassain M (1973) Capture d'un insecte Collembola
par deux Myxomycetes. Documents mycologiques
Lille 8, 37-38.

Christiansen K (1964) Bionomies of Collembola
Annual Review of Entomology 9, 147-178,

Prederick L (1990) Phylum Plasmodial Slime Molds
Class Myxomycota, In Handbook of Protoctista! the
structure, cultivation, habitats and life eycles of the
eukaryotic microorganisms and their descendants
exclusive of animals, plants and fungi, A guide to the
algae, ciliates, Foraminifera, sporazou, water molds,
slime molds and the other protoetists, pp. 467-483. J.
Eds O Corliss, M Melkonian and DJ Chapman.
(Jones and Bartlett Publishers: Boston)

Greenslade P (1991) Notes on Australian
Uchidanurinae (Collembola: Neanuridae). In
Advances in Management and Conservation of Soil
Fauna, pp. 63-65, Eds GK Veeresh, D Rajagopal and
CA Viraktamath. (Oxford/IBH Publishing Co. Pvt.
Lid: New Delhi)

Greenslade P (1993) Collembola, In 7nsects o/
Australia, pp. 252-264. Ed C SIRO. (Melbourne
University Press: Melbourne)

Hopkins $ (1997) Biology of the Springtails (Insecta:
Collembola). (Oxford University Press: Oxford)

Ing B (1967) Myxomycetes as food for other organsisms.
Proceedings of the South London Entomological and
Natural History Society 1967, 18-23,

Lawrence JF and Milner RJ (1996) Associations
between arthropods and fungi. In Fungi of Australia
Volume 1B. Introduction -= Fungi in the Environment,
pp. 137-200. Eds K Mallett and € Greurinoyie.
(Australian Biological Resources Study: Canberra)

Rusek J (1998) Biodiversity of Collembola and their
functional role in the ecosystem. Biodiversity and
Conservation 7, 1207-1219,

Stephenson SL. and Stempen H (1994) Myxomycetes: A
Handbook of Slime Moulds. (Timber Press: Portland,
Oregon)

Wheeler Q (1984) Evolution of slime mold feeding in
lviodid beetles, In F'ungus-Insecr relationships
Perspectives in Ecology and Evolution, pp. 446-477.
Eds Q Wheeler and M Blackwell. (Columhia
University Press; New York)

223

Contributions

Butterfly Management Uses Pollard Walk

Bryan T Haywood' and Christopher J Wilson

Abstract

South Australia's first set of butterfly walks was conducted in the Penambol Conservation Park from
13 November 2000 to 25 April 2001, A simple survey technique devised by Pollard (1977) was used
to record the changes in abundance of common and rare butterflies during a weekly transect walk,
l'en butterfly species were recorded during the survey period with 4 species being regionally threat-
ened species. The survey technique has great relevance for the survey and management of threatened
species in many terrestial environments with the assistance of volunteers also important for its suc-
cess. The presence and absence of particular species in specific habitats can also be objectively used
as a guide to overall habitat health and condition. (The Victorian Naturalist 119 (3), 2002. 224-226)

Introduction

A simple survey technique to observe and
record the abundance in butterfly fauna was
devised by Pollard (1977) as a method for
assessing the changes in the abundance of
butterflies, The method was first developed
at Monks Wood Experimental Station (UK)
to enable a person (with relatively little
experience) to assess the changes in abun-
dance of butterflies in their locality.
Endersby (1994) and Brooks (1993) high-
lighted the use of the Pollard walk for
counting insects, especially dragonflies,
suggesting minor modifications to reflect
local conditions but overall finding the tech-
nique repeatable and very effective. Staff of
the Department for Environment and
Heritage (DEH) in South Australia, with the
assistance of local volunteers, want to fur-
ther highlight the suitability of the Pollard
walk as a tool for the management of butter-
fly species and as a component in assessing
remnant vegetation health.

The Pollard walk, ‘the first of its kind in
South Australia,’ was initiated in 2000/01
and is being used for surveying butterflies
in the Penambol Conservation Park, South
Australia, a 178 hectare reserve situated 25
km south-east of Mount Gambier in an area
more commonly known as Caroline Forest.

The aim of the survey was to:

* highlight the simplicity of monitoring a
site for butterflies using the Pollard walk;

* commence the first annual Butterfly walk
for South Australia;

' Department for Environment and Heritage, PO Box
1046, Mount Gambier, South Australia 5290

Email Haywood, Bryan(@saugoy.sa.gov.au

' Wexford Wildfowl Reserve, North Slob, Wexford,
Ireland, Email ewilson(@ealga,ie

224

*record the numbers and species of
Lepidoptera present in the Penambol
Conservation Park;

*encourage loeal
involved;

* use the information collected on butterfly
presence as a guide to the assessment of a
habitat's health and condition.

Fisher (1983) and Grund and Hunt (1997,
2000) have previously recorded the butter-
fly fauna in the Lower South East includ-
ing the Penambol Conservation Park area.
They determined the locality as one of
great potential for the discovery of various
species occuring in Victoria but rare in this
region. This is principally due to the fact
that the Lower South East of South
Australia is the westerly extension of habi-
tats (and the range of flora and fauna
species) occuring throughout south-eastern
Australia.

The habitat throughout the transect includ-
ed Messmate Stringybark Eucalyptus obli-
qua on deep sandy soil, Swamp Gum Æ.
ovata on red loam over limestone, open
cleared ground of Bracken Pteridium escu-
lentum with emergent E. obliqua over intro-
duced grasses and weeds, and Blackwood
Acacia melanoxylon over Mat Rush
Lomandra longifolia.

Method

The Pollard walk suitably should be
undertaken from Spring to Autumn each
year (Pollard 1977). As this was the first
year of the walk the authors began the sur-
vey from 13 November 2000 and ended on
25 April 2001. Each week during this peri-
od a walk was undertaken along a fixed
and segmented transect (between 1100 and

volunteers to be

'The Victorian Naturalist

1600 hours). Pollard (1977) stressed the
importance of undertaking a walk during
optimum conditions. The weather can have
considerable effect on the numbers of but-
terflies seen. Therefore before each survey
the temperature, approximate wind speed
(Beaufort Scale) and direction and cloud
cover (sunshine) were recorded. Any day
during the allocated week could be chosen
for the Pollard walk to allow for the best
conditions. Each walk was completed at a
slow pace and took on average 25 min.
The walk was divided into 7 sections with
each section reflecting a change in habitat.
All butterfly species seen were identified
and counted at a distance up to 10 m in
each of the seven sections. Any un-identi-
fiable butterflies were recorded as such
and/or caught and later identified using
Braby (2000) and/or Fisher (1978). Some
uncommon and rare species were taken as
specimens, mounted, labelled and stored at
the Mount Gambier DEH office inverte-
brate collection for future reference. Notes
were also taken relating to general obser-
vations, feeding plants and other fauna
sighted during each walk. All data record-
ed were entered into a spreadsheet and for-
warded to the Biological Survey and

Contributions

Research section of the Department for
Environment and Heritage.

The habitat was assessed using a habitat
assessment technique developed for the
Property Management Planning program
(Primary Industries South Australia 1997).
The assessment process was to answer a
number of simple questions as to the pres-
ence and/or absence of various natural fea-
tures of a habitat of that type and associat-
ed wildlife. The assessment included the
allocation of points for positive and nega-
tive features contained within the habitat.
For example the presence of understorey
plants especially grasses would receive 5
points and the presence of weeds would
receive 5 minus points and so on. The tally
at the end of the exercise would provide an
assessment of the habitat depending on the
score.

Results

In 2000/01 a total of 10 species was
recorded along the weekly walks (Table 1).
The most commonly recorded butterflies
were the Common Brown /Heferonympha
merope merope, Common Grass Blue
Zizina labradus labradus and the Klug’s
Xenica Geitoneura klugii. The graph

Table 1. Butterfly species recorded including conservation status for South Australia (South
Australian Butterflies 1998). SA conservation ratings: U, uncommon; R, rare.

Common Name Scientific Name

Vanessa itea

Vanessa kershaw
Dispar compacte
Signeta flammea

Australian Admiral
Australian Painted Lady
Barred Grass-skipper
Bright Shield-skipper
Cabbage White
Common Brown
Common Grass Blue
Klug's Xenica
Shouldered Brown
White-banded Grass-dart

Heteronympha n

Heteronympha p

Table 2. Habitat assessment for each section of the

Taractrocera papyria papyria

Pieris rapae rapae

Zizina labradus labradus
Geitoneura klugii

Conservation
Status

i
1 u
ta R
"rope merope
enelope penelope U

R

walk showing the presence of rare and common

species and total butterfly numbers recorded for each section during survey period.

Section Habitat Value Number of Number of Total
rare species common species numbers
1 High 3 5 517
2 Low | 5 86
3 Moderate 3 5 230
4 Low 2 6 93
5 Low 0 2 106
6 Moderate l 5 155
1 High 2 5 CE,

Vol. 119 (5) 2002

225

Contributions

160 "s
= = Common Brown
r
4 — Klup's Xenica
PA
E L —— Grass Blue
120 ' —————
k '
u m
£4 i r
8 z p` ` ry
v E T y ty k t
[P »
eg ' " ; "
ps9 80 G p `
2g P : d 1 ww *
ES " LI , ^, d
25 " p ' `
EN " A ' E
& 40 t i «
S ` ^'
E i
à
\
0
ET
c

Weekly dates of transect walks

Fig. 1. The abundance of the three most common species recorded throughout the Pollard walk dis-

playing the total for each week.

shows the abundance of these three species
each week during the walk (Fig. 1).

The sections of the walk that received a
low habitat score had fewer butterflies
and/or rare species recorded. Those com-
mon species recorded in these sections
were normally seen in open impoverished
habitats. The areas with moderate/high
habitat value recorded a greater diversity
and abundance including most species of
conservation significance (Table 2).

Conclusion

Initial findings indicated that data collected
using the Pollard walk could assist in identi-
fying variations in the presence of
rare/threatened populations of butterflies at a
site. The long term trends of any monitoring
program are critical to the actions undertak-
en for rare/threatened species especially if
the technique for the collection of data can
be simplified. Pollard (1977) offers such a
technique for scientific authorities and. pro-
fessionals, amateur entomologists or nature
enthusiasts. The contribution and assistance
offered by volunteers is also a critical com-
ponent to the success of a wildlife monitor-
ing program like the Pollard walk.

The presence of rare species in some sec-
tions of the walk indicates that the habitat is
of moderate/high biodiversity value. The
areas of the walk in which few butterfly
species were found could be an indication
that the habitat is impoverished and needs
attention, However with only one year's
data this conclusion cannot be substantiated,

226

Acknowledgements

The authors wish to thank the following individ-
uals who assisted or participated in weekly
walks, identification of butterflies, signage
along the trail, maps for a Park brochure and
slashing of walking trail: Jason Bonney, Katrina
Eggleston, Zoe Griffiths, Roger Grund, Torsten
Gustavsson, Jean Haywood, Toni Haywood,
Brad Lay, Paul Steed, Greg Wandin, and Ann
Wilson.

References and Bibliography

Braby MF (2000) Butterflies of Australia, Volumes |
and 2. (CSIRO: Melbourne)

Brooks SJ (1993) Review of a method to monitor adult
dragonfly populations. Journal of the British
Dragonfly Society 9, 1-4.

Endersby ID (1994) Counting insects while doing the
Pollard Walk. Victorian Entomologist 24, 134-135.

Fisher RIT (1978) Butterflies of South Australia
Lepidoptera: Hesperioidea, Papilionoidea,
(Government Printer; Adelaide)

Fisher RH (1983) Butterflies of the South East. In
Natural History of the South East, pp 215-222. Eds
MJ Tyler, CR Twidale, JK Ling and JW Holmes.
(Royal Society of South Australia: Adelaide)

Grund R and Hunt L (1997) New butterfly records and
hew range extensions, food plant recordings and biol-
ogy for some butterflies from the lower south east
region of South Australia, Victorian Entomologist 27,
50-33,

Grund R and Hunt L (2000) Butterfly Conservation in
the Lower South East Region. (National Parks
Foundation Inc: Adelaide)

Pollard E (1977) A method for assessing changes in the
abundance of butterflies. Biological Conservation 12,
115-134.

Primary Industries South Australia (1997) How healthy
is your patch of native vegetation — worksheet. In
Property Management Planning — Extension
Resource Manual. (PISA; Adelaide).

South Australian Butterflies website (1998).
www.chariot.net.au/-rgrund

'The Victorian Naturalist

Contributions

First Record of the Asian Freshwater Leech Barbronia weberi
(Blanchard, 1897) (Euhirudinea: Erpobdellidae) in Australia

Fredric R Govedich', Bonnie A Bain' and Ronald W Davies"

Abstract

Two individual Barbronia weberi, an invasive species from India and southeast Asia, were collected
from Common Waterweed Hydrilla verticillata purchased in June, 2001 from an aquarium supplier
in Melbourne, Australia. Adult leeches were fed Tubifex sp. ad libitum and successtully reproduced
under laboratory conditions (21°C). This invasive species has the potential to adversely affect the
biodiversity of native Australian freshwater invertebrates and the extent of this invasion is not clear-
ly known. (The Victorian Naturalist 119 (5), 2002, 227-228)

Introduction

Members of the family Erpobdellidae are
typically freshwater predators feeding on a
range of invertebrate prey including crus-
taceans, insects, molluscs, and annelids.
Unlike their bloodsucking cousins, these
leeches lack both a proboscis and jaws and
often feed by swallowing their prey whole.
Erpobdellids are well known for being
important sit-and-wait predators in aquatic
ecosystems, with a few species being
pelagic predators in the absence of fish
(Blinn et al, 1987). In addition, erpobdel-
lids may be important prey for vertebrate
and invertebrate predators (Sawyer 1986;
Davies and Govedich 2001).

Barbronia weberi (Blanchard, 1897) was
originally described from India and is native
to south-east Asia, Java, Sumatra, Celebes,
Borneo, Philippines and Afghanistan
(Sawyer 1986; Sods 1966; Nesemann and
Sharma 1996). It has been introduced to
New Zealand, England, Brazil, and the
United States of America, apparently
through the aquarium trade (Mason 1976;
Sawyer 1986; Pamplin and Rocha 2000;
Rutter and Klemm 2001). Barbronia weberi
is ofien associated with aquatic vegetation,
using aquatic plants as a substrate for
attachment and for the attachment of
cocoons, and it is likely this species has
been transported with common aquarium
plants, Hydrilla verticillata (Linnaeus,
1782) and Elodea sp.

Methods
Two reproductively mature leeches were
found on Hydrilla verticillata purchased

School of Biological Sciences, Monash University,
Clayton, Victoria 3800

Department of Biological Sciences, University of
Calgary, 2500 University Drive NW, Calgary, Alberta,
Canada TIN 1N4

Vol. 119 (5) 2002

(June, 2001) from an aquarium supplier in
Melbourne, Australia. The adult leeches
were fed Tubifex sp. ad libitum and main-
tained in 200 ml of artificial pond water (pH
= 6.96, conductivity = 31.6 uS, 21°C) for 3
months. Following the production of
cocoons the adults were preserved in 7096
ethanol and examined using an Olympus
SZH-10 dissecting microscope to determine
their identity.

Results and Discussion

Barbronia weberi is distinguishable from
two superficially similar Australian erpob-
dellids, Vivabdella arcana Richardson,
1970 and Dineta cylindrica Goddard, 1908.
Both B. weberi and V. arcana have three
pairs of eyes and accessory copulatory
pores (Fig. 1d) distinguishing them from D,
cylindrica which has four pairs of eyes and
lacks accessory pores (Fig. 1c). Barbronia
weberi can be distinguished from F. arcana
(Fig. la, f) based on the presence of pha-
ryngeal stylets (Fig. 1b) and the absence of
simple postcaeca (Fig. le).

Each adult Barbronia weberi produced
between 8 and 10 cocoons during a two
week period and each cocoon was attached
to solid substrates (aquatic vegetation,
rocks or the sides of aquaria). Cocoons
were produced every other day over the
two weeks and each cocoon contained two
or three small eggs surrounded by irans-
parent fluid. Upon hatching the young
were 5.6 + 0.6 mm long and 0.7 + 0.2 mm
wide (relaxed), capable of feeding inde-
pendently on Tubifex sp. and became
reproductively mature within four months
of hatching (n = 7). At reproductive matu-
rity, leeches had a relaxed length of 21.5 +
3 mm and width of 2.3 £0.3 mm (n = II).

227

Contributions

Adult Barbronia weberi were found
attached to Hydrilla verticillata, a weedy
aquatic plant that has been known to
invade freshwater ecosystems. The leeches
were usually attached near the base of the
leaf whorls or along the stem, making
them difficult to see. While most of the
individual cocoons were attached to the

bottom or sides of the container, some of

the cocoons were laid on individual leaves.
All cocoons, regardless of attachment site,

eventually hatched. Both the behaviour of

the parent and the attachment of cocoons
to H, verticillata allow this species to be
transported as both an adult and as a
cocoon. This close association, at both the
cocoon and adult stages, has allowed B.
weberi to successfully invade many new
areas via the aquarium trade (Mason 1976;
Sawyer 1986; Pamplin and Rocha 2000;
Rutter and Klemm 2001).

The H. verticillata which contained the B.
weberi had been part ofa shipment that orig-
inated from a pool of locally (Melbourne)
and nationally grown plants making it diffi-
cult to identify the source of the B. weberi.
However, all stocks of the Æ. verticillata
were from Australian sources, indicating
that B. weberi has been and is being main-
tained in the Australian aquarium trade.
When placed into an aquarium with a gravel
base, individual leeches were observed to
burrow into the gravel, making them virtual-
ly undetectable, Like other erpobdellid
leeches, B. weberi is an aquatie predator that
swallows its invertebrate prey (insects,
worms, etc.) whole. Being a predator with
rapid development and the ability for adults
and cocoons to be transported in aquatic
vegetation, allows B. weberi to invade new
systems where they have the potential to
compete with or feed on native invertebrate
species. The introduction of this invasive
species could potentially have long term
consequences for the biodiversity and sur-
vival of native Australian freshwater fauna
and the ecosystems they inhabit.
Unfortunately it is not clear how many B.
weber? are in Australia, or the extent of the
invasion of Australian aquatic systems, and
further studies need to be completed to
determine the extent and effect this species
is having in Australia.

228

out

1,25mm.

ti crop

[]
b
=

postegeen

dri some

d ana
n t

Fig. 1. Morphological features of Barbronia
weberi and Vivabdella arcana, Top left:
Dissected view of the buccal cavity of V.
arcana (a), without stylets and B, weberi (b)
containing stylets. Bottom left; External view
showing the gonopores of a typical erpobdellid
(c) and the gonopores and accessory copulatory
pores of B. weberi and V. arcana (d). Right:
Digestive system of B. weberi (e) without post-
caeca and 7. arcana (V) with posteaeca.

References

Blim DW, Davies RW and Dehdashti B (1987)
Specialized pelagic feeding by Erpobdella montezu-
ma (Hirudinea). Holaretic Ecology 1, 235-240.

Davies RW and Govedich FR (2001) Annelida:
Euhirudinea and aeanthobdellids, In Ecology and
Systematics of North American Freshwater
Inverfebrates, 2nd ed, pp 465-504. Ed JH Thorp and
AP Covich. (Aeademic Press: San Diego)

Mason J (1976) Studies on the freshwater and terrestrial
leeches of New Zealand, 2. Orders Gnathobdelliformes
and Pharyngobdelliformes. Journal of the Royal
Society of New Zealand 6, 255-276.

Nesemann H and Sharma S (1996) Contribution to the
knowledge of the leeches of Nepal (Annelida:
Hirudinea). deta Zoologica Academiae Scientiarum
Hungaricae A42, 231-249,

Pamplin P and Rocha O (2000) First report of
Barbronia weberi (Hirudinea: Erpobdelliformes:
Salifidae) from South America, Revista de Biologia
Tropical 48, 723.

Rutter RP and Klemm DJ (2001) The presence of an
Asian leech, Barbronia weberi, in à home aquarium
in south Florida (Hirudinea: Salilidae). /"lorida
Scientist 64, 216-218,

Sawyer RT (1986) Leech Biology and Behaviour
Feeding Biology, Ecology and Systematics, Volume
IL. (Oxford University Press: Oxford)

Soós A (1966) Identification key to the leech
(Hirudinoidea) genera of the world with a catalogue
of the species. I1, Family: Erpobdellidae, Acta
Zoologica Academiae Scientiarum Hungaricae 12,
371-407.

The Victorian Naturalist

Contributions

Dragonfly Nymphs with Dipteran *Hitch-hikers': an Example
of Phoresy Found in Dandenong Creek

Anneke Veenstra-Quah'

Abstract

A phoretic association was observed between larvae of Rheotanytarsus juliae (Diptera:
Chironomidae) and larvae of Austroaeschmna unicornis unicornis (Odonata: Aeshnidae) collected
from one site near the headwaters of Dandenong Creek, Victoria. (The Vierorian Naturalist 119 (5), 2002,

229-231)

Introduction

Phoresy, from the Greek phoresis, ‘a car-
rying’, is a phenomenon where an individual
of one species is transported by an individ-
ual of another species (Gullan and Cranston
1994). Phoresy has been observed in fresh-
water ecosystems where small or slow-mov-
ing animals rely on larger more mobile
species for transport. There have been
numerous overseas reports of dipteran lar-
vae, particularly non-biting midges from the
Family Chironomidae, attaching themselves
to several insect orders including
Ephemeroptera (White et al. 1980; Tokeshi
1986), Plecoptera (Bottorff and Knight
1987), and Odonata (Rosenberg 1972;
White ef al. 1980; Dudgeon 1989), Hawking
and Watson (1990) reported the first associ-
ation between larvae from the chironomid
genus Rheotanytarsus and the dragonfly
Austroaeschna atrata in Australian waters.

Chironomid larvae are popular prey items
for many predators, and increasing their
effective size by attaching to a larger organ-
ism like a dragonfly nymph can decrease the

potential number of predators capable of

feeding on them (Tokeshi 1995). A phoretic
association has now been observed between
Rheotanytarsus juliae (Diptera: Chirono-
midae) and 4ustroaeschna unicornis unicor-
nis (Odonata: Aeshnidae) collected from the
headwaters of Dandenong Creek,
Rheotanytarsus is a widespread, tube-
building genus found in flowing water
(Cranston 1996). Their distinctive non-
transportable cases constructed of detrital
and algal material, have arm-like extensions
on which the larva spins a silken web for
filtering fine food particles carried by the

Faculty of Science and Techonology, Deakin
University, Burwood Campus 221 Burwood Highway,
Burwood, Victoria 3125. Email yquah@ideakin.cdu.au

Vol. 119 (5) 2002

current, Rheotanytarsus juliae is the most
abundant species in cool temperate regions
of south-eastern Australia (Cranston 1997),

Austroaeschna unicornis unicornis
nymphs are ambush predators, inhabiting
mud or gravel substrates, crevices in rocks
and logs, in both montane and lowland
streams in south-eastern Australia
(Hawking 1986). dustroaeschna nymphs
have slow growth rates, hard cuticles with
many sites suitable for attachment, and they
remain stationary for long periods in mud
and detritus, habitats favoured by chirono-
mids (Hawking and Watson 1990).

Methods

The headwaters of Dandenong Creek
upstream of Edgar Track in Doongalla
State Forest 37° 51' S, 145?20' E were sam-
pled using the rapid bioassessment method
of Tiller and Metzling (1998) on 30 August
2001. At this site, the streambed was com-
prised of small boulders and cobbles with
large amounts of detritus present. Tree
ferns and eucalypts dominated the riparian
zone vegetation (Fig. 1).

Invertebrates were collected by kick sam-
pling for five minutes using a collecting
net with mesh size 250 um, live sorted in
the field, preserved in 70% ethanol, and
initially identified to family level in the
laboratory using published keys. Dragonfly
nymphs were later identified to species
level using Hawking (1986) and chirono-
mids to species level using Cranston
(1996, 1997).

Results

Of the five 4. unicornis unicornis nymphs
collected from this site on Dandenong
Creek, two had larval R. juliae attached,
Two chironomid larvae were associated with
the largest dragonfly nymph — one attached

229

Contributions

Doongalla
.,- .. State Forest
cd T J

j A x
2 |
ous J
I! f

2>

WC sampling
: site
| |
Dandenong”
Creek
T
1 ap
Port Phillip \ d

Bip” AL. (Carrum

Fig. 1. Location and photograph of the sampling
site near the headwaters of Dandenong Creek
upstream of Edgar Track in Doongalla State
Forest.

horizontally to the tibia of the hind leg and
the other near the wing pad margin (Fig. 2).
The other nymph had a single case attached
laterally to the first abdominal segment.

Discussion

The phoretic association between aeshnid
dragonfly nymphs and chironomid larvae
is thought to be quite common (JH
Hawking 2001 pers. comm.). The headwa-
ters of Dandenong Creek have been sam-
pled since the late 1970s (Campbell 1978;
Nuttall 1982: Ferdinands er al. 1995) and
dragonfly nymphs from the family
Aeshnidae have often been collected in the
vicinity of the site sampled in August
2001. However, there are no published
records of chironomids attached to aeshnid
nymphs. This may be because the associa-
tion was not noticed when samples were
sorted due to the fact that the nymphs are
dark brown in colour, often covered in
detritus and the brown Rheotanytarsus
cases may be easily overlooked.

It is interesting to note that of the 41 A.
atrata nymphs collected by Hawking and
Watson (1990) from two sites in the upper
catchment of the Kiewa River, only two had
Rheotanytarsus attached, compared with five
A. unicornis unicornis nymphs collected
from the headwaters of Dandenong Creek
and two of which had Rheotanytarsus juliae
attached.

230

Fig. 2. Two Rheotanytarsus juliae larvae asso-
ciated with Ausiroaeschna unicornis unicornis
the largest dragonfly nymph collected - one
attached horizontally to the tibia of the hind leg
(black arrow and inset) and the other near the
wing pad margin (white arrow).

Acknowledgements

| would like to thank Helen Madden for the
excellent dragonfly nymph portrait, Suzanne
Kiss von Soly, for her kick-sampling expertise,
Rebecca Joyce, an observant student who
noticed that there was something unusual about
one dragonfly nymph, and other students from
the School of Ecology and Environment
(Deakin University) Freshwater Biology class —
Louise Biggs, Katie Christopher, Rebecca
Pretty, Jacqueline Rosewall and Haylee Weaver,

References

Bottorff RL and Knight AW (1987) Ectosymbiosis
between Nanocladius downesi (Diptera:
Chironomidae) and Acroneuria abnormis
(Plecoptera: Perlidae) in a Michigan stream, USA.
Entomologia Generalis 12, 97-113.

Campbell IC (1978) A biological investigation of an
organically polluted urban stream in Victoria.
Australian Journal of Marine and Freshwater
Research 29, 275-291.

Cranston PS (1996) Identification Guide to the
Chironomidae of New South Wales. AWT
Identification Guide Number 1, (Australian Water
Technologies Pty Ltd: West Ryde, NSW)

Cranston PS (1997) Revision of Australian
Rheotanytarsus Thienemann & Bause (Diptera:
Chironomidae) revised, with emphasis on the imma-
ture stages. Invertebrate Taxonomy 11, 705-734.

Dudgeon D (1989) Phoretic Diptera (Nematocera) on
Zygonyix iris (Odonata: Anisoptera) from a Hong
Kong river: incidence, composition and attachment
sites. Archiv für Hydrobiologie 115, 433-439.

Ferdinands K, Vertessy D and Smith T (1995)
Biological Survey of Dandenong Creek and
Associated Tributaries, Report No 8/95, (Water
Ecoscience: Melbourne)

Gullan PJ and Cranston PS (1994) The Insects: An
Outline of Entomology, (Chapman and Hall: London)

The Victorian Naturalist

Hawking JH (1986) Dragonfly Larvae of the River
Murray System: a Preliminary Guide to the
Identification of Known Final Instar Larvae of
South-eastern Australia. Technical Report No 6,
Albury-Wodonga Development Corporation:
Albury,

Hawking JIL and Watson JAL (1990) First Australian
record of chironomid larvae epizoic on larval
Odonata. Aquatic Insects 12, 214-245.

Nuttall PM (1982) 4 Biological Assessment af Water
Quality in Dandenong Creek and its Major
Tributaries, Victoria. (Dandenong Valley Authority:
Dandenong)

Rosenberg D (1972) A chironomid (Diptera) larva
attached to a libellulid (Odonata) larva. Quaestiones
Entomologicue 8, 326-336.

Contributions

Tiller D and Metzling L (1998) Rapid Bioussessment of
Victorian Streams: The Approach and Methods of the
Environment Protection Authority. Publication 604.
(Environment Protection Authority: Melbourne)

Tokeshi M (1986) Population ecology of the commensal
chironomid Epoicocladius flavens on the may fly host
Ephemera danica. Freshwater Biology 16, 23 243.

Tokeshi M (1995) Species interactions and community
structure. In The Chironomidae: the Biology and
Ecology of Non-biting Midges, pp 297-335. Eds P
Armitage. PS Cranston and LCV Pinder (Chapman
and Hall; London)

White TR, Weaver JS Hl and Fox RC (1980) Phoretic
relationships hetween Chironomidae (Diptera) and
benthic macroinvertebrates. Entomological News 91,
69-74.

Acacia obtusifolia — Introduction and Spread in Native Bush

David Cheal'

Abstract

Acacia obtusifolia has reproduced and expanded from a single introduction in the upper Yarra Valley
in native bush as a result of inadvertent introduction of seed on earth-moving equipment. Initial estab-
lishment was dependent on soil disturbance, but subsequent spread has been episodic and dependent
on fire-promoted germination and establishment. There is no evidence of seed germination and estab-
lishment in the absence of fires, The next fire may lead to a dramatic increase in both the number of
A. obtusifolia plants and their local distribution. (The Victorian Naturalist 119 (5), 2002, 231-232)

Many native species have spread from
their former habitats following European
settlement. In south-eastern Australia some
of the most troublesome include Acacia
longifolia (Jameson 1994), Leptospermum
laevigatum (Burrell 1981; Molnar ef al.
1989; Offor 1990) and Pittosporum undu-
latum (Gleadow and Ashton 1981;
Gleadow 1982), but many others have
been occasionally recorded as weeds or as
the product of recent human-assisted
spread (see Walsh and Entwisle 1994;
Walsh and Entwisle 1996: Low 1999;
Walsh and Entwisle 1999).

Acacia obtusifolia is a tall wattle character-
istically found on the slopes and low ridges
of drier East Gippsland forests (Walsh and
Entwisle 1996). However, the Flora

Information System of the Department of

Natural Resources and Environment,
aise tel has an isolated record from the

Upper Yarra Valley, much closer to
Melbourne and grpeor innately 160 km from
the nearest Gippsland record (see Walsh and
Entwisle 1996: 597). This small stand is an
example of recent, human-assisted spread

Arthur Rylah Institute for Environmental Research,
123 Brown Street, Heidelberg, Victoria 3084

Vol. 119 (5) 2002

rather than a natural outlier. The site is at
5511CU975628 on the 1:100000 Healesville
Mapsheet.

In 1977 a bulldozer was hired by local
residents to make an access track along à
ridge approximately | km north-east of
MeMahons Creek and through otherwise
undisturbed forest typical of the region
(Eucalyptus obliqualEucalyptus radiata
dives/scattered Eucalyptus sieberi, above
shrubs including Acacia mucronata,
Banksia spinulosa, Hakea decurrens,
Kunzea ericoides and Leptospermum conti-
nentale; nomenclature follows Walsh and
Entwisle 1996), The bulldozer had been
most recently used in fire prevention works
near Bruthen in East Gippsland and was
brought to the Upper Yarra Valley on the
tray of a large lorry.

In the first year after the track was con-
structed an unusual wattle was noted grow-
ing in the disturbed earth adjacent to the
track. That single shrub turned out to be an
Acacia obtusifolia (Figs | and 2). In subse-
quent years it grew to approximately 3-4 m
tall, flowered and set seed. Despite annual
searches over the next 4 to 5 years, no other
A. obtusifolia was found anywhere in the

231

Contributions

Fig. 1. Acacia obtusifolia at MeMahons Creek,
February 2001.

region (the seed that had been dropped
from this one specimen did not germinate
and establish in those first few years).

The ridge, including this one wattle shrub
(by now a small tree), was burnt in the Ash
Wednesday fires of March 1983. The fire
was of very high intensity (no nearby
unburnt refuges and all above-ground veg-
etation burnt). However, in the following
years regeneration was prolific and some
formerly less common shrubs increased in
abundance (notably B. spinulosa and E.
sieberi). The single A. obtusifolia present
before the fire was killed, but a few
seedlings established nearby after the fire.

In February 2001 the site was revisited.
There were now 14 A. obtusifolia individu-
als, all growing within 4 m of the original
(now dead) plant. The tallest was over 4 m
tall and most of them had flowered. A few
retained healthy seed in pods from the pre-
vious season (seed viability was not test-
ed). There were many Acacia seeds in the
soil around this small stand of 4, obtusifo-
lia. As A. mucronata was also common at
the site, it is not known what proportion of
these seeds were A. obtusifolia. The next
fire may lead to a dramatic increase in both

232

Bl" p as 3
Fig. 2. A closer view of Acacia obtusijolia at
McMahons Creek.

the number of 4. obtusifolia plants and
their local distribution.

References

Burrell JP (1981) Invasion of Coastal Heaths of
Victoria by Leptospermum laevigatum (J. Gaertn.) F.
Muell. 4ustralian Journal of Botany 29, 747-764.

Gleadow RM (1982) Invasion by Pittosporum undula-
tum of the forests of Central Victoria IH. Dispersal,
germination and Establishment. Australian Journal
of Botany 30, 185-198.

Gleadow RM and Ashton DH (1981) Invasion by
Pittosporum undulatum of the forests of Central
Victoria |. Invasion patterns and plant morphology.
Australian Journal of Botany 29, 705-720.

Jameson G (1994) A case of your Longyfolia. The
Victorian Naturalist 141, 145-150.

Low T (1999) Feral future. (Viking: Ringwood)

Molnar CD, Fletcher D and Parsons RF (1989)
Relationships between heath and Leptospermum lae-
vigatum serub at Sandringham, Victoria, Proceedings
of the Royal Society of Victoria 101, 77-88,

Offor T (1990) What future for the sandy heaths of
Wilson's Promontory. The Victorian Naturalist 107,
120-123.

Walsh NG and Entwisle TJ (1994) Flora of Victoria
Volume 2. Ferns and Allied Plants, Conifers and
Monacotyledons, (Inkata Press: Melbourne)

Walsh NG and Entwisle TJ (1996) Flora of Victoria
Volume 3. Dicotyledons Winteraceae to Myrtaceae.
(Inkata Press; Melbourne)

Walsh NG and Entwisle TJ (1999) Flora of Victoria
Volume 4. Dicotvledons Cornaceae to Asteraceae.
(Butterworth-Heinemann: Port Melbourne)

The Victorian Naturalist

Contributions

The Discovery of Leadbeater’s Possum Gymnobelideus lead-
beateri along the Woori Yallock Creek, Yellingbo

Dan Harley'

Abstract

A single Leadbeater's Possum Gyrminobelideus leadbeateri was discovered denning in a nest box sit-
uated along the Woori Yallock Creek, Yellingbo Nature Conservation Reserve, This is the first
record of the species from this section of the Reserve, which is more than a kilometre from other
sites at which the possum occurs at Yellingbo. It is likely that the individual dispersed from the
Cockatoo Creek across unfavourable habitat. (The Vietorian Naturalist 119 (5), 2002, 233-235)

In 1986, a small, outlying population of
the endangered Leadbeater's Possum
Gymnobelideus leadbeateri was discov-
ered inhabiting lowland swamp forest at
Yellingbo Nature Conservation Reserve
(Smales 1994). Significantly, it is the only
known extant lowland population,
although there are historical records of the
species in similar habitats from low eleva-
lions in south-west Gippsland (McCoy
1867; Brazenor 1932, 1946).

Yellingbo is located approximately 50 km
east of Melbourne (37° 47' S, 145° 32' E).
The reserve encompasses 591 ha of remnant
riparian forest and comprises three separate
sections that span stretches of four water-
courses, the Woori Yallock, Sheepstation,
Cockatoo and Macclesfield Creeks. The
nearest records of Leadbeater's Possum to
Yellingbo are from Ben Cairn, approxi-
mately 17 km to the north-east (Owen 1963;
Lindenmayer er al. 1989), and Mt Beenak,
approximately 18 km to the east (Loyn and
MeNabb 1982), Mountain Ash Eucalyptus
regnans forest, typical of that inhabited by
the species throughout its core range in the
Victorian Central Highlands, occurs at both
of these sites.

Since the rediscovery of Leadbeater's
Possum in 1961 (Wilkinson 1961), most
records have come from montane ash forest
above 600 m ASL. (Lindenmayer et al.
1991). Yellingbo is situated at 120 m Ast.
The sites inhabited by the species at
Yellingbo are dominated by Mountain
Swamp Gum Eucalyptus camphora with a
dense middlestorey in places, comprising
Woolly Tea-tree Leptospermum lanigerum,
Prickly Tea-tree L. juniperinum or Scented
School of Biological Sciences, PO Box 18, Monash
University, Victoria 3800.

Email dkpharley@hotmail.com

Vol. 119 (5) 2002

Paperbark Me/aleuca squarrosa (Harley
unpubl. data). Connecting vegetation in
either the canopy or middlestorey is an
important attribute of Leadbeater's Possum
habitat, as the species rarely descends to the
ground (Harley unpubl. data). Notably, aca-
cias such as Silver Wattle Acacia dealbata, a
key food source for the possum in montane
ash forest (Smith 1984), are absent from the
sites utilised by Leadbeater's Possum for
foraging at Yellingbo (Harley unpubl. data).

In order to study the ecology of
Leadbeater's Possum in lowland swamp
forest, 150 nest boxes were installed at
Yellingbo between 1995 and 2000. Coupled
with the results of stagwatching surveys, the
nest box results suggested that the
Yellingbo population of Leadbeater's
Possum is restricted to a single section of
the reserve along the Cockatoo and
Macclesfield Creeks (Harley unpubl. data).
The possum's distribution there extends
over a long, narrow corridor of forest,
approximately 6 km long and typically less
than 200 m wide. In contrast, Lindenmayer
et al. (1993) detected the species at only one
of 49 linear corridors surveyed in montane
ash forest. Many of these sites contained
habitat considered to be suitable for the
species, leading the authors to conclude that
the possum does not utilise linear corridors
of forest in montane ash forest.

As part of the survey to establish the dis-
tribution of Leadbeater's Possum at
Yellingbo, five nest boxes were installed
along the Woori Yallock Creek in
February 1999, They were positioned on
eucalypts at a height of 3.5 m, with a
south-easterly aspect. Each nest box had
internal dimensions of 236 x 195 x 356
mm, 19 mm thick walls and an entrance
hole of 50 mm diameter. They were locat-

233

Contributions

ed at sites with habitat most resembling
that occupied by Leadbeater's Possum
along the Cockatoo and Macclesfield
Creeks (e.g. presence of & camphora with
a tea-tree middlestorey). The five nest
boxes were inspected in November 1999,
eight months after installation, at which
lime there was no sign of use by
Leadbeater's Possum,

During the course of monitoring the
Yellingbo nest boxes in April 2001, a sin-
gle Leadbeater's Possum was found den-
ning in one of the nest boxes located along
the Woori Yallock Creek. This is the first
record of the species along this creek, The
animal was an adult male weighing 121 g
and it had built a nest of shredded bark
inside the nest box. It is unclear how long
the individual had been resident at the site.
No evidence of the species (e.g. nesting
material) was found in the other four nest
boxes positioned along the Woori Yallock
Creek, all of which occur to the north of
where the possum was discovered.

On 29 December 2001, one hour after
dusk, the individual was observed approxi-
mately 30 m north of the nest box in which
it was first discovered, eight months earlier.
It was detected through imitation of
Leadbeater’s Possum calls and approached
to within one metre, This observation indi-
cates that the individual was still resident at
the site, presumably having established its
territory in the immediate area.

The riparian forest along the Woori
Yallock Creek is dominated by Manna
Gum Æ. viminalis, with an understorey of
Blackwood A. me/anoxylon and pockets of
Prickly Tea-tree. Much of the forest is
quite open in structure. However, there are
a number of small, swampy areas along the
creek, where the vegetation more closely
resembles that found along the Cockatoo
and Macclesfield Creeks. The habitat at the
site where the possum was discovered is
such an area, The canopy is dominated by
Mountain Swamp Gum with an under-
storey of Woolly Tea-tree.

More than a kilometre of partially cleared
farmland separates the Cockatoo and
Macclesfield Creeks from the site where
the possum was discovered on the Woori
Yallock Creek. The habitat that remains
between the two sites is not suitable for the
species, as it consists of a ridge dominated

234

by Messmate E. obliqua, and is open in
structure. It is very different to the flood-
plaín forest inhabited by Leadbeater's
Possum along the Cockatoo and
Macclesfield Creeks.

Two likely scenarios could explain the ori-
gin of the animal found on the Woori
Yallock Creek. Either it had dispersed from
Cockatoo Creek or a small, remnant popula-
tion of Leadbeater’s Possum exists along the
Woori Yallock Creek. The dispersal sce-
nario would require the possum to have trav-
elled more than 1100 m through partially
cleared farmland. While seven dispersal
events of this magnitude (i.e. 870-1120 m)
have been detected at Yellingbo, the average
dispersal distances for Leadbeater's
Possums through favourable habitat along
the Cockatoo and Macclesfield Creeks are
480 m for males and 450 m for females
(Harley unpubl. data).

At sites with few tree hollows, nest boxes
can be a valuable survey technique for
cryptic, arboreal mammals (Traill and
Coates 1993; Soderquist et al. 1996; Ward
2000). The discovery of a Leadbeater's
Possum along the Woori Yallock Creek
highlights their value as a long-term sur-
vey technique. One of the main benefits of
nest boxes is that they can be left on site
for extended periods of time, in marked
contrast to the short duration of trapping
surveys. They are particularly valuable in
surveying for Leadbeater's Possums,
because the species’ presence in an area
can be established from the possums’ dis-
tinctive nesting material, with no necessity
for the animals to be denning in the nest
box on the day of inspection.

Between 1995 and 2001, twenty-seven
different Leadbeater's Possum colonies
deposited nesting material in [11 of the
131 nest boxes (85%) installed along the
Cockatoo and Macclesfield Creeks (Harley
unpubl, data). This result suggests that, at
Yellingbo, if Leadbeater’s Possums are
resident at a site, there is a strong likeli-
hood that they will deposit nesting material
in nest boxes located within their territory.
Thus, the absence of nesting material from
the four northernmost nest boxes, located
along the Woori Yallock Creek, suggests
that the species probably does not occur in
this part of the reserve. However, it is pos-
sible that more individuals occur in the

The Victorian Naturalist

immediate vicinity of the southernmost
nest box that contained the possum, as an
area of suitable habitat occurs here on pri-
vate property adjacent to the reserve.
Further surveys will be conducted along the
Woori Yallock Creek to determine which
of the two possible scenarios is correct.

Acknowledgements

Nest boxes were purchased with funds from the
Holsworth Wildlife Research Fund and
Australian Geographic. Barry Traill and
Malcolm Macfarlane kindly donated nest boxes
to the study. Financial and field support for the
2001 nest box monitoring was provided by
Parks Victoria (Woori Yallock). | would partic-
ularly like to thank lan Roche and Joanne
Antrobus for their support of the project, and the
Parks Victoria staff that assisted in the field. My
thanks also go to Terry Harley, Marianne
Worley and Melinda Collinson, all of whom
assisted with the installation and inspection of
the Yellingbo nest boxes. Marianne Worley,
Alan Lill and an anonymous reviewer provided
valuable comments on the manuscript. The pro-
ject was conducted under Victorian Department
of Natural Resources and Environment Wildlife
Permit Number 10001246.

References

Brazenor CW (1932) A re-examination of
Gymnobelideus leudbeateri McCoy. Australian
Zaologist 7, 106-109,

Brazenor CW (1946) Last chapter to come: a history of
Victoria's rarest possum. Mild Life 8, 382-384.

Lindenmayer DB, Smith AP, Craig SA and Lumsden
LF (1989) A survey of the distribution of
Leadbeater's Possum, Gyminobelideus leadbeateri
McCoy in the Central Highlands of Victoria. The

Contributions

Victorian Naturalist 106, 174-178.

Lindenmayer DB, Cunningham RB, Tanton MT, Nix
HA and Smith AP (1991) The conservation of arbo-
real marsupials in ihe Montane Ash Forests of the
Central Highlands of Victoria, South-East Australia:
IIT. The habitat requirements of Leadbeater’s Possum
Gymnobelideus leadbeateri and models of the diver-
sity and abundance of arboreal marsupials. Biological
Conservation 86, 295-315.

Lindenmayer DB. Cunningham, RB and Donnelly, CF
(1993) The conservation of arboreal marsupials in the
Montane Ash Forests of the Central Highlands of
Victoria, South-East Australia, IV. The presence and
abundance of arboreal marsupials in retained linear
habitats (wildlife corridors) within logged forest.
Biological Conservation 66, 207-221.

Loyn RH and MeNabb EG (1982) Discovery of
Leadbeater’s Possum in Gembrook State Forest. The
Victorian Naturalist 99, 21-23,

McCoy F (1867) On a new genus of Phalanger. Annals
and Magazine of Natural History 3, 287-288.

Owen WH (1963) Further sight records of Leadbeater's
Possum. The Victorian Naturalist 79, 292-293.

Smales IJ (1994) The discovery of Leadbeater’s
Possum, Gymnobelideus leadbeateri McCoy, resi-
dent in lowland swamp woodland. The Victorian
Naturalist 111, 178-182.

Smith A (1984) Diet of Leadbeaters Possum,
Gymnobelicdeus leadbeateri (Marsupialia). Australian
Wildlife Research 11, 265-273.

Soderquist TR, Traill BJ, Faris F and Beasley K (1996)
Using nest boxes to survey for the Brush-tailed
Phascogale Phascogale tapoatafa. The Victorian
Naturalist 113, 256-261,

Traill BJ and Coates TD (1993) Field observations on
the Brush-tailed Phascogale Phascogale rapoatafa
(Marsupialia: Dasyuridae). Australian Mammalogy
16, 61-65,

Ward SJ (2000) The efficacy of nestboxes versus spol-
lighting for detecting feathertail gliders, Wildlife
Research 27, 15-79.

Wilkinson HE (1961) The rediscovery of Leadbeater's
Possum, The Victorian Naturalist 78, 97-102.

Ooline Cadellia pentastylis F. Muell.: a Survivor

Ron Fletcher!

Some plants lead adventurous lives. They
manage to survive flood, fire, famine and
even the attentions of the taxonomists. A
chance meeting with one such was recently
made in the small Tregole National Park, a
few kilometres south of Morven on the
Warrego Highway in south-central
Queensland.

Cadellia pentastylis is a large, attractive
tree, variously described as 15-22 m tall
with some as little as 5 m and the tallest at
28 m, with grey bark described as fissured
or tesselated (Figs. | and 2) . It apparently
flowers only spasmodically and little is
known about the viability of the seed.

2% Marjorie Avenue, Belmont, Victoria 3216

Vol. 119 (5) 2002

Reproduction is often achieved by sucker-
ing, regrowth from rootstock or coppice
growth from stumps. The populations are
widely scattered through northern New
South Wales and south-central Queens-
land, and the species is registered on the
rare or threatened Australian plant list.
Some existing stands are on private prop-
erty where cooperation is sought in pre-
serving populations but where there is
pressure from clearing and grazing, Others
are more secure in reservations in both
New South Wales and Queensland
(Benson 1993).

Cadellia pentastylis is typical of relict
populations of several species of plants
such as Nothofagus, Livistonia, Casuarina,

235

Contributions

ig. 1. Ooline Cadellia pentastylis, Tregole
National Park, July 2001.

Acacia and so on, which are scattered
throughout Australia, and many of them
are survivors from warmer and wetter
times when rainforest covered much more
of our continent. In some cases they have
found niches that have some resemblance
to earlier conditions and in others they
have modified their habits to keep up with
changing climatic conditions (White
1994). Cadellia pentastylis would appear
to be one of the latter and survives in what
has been described with the curious term
‘semi-arid rainforest’.

Taxonomically, it has survived several
changes, being first described by Ferdinand
von Mueller in 1860 from a specimen col-
lected near Tenterfield in Northern New
South Wales (APNI), and placed in the fam-
ily Malpighiaceae. Later, Jadin (1901)
included C. pentastylis in the Simarou-
baceae along with C. monostylis. Engler
(1931) split Guilfoylia monostylis from
Cadellia and established the monotypic
genera Cadellia, Suriana and Guilfoylia.
Cronquist (1981) placed Cadellia, Suriana,
Guilfoylia and Stylobasium in a new family,
the Surianaceae, and this is the classifica-
tion followed in the Flora of Australia
(Benson). The monotypic genus Cadellia
takes its name from Francis Cadell, a pio-

236

Fig. 2. Foliage of Ooline Cadellia pentastylis,
Tregole National Park, July 2001,

neer river-boat captain on the Murray-
Darling system during the late 1850s. His
shipping company was responsible for the
transport of about 16,000 specimens, botan-
ical and zoological, from Mildura to
Adelaide in 1858, from where they were
sent on to Melbourne. Mueller had of
course been actively interested in the explo-
ration of the Murray-Darling Basin and vis-
ited the area in 1853-54, soliciting assis-
tance and making extensive collections
which resulted in many specimens being
shipped from the region in the subsequent
decade (Orchard 1997).

Acknowledgements

The author is grateful to Neville Walsh, Helen
Cohn and Sheila Houghton for direction to basic
references, and to Linden Gillbank for an intro-
duction to Francis Cadell and the Australian
Plant Name Index.

References and Further Reading

Benson J (1993) Biology and Management of Ooline
(Cadellia pentastylis) in New South Wales. New
South Wales Parks and Wildlife Service, Hurstville,
NSW.

Chapman, AD (1991) Australian Plant Name Index, 4
vol. (Australian Government Publishing Service:
Canberra)

Orchard K (1997) The scientific savant in nineteenth
century Australia. Mistorical Records of Australian
Science 11, 389-405.

White ME (1994) After the Greening. (Kangaroo Press:
Kenthurst)

The Victorian Naturalist

Contributions

Foraminifera from Lake Corangamite, Victoria

Ken N Bell'

Abstract

Surface sediment samples collected at Lake Corangamite, Victoria, contained dead specimens of the
calcareous foraminiferan Elphidium excavatum excavatum (Terquem, 1875). (The Victorian Naturalist

119 (5), 2002, 237)

Foraminiferans are usually restricted to
fully marine or marine brackish waters.
They have been recorded from non-marine
saline waters in many parts of the world
(Resig 1974, and references therein). In
Victoria, specimens of an unidentified, liv-
ing foraminiferan were reported from a salt
lake near Douglas, Western Victoria (Parr
1942) and two agglutinated species
(Trochamminita irregularis Cushman and
Bronnimann, Miliammina fusca (Brady ))
have been found in freshwater (Bell 1983).
Cann and DeDeckker (1981) reported a
fauna of two species (E/phidium sp.,
Trochammina sp.) from the Coorong,
South Australia.

Lake Corangamite, which lies to the west
of Colac, is the largest natural lake in
Victoria. It is about 32 km long and ranges
in width from 2 to |! km, giving a total area
of about 250 km’, In recent geological times
the lake was much larger, having an area of
perhaps 1700 km' and being markedly deep-
er (Currey 1964). The lake is shallow,
although this, of course, varies with seasonal
changes; Skeats and James (1937) give the
depth as less than 1.5 m whilst Bayly and
Williams (1973) give the maximum depth
during the great floods of 1953 as 3.4 m.
Because Lake Corangamite is a basin of
internal drainage its salinity can be high, but
it shows significant fluctuations with values
of three times seawater (Skeats and James
1937) to 12.5 ppt in 1953 and 6.5 ppt in
1960 (Bayly and Williams 1973).

The samples which contained the
foraminiferans were collected (May 2001)
on the northwest side of the lake near
Foxhow. The calcareous benthic species
Elphidium excavatum excavatum (Terquem)
was present. No specimens were alive when
collected and none had traces of protoplasm
present. Specimens were well preserved with
only slight surface frosting in a few cases

120 McCallum Road, Inverleigh. Victoria 332]

Vol. 119 (5) 2002

and were present in infrequent numbers,
about 10 per 200 g of sediment. This species
is usually restricted to brackish or marginal
marine environments such as estuaries or
brackish lagoons. There seems little possibil-
ity of the specimens being eroded from geo-
logical sediments; the COLAC Geological
Map (1973) shows no Tertiary sediments
outcropping around the lake edges (although
Skeats and James (1937) mention that such
outcrops do occasionally occur, they give no
locations), and the Pliocene Moorabool
Viaduct Sands, which underlie the recent
soils and lunettes in the area, consist of non-
marine sands and gravels. The only recorded
fossil occurrences of this taxon are two
records in the New Zealand Pliocene
(Hayward ef al. 1997). One would also
expect more species to be present if the spec-
imens were geologically derived. Whether or
not these specimens represent a living popu-
lation of foraminiferans in Lake
Corangamite or are representatives of a tran-
sient, introduced population will require
more study.

References

Bayly LAE and Williams WD (1973) Inland Waters
and their Ecology. (Longman: Melbourne)

Bell KN (1983) Foraminifera in some Victorian

Freshwater Streams. The Victorian Naturalist 100,
159-162.

‘ann JH and DeDeckker P (1981) Fossil Quaternary

and living foraminifera from athalassic (non-marine)

saline lakes, Southern Australia, Journal of

Paleontalogy 55, 660-670.

COLAC Geological Map (1973) 1:250000, | ed.
Geological Survey of Victoria, Sheet 154-12.

Currey DT (1964) The former extent of Lake
Corangamite. Proceedings of ihe Royal Society of
Victoria 77, 377-386.

Hayward BW, Hollis CI and Grenfell HR (1997)
Recent Elphidiidae (Foraminiferida) of the South-
west Pacific and fossil Elphidiidae of New Zealand.
Institute af Geological and Nuclear Sciences mono-
graph 16,

Parr WJ (1942) Foraminifera. The South Australian
Naturalist 21, 1-9.

Resig JM (1974) Recent foraminifera from a Hawaiian
Lake, Journal of Foraminiferal Research 4, 69-76.
Skeats EW and James AVG (1937) Basaltic barriers
and other surface features of the Newer Basalts of
Western Victoria, Proceedings of the Raval Society

of Victoria 49, 245-292

^

237

Naturalist in the Mountains

Winter Keeps Us Warm

The lines from The Waste Land by TS
Eliot,
‘Winter kept us warm, covering
Earth in forgetful snow..."

have always appealed to me for a couple of

reasons, Apart [rom my enjoyment of the
apparent incongruity, they have so much
additional meaning to an alpine ecologist,
The natural assumption about mountain
environments is that they are especially
harsh on animals, particularly in winter,
This is more true of countries such as
Australia where the fauna occupying, the
mountains are not pre-adapted to the envi-
ronment by reason of living at high latitude
(with the attendant benefit of experiencing
harsh conditions at lower altitude),

Fig. 1. Antechinus swainsonii Dusky Antechinus

However, one species gives us a good clue
as lo when the weather is most harsh. The
Dusky Antechinus Antechinus swainsonii
(Fig. 1), along with others of its genus, is
well known for the male die-ofT immedi-
ately after breeding. Yet looking at my
records gathered over the years in the
mountains, and at what foxes pick up in
their foraging, autumn seems by far the
most dangerous time, Spring has its male
die-off admittedly, and winter is a time
when you would expect an insectivorous

238

mammal in the snow to do it tough, but all
is not quiescent beneath the snow (see
Green 1997 for what goes on with insects
beneath the snow).

I was doing a small mammal radio track-
ing project some years ago, looking at what
changes in home range were caused by
snow cover, and I was alarmed at the mor-
tality among my subject animals. This was
irrespective of who collared the animals,
whether it was | or a more experienced col-
league. | looked back on the haleyon days
of my PhD to when I had zero mortality in a
winter radio tracking program. What | now
know is that the big die-off in mountain
Dusky Antechinuses is in autumn. | was
merely collaring already doomed animals.
In a paper published last year (Green 2001),
I| looked at autumnal reduction in body
weight in the Dusky Antechinus. This phe-
nomenon, of a reduction in lean mass rather
than a metabolisation of fat reserves
(Dehnel’s phenomenon), was known previ-
ously only from northern hemisphere
rodents and insectivores. However, above
the snowline in the Snowy Mountains, the
average weight of individual Dusky
Antechinuses falls by over 12% in the short
period from April to May. To go and lose
all this body weight put on at so much
expense in foraging effort in late summer
seemed inexplicable at first — after all, if
there was an advantage to being small in
winter, why go to so much trouble to get big
in summer? The reason for this loss of
Weight in autumn is not inextricably linked
with the availability of food. This is appar-
ent when you look at animals that have lost
weight in autumn and revisit the same ani-
mals in winter to find that they have put the
weight back on again under the cover of
snow, despite the fact that there is no
increase in the amount of food available.

What is happening is that the animals are
undergoing a period of great thermal stress
in a harsh microclimate when they are
exposed to the frost and cold rain and sleet
of autumn but are less stressed in winter
when the snow cover keeps them warm.
The evidence for this can be seen in the fact
that the heavier animals (that lose a bit of
weight in autumn) can be found throughout

The Victorian Naturalist

winter whereas the lighter animals disap-
pear from the population. It is an advantage
to be a lightweight under the winter snow
(the less you weigh the less you have to eat
to keep going and therefore the less you
have to expose yourself to the elements by
foraging) whereas in autumn it pays to be a
heavyweight so that you have something in
reserve, This autumnal thermal stress was
what was killing my animals, Foxes benefit
from this. You would expect that consump-
tion of antechinuses would peak in spring,
with foxes scavenging all the dead males.
However, this is not the case. In a study I
have just completed (Green in press),
remains of antechinuses in scats peaked in
autumn at both alpine and subalpine alti-
tudes, in all three years of the study. This
high autumnal mortality of small mammals
due to severe thermal stress is not confined
to Australia but is well known overseas, as
are the ameliorating effects of snow cover.
Eliot seems to have known it too.

Winter kept us cool? Some species, how-
ever, are more in need of keeping cool —
2*C to be exact. This is the preferred winter
temperature of the Mountain Pygmy-pos-
sum Burramys parvus. 1f the temperature
goes above or below this then the shorter
bouts of torpor (or rather the longer periods
of arousal) inerease energetic demands and
hence use up valuable fat reserves that have
to support the animal through the winter.
This is particularly the case when tempera-
tures rise, causing increased body tempera-
ture and hence metabolic rate.

Other small mammals in the mountains,
whilst not hibernating, do restrict their activ-
ity. The Broad-toothed Rat Mastacomys fus-
cus restricts its movements to a minimum in
winter and builds large stores of grass to
feed on when foraging is restricted. It also
builds a large ball nest of grass on the soil

Naturalist in the Mountains

surface (but beneath the snow) where it
takes advantage of a double layer of insula-
tion (Green 1998). In contrast, the Bush Rat
Rattus fuscipes tends to dig deeper burrows,
and the newly excavated fans of soil coming
from a rat hole are a feature of autumn
beneath the bushes in the mountains. The
Bush Rat and the two species of antechinus
are particularly active in winter. The Agile
Antechinus 1. agilis, also active under the
snow, is less common above the treeline. It
builds a nest in a hollow tree or rock and
insulates it with leaves. In one nestbox I
placed in a tree for this purpose, | counted
over 1000 dry Snowgum leaves that had
been carried the three metres or so up the
tree trunk from the woodland floor by the
resident Agile Antechinuses.

The rest of the animals above the snow
(such as ourselves), not Kept warm by the
snow, just have to rug up for the winter or
fluff up like birds (although thornbills and
scrubwrens will roost beneath the snow at
times). But for the small mammals an early
snowfall such as we had this year was a
blessing, reducing autumn to a short period
and covering Earth in forgetful snow.

References

Green K (1997) Inter-annual, seasonal and altitudinal
differences in invertebrate activity in the Snowy
Mountains. The Vieforian Naturalist 14, 222-229,

Green K (1998) A winter niche: the subnivean space.
In Snow: A Natural History; an Uncertain Future, pp
125-140, Ed K Green, (Australian Alps Liaison
Committee: Canberra.)

Green K (2001) Autumnal body mass reduction in
Antechinus swainsonii (Dasyuridae) in the Snowy
Mountains. Australian Mammalogy 23, 31-36,

Green K (in press) Altitudinal and temporal differences
in the food of foxes (Fulpes vulpes) at alpine and
subalpine altitudes in the Snowy Mountains, Wildlife
Research,

Ken Green

National Parks and Wildlife Service
PO Box 2228, Jindabyne, NSW 2627
Email ken,green(@inpws.nsw.goy au

Special Issues
The Victorian Naturalist
Murray River Special Issues, Volume 119 (3 and 4) 2002

Frederick MeCoy Special Issues, Volume 118 (5 and 6) 2001
Fungi Special Issue, Volume 118 (2) 2001

Copies are available for purchase from FNCV Office,
Locked Bag 3, Blackburn 3130, Victoria.

Price: $5 for Murray River and Fungi Special Issues; $8.50 for McCoy (includes GST)

Vol. 119 (5) 2002

239

Naturalist in the Mountains

Snow in Spring

In the short days of winter it is some-
times difficult to find any evidence of ani-
mal activity in the Snowgum woodlands,
particularly on a windy day when every
movement is disguised by blowing snow,
flapping bark or falling leaves. The winter
just past, with its early, deep snow meant
that the few resident birds were easy to
see, forced up out of the shrubs as these
became buried in the best early snow for

some years, One day, | managed to tick off

ten species out of the 12 or so I could have
hoped to record, by sight, sound or other
evidence. But even on that day the wood-
lands seemed quiet. Come spring, even
with snow on the ground, the Snowgum
woodlands come alive with birds, particu-
larly once the Royal Grevillea Grevillea
vietoriae comes into flower. It is quite

interesting to follow the regular pattern of

bird arrivals in this spring migration.
Everything is not fixed in time though,

and the timing, if not the pattern of

arrivals, has changed over the years. There
has been about a 30% reduction in snow
cover in the Snowy Mountains over the
past 45 years. This has led to interesting,
responses from the birds migrating back to
the mountains. There is a pattern of earlier
arrival in the 1980s and/or 1990s com-
pared to the 1970s. For 11 bird species for
which there are sufficient data, the earliest
record for nine of them was in the 1990s
and the 1980s (the latter generally differ-
ing by only a few days from the earliest
date for the 1990s) with the time of influx
into the mountains sometimes more than a
month earlier than in the 1970s,
Interestingly, two species showed virtual-
ly no change in arrival date across the three
decades. The Grey Fantail Rhipidura fuligi-
nosa is probably constrained by the need
for flying insects and Silvereyes Zosterops
lateralis — well, who knows what local
migration cues they follow? In some earlier
work that | did with Will Osborne, we
looked at the cues that brought in the birds.
The birds that we recorded as arriving earli-
er included honeyeaters that depend on the
flowering of shrubs which pop out from
under the snow, and birds that are largely
dependent upon snow-free ground for for-

240

aging such as Australian Kestrel Falco
cenchroides, Flame Robin Petroica
phoenicea and the Australian Pipit Anthus
australis. We built up that work from years
of accumulated records, but to look more
closely at the impacts of snow depth, cover,
and shrub exposure and flowering | have
had to undertake a weekly survey from
about mid-August until the last of the strag-
glers (Grey l'antails) arrive.

It is interesting to make these weekly bird
observations and to see these patterns fol-
lowed year after year. OF course some unex-
pected effects are unearthed. In spring,
Flame Robins forage in the open alpine
snowfields (above treeline). This means that
lor a few weeks the most commonly seen
robin on my rounds of the woodlands is the
Pink Robin Petroica rodinogaster and
another common species at this time is the
Olive Whistler Pachvcephala olivacea.

Other movement patterns in the mountains
are of interest in spring, including the arrival
of insects on the snow. These are not the
annual migrants, but insects swept up into
the mountains by weather patterns in which
they would have been better advised not to
fly. Last spring, half an hour after crossing a
snow-filled pass on foot, I retraced my steps
and counted the insects that had accumulat-
ed in my footprints. The insects were rolling
and tumbling, across the pass at a rate of at
least 135 per hour (rolling and tumbling
because if they had been airborne they
wouldn't have accumulated in my foot-
prints). Some migratory insects do get
caught; the Caper White Butterfly
Anaphaeis java which includes the Snowy
Mountains region in its strange southerly
looping migratory path is often trapped,

Then, of course, there is the Bogong Moth
Agrotis infusa. This appears on the snow in
spring and is quickly taken up by Little
Ravens Corvus mellori and Richard's
Pipits. | have sometimes wondered where
the moths hide during the day whilst wait-
ing for their boulder pile aestivation sites to
thaw out. In the classic 1950s work on the
species, lan Common found temporary
camps in locations such as under logs in the
woodlands surrounding his study site on Mt
Gingera (on the ACT/NSW border). That's

The Victorian Naturalist

all very well, and the moths do a similar
thing in woodlands in the Snowy
Mountains, but so many of the aestivation
sites are well above the treeline, so where
are the moths hiding so early? The clue
came to me as | sat in my tent while watch-
ing ravens forage in the grass and shrubs
that were exposed in patches in the snow
cover. Despite days of searching for what
they were feeding on | could find only a
few dead arctiid moth larvae, the spotted
grasshoppers Monistria concinna which the
birds seem to ignore, and ants. It was later
while catching Bogong Moths on a cold
night for studies on arsenic transport that I
got confirmation of my guess. | released the
moths that were excess to my needs onto
the grass; they disappeared and within sec-
onds I could find none. Of course, being
grown-up cutworms they are probably used
to hiding in grass, but | was amazed at how
quickly and completely they were hidden
from my searching fingers — but obviously
not from the raven beaks!

All through spring time the snow is slowly
retreating. One thing slower to retreat is the
ice on Blue Lake, the largest and deepest of
the mainland glacial lakes. The ice gets to
quite a reasonable thickness; | had to drill
1.4 metres to get through one September.
Eventually, with the warmer weather and
the pressure of incoming water from the
thawing snow, the ice cover breaks up and
the floating bits wash away or slowly melt.
In 1970 and 1971 the ice didn’t break up
until late November and there was still ice
on the lake in December, whereas in 1999 it
was all gone by early October.

Last spring, | was on the middle of Blue
Lake on 23 October and the ice was still a
good 50 cm thick. In places there were
patches of pink — snow algae. I didn’t think
much more about it until I was in the car
on my way home. Then ! recollected that
Harvey Marchant's work on snow algae
stated that the spores came from the soil
and moved up into the snowpack once
there was free liquid water, returning to the
soil as soon as the snow melted. So how

Naturalist in the Mountains

did they get to the middle of the lake? I
rang Harvey. Snow algae hadn't been
recorded on Australian lake ice before —
could | get a sample? The next day was
thundery so | gave it a miss and went back
on the 25". The lake was now either open
water or watery-looking ice too thin to
support me. Only about 20% of the lake
could support a person on skis — this just
two days after | had lunched in the middle!
I was able to get out 60 m to get a sample,
but it was useless for identification. Still,
maybe this month ...?

Reading list

Common IFB (1954) A study of the ecology of the adult
bogong moth Agrotis infusa (Boisd.) (Lepidoptera:
Noctuidae), with special reference to its behaviour
during migration and aestivation, Australian Journal
of Zoology 2, 223-203,

Green K and Pickering CM (2002) A scenario for
mammal and bird diversity in the Australian Snowy
Mountains in relation to climate change. In Mountain
Biodiversity: a Global Assessment, pp 241-249. Eds
C Körner and EM Spehn, (Parthenon Publishing,
London)

Marchant HJ (1998) Life in the snow: algae and other
microorganisms. In Snow: a Natural History; an
Uncertain Future, pp 83-97. Ed K Green. (Australian
Alps Liaison Committee, Canberra)

Osborne WS and Green K (1992) Seasonal changes in
composition, abundance and foraging behaviour of
birds in the Snowy Mountains. Emu 92, 93-105.

Note: The trips to Blue Lake referred to above
are to build up a data base of ice breakup over
time as a means of monitoring regional evidence
of global warming. | am interested in obtaining
photographs or recollections of when ice breakup
occurred in the past, A photograph of the lake
with full ice on it is of no use (unless it is very
late in the spring/summer), nor is a photograph
of open water (unless it is very early in the
spring). However, if anyone has photos of Blue
Lake (with year and month recorded) with ice
already breaking up then | will be able to add
another year to the list, | am particularly interest-
ed in 1982 (the cold year when there was not
much snow) and anything pre-1970, Therefore if
you, or anyone you know, has relevant dated
photos | would be delighted to hear from you. 1
can he contacted at the address below.

Ken Green

National Parks and Wildlife Service
PO Box 2228, Jindabyne, NSW 2627
Email ken.green(a npws.nsw.pov.gu

For assistance in preparing this issue, thanks to Kate Smith (desktop publishing), Ann

Williamson (label printing) and Dorothy Mahler (administrative assistance).

Vol. 119 (5) 2002

241

Tribute

William Nigel Balcombe Quick

6 June 1928 — 4 April 2002

Nigel Quick (often known as ‘Nige’) was
born in Melbourne and grew up in the inner

suburb of Toorak, He was the only child of

Hilary and William Balcombe Quick, an
honorary surgeon and head of the Alfred
Hospital Board at Melbourne. Nige was the
great great grandson of Lieutenant Colonel
William Balcombe, Napoleon's jailer on
the small oceanic island of St Helena in the
southern Atlantic, and East India Company
Representative charged with provisioning
the Company's ships.

Nige was educated at Melbourne Grammar
School from 1938 until 1946 and it was dur-
ing this time that his interest in entomology
began. He made contact with Alex Burns,
curator of Entomology at the National
Museum of Victoria, and through this asso-
ciation met other keen ‘youngsters’ and
experienced amateurs who provided encour-

agement, guidance and local knowledge of

the fauna of the Melbourne environment.
Nige went on to study science at
Melbourne University but never completed
his degree, instead taking employment at the
engineering firm MePhersons where over an
eight year period he acquired exceptional
knowledge of tools and their practical use.
During this period Nige, often accompa-
nied by Charles MeCubbin, travelled far
and wide throughout the State to collect
butterflies and other insects, fish and
plants, making, trips to the poorly surveyed
areas of the north-west (Big and Little
Deserts) and far east (East Gippsland).
Around 1956, Nige left McPhersons and
started his own business, setting up a plant
nursery, *Dargon Hill’, which he estab-

lished on an elevated site on the edge of

the suburb of Glen Waverley. The nursery
specialised in native plants, all of which
Nige grew from seed and cuttings collect-
ed on field trips. Most of the plant material
came from Victoria but he made an
extended trip to WA specifically to collect
samples of the rich endemic flora. Nige
was a pioneer in the propagation and sell-
ing of native plants.

In 1960, he married Gwenda Gerraty.
They had two children, David and Lucinda.

242

During the 1970s the family opened a shop
called ‘Gem Minerals in Syndal’. The busi-
ness reflected a long interest Nige had
developed in gemstones and minerals over
the years while collecting plant seed.

Nige joined the Entomological Society of
Victoria when it was reformed in 1961,
and was Vice-President from 1973 to
1976, In 1971 he began writing articles for
the Victorian Entomologist, the Society’s
journal. Over the ensuing 28 years he con-
tributed 51 articles. He also published
three papers in The Victorian Naturalist,
Queensland Naturalist and Australian
Journal of Entomology. Wis most produc-
live period was during the early and mid-
1970s when he handled the publication of
the journal and was assistant editor for sev-
eral issues. Most publications dealt with
the natural history of butterflies, novel
methods of setting, preserving and rearing
specimens, and technical aspects of the
ENTRECS scheme, a computerised distri-
bution database that he pioneered in the

di M t

[^ | = »
Nigel Quick at Garfield North in the late 1990s.

The Victorian Naturalist

mid-1970s (well before most people had
even heard of computers!). However, he
also wrote on a range of other topics,
including species protection and conserva-
tion, book reviews and field trip reports.

In 1979, he was made an Honorary mem-
ber of the Entomological Society of
Victoria in recognition of his pioneering
work on the ENTRECS scheme. However,
he subsequently resigned from the Society,
mainly because of poor health, and con-
tributed very few articles after 1985.

In 1980, Nige and Gwen lived at Kuranda
near Cairns where Nige studied the local
butterflies, plants and wildlife before return-
ing to Victoria in 1983.

Nige served on the advisory committee
of the Melbourne Zoo butterfly house in
1984, and provided advice on suitable
species, breeding requirements, and the
conditions needed to sustain tropical insect
populations. In 1987, 4023 specimens of
Australian butterflies from Nigel's private
collection were incorporated with the DF
Crosby collection and donated to the
Australian National Insect Collection. This
collection includes much important histori-
cal material from areas where populations
of many species are no longer extant, espe-
cially in Victoria.

Nige had a very wide interest in natural
history, including botany, geology,
ornithology and the ecological associations
between animals and plants. During the
early years he became an expert on
Victorian native freshwater fish, and col-
lected many species throughout the State.
However, entomology, particularly butter-
flies, remained a major preoccupation
throughout his life. Nige made a signifi-
cant contribution to entomology in
Australia and was widely known and
respected by professional entomologists.
He corresponded frequently with other
entomologists, assisted colleagues, stu-
dents and newcomers to the field, and was
very generous and helpful in passing on his
knowledge, particularly in his later years.
He thought deeply about his subject and
was never short of scientific ideas to

Vol. 119 (5) 2002

Tribute

explain natural phenomena. Nige was an
exceptionally gifted naturalist and a per-
fectionist in all aspects, from making care-
ful field observations, rearing the early
stages, to preparing specimens for the col-
lection, which he did with meticulous pre-
cision. He was a proficient photographer
and had expert working knowledge of
cameras and their lenses. But he was also
an excellent draughtsman and à competent
wildlife artist. He painted quite a few
transparent watercolour paintings of
insects, and also some landscapes in oil.

His current private collection, compris-
ing about 40 cabinet drawers and 15 store-
boxes (representing a diverse array of tem-
perate and tropical butterflies, macro
moths, cicadas and jewel beetles), along
with his vast personal photographic
library, has kindly been donated by
Nigel's descendants to the Australian
National Insect Collection where research
workers will have ready access to study
his material.

Nige was courteous, respectful and elo-
quent with a witty sense of humour. He
was a thorough gentleman, although at
times temperamental. He died quite sud-
denly at the age of 73 soon after being
diagnosed with cancer. A funeral service
was held on 12 April 2002 at the St
Andrews Uniting Church in Bunyip, fol-
lowed by a graveside commital in the
Bunyip Cemetary.

Those of us who knew or were close to
Nige were the beneficiaries of his enor-
mous talent and intellect, He was the last
of the older generation of lepidopterists in
Victoria still active in the field. Nige will
be greatly missed in the future, but his
contribution to entomology will live on
for a very long time.

Michael F Braby' and
Charles MeCubbin*

! School of Botany and Zoology
Australian National University
Canberra, ACT 0200

' PO Box 248

Sale, Victoria 3850

243

Book Reviews

Butterflies of A1

stralia: their Identification, Biology and

Distribution

by Michael F Braby

Publisher: CS/RO Publishing, Collingwood, 2000, 1008 pp, 2 hardback volumes,
70 colour plates. ISBN 0643065911. RRP $195

l'or centuries enthusiasts have found
Lepidoptera to be popular and accessible

insects for study, Butterflies in particular

depict, with their irresistible, exquisite
wing. patterns and complex life histories,
an interesting subject of beauty and curios-
ity, They are also of scientific importance,
being excellent indicator species of biodi-
versity and useful for genetic research or
studies in ecology and evolution.

Butterflies of Australia by Michael Braby
has followed the similarly titled classic
publications by IFB Common and DE
Waterhouse (1972 to 1981). Two volumes
of this latest work represent an indication
of the massive increase in information that
has occurred in the last twenty years, Braby
acknowledges that much of this new data
has been provided by amateur researchers.

Volume One begins with an introduction
to the history, classification, morphology
and distribution of Australian butterflies and
also includes an important section on their
conservation, Then follows a chapter (most-
ly for amateurs) on the collection, preserva-
tion and study of these insects. Following a
checklist of species and subspecies recorded
from the Australian region, the species
accounts begin, These include three fami-
lies; Hesperiidae (Skippers), Papilionidae
(Swallowtails) and Peridae (Whites and
Yellows), Then follows colour plates (with
label data) of all species (400 plus) and
many subspecies found in the Australian
faunal region and peripheral political
boundaries. Volume One concludes with a
selection of field photographs of adult and
juvenile butterflies,

Volume. Two has species accounts of the
remaining two families; Nymphalidae
(Browns, Nymphs and Danaines) and
Lycaenidae (Blues, Coppers, Hairstreaks
and Metalmarks). The volume is completed

with a series oF appendices including lists of

larval food plants, attendant ants and a use-

244

ful glossary, A comprehensive list of refer-
ences is given only in the second volume.
All chapters are evenly presented with
accurate information included (where
known) for all families, subfamilies, gen-
era and species. The systematic placement
of most genera is well researched and the
description of each species and genus is
generally accurate. A distribution map is
given for each species, and includes
boundaries delimiting the subspecies.
Some important additions have been
inade since the publications of Common
and Waterhouse, These include the use of
hew common and recently established or
corrected scientific names and up-to-date
informative data on butterfly life histories,
behaviour and distribution. Hach species’
text is accompanied by black and white
photos with bar indicators emphasising
specifie points of identification.
Significantly, several subspecies have
been ‘sunk’. With such an action, which
may or may not be productive, an illustra-
tion of specimens (or similar specimens)
from the type locality is essential. This has
not been done for species such as Zrapezites
lutea in Volume One and may be mislead-
ing. Similar difficulties may occur for over-
seas readers, lor instance, where an
Australian representative species, that may
have numerous subspecies in South East
Asian or Pacific Islands, cannot be identi-
fied at the subspeeifie level in the text until
the distribution and habitat section is seruti-
nised, This is somewhat rectified by includ-
ing the Australian subspecies in the check-
list in Volume One, Inevitably, in such a
large two-volume publication, referral
between the volumes ean be cumbersome.
Amateur enthusiasts would probably
have expected more coloured figures of
aberrations, forms and subspecies. There is
certainly room on some coloured plates for
this. It appears to be an oversight that the

The Victorian Naturalist

distinctive female of that great Australian
icon, the ‘Ulysses Swallowtail’ is omitted,
wheras the equally spectacular two species
of Birdwing are well accommodated, shar-
ing two plates. The species in some colour
plates (e.g. 25 and 34) are reduced a little
too far for accurate identification,

These inconsistencies do not detract from
the value of the publication. There are few
typographical errors and the text informa-
tion and photographie presentation is of a
high standard. Michael Braby has produced

a masterful work, which in this world of

information and technology, sets the stan-
dard for all other proposed publications on
the butterfly fauna of this planet. The excel-

Book Reviews

lent quality is seen foremost as a reflection
of the abilities and perseverance of the
author. There are also other fine technical
contributions made by ED Edwards, MS
Upton and Professor JFR Kerr.

Butterflies of Australia recently (and
deservedly) won the prestigious 2001
Whitley Medal. It is a publication worthy
of the bookshelves of all naturalists, butter-
fly enthusiasts and researchers. It is to be
hoped that a smaller *glove-box' edition
will follow.

Andrew Atkins

9 Bathurst Street
Dudley, NSW 2290

Dragonflies of the World

by Jill Silsby

Publisher: CSIRO Publishing, 2001. Hardback, 224 pages, colour illustrations.
ISBN 0643065121. 859.95

This book is an excellent introduction to
the dragonfly fauna of the world and will
be a valuable addition to the odonate liter-
ature, This wonderfully presented colour
book will interest amateur collectors
through to the professional odonatologists.
The author has written the book in a
relaxed, easily understood, semi-technical
style that makes it a delight to read. The
book contains superb colour photographs
of over 300 dragonfly species, some as lar-
vae, and many of their habitats. It is testa-
ment to the 20 years of hard work by the
author photographing and studying drag-
onflies in many countries.

The layout and production is very pleas-
ing, with colour photographs on almost
every page. However, it is not just a colour

book, as the chapters provide a wealth of

general information on dragonflies, ranging
from evolution to ecology, to conservation,
The science is strengthened by having many
leading odonatologists write or collaborate
on many chapters. The book is well struc-
tured, with 12 chapters, 11 short, covering
the Introduction, Today’s Odonata, Life
Cycle, The Perfect Hunting Machine, Lords
of the Air, Colour Polymorphism, Territory

Vol. 119 (5) 2002

and Reproduction, Habitat and Refugia,
Evolutionary Riddles, Artificial Rearing,
and Conservation, and a large chapter of
114 pages on Odonata Around the World.
This large chapter is the backbone of the
book and is where the reader is provided
with an overview of the world's dragonfly
fauna to the subfamily level. A general
overview of each subfamily is given, as
well as the ecology of the most distinctive
and interesting representatives.

The book also includes a very informa-
tive glossary, which will be valuable for
readers unfamiliar with dragonfly termi-
nology. At the back is included a list of the
World Dragonfly Societies, a helpful list
of the bibliographies used, and indexes to
both the species and to general terms.

Australian odonatologists will find the
many excellent photographs of 36 species
found in Australia very useful. The choice
of a large photograph of the dazzling
Rhyothemis graphiptera on the front cover
also highlights the beauty of Australia's
dragonflies. The Australian fauna is well
detailed and with the many photographs
provides the reader with a good insight
into the diversity of the Australian species.

245

Book Reviews

There are a few minor errors, which is
understandable when trying to cover such
a large fauna. The distribution information
of the caption *Tasmania's very rare
Ictinogomphus dobsoni Dobson's Tiger
wingspan 75 mm" (the lower photograph
on page 140), is incorrect, as 7. dobsoni
only occurs in the north-west of Western
Australia (Watson ef al. 1991). The central
photograph on page 153 (identified as a
male Eusynthemis nigra) is a male
Hemicordulia australiae. The third sen-
tence of the first paragraph on page 190
states that ‘A second is to break up the
small family Neopetaliidae, with one
species to remain in Aeshnoidea while
seven are transferred to Libelluloidea’ is
incorrect, It should read that one species
remains in the family Neopetaliidae, which
is moved to the Libelluloidea, while the
remaining, eight species are placed in the
family Austropetaliidae and remain in
Aeshnoidea (Carle and Louton 1994, Carle
1995), Also on page 190, paragraph. four

heavily criticises the classification system
of only Bechly and not other phylogeneti-
cists, which is unnecessary, as the jury is
still out in the classification case.

| would recommend this wonderful
colour book as a ‘must buy’ for all people
interested in dragonflies.

John Hawking

Murray Darling Freshwater Research Centre
Cooperative for Freshwater Ecology
CSIRO Land and Water

PO Box 921, Albury, NSW 2640

References

Carle PL (1995) Evolution, taxonomy, and biogeogra-
phy of ancient Gondwanian Libelluloides, with eom-
ments on Anisopteroid evolution and phylogenetic
systematics (Anisoptera: Libelluloidea).
Odonatologlea 24, 383-424.

Carle FL and Louton JA (1994) The larva of Neopetalia
punctata and the establishment of Austropetaliidae
faim, nov, (Qdonata), Proceedings of the Entomological
Societv af Washington 96, 147-155.

Watson JAL, Theischinger G and Abbey HM (1991)
The Australian Dragonflies; a Guide ta the
Identification, Distribution and Habitats of
lustralian Odanata. (CSIRO: Canberra and
Melbourne)

Possums: The Brushtails, Ringtails and Greater Glider

by Anne Kerle

Publisher: University of New South Wales Press 2007, 128 pp.
ISBN 0868404195, RRP $39.95

This is the twelfth book in the Australian
Natural History Series, a series to be com-
mended for bringing well-written species
accounts to the general public. For the past
few decades, natural history writing for the
general audience has consisted of books for
identification of the various taxa, with the
occasional travelogue through the natural
history world, or books focussed on particu-
lar regions. The Australian Natural History
Series from the University of New South
Wales Press was a natural, and most wel-
come, extension of this. With this series the
facts about a species or group of related
species are available to those with limited
access to the scientific literature. Unlike
some single-author views of a species,
where the veracity of an opinion cannot be
cheeked, this series employs the scientific

tradition of acknowledging the sources of

246

the information, albeit in a more reader-
friendly way than the usual scientific nota-
tion. This makes the series useful to the pro-
fessional as well as to the casual reader.

The latest offering is from Anne Kerle
who grew up geographically and profession-
ally with possums, starting with her honours
project on Mountain Pygmy Possums (cov-
ered in this series by lan Mansergh and
Linda Broome) before moving on to north-
ern Brushtail Possums for further research.
The breadth of knowledge, gained from her
own research, from the scientifie work of
others and from indigenous people, ts
brought together in this book to give a high-
ly readable and fact-filled account.

The first mainland marsupial to be recog-
nised as a marsupial was not a kangaroo but
a possum. Cook and Banks likened the ani-
mal they saw to the Brazilian Opossum (the

The Victorian Naturalist

first marsupial known to European science)
and the name stuck - minus the ‘O°. Apart
from noises on the roof or depredations to
fruit and rose bushes what do we know
about possums? Who would have thought
that the Common Brushtail Possum, so
adept at living with humans in our cities and
as a pest in New Zealand could be declining
here in Australia? It has disappeared from
half of its pre-European range. Some of the
threats to possums are covered in this book.
The fur trade took four million pelts in
1906, but today the threats are more from
the usual suspects: habitat clearance and
ferals, including one not generally thought
of, the honey bee that in one location occu-
pied 25% of hollow trees making them
unsuitable for possums.

This book covers the thirteen species of
large Australian possums in two families,
the Phalangeridae (five species) and the
Pseudocheiridae (eight species). Four fam-
ilies are not covered and, apart from one
single species treatment already published
there is room for another book in this
series. This book examines what is a pos-
sum and how the group evolved, has gen-
eral accounts of the thirteen species includ-
ing distribution, scats, skulls and footprints
of the better-known ones, habitat prefer-
ences, food, breeding, behaviour and final-
ly their interactions with humans.

I had few quibbles. Figure 1.1 has
Vombatidae as a suborder (rather than

Book Reviews

Vombatiformes) thus on p. 16 the Koala
(family Phascolarctidae) is included in the
Vombatidae (the wombats). On p. 19 the
flying term ‘attitude’ (meaning the angle of
its axes relative to the airflow) has morphed
into altitude (height above sea level).
Additionally (a quibble from a mountain
ecologist and a lapse in an author who has
studied the Mountain Pygmy Possum) on
p. 52 Common Brushtail Possums have
been found ‘in sub-alpine woodlands and
above the snowline’ (as if sub-alpine wood-
lands occur anywhere else). | also had trou-
ble working out what were the units on the
Y axis in Fig. 5.2.

There has been some debate in wildlife
circles about immunocontraception to con-
trol reproduction in pest animals. This is a
potential pathway in Australia to reduce
numbers of foxes (a long way from where
there are native populations of this species).
The possibility of this technique becoming
widespread in New Zealand to control pos-
sums (relatively close to source native pop-
ulations) may well widen the debate.

In all, the book was informative and
enjoyable to read and I for one will be dip-
ping into it to learn more about the animals
that manage to raid my grape vine the night
before the grapes are ready for picking.

Ken Green

National Parks and Wildlife Service
PO Box 2228, Jindabyne, NSW 2627

One Hundred Years Ago |

Notes on a Rare Victorian Shark
By Jäs. A. KERSHAW, F.E.S., Curator of the Zoological Dept., National Museum
(Read before the Field Naturalists’ Club of Victoria, 14th July, 1902.)

On the 2nd May last a large shark was captured in Hobson's Bay, off Williamstown, which
was quite unknown to any of the fishermen and others who saw it. It was captured through
becoming entangled in the nets of some local fishermen, which it damaged considerably, and
unfortunately bruised itself a good deal in its powerful struggles to escape.

Its captors immediately disembowled it, and had it conveyed to the city for exhibition purpos-
es, where it was secured for the Museum.

It proved to be a medium-sized male specimen of the Basking Shark, Cetorhinus maximus,
Gunner, and measured a total length of 12 feet 11 inches.

The only previous record of its occurrence in Victorian waters is that given by Sir Frederick
M*Coy in the Prodromus Zoology Victoria, vol. ii, where he describes and figures a large speci-
men measuring 31 feet 6 inches, which was captured in the fishermen’s nets at Portland in
November 1883. This specimen was exhibited in the city for some days, but, being hot weather,
it was rendered useless for museum purposes.

| From The Victorian Naturalist XIX, p 62, August 7, 1902

Vol. 119 (5) 2002 247

The Field Naturalists Club of Victoria Inc.

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ai E

Victorian —
Naturalist

Volume 119 (6) December 2002

Published by The Field Naturalists Club of Victoria since 1884

From the Editors

Authors are asked to carefully read the guidelines and consult the most recent issue of the
journal when preparing a manuscript for publication.

The Victorian Naturalist would not be successful without the enormous amount of time
and effort given voluntarily by a large number of people who work behind the scenes.

One of the most important editorial tasks is to have papers refereed. The Editors would
like to say thank you to those people who refereed manuscripts published during 2002:

Robyn Adams John Hawking Glen Saunders
Andrew Bennett Paul Humphries John Seebeck
Dave Britton Laurie Laurenson Rick Shine
Rhonda Butcher Richard Loyn Russell Shiel
Malcolm Calder Brian Malone Letitia Silberbauer
Rohan Clarke Richard Marchant Dianne Simmons
John Dean Peter Menkhorst John White
Kelvyn Dunn John Neil Alan Yen

lan Endersby Tim New

Linden Gillbank Peter Robertson

The Victorian Naturalist publishes articles for a wide and varied audience. We have a
team of dedicated proof-readers who help with the readability and expression of our
articles.

Ken Bell Pat Grey Geoffrey Paterson

Andrew Bennett Murray Haby John Seebeck

Arnis Dzedins Virgil Hubregtse Rob Wallis

lan Endersby Michael McBain Gretna Weste

Maria Gibson Tom May Jenny Wilson

Ken Green lan Mansergh Alan Yen

Sincere thanks to our book reviewers for 2002 who provided interesting and insightful
comments on a wide range of books and other materials:

Andrew Atkins Ken Green Cheryl O'Dwyer
Margaret Corrick John Hawking Gary Presland
Stuart Dashper Simon Lewis Gerry Quinn
Gwen Elliot Angus Martin Melody Serena
Rodger Elliot Sara Maroske Jenny Tonkin
Cecily Falkingham Tim New

As always we particularly thank our authors who provide us with excellent material for
publication.

Special thanks to Ken Green for his ‘Naturalist in the Mountains’ series to commemorate
The United Nations International Year of Mountains 2002.

Our editorial advisory team continues to provide valuable advice and assistance: lan
Endersby, lan Mansergh, Tom May, and John Seebeck.

On the production side, thank you to:

Ken Bell, who prepares the annual index,

Dorothy Mahler for administrative assistance,

Kate Smith, for desktop publishing,

Ann Williamson, for printing the mailing labels, and
Printers, BPA Print Group, especially Steve Kitto.

The

Victorian

K
Naturalist
f
Volume 119 (6) 2002 Aq December

F.N.C V,

Executive Editor: Merilyn Grey
Editors: Alistair Evans and Anne Morton

Research Reports A Population Study of Eastern Grey Kangaroos Macropus
giganteus at Tower Hill State Game Reserve, South-western
Victoria, by Eve McDonald-Madden and Graeme Coulson .......252

Potential Dispersal Agents of Higher Plants in the Kosciuszko
Alpine Area, by Karen C Rudkin and Catherine M Pickering ...258

The Red Fox Vulpes vulpes L. in the Dandenong Creek Valley:
Impacts on Wildlife and an Integrated Program for their

Control, by A Morton, D Tagg, R Wallis and J White ................ 269
Naturalist in the Flying Spiders and Crawling Flies, by Ken Green ........................ 276
Mountains
Book Reviews Grassed Up: Guidelines for Revegetating with Australian
Native Grasses, hy Cathy Waters, Wal Whalley and
Charles Huxtable, reviewed by Cheryl O "Dwyer ............ ss 278

Platypus: the Extraordinary Story of How a Curious Creature
Baffled the World, by Ann Moyal, reviewed by Melody Serena.279
Working on Country: Contemporary Indigenous Management
of Australia's Land and Coastal Regions, edited by Richard
Baker, Jocelyn Davies and Elspeth Young, reviewed by
Gary EP ESIAMA a. Ia. ocean Ac Etre Lr eoru nae 280

Gardening With Australian Rainforest Plants, by Ralph Bailey
and Julie Lake, reviewed by Gwen and Rodger Elliot ................28

Guidelines for Authors at the end of this issue

ISSN 0042-5184

a
Cover: Eastern Grey Kangaroo Macropus giganteus. See Research Report on p. 252. Photo
by Keith Dempster.

Web address: http://www.vicnet.net.au/~fnev/
Email fnev@vicnet.net.au

Research Reports

A Population Study of Eastern Grey Kangaroos
Macropus giganteus at Tower Hill State Game Reserve,
South-western Victoria

Eve McDonald-Madden! and Graeme Coulson!

Abstract

Overabundance in native species has recently become an important focus for wildlife management.
The negative effects of overabundant macropod populations include threats to human life or liveli-
hood, depression of the density of favoured species, decline in body condition and reproduction, and
a loss of equilibrium between fora and fauna, This study was undertaken at Tower Hill State Game
Reserve in south-western Victoria, The population of Eastern Grey Kangaroos Macropus giganteus
in the reserve is derived from a number of individuals released in 1964, and this study established
that the current population totals about 150 individuals. The adult sex ratio was at parity and 71 % of
females in the population were breeding, suggesting that this population has the potential for rapid
increase, Further study into the population’s growth trajectory and the development of a manage-
ment plan is needed to avoid problems of over-grazing and oyer-browsing. (The Victorian Naturalist

119 (6), 2002, 252-258)

Introduction
Overabundance in native species has

recently become an important focus of

wildlife management (Garrott e? al. 1993;
Rodger 1998). In parts of North America,
overabundance of ungulates, particularly
White-tailed Deer Odocoileus virginianus
and Elk Cervus elaphus, has become a
major management issue in national parks
and on private land (e.g. Healy er al. 1997;
McShea and Rappole 1997). Some
Australian species seem to be undergoing
parallel growth rates to those witnessed in
North America. In Australia, the Koala
Phascolarctos cinereus (Martin and
Handasyde 1999) and a number of macro-
pod species have been identified as prob-
lematie (Coulson 1998).

Overabundant macropod species have
been shown to have a negative effect on
agricultural yields (Hill eż aZ. 1988) and can
also have deleterious effects on biota in
areas where grazing pressure is concentrat-
ed, such as nature reserves (Neave and
Tanton 1989; Coulson 1997). In particular,
problems arise in populations that are closed
to emigration, either by impenetrable habitat
or by fencing. In the past the deliberate
introduction of kangaroo populations into

! Department of Zoology, University of Melbourne,
Victoria 3010

Current address: Department of Natural Resourees
and Evnironment, Arthur Rylah Institute for
{ nvironmental Research, PO Box 137, Heidelberg,
Victoria 3084
Email evememadt(ahotmail com

252

closed areas has resulted in problems due to
overpopulation (see Coulson in press).

This study was undertaken at Tower Hill
State Game Reserve, a small reserve of
native habitat, isolated from other native
vegetation by expanses of cleared pasture.
Over the years, macropod species includ-
ing Eastern Grey Kangaroos Macropus
giganteus, Black Wallabies Wallabia
bicolor and Red-necked Wallabies
Macropus rufogriseus (introduction unsuc-
cessful), have been released into the
reserve (Goldstraw 1986), In November
1964, Eastern Grey Kangaroos were rein-
troduced into the reserve (Goldstraw
1986). They were initially introduced to
two enclosures planted with open forest
vegetation, designed to acclimatize macro-
pod breeding stock before ultimate release.
The initial numbers of kangaroo released
seem to be unrecorded; however anecdotal
observations of releases and deaths [rom
1979 to 1985 are available. Records show
that 19 kangaroos were released intermit-
tently over six years, with nine of those
released animals dying in their enclosures
before release into the reserve itself.
Records of observations from outside the
enclosures over the same period show a
maximum number of 36 animals,

Eastern Grey Kangaroos have been
shown to inhabit ecotones of ample forest
cover and open grassy feeding grounds
(Hill 1981; Short and Grigg 1982). Due to

'The Victorian Naturalist

their need for cover these kangaroos are
highly restricted to areas of lateral vegeta-
tion, and are reluctant to cross large areas
lacking cover. The population at Tower
Hill is thus effectively closed to natural
emigration and immigration, owing to the
expanse of surrounding open habitat.

An overabundance of Eastern Grey
Kangaroos at Tower Hill has the possibility
to fulfill all of the problems of overabun-
dance outlined by Caughley (1981). Tower
Hill is an ecotourist destination, and there
is concern about the possibility of injury to
visitors. In addition, increased grazing pres-
sure by overabundant kangaroos is likely to
affect other ground dwelling herbivores and
also have a negative effect on native flora,
leading to decreased species diversity with-
in the reserve. As has already been wit-
nessed at Tower Hill with the Koala popu-
lation, overabundance and the inability of
animals to emigrate out of the system to
find food may see an increase in mortality
due to starvation. Recently, Tower Hill
managers and *Friends of Tower Hill’ have
become concerned about the number of
kangaroos present in the reserve. Despite
this, the population has never been properly
surveyed. Available records are sporadic
and do not account for the possibility of
double counting; therefore, estimates of
population size are unreliable for manage-
ment assessment.

The objectives of this study were firstly,
to obtain an accurate population count of
Eastern Grey Kangaroos in the reserve;
secondly, to determine population structure
with the aim of establishing the likely
growth trajectory of the population; and
thirdly, to assess habitat usage. Overall, the
aim of this study was to provide informa-
tion for the management of the population
of Eastern Grey Kangaroos at Tower Hill
State Game Reserve.

Study area and methods

Tower Hill State Game Reserve is located
about 15 km northwest of Warnambool,
Victoria. Tower Hill is an extinct volcanic
crater of approximately 3 km diameter
(Morgan 1997). The total land area of the
study site is 264 ha, partitioned into the
Main Island (125 ha), Fairy Island (18 ha),
Hat Island (1 ha) and the crater rim (120 ha).

This small reserve is isolated from other

Vol. 119 (6) 2002

Research Reports

areas of native vegetation by farmland pas-
ture and a steep crater rim. By the end of
the 19" century it was completely cleared
for agriculture. In 1961, the decision was
made to regenerate this area as a State
game reserve. Regrowth throughout the
reserve can be classified into five major
vegetation types: Eucalypt forest, predomi-
nantly Eucalyptus viminalis; Eucalypt
woodlands, dominated by E. viminalis and
E. ovata; Allocasuarina woodlands, domi-
nated by A. verticillata and A. stricta;
shrublands dominated by Acacia mearnsii,
A. decurrens and A, melanoxylon; and
grasslands.

This study was conducted during May
and June 1999, Sampling of the kangaroo
population was undertaken using à grid
system of 15 transect lines running east-
west across the main island of the reserve
(Fig. 1; Morgan 1997) and by 4WD along
the tracks around the outer rim of the
reserve. Transects were mapped out by the
authors, 160 m apart (as in the Koala sur-
vey of Morgan 1997) and, before data col-
lection, transects were walked on the rele-
vant east or west compass bearing, and
marked using flagging tape.

Over the course of this study, each line
was walked three times, always between
1000 and 1600 hours. Odd lines were
walked in succession, followed by evens,
to minimise the possibility of double
counting created when animals moved
between lines, The observer set off from
the beginning of each transect and walked
at a constant speed. When a kangaroo
group was observed in front of the observ-
er, the following data were recorded: the
number of kangaroos in the group; the dis-
tance from the observer to the centre of the
group (measured with a rangefinder); the
compass bearing from the observer to the
centre of the group; the sex, if deter-
minable, of the individuals within the
group, along with the reproductive status
of females; and the habitat type in which
the group was observed.

The outer rim was sampled by observa-
tion from a vehicle. Established tracks
were driven in a 4WD vehicle between
1630 and 1730 hours. When a group was
observed, the number of individual kanga-
roos was recorded along with sex and
reproductive status of females, in the hope

253

Research Reports

Å 0

200 400
N remm
| Metres

Hat
Island

Main Track

Transects/ Management Tracks ----

Fairy Island

Tower Hill State Game Reserve

Fig. 1. Map of Tower Hill showing grid system of transects (adapted from Morgan 1997).

of establishing a maximum stable number
for the outer rim population. If animals
moved extensive distances out of sight,
they were not recorded. This was done to
minimise the chance of double counting.

The percentage of each habitat type with-
in the reserve was estimated from an aerial
photo. Using | mm graph paper, habitat
percentages were calculated from the num-
ber of squares within the reserve occupied
by each habitat.
Results

The population density (kangaroos/km")
was estimated using the computer program
MNPS (see Morgan 1986). This program

254

estimates population density and three other
parameters from a set of frequency distribu-
tions: a coefficient of conspicuousness (in
this study 12.91 + 1.724 m), the lateral veg-
etation cover (0.05317 50.007573), and the
maximum direct-line distance at which
species recognition is possible (dmax).

The minimum detection distance was 8
m, whilst the maximum (dmax) was 300
m. The highest frequency of kangaroo
groups was observed at 23 m. The sighting
distance and the curve of the model fitted
to those data are shown in Fig. 2.

From this set of data, a density of 81.4 +
3.54 kangaroos/km' (mean + SE) was esti-
mated. The population for the main island

The Victorian Naturalist

25

20

Frequency
a

za,
o

Research Reports

A

\
L3 A A A
0 ^ 4 S22
0 50 100 150

yet 2 —18—9—42— tt th tt tt 9 tt

200 250 300 350

Distance (m)

Fig. 2. Frequency distribution of sighting distances from the observer to the kangaroo groups.
Triangles indicate actual data points and the line indicates the curve of best fit using the computer

program MNPS (Morgan 1986).

was calculated by multiplying the estimated
density by the area of the main island, giv-
ing an estimate of 117 individuals (Table
1). The stable number observed for the
outer rim of the reserve, which was taken
as the maximum number observed over the
four counts, was 34 (Table 1). The overall
estimate for the reserve was 151, with nar-
row confidence intervals (Table 1).

The minimum percentage of adult
females breeding in the population was
71%. The percentage of males within the
sample was 54%. A chi-squared test found
that the sex ratio of adults in the popula-
tion did not differ from parity (y^ = 0.00, p
7 0.05, df — 94).

The pattern of percentage use of the five
defined habitats by this population can be
seen in Fig. 3. A chi-squared test was done
comparing raw frequencies observed in
each habitat with expected frequencies cal-
culated from the availability of each habitat
(Set | - fine conditions: x^ = 9.63, p < 0.05,
df = 4; Set 2 - wet-windy conditions: X^ =
21.98, p < 0.05, df = 4; Table 2). The null
hypothesis assumes that habitats were used
in proportion to availability. Both analyses
rejected the null hypothesis, and show that
kangaroos were not using habitats in pro-

Vol. 119 (6) 2002

Table 1. The population estimates (+ standard
error) and 95% confidence intervals for Eastern
Grey Kangaroo Macropus giganteus at Tower
Hill State Game Reserve.

Estimated 95% CI

Population
Main Island 1175 107 to 127
Maximum stable
number for outer
rim population 34 -
Overall (for outer rim

151 141 to 161

and main island)

portion to their availability. Differences
between the two sets are observed within
the selection of woodlands and shrublands.
Set 1 shows an observed usage of shrub-
lands less then expected and an increased
usage of Allocasuarina woodlands in fine
conditions, In comparison, set 2 shows an
increased usage of shrublands and decrease
in the use of eucalypt woodlands on days of
wet and windy conditions (Table 2).

Discussion

This study shows that the population of
Eastern Grey Kangaroos at Tower Hill
State Game Reserve has increased from a
minimum of 19, between 1979 and 1985,

285

Research Reports

45

40

Percentage

0 —
Eucalypt
Woodlands

Allocasuarina
Woodlands

Shrublands

Grasslands Eucalypt Forest

Habitat type

Fig. 3. The percentage of the five main habitats within the reserve (shaded) and the percentage of
Eastern Grey Kangaroos Macropus giganteus observed within these habitats (unshaded).

to an estimate of 151 in 1999, This esti-
mate suggests that the population has
increased dramatically. Along with this
growth witnessed over the past 14 years
the demographics of the population at
Tower Hill suggest that there is potential
for continued growth in the future. The
rate of increase of a population depends
upon its current age distribution and sex
ratio (Bayliss 1985; Caughley et al, 1987).
l'his population showed no apparent sex-
biased mortality in adult individuals with
the sex ratio of adults being at parity.
Norbury et al. (1988) noted a female bias
in Western Grey Kangaroos M. fuliginosus
melanops after drought, resulting in a
decreased growth trajectory, However,
parity or male biased populations are
thought to be indicative of growing popu-
lations (Norbury eż al. 1988).

The population of kangaroos at Tower
Hill also had a high percentage of breeding
females, with a minimum of 71% of adult
females carrying young. Females with very
small pouch young are extremely difficult
to identify as breeding individuals; conse-
quently, this figure is likely to underesti-
mate the breeding potential of the popula-
tion. Kangaroos may respond to changes in
rate of population inerease by adjusting
their rates of reproduction (Bayliss 1985;
Caughley eż al. 1987). In harsh conditions,

256

kangaroos have been shown to have a dra-
matically reduced fecundity (Norbury ef
al. 1988). Continued growth of the Tower
Hill population can be expected, as a high
percentage of the females are breeding.

Overabundance of any grazing or brows-
ing species has the potential to be devastat-
ing to the vegetation within any reserve.
Research at other sites has shown that
overgrazing by kangaroos can have a nega-
tive effect on the structure and floristics of
ground cover and on soil invertebrates and
ground-dwelling birds (e.g. Neave and
Tanton 1989; Coulson 1998). In these
areas, removal of kangaroos had a marked
effect on the regeneration of native ground
vegetation and dicotyledon species (Neave
and Tanton 1989; Coulson 1998), Since
Tower Hill’s classification as a State Game
Reserve in 1961, a great deal of effort has
gone into the re-establishment of indige-
nous vegetation on this once cleared site.
Overgrazing by Eastern Grey Kangaroos
may threaten the continued success of this
revegetation effort,

In this study, kangaroos actively selected
habitats, although the preferences shifted
depending on the environmental condi-
tions. Selection, it seems, was affected by
the need for shelter from the elements. For
example, when environmental conditions
were harsher, more time was spent in

The Victorian Naturalist

Research Reports

Table 2. The frequency of kangaroos observed in each habitat and the expected frequency according
to the proportion of the reserve occupied by each habitat type.

Habitats Eucalypt Allocasuarina Shrubland Grassland Eucalypt
Woodland Woodland Forest

Set | (fine conditions)

Observed 23 13 10 14 0

Expected 21 8 20 11 0.5

Set 2 (wet-windy conditions)

Observed 7 9 37 2 0

Expected 23 8 22 2 0.5

shrublands of greater density where there is
more cover than in open woodlands, Such
results are important to consider when con-
templating management of the reserve.
Time and money should be invested in the
rehabilitation and protection of areas of the
reserve at greatest risk. For example, in
cooler months, increased management of
over-used shrubland areas may reduce the
impact of kangaroo grazing in the reserve.
It appears that the population of kanga-
roos at Tower Hill State Game Reserve has
increased dramatically and will continue to
increase. The first step in assuring the
effective management of the Tower Hill
population should be to outline the issues
within the reserve that are of management
priority. With management priorities clear-
ly stated, an appropriate management plan
should be designed to effectively maintain
these goals. The management plan should

determine an optimal population size of

kangaroos within the reserve. Appropriate
methods of control should be investigated,
and consideration given to use of concur-
rent methods such as exclusion fencing,
culling and fertility control.

This study has been the first comprehen-
sive study completed on the kangaroo pop-
ulation at Tower Hill since their reintro-
duction in 1964. Results have shown that
numbers in the reserve have increased dra-
matically and in light of the population
demography, have the potential to increase
further, thereby threatening the biodiversi-
ty of the reserve. Further study, and devel-
opment of management practices, are
essential to ensure that efforts already put
into the reserve by staff, Friends and the
community have not been in vain.
Acknowledgements
We would like to thank: the Rangers of Tower

Hill, Kerry Murphy and Jodie Honan, for their
assistance with this project; David Morgan and

Vol. 119 (6) 2002

Dustin Marshall for statistical advice; Mick
Wright for his unwavering support and assistance
in laying out transect tape; and Jill McDonald for
her great help in all facets of this project, includ-
ing the dreaded removal of transect tapes. Also
thanks to Sue, Alex and Joe MeCulloch for pro-
viding accommodation during this study. Our
thanks must also go to Friends of Tower Hill for
their financial support,

All the hard work that went into this study and
into this final report is dedicated to John
MeCulloch.

References

Bayliss P (1985) The population dynamics of red and
western grey kangaroos in arid New South Wales,
Australia. If, The numerical response function.
Journal of Animal Ecology 54, 127-135.

Caughley G (1981) Overpopulation. In Problems in
Management of Locally Abundant Wild Mammals, pp
7-19. Eds PA Jewell, S Holt and D Hart. (Academic
Press: New Yark)

Caughley G, Shepherd N and Short J (1987)
Kangaroos; their Ecology and Management in the
Sheep Rangelands of Australia. (Cambridge
University Press; Cambridge)

Coulson G (1997) Woodlands Historic Park Kangaroo
Management Plan; a report to Parks Victoria.
Department of Zoology, The University of
Melbourne.

Coulson G (1998) Management of overabundant
macropods ~ are there conservation benefits? In
Managing marsupial abundance for conservation
benefits: Proceedings of a Symposium held at the
Society for Conservation Biology Conference,
Svdney, Australia, July 1998,

Coulson G (in press) Overabundant kangaroo popula-
tions in southeastern Australia. In Wildlife, Land, and
People: Priorities for the 21st century. Proceedings
of the Second International Wildlife Management
Congress. Eds R Field, RJ Warren, H Okarma and
PR Sievert, (The Wildlife Society: Maryland)

Garrott RA, White PJ, and White CAV (1995)
Overpopulation: An issue for conservation biolo-
gists? Conservation Biology 7, 946-949,

Goldstraw P (1986) Department of Natural Resources
and Environment internal file.

Healy WM, Decalesta DS and Stout SL (1997) A
research perspective on white-tailed deer overabun-
dance in the north eastern United States, Wildlife
Society Bulletin 25, 259-263.

Hill GJE (1981) A study of the habitat preferences in
the Grey Kangaroo. Australian Wildlife Research 8,
245-254.

Hill GJE, Barnes A and Wilson GR (1988) The use of
wheat crops by kangaroos, Macropus giganteus, in
southern Queensland, Australian Wildlife Research
15, 111-117.

257

Research Reports

Martin R and Handasyde K (1999) The Koala: Natural
History, Conservation and Management. (1 Iniversity
of New South Wales Press: Sydney)

MeShea WJ and Rappole JH (1997) The science and
politics of managing deer within a protected area.
Wildlife Society Bulletin 25, 443-446.

Morgan DG (1986) Estimating vertebrate population
densities by line transect methods. Me/bourne
College of Advanced Education Occasional Papers
No 11, March 1986

Morgan DG (1997) Tower Hill State Game Reserve
Koala Population, April 1997: Report for the
Department of Natural Resources and Environment,
Victoria. Department of Zoology, The University of
Melbourne,

Neave HM and Tanton MT (1989) The effects of graz-
ing by kangaroos and rabbits on the vegetation and

ihe habitat of the fauna in the Tidbinbilla Nature
Reserve, Australian Capital Territory. Australian
Wildlife Research 16, 337-381.

Norbury GM, Coulson GM and Walters BL (1988)
Aspects of the demography of the Western Grey
Kangaroo, Macropus fuliginosus melanops, in semi-
arid north-west Victoria. Australian Wildlife
Research 15, 257-566,

Rodger JC (1998) Overabundance: a new issue in con-
servation. In Managing marsupial abundance for
conservation benefits; Proceedings of a Symposium
held at the Society for Conservation Biology
Conference, Svdney, Australia, July 1998.

Short J and Grigg GC (1982) The abundance of kanga-
roos in suboptimal habitats: wheat, intensive pastoral,
and mallee. Australian Wildlife Research 9, 221-227.

Potential Dispersal Agents of Higher Plants in the Kosciuszko
Alpine Area

Karen C Rudkin' and Catherine M Pickering"

Abstract

Morphology of higher plants in the Kosciuszko alpine area was investigated in order to determine
probable dispersal agents. Attributes including seed and fruit morphology, presentation mechanisms
and habitat for each plant were matched to ecological syndromes representing four primary dispersal
agents: wind, water, animals and the plant itself, The most common probable dispersal agent was
wind, followed by animal, water and parent dispersal. (The Vietorian Naturalist 119 (6), 2002, 258-268)

Introduction

Dispersal mechanisms in plants have
been the subject of extensive theoretical
and empirical research, much of which has
concentrated on the anatomical structure of
the dispersal unit or diaspore i.e. the seed,
fruit, spores and/or vegetative parts (see
reviews by van der Pijl 1982; Murray
1986). Several criteria can be used to clas-
sify dispersal methods (Dansereau and
Lems 1957, cited in van Rheede van
Oudtshoorn and van Rooyen 1999),
although the system most commonly
applied is that of Ridley (1930) using the
agent of transport as the criterion (e.g. van
Rheede van Oudtshoorn and van Rooyen
1999). In this system, dispersal traits repre-
senting ecological ‘syndromes’ are
matched to the most effective dispersal
agent (Ridley 1930; van der Pijl 1982).
This may involve a single trait, although in
most cases it is a combination of character-
isties (van der Pijl 1982). For example,
plants growing. in open habitats with cap-
! Griffith University, School of Environmental and
Applied Sciences, PMB 50 Gold Coast Mail Centre,

Queensland, 9726
‘Email ce; piekeringicmailbox.gu,edu.au

258

sules set upon elongated stalks that open to
release small mature seeds when humidity
is low are considered to be wind dispersed
(van der Pijl 1982).

Principle transport agents of plant dias-
pores are animals, wind, water and the par-
ent plant itself, with the resulting dispersal
mechanisms termed zoochory, anemo-
chory, hydrochory and autochory respec-
tively (van Rheede van Oudtshoorn and
van Rooyen 1999), Wind is considered the
"basic? dispersal mechanism (Sernander
1927, cited in van der Pijl 1982), although
in ferns and conifers, animal dispersal was
found to be dominant (van der Pijl 1982),
Wind is a common dispersal mechanism in
open habitats and regions characterized by
severe climates and limited numbers of
animal vectors (van der Pijl 1982). In
regions with few animal dispersal agents,
the flora may show a range of traits for
wind and water dispersal (Ridley 1930). In
mountainous areas, mammal, bird and
insect diversity tends to decline as altitude
increases (Meyer and Thaler 1995),
Around Mt Kosciuszko, the largest contin-
uous alpine area in Australia, there appear

The Victorian Naturalist

Research Reports

'Table 1. Animals of Kosciuszko alpine area with potential to act as dispersal agents for flora. R,

alpine resident; S, reside in the alpine area during the

warmer months only; * introduced species.

Taxon Status Habits/Diet References
Mammals
Burramys parvus R Hibernates during winter, diet includes Mansergh et al. 1990;
Mountain Pygmy- cached seeds eg. Alpine Hovea Hoved Green and Osborne
possum montana and Mountain Plum Pine 1994
Podocarpus lawrencei
Antechinus spp. R Small marsupial, insectivore, occurs in Dimpel 1976; Green
Brown and Dusky moister areas where adequate ground cover — 1989
Antechinus of shrubs exist
Mastacomys fuscus R Mainly herbivorous, takes small amounts of Carron e/ al. 1990
Broad-toothed Rat seeds, builds nests from grass
Rattus fuscipes R Found in woodland, heath and grassland, Carron ef al. 1990
Southern Bush Rat in winter moves to alpine herbfield, Diet
diverse including seeds, dicots taken year
round, monocots in late autumn and winter
Vulpes vulpes* R Diet: insects, mammals and minimal Green and Osborne
Red Fox vegetable matter 1981
Birds
Anas superciliosa S Found near water, feeds mainly on seeds Frith 1986
Pacific Black Duck and vegetable matter
Gallinago hardwickii S Feeds in fens, bogs and wet grassland on Green and Osborne

Latham's Snipe

Anthus novaeseelandiae S

Richard's Pipit

seeds and invertebrates

Tall alpine herbfield and tussock grassland,
eats insects and small seeds, grass lined
nest situated in thick vegetation e.g.
Richea continentis

Nest of grass, feeds on insects, nectar

Nest of grass, feeds on insects and plant
foods on the ground or in trees

Nests on the ground in grass lined
depression, feeds on seeds, green grass

Zosterops lateralis S

Silvereye and berries
Sturnus vulgaris* S

Common Starling
Coturnix pectoralis S

Stubble Quail

and insects

Ants
Iridomyrmex spp. R

Omnivorous, can be found in dry and wet
habitats eg. bogs, tracks, herbfield and
grassland

1994
Green and Osborne
1994

Green and Osborne
1994

Green and Osborne
1994

Cayley 1975

Green and Osborne
1994

OOO eee em

to be few animal agents for seed dispersal
(Table 1). In contrast, much of the area
consists of wind exposed slopes or valley
floors with creeks, fen or bogs (Costin ef
al. 2000), providing opportunities for dis-
persal by wind and water. In addition, par-
ticularly in harsh environments, optimal
dispersal may not require maximizing dis-
tance, as the most suitable habitat may be
more readily available close to the parent
plant (Ellner and Shmida 1981).

To better understand the role of seed dis-
persal in the ecology of the Kosciuszko
alpine zone, this review identifies morpho-
logical dispersal characteristics for all seed
plants recorded in the region and matches
them to potential dispersal agents.

Vol. 119 (6) 2002

Methods
Study area

The Kosciuszko alpine area extends from
the treeline at around 1830 m above sea
level to the top of Australian’s highest
mountain (Mt Kosciuszko: 2228 m ASL) an
area of approximately 100 km’ (Costin ef al.
2000). This area contains the most diverse
alpine environment on the mainland with
204 native angiosperm taxa from 40 fami-
lies, one gymnosperm species and fourteen
introduced species of angiosperms (Costin
et al, 2000). Plants exhibit a range of
growth forms including shrubs, grasses,
sedges, rushes, herbs, mats and cushions,
with herbs the most dominant.

259

Research Reports

Table 2. Dispersal syndrome used for alpine flora of Kosciuszko National Park for the four primary

dispersal agents for seed bearing plants.

Poaceae

[airs present on lemma and/or palea
Bearded callas

Dispersal Dispersal syndrome characteristics
syndrome General
Anemochory — Dust-like seeds

(wind) Fruit on long peduneles

Peduncles elongate during fruit maturation

Dry fruits that dehisce in dry air

Awn present
Plumose rachilla

Predominantly small herbaceous species

Small (- Imm), plumed, winged or ribbed

sceds

Hydrochory
(water) Dry fruits that dehisee in moist air
Floating utricle or loose seed coat

Seed coat to prevent germination during

transport
Moist or low-lying habitat
Low-growing,
Zoochory Fruit berry, drupe or nut
(animal)

Elaiosome present on seed

Small (^1 mm) and/or scabrous diaspore

Moist habitat

Autochory
(parental plant)

Diaspores without other mechanisms (Imm)

Fruit or seed colour red, black or purple
Burred, scabrous, hooked, or sticky diaspore

Open habitats

Glabrous lemma, palea and/or callas

Bristled rachilla, habitat near water-
course

Spikelets droop at maturity, fine hairs
on lemma

Absence of awn and/or rachilla

Barbed callas

Spiked or scabrous lemma and/or
palea

Presence of muero

Large floret (75mm), glabrous
lemma

Short rigid awn, bristled rachilla

Explosive dehiscence of capsule or follicle

Classification of dispersal syndrome

Due to the number of taxa to be studied
and diversity of structures associated with
dispersal, classification by agent dispersal
syndromes was limited to the four principal
agents of dispersal (after Ridley 1930;
Table 2). Criteria used to classify taxa were
based on (1) seed morphology including
structures attached to or enclosing, seed, (2)
plant habitat, and (3) seed presentation
(where on parent plant). Some difficulties
were encountered in applying the criteria to
the specialized structures of grasses
(Poaceae); therefore, these taxa were con-
sidered separately (Table 2). Morphological
characteristics for the seed plants of
Kosciuszko alpine area were identified from
taxonomic descriptions by Costin e/ al.
(2000), Harden (2000) and Walsh and
Entwisle (1999), Native flora and intro-
duced weed species were categorized sepa-
rately. When attributes of a species could be
attributed to more than one agent syndrome,
a subjective decision was made as to the
"most likely" dispersal agent based on the
combination of attributes.
Results

The major potential dispersal agent was
identified for each of the 205 native taxa

260

(Appendix 1) and 14 introduced species
(Appendix 2). For the native taxa, the most
common agent was wind (anemochory,
114 taxa), followed by animals (zoochory,
53), water (hydrochory, 34) and the parent
plant (autochory 4; Fig. 1). The dominance
of wind dispersed seeds was due to the
prevalence of herbs with small plumed,
winged or ribbed seeds (Fig. 2), which
together accounted for 73 (35%) of the
total native taxa.

Growth forms typically associated with
wet habitats such as sedges and rushes
were characterized by water and animal
dispersal (Fig. 2) while low-growing eush-
ions and mat/herbs were wind dispersed.
Taxa with shrub/subshrub growth forms
(Epacridaceae) were also wind dispersed.
Of the grasses, 47.8% are likely to be wind
dispersed, with the remaining 52.2% divid-
ed evenly between both water and animal
dispersal. Animal dispersal was the deter-
mined agent for the three dwarf shrubs
Alpine Ballart Exocarpus nanus (Santal-
aceae), Alpine Kelleria Ke/leria dieffen-
bachii (Vhymelaeaceae) and Alpine Rice-
Flower Pimelea alpina (Thymelaeaceae).

The most probable dispersal agent of the
single representative of Gymnospermae,
Mountain Plum Pine Podocarpus lawrencei

The Victorian Naturalist

son

40% 7

30% 7

2097

Percent taxa

10" =

g ' ' P
Anemochory — Hydrochory Zoochory Aulochnry

Fig. 1. Native taxa recorded in the Kosciuszko
alpine area attributed to the four probable dis-
persal agents. Anemochory = wind, hydrochory
= water, zoochory = animal, autochory =
parental plant. n = 205.

was found to be zoochory. Seeds of this
species are included in the diet of the
Mountain Pygmy Possum Burramys
parvus, known for their habit of caching
seeds in and around nests (Table 1). Within
angiosperms, seed dispersal by animals was
found in four of the six monocot families
(40% of taxa) and thirteen of the thirty-two
dicot families (20% of taxa). Of the native
taxa likely to have diaspores dispersed by
animals, 57% had characteristics indicating
dispersal that involved viable seed being
eaten and excreted undamaged. The
remaining 43.4% had sticky, hooked or
burred seeds likely to be dispersed by ani-
mals externally (e.g. carried on fur; Fig. 3).

An alternative method for animals to
carry seed externally was identified by
Ridley (1930) where the small otherwise
un-mechanized seeds of Juncaceae were
transported from one area to another in
mud sticking to the feet of waterfowl. This
may be the method of seed dispersal for
Juncus antarcticus, J. faleatus and J.
thompsonianus in the Kosciuszko alpine
area, as the plants invariably grow in
marshy spots where they may come into
contact with the feet or plumage of the
Pacific Black Duck Anas superciliosa
residing in the area over summer (Table 1).
However, the small seed size, presentation
mechanism and growth form of these taxa
also suggests dispersal by wind.

The six remaining taxa of the family
Juncaceae are comprised of the genus
Luzula, many of which were also found near
water, and their larger seed size (1.0-1.8
mm) indicates water dispersal. Interestingly,
many species of Luzula, including one

Vol. 119 (6) 2002

Research Reports

Agent
100 4
E autochory
04 —
80 ISSSVzoochóry
m Hydrachory
EE A nemochory

Number of taxa

Subsarub
Shrub 'subshrub
Ma

Herb’
rub

= 3

Herth/cushion

Growth Porm

Fig. 2. Probable dispersal agents of native taxa
(in terms of growth forms) of the Kosciuszko
alpine area (Costin er al. 2000).

species found in this region, Luzula acutifo-
lia subsp. nana, have a small fleshy out-
growth of the seed coat (caruncle). These
structures on Luzula multiflora were found
to be attractive to ants (Ridley 1930), which
transport the seeds short distances. Ridley
(1930) also recorded finding Luzula inflores-
cences, with seed, used as nesting material
by sparrows; however, this has yet to be
determined for the species considered here
and is unlikely to account for the wide distri-
bution of these plants as there are few bird
species, and no sparrows.

Herbs comprised 55.6% of all native taxa
(Costin er al. 2000). Taxa from common
herb families such as the Asteraceae and
Apiaceae were determined to potentially be
wind dispersed, as were those of
Orchidaceae, Crassulaceae, Geraniaceae,
Onagraceae, Gentianaceae, Scrophu-
lariaceae, Plantaginaceae, Campanulaceae
and Stylidiaceae (Fig. 4). Herbs also
accounted for all species classed as auto-
chores (mechanical self dispersal) compris-
ing the three Cardamine spp. (Brassicaceae)
and the Violet Viola betonicifolia subsp.
betonicifolia, each characterized by explo-
sive seed capsules,

There were fourteen established introduced
weed species in five families, and all but one
were classified as wind dispersed (Fig. 5).
Timothy grass Phleum pretense (Poaceae)
was classified animal dispersed primarily
due to the thin and dry outermost bract of
the floret lemma extending as a point.
Ridley (1930) cited several examples of
viable seeds of this species being dispersed

261

Research Reports

Animal Agent
___|Exozoochory
| Endozoochory

141
Monocotyledoneae
23

12 ie
Dicotyledoneae

10
9

84

23
6*4

Number of plant taxa

Poaceae
Juncaceae
Lilliaceae

Cyperaceae
Santalaceae
Winteraceae 4

Podocarpaceae

$-- Tu umen camelis 3
8 d € B m S os 8
E! g m] Z E 5 B
E cs of d EROS 5
= w - E 50 ed vd
m get EL EI aM E, 3
* CX XA a
E É u a [v]
N

Family

Fig. 3. Number of taxa in each family exhibiting characteristics that indicate seed dispersal by ani-
mals. Endozoochores transport seeds internally, excreting them undamaged. Seed transport on the

fur or plumage of animals are class

ified exozoochores. Numbers at the top of each column indicate

the total number of native taxa in each family recorded in the Kosciuszko alpine area.

401

Monocotyledons

201

Number of plant taxa

101

OLS COMMO
o0 9 o 9 v 9
ss 3 8 SS 3 g
Akan iz. MAI VD Dum gx 8
E bese 2 TPA ature
S d$ S Ww s d"
= LL ucc Mar epic
=EEsS eyes SI
RAR ee ope? eet
E SO E S or
Gud ES. [el o
PD
E:
[9]

Fig. 4. Number of native herbs by family recorded in the

q o y 90 o9 o O q 1
S 89 $ s 84 9 9 E 2 2 E
o 0 02070 00 o O OD oO d o
D E ONES ES gery I MC SL I2: S8
BU ORIS UTE ces resti CR GT ue es rd =
bre B gfe & oS g rc g
S Ep sD ees Se "BD Vice MS a D
B oO "$$ S S x MS 3 2
6 3 O E B S e s <
a SIS UE B S
T ao x i=! [G)
ARA oes IS
Family m

Dicotyledons

Agent

Autochory

Zoochory
ES Hydrochory

E Anemochory

Kosciuszko alpine area (Costin er a/. 2000)

attributed to the four probable dispersal agents. Numbers at the top of each column indicate the total
number of native taxa in each family recorded in the Kosciuszko alpine area.

by cattle and horses, both of which were pre-
sent in the alpine area until recently.

Discussion

The diversity of alpine diaspore structures
and presentation at maturity presumably
increase the chance of seeds being
favourably dispersed (Ridley 1930).

262

However, many of the species studied in
the Kosciuszko alpine area had little or no
apparent adaptations for diaspore dispersal
based on morphological descriptions of
seed in Floras. More detailed examination
of material and field work may clarify this.
Complications also arose in seeds adapted

The Victorian Naturalist

T

T

1

| | P Pr n
Cee Carpi | PUMP. ces Fabaceae mm

Famil

Fig. 5. Number of introduced weed taxa by
family recorded in the Kosciuszko alpine area
(Costin et al. 2000) attributed to the probable
dispersal agents. n = 14.

for several agents of dispersal, or for a suc-
cession of agents (Ridley 1930; van Rheede
van Oudtshoorn and van Rooyen 1999).
For example, feathery hairs on seed can be
dispersed by wind, provide buoyancy in
water, facilitate adherence to animals, and
assist with anchorage to the ground during
germination (Ridley 1930). It has also been
demonstrated that the awn of barley is an
important photosynthetic structure, sug-
gesting the diaspores may have functions
other than just dispersal (van der Pijl 1982).
Therefore by assigning seed to a single
agent, we may have underestimated animal,
water and plant dispersal mechanisms.
Conditions favorable for growth in alpine
environments are often restricted to small
micro-habitats, Seed morphologies that
restrict dispersal to the vicinity of the parent
plant may therefore be advantageous (van
der Pijl 1982). Zohary (1962, cited in van
Rheede van Oudtshoorn and van Rooyen

1999) distinguished two major categories of

localized dispersal; 1) restriction of dispersal
in space and 2) restriction of dispersal in
time. The former includes burying of seed or
formation of fruits immediately above the
ground, limiting separation of seed, and the
presence of sticky material (mucilage), burrs
or other anchorage mechanisms on the seed.
Inhibitory mechanisms of the latter include
postponement of dispersal until the following
season, and opening or closing of the disper-
sal organ (e.g. capsule or follicle) as a func-
tion of moisture. The taxonomic descriptions
available did not include enough information
to make determinations as to the presence or
absence of these types of mechanisms, nor
did they provide post-dispersal details e.g.
weight, buoyancy, absorbency of seed coat,
hydrophilic nature of hairs, etc.

Vol. 119 (6) 2002

Research Reports

This study provides information about
potential dispersal mechanisms. Observations
of species growing in the natural habitat are
required to confirm the potential dispersal
mechanisms proposed. This also includes
examining the role of vegetative dispersal
mechanisms such as the transport of torn-off
rhizomes, stems and stolons. Based on the
data presented here, however, we can con-
clude that wind appears to be the most com-
mon type of dispersal mechanism for the
Kosciuszko alpine flora.

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263

Research Reports

Appendix I. Native gymnosperm and angiosperm species, subspecies and varieties recorded from the Kosciuszko
alpine area including growth form and habitats as recorded by Costin er al. (2000), Habitat abbreviations: B, bog;
F, fen; FMa, feldmark (Epacris-Chionohebe alliance); FMb, (Coprosma-Colobanthus) alliance; H, heath; STG, sod
tussock grassland; SAH, short alpine herbfield; TAHa, tall alpine herbfield (Celmisia-Poa alliance); TAHDb, tall
alpine herbfield (Brachyscome-Austrodanthonia alliance). Zoochory* indicates endozoochory; Zoochory** indi-

cates exozoochory; « endemic to Kosciuszko alpine area,

Species Common Name Growth
Form
Gymnospermae
Podocarpaceae
| Podocarpus lawrencei Mountain Plum-pine Shrub
Angiospermae: Mononcotyledoneae
Poaceae
2 Agrostis meionectes Alpine Bent Grass
3 Agrostis muelleriana Mueller's Bent Grass
4 Agrostis parviflora Hair Bent Grass
5 Agrostis venusta Graceful Bent Grass
6 Agrostis sp. Swamp Bent Grass
7 Australopyrum velutinum Mountain Wheat-grass Grass
8 dusirodanthonia alpicola Crag Wallaby-grass Grass
9 Chionochloa frigida e Ribbony grass Grass
10 Deschampsia caespitosa Tufted Hair-grass Grass
ll Deyeuxia affinis Dwarf Bent-grass Grass
12 Deyeuxia carinata Slender Bent-grass Grass
13. Deyeuxia crassiuscula Coarse Bent-grass Grass
l4 Deyeuxia monticola var. Mountain Bent-grass Grass
monticola
15 Hierochloe submutica Alpine Holy-grass Grass
16 Poa costiniana Prickly Snow-grass Grass
17 Poa fawcettiae Smooth-blue Snow-grass Grass
18 Poa hiemata Soft Snow-grass Grass
19 Poa saxicola Rock Poa Grass
20 Rytidosperma australe Snowpatch grass Grass
2] Rytidosperma nivicola Snow Wallaby-grass Grass
22 Rytidosperma nudiflorum Alpine Wallaby-grass Grass
23 Rytidosperma pumilum Feldmark grass Grass
24 Trisetum spicatum subsp. Bristle-grass Grass
australiense
Cyperaceae
25 Carex archeri Archer’s Sedge Sedge
26 Carex breviculmis Short-flower Dryland Sedge Sedge
27 Carex canescens Yellow-headed Sedge Sedge
28 Carex cephalotes Button Sedge Sedge
29 Carex echinata Star Sedge Sedge
30 Carex gaudichaudiana Tufted Sedge Sedge
31 Carex hebes Dryland Sedge Sedge
32 Carex hypandra Dark Fen-sedge Sedge
33 Carex jackiana Short-flower Swamp Sedge Sedge
34 Carpha alpina Small Flower-rush Sedge
35 Carpha nivicola Broad-leaf Flower-rush Sedge
36 Isolepis aucklandica Slender Club-rush Sedge
37 Isolepis crassiuscula Alpine Club-rush Sedge
38 lsolepis habra Tufted Club-rush Sedge
39 Isolepis montivaga Mountain Club-rush Sedge
40 Isolepis subtilissima Dwarf Club-rush Sedge
41 Oreobolus distichus Fan Tuft-rush Sedge
42 Oreobolus pumilio subsp. Alpine Tuft-rush Sedge
pumilio
43 Schoenus calvptratus Alpine Bog-rush Sedge
44 Uncinia compacta Compact Hook-sedge Sedge
45 Uncinia flaccida Mountain hook-sedge Sedge
46 Uncinia sinclairii Sinclair's Hook-sedge Sedge
47 Uncinia sulcata Furrowed Hook-sedge Sedge
Restionaceae
45 Empodisma minus Spreading Rope-rush Rush
Juncaceae
49 Juncus antarcticus Cushion Rush Rush
50. Juncus faleatus Sickle-leaf Rush Rush
5| Juncus thompsonianus Thompson's Rush Rush
52 Luzula acutifolia subsp. nanas Dwarf Woodrush Rush
53 Luzula alpestris Mountain Woodrush Rush

Habitat Probable
Dispersal
Agent

H Zoochory*
STG, SAH, F, B Anemochory
TAla, STG, F, B. FMa Hydrochory
? Anemochory
TAHa, STG Anemochory
STG, F, B, TAHa Hydrochory
TAHa, STG Zoochory**
TAHb Anemochory
TAHa Anemochory
STG Anemochory

SAH Zoochory**
STG, F, B, H Zoochory**
TAHa, STG Zoochory**
STG, TAHa Zoochory**
TAHa Hydrochory
STG, F, B, TAHa Hydrochory
TAHa, STG Anemochory
TAHa Anemochory
TAHa Zoochory*
SAH Hydrochory
STG, TAHa, F Anemochory
STG, TAHa, F Anemochory
FMa Hydrochory
TAHa, STG Anemochory
B Hydrochory
TAHa, STG Hydrochory
F, B, TAHa, STG Hydrochory
F, B. SAH, STG Hydrochory
SAH, TAHa, B Hydrochory
F, B, STG Hydrochory
TAHa, STG Hydrochory
F, B Hydrochory
SAH, B, F, STG Hydrochory
STG, F, B Hydrochory

F, B Hydrochory
SAH, B, F Zoochory*
SAH, TAHa Zoochory*
B Zoochory*

SAH, F, STG Zoochory*
SAH Zoochory*

B, SAH, TAHa Zoochory*
SAH, F Zoochory*
SAH Zoochory**
TAHa, STG Zoochory**
TAHa, STG Zoochory**
STG Zoochory**
TAHa, STG Zoochory**
B, STG, TAHa Iydrochory
SAH, B Zoochory**
PB Zoochory**

B, F, SAH Zoochory**
SAH Hydrochory

STG, TAHa Hydrochory

264

The Victorian Naturalist

Research Reports

Appendix 1 continued.

Species Common Name Growth Form Habitat
54 Luzula atrata Slender Woodrush Rush B, F, SAH
SS om australasica subsp. — Feldmark Woodrush Rush Fma
durae
56 Luzula modesta Bog Woodrush Rush B
57 Luzula novae-cambriae Roek Woodrush Rush TAHa, H
Liliaceae
58 Astelia alpina var. Silver Astelia Herb B, TAHa
novae-hollandiae
59 Astelia psychrocharis e Kosciuszko Pineapple-grass Herb B, TAHa
60 Dianella tasmanica Tasman Flax-lily Herb TAHa, H
61 Herpolirion novae-zelandiae Sky Lily Herb STG
Orchidaceae
62 Caladenia alpina Alpine Caladenia Herb B
63 Prasophyllum alpestre Highland Leek-orchid Herb B, TAHa, STG
64 Prasophyllum tadgellianum | Tadgell's Leek-orchid Herb B, TAHa, STG
Angiospermae: Dicotyledoneae
Proteaceae
65 Grevillea australis Alpine Grevillea Shrub H
66 Grevillea victoriae
subsp. nivalis Royal Grevillea Shrub H
67 Orites lancifolia Alpine Orites Shrub H
Santalaceae
68 Exocarpos nanus Alpine Ballart Dwarf Shrub H, TAHa
Portulacaceae
69 Neopaxia australasica White Purslane Herb SAH, TAH a
Caryophyllaceae
70 Colobanthus affinis Alpine Colobanth Herb VAHa, FMa
71 Colobanthus nivicola e Snowpatch Cushion Cushion PMb, SAH, TAHa
72 Colobanthus pulvinatus Feldmark Cushion Cushion FMa, TAHa
73 Scleranthus biflorus Twin-flower Knawel Mat/Cushion LAHa, STG

74 Scleranthus brockier

75 Scleranthus singuliflorus

76 Stellaria multiflora
Ranunculaceae

77 Caltha introloba

78 Ranunculus acrophilus e

79 Ranunculus anemoneus e

80 Ranunculus dissectifolius e

81 Ranunculus graniticola

32. Ranunculus gunnianus

83 Ranunculus millanii

84 Ranunculus muelleri

85 Ranunculus niphophilus e
Winteraceae

86 Tasmannia xerophila subsp.

xerophila

Brassicaceae

87 Cardamine astoniae

88 Cardamine lilacina s.l.

89 Cardamine robusta e

Droseraceae

90 Drosera arcturi
Crassulaceae

91 Crassula sieberiana s.l.
Rosaceae

92 Acaena sp.

93 Alchemilla xanthochlora auct.
Fabaceae

94 Fovea montana

95 Oxylobium ellipticum

96 Podolobium alpestre
Geraniaceae

97 Geranium antrorsum

98 Geranium potentillaides

var, abditum

99 Geranium potentilloides var.

100 Pelargonium helmsii
Rutaceae

101 Phebalium ovatifolium e

Vol. 119 (6) 2002

Brockie's Knawel Mat/Cushion TAHa
One-flowered Knawel — Herb/Cushion TAHa, STG, FMa
Rayless Starwort Herb TAHa
Alpine Marsh-marigold Herb SAH
Feldmark Buttercup Herb FMa
Anemone Buttercup Herb FMb, TAHa, SAH
Feather Buttercup Herb TAHa, STG, H
Granite Buttercup Herb TAHa, STG
Gunn’s Alpine Buttercup Herb TAHa, STG
Dwarf Buttercup Herb STG, F, B
Felted Buttereup Herb TAHa, STG
Snow Buttercup Herb SAH, TAHa
Alpine Pepper Shrub H
Aston's Bitter Cress Herb B
Nalive Bitter Cress Herb STG, TAHa
Snow Bitter Cress Herb SAH, TAHab
Alpine Sundew Herb B, SAH
Austral Stonecrop Herb TAHa
Bidgee-widgee Subscrub TAHa, H, STG
Lady's Mantle Herb TAHab
Alpine Hovea Shrub H
Common Shaggy Pea — Shrub/Subshrub IH
Alpine Shaggy Pea Shrub/Subshrub NH
Rosetted Crane's-bill Herb TAHa, STG, B
Mountain Crane’s-bill Herb TAHa, STG, H
Alpine Swamp Crane's-bill — Herb TAHa, STG
Alpine Stork’s-bill Herb TAHa
Ovate Phebalium Shrub H

Dispersal

Hydrochory
Hydrochory

Hydrochory
Hydrochory

Zoochory *

Zoochory*
Zoochory *
Zoochory *

Anemochory
Anemochory
Anemochory

Anemochory

Anemochory
Anemochory

Zoochory*
Hydrochory

Anemochory
Anemochory
Anemochory
Hydrochory
Hydrochory
Hydrochory
Anemochory

Hydrochory
Zoochory **
Zoochory **
Zoochory **
Zoochory**
Zoochory**
Zoochory**
Zoochory**
Zoochory**

Zoochory*
Autochory
Autochory

Autochory
Hydrochory
Anemochory

Zoochory**
Hydrochory

Anemochory
Anemochory
Anemochory

Anemochory
Anemochory

Anemochory
Anemochory

Anemochory

265

Research Reports

Appendix 1 continued,

Species

Common Name

Stackhousiaceae
02 Stackhousia pulvinaris
Violaceae
03 Melicytus sp. (Hymenanthera
auct.)
04 Viola betonicifolia subsp.
hetonicifolia
Thymelaeaceae
05 Kellaria dieffenbachii
106 Pimelea alpina
07 Pimelea axiflora subsp. alpina
08 Pimelea ligustrina subsp.
ciliata
Myrtaceae
109 Baeckea gunniana
110 Baeckea utilis var. utilis
HE Kunzea muelleri
Onagraceae
112 Epilobium gunnianum
113 Epilobium sarmentaceum
14 Epilobium tasmanicum
Haloragaceac
15 Gonocarpus micranthus
subsp. micranthus
16 Gonocarpus montanus
17 Myriophyllum pedunculatum
subsp. pedunculatum
Apiaceae
18 Aciphylla glacialis
19 Aeiphylla simplicifolia
20 Dichosciadium ranuncula-
ceum* var, ranunculaceum e
121 Diplaspis nivis
22 Gingidia algens e
123 Oreomyrrhis brevipes
124 Oreomyrrhis ciliata
125 Oreomyrrhis eriopoda
26 Oreamyrrhis pulvinifica
27 Oschutzia cuneifolia
28 Schizeilema fragoseum
Epacridaceae
29 Epacris glacialis
30 Epacris microphvila s.A.
31 Epacris paludosa
32 Epacris petrophila
33 Leucopogon montanus
34 Pentachondra pumila
35 Richea continentis
Gentianaceae
136 Chionogentias muelleriana
subsp. alpestris e
Boraginaceae
137 Myosotis sp.
Lamiaceae
138 Prostanthera cuneata
Scrophulariaceae
139 Chionoliebe densifolia
140 Euphrasia alsa
141 Euphrasia collina var.
diversicolor
142 Euphrasia collina subsp.
glacialis e
143 Euphrasia collina var.
lapidosa e
144 Veronica serpyllifolia s.l.
Plantaginaceae
145 Plantago alpestris
146 Plantago euryphvlla
147 Plantago glacialis
148 Plantago muelleri

Alpine Stackhousia Mat

Woody Violet Subshrub

Showy Violet Herb
Alpine Kellaria Dwarf Shrub
Alpine Rice-flower Dwarf Shrub
Alpine Bootlace-bush Subshrub
Kosciuszko Rose Shrub
Alpine Baeckea Shrub

Mountain Baeckea
Yellow Kunzea

Gunn's Willow-herb Herb
Mountain Willow-herb Herb
Snow Willow-herb Herb
Creeping Raspwort Herb
Mat Raspwort Herb
Mat Water-mil foil Herb
Mountain Celery Herb
Mountain Aciphyll Herb
Wreath Penny wort Herb
Snow Penny wort Herb
Kosciuszko Aniseed Herb
Rock Carraway Herb
Bog Carraway Herb
Australian Carraway Herb
Cushion Carraway Herb
Wedge Oschatzia Herb
Alpine Pennywort Herb

Bog Heath
Coral Heath
Swamp Heath
Snow Heath

Shrub

Snow Beard-heath Shrub
Carpet Heath Mat/Subshrub
Candle Heath Subshrub
Mueller’s Snow-gentian Herb
Forget-me-not Herb
Alpine Mint Bush Shrub
Feldmark Snow Hebe Mat/Subshrub
Dwarf i Herb
Variable Eyebright Herb
Snow Eyebright Herb
Feldmark Evebright Herb
Thyme Speedwell Herb
Veined Plantain Herb
Broad Plantain Herb
Small Star Plantain Herb
Star Plantain Herb

Growth Form

Shrub/ Subshrub
Shrub/ Subshrub

Shrub/Subshrub
Shrub/Subshrub

Shrub/Subshrub

Habitat Dispersal
STG, TAHa Zoochory*
H Zoochory*
TAHa, STG Autochory
FMa Zoochory*
TAHa, STG, H Zoochory*
H, TAHa, STG Zoochory*
H Zoochory*
B, H Anemochory
H, B Anemochory
H, STG Anemochory
TAHa, STG, B, H Anemochory
lAHa, STG Anemochory
SAT, FMb Anemochory
F, B, STG Ilydrochory
TAHa Hydrochory
F Hydrochory
TAHa Anemochory
TAHa, STG Anemochory
SAH, B, TAHa Anemochory
SAH, B, TAHa Anemochory
TAHa Anemochory
TAHa, FMa Anemochory
B Anemochory
TAHa, H Anemochory
SAH, TAHa Anemochory
TAHa, B Anemochory
TAHa Anemochory
H, B, STG, TAIHa Anemochory
FMa, H, B Anemochory
B, H Anemochory
FMa Anemochory
H Zoochory*
H, TAHa, STG, FMa Zoochory*
B Anemochory
STG, TAI a Anemochory

TAHab Zoochory*

H Zoochory*
FMa Anemochory
FMa, STG, TA Ha Anemochory
TAHa, STG, H Anemochory
STG, SAH, F, TAlla — Anemochory
FMa Anemochory

STG, B Anemochory
STG, TAHa Anemochory
STG, TAHa Anemochory
SAH, B Anemochory
SAIL B Anemochory

266

The Victorian Naturalist

Research Reports

Appendix 1 continued.

Species Common Name Growth Form Habitat Dispersal
Rubiaceae
149 Asperula gunnii Mountain Woodruff Herb TAHa, STG, H Zoochory *
150 Asperula pusilla Alpine Woodruff Herb FAHa, STG, H Zoochory*
151 Coprosma niphophila Snowpatch Coprosma — Mat/Subshrub FMb Zoochory *
152 Nertera sp. Matted Nertera Herb STG.F, B Zoochory*
Campanulaceae i
153 Wahlenhergia ceracea Waxy Bluebell Herb TAHa, STG Anemochory
154 Wahlenbergia gloriosa Royal Bluebell Herb TAHa Anemochory
Lobeliaceae
155 Pratia surrepens s.l. Mud Pratia Herb PF, B, STG Zoochory*
Goodeniaceae
156 Goodenia hederacea subsp. Ivy Goodenia Herb VAHa Zoochory**
alpestris
Stylidiaceae
157 Stvlidium sp. (alf. Alpine Trigger Plant Herb TAHa, STG, H, B Anemochory
graminifolium)
Asteraceae
158 Abrotanella nivigena Snow-wort Mat/Herb SAT Anemochory
159 Brachyscome nivalis Snow Daisy Herb TAHb Anemochory
160 Brachyscome obovata Baw Baw Daisy Herb LAHa Anemochory
161 Brachyscome scapigera Tufted Daisy Herb I AHa, STG, H, B Anemochory
162 Brachyscome spathulata Spoon Daisy Herh TAHa, STG, FMa Anemochory
subsp. spathulata
163 Brachyscome stolonifera e — Gwenda's Daisy Herb SAH, TAHa, STG Anemochory
164 Brachyscome sp. (alf. Creeping Daisy Herb SAH, TAHa Anemochory
tadgellii)
165 Brachyscome sp. (aff. Mountain Daisy Herb STG, TAHa Anemochory
fenuiscapa)
166 Celmisia costiniana Herbfield Celmisia Herb TAHa, STG Anemochory
167 Celmisia pugioniformis Dagger-leaf Celmisia Herb TAHa, STG Anemochory
168 Celmisia tomentella Bog Celmisia Herb B Anemochory
169 Cotula alpina Alpine Cotula Herb B, TAHa, STG Anemochory
170 Craspedia alba Dwarf Billy-button Herb STG Anemochory
171 Craspedia aurantia Orange Billy-button Herb H, TAHa Anemochory
172 Craspedia costiniana e Hairy Billy-button Herb TAHa, STG Anemochory
173 Craspedia jamesii James's Billy-button Herb lAHa, STG Anemochory
174 Craspedia lamicola e Shiny-leaf Billy-button Herb TAHa, STG Anemochory
175 Craspedia leucantha Pale Billy-button Herb SAH, TATa Anemochory
176 Craspedia maxgrayi Woolly Billy-button Herb TAHa, STG Anemochory
177 Craspedia sp. B Sticky Billy-button Herb TAHab, FMa Anemochory
178 Erigeron bellidiodes Violet Fleabane Herb STG, TAIHa Anemochory
179 Erigeron nitidus Sticky Fleabane Herb STG, TAHa Anemochory
180 Erigeron paludicola Bog l'leabane Herb B Anemochory
181 Erigeron setosus » Dwarf Fleabane Merb SAH Anemochory
182 Euchiton argentifolius Silver Cudweed Mat/Herb TAHa, STG Anemochory
183 Euchiton fardianus Ford's Cudweed Herb VAHa, STG Anemochory
184 Euchiton nitidulus Shining Cudweed Mat/Ierb STG Anemochory
185 Euchiton poliochlorus Swamp Cudweed Herb B Anemochory
186 Euchiton umbricola Cliff Cudweed Herb TAHb Anemochory
187 Ewartia nubigena Silver Ewartia Mat/Subshrub FMa, TAHa Anemochory
188 Helichrysum scorpioides Button Everlasting Herb TAHa Anemochory
189 Lagenophora stipitata Blue Bottle-daisy Herb STG, TAHa Anemochory
190 Leptorhynchos squamatus s.|. Scaly Buttons Herb STG, TAHa, FMa Anemochory
191 Leucochrysum albicans Alpine Sunray Mat/Herb 'TAHa, FMa Anemochory
subsp. alpinum
192 Microseris lanceolata Native Dandelion Herb TAHa, STG Anemochory
193 Olearia algida Alpine Daisy-bush Shrub H Anemochory
194 Olearia phlogopappa var. Dusty Daisy-bush Shrub H Anemochory
flavescens
195 Olearia phlogopappa var. Dusty Daisy-bush Shrub H Anemochory
subrepanda
196 Ozothamnus alpinus Alpine Everlasting Shrub B, H Anemochory
197 Ozothamnus sp. (aff. hooker?) Kerosene Bush Shrub B, H Anemochory
198 Ozothamnus secundiflorus Cascade Everlasting Shrub H Anemochory
199 Parantennaria uniceps Snowpatch Daisy Mat/Herb SAH, STG, B Anemochory
200 Picris angustifolius subsp. Mountain Picris Herb TAHa Anemochory
merxmuelleri
201 Padolepis robusta Alpine Podolepis Herb VAHa, STG Anemochory
202 Rhodanthe anthemoides Chamomile Sunray Herb TAHa Anemochory
203 Senecio gunnii Gunn’s Groundsel Herb TAHa Anemochory
204 Senecio pinnatifolius var. Highland Groundsel Herb TAHa, STG Anemochory
pleiocephalus
Alpine Groundsel Herb TAHa, FMa Anemochory

205 Senecio pectinatus var. major

Vol. 119 (6) 2002

267

Research Reports

Appendix 2. Introduced species well established in the Kosciuszko alpine area. Taxa from Costin ef

al. (2000).

Species Common Name Probable Dispersal Agent
Poaceae

Agrostis capillaris Brown-top Bent Anemochory
Dactylis glomerata Cocksfoot Anemochory
Festuca rubra Red Fescue Anemochory
Phleum pratense Timothy Grass Zoochory

Poa annua Annual Meadow-grass Anemochory

Poa pratensis Kentucky Blue-grass Anemochory

Caryophyllaceae
Cerastium vulgare (syn. Cerastium
Jontanum subsp. trivale)

Common Mouse-ear Chickweed

Anemochory

Spergularia rubra Red Sand-spurrey Anemochory

Polygonaceae

Acetosella vulgaris (syn. Rumex Sheep Sorrell Anemochory
acetosella)

Fabaceae

Trifolium ambiguum Pellet Clover Anemochory

Trifolium repens White Clover Anemochory

Asteraceae

Achillea millefolium Milfoil, Yarrow Anemochory

Hypochoeris radicata Cat's-ear Anemochory

Taraxacum officinale sp. agg. Dandelion Anemochory

One Hundred Years Ago

On The Fertilization of Phanerogams
I.—DISPERSION OF POLLEN BY THE WIND.
By G. WEINDORFER
(Read before the Field Naturalists’ Club of Victoria, \3th July, 1902.)

For the conveyance of pollen between flowers there exist two main agents, viz., the wind and
insects, Phanerogamous plants have been separated by botanists into "anemophilae," or wind-fer-
tilized, and “entomophilz,” or insect-fertilised plants.

As would be naturally expected, it is, speaking generally, only pollen of a dusty consistency
which is transported by the wind; but the pollen of some flowers is occasionally torn away from
the anthers, in the form of sticky masses, and conveyed to the stigmas of neighbouring flowers
by the wind, but the occurrence can only be looked upon as accidental, and would happen in the
rarest instances. Still more remarkable is the fact that in certain water plants the pollen, though
cohering in sticky masses, is blown by the wind in a kind of little boat to the stigmas, which are
raised above the surface of the water.

From The Victorian Naturalist XIX, p 98, November 6, 1902

Vale

Club Members will be saddened to learn of the recent death of the follow-
ing members:

Natalie Smith in July

Elsie Costermans in August

Eric Allen in September

Each made a great contribution to natural history and individual tributes
will appear in later issues.

268 The Victorian Naturalist

Research Reports

The Red Fox Vulpes vulpes L. in the Dandenong
Creek Valley: Impacts on Wildlife and an Integrated Program
for their Control

A Morton', D Tagg', R Wallis? and J White!

Abstract

Foxes in the Dandenong Creek Valley were found to consume many native vertebrate species,
including three mammals, thirteen birds and at least one lizard. As well, foxes spread weeds by dis-
persing viable Blackberry seeds in their scats. A Co-ordinated Fox Control Committee representing
eight landholders in the study site oversees the continuing project on land which includes many
parks, four golf courses. a freeway easement, a water retarding basin, a quarry, two waste transfer
stations and small blocks used for agriculture and stock agistment, Recommended control methods
include den fumigation and removal of diurnal weed shelter. (The Victorian Naturalist 119 (6), 2002, 269-

275)

Introduction

The Red Fox Vulpes vulpes L. is an intro-
duced canid predator that is now wide-
spread across the southern half of the
Australian continent, Fox predation is con-
sidered a major threat to the conservation of
native Australian fauna, especially ground
nesting birds and those mammals in the crit-
ical weight range of between 35 and 5500 g
(Burbidge and McKenzie 1989; Saunders e/
al. 1995). Predation by foxes is listed as a
key threatening process under both Federal
and Victorian Government legislation and
appropriate plans have been devised to
address the threat to wildlife (Mansergh and
Marks 1993; Environment Australia 1999).
Foxes can also be serious pests in agricul-
tural areas, preying heavily on lambs and
poultry in particular and are a potential vec-
tor for the transmission of rabies (Saunders
etal. 1995).

Foxes are opportunistic in their diet, tak-
ing a variety of vertebrates, insects and
vegetation. A number of dietary studies
have been conducted in rural Australia in
both forested and agricultural settings (see
Brunner and Wallis 1986), but there have
only been a few reports published on fox
diet (Wallis et al. 1996) or on fox densities
in urban environments in Australia (Marks
and Bloomfield 1999).

In this report we describe a co-ordinated
fox control program conducted by eight
land manager/landholder groups in the
Dandenong Creek Valley in Melbourne's

School of Ecology and Environment, Deakin
University, Melbourne Campus, 221 Burwood
Highway, Burwood, Victoria 3125

School of Ecology and Environment, Deakin
University, Warrnambool, Victoria 3280

Vol. 119 (6) 2002

eastern suburbs. The program relied on the
establishment of a committee that oversaw
a study into the diet, habitat use and con-
trol of foxes conducted by staff and stu-
dents from Deakin University's School of
Ecology and Environment. We also report
on ‘pre-control’ data gathered to date in
this four-year program and an outline of
planned future activities.

Methods
Study site

The study site in the Dandenong Creek
Valley is approximately 13 km long and 1-3
km wide. Situated in Melbourne's eastern
suburbs, it extends from north of Boronia
Road, Wantirna, to Stud Road, Rowville, in
the south (Fig. 1). Dandenong Valley
Parklands, managed by Parks Victoria,
forms the largest portion of the study site
and combines 1325 ha of popular recreation
and parkland settings with conservation
areas and farmland. Managed parks include
Koomba Park, Bushy Park Wetlands,
Nortons Park, Shepherds Bush, the Linear
Trail, Jells Park and Chesterfield Farm.

Other agencies that manage land in the
study site and are members of the Co-ordi-
nated Fox Control Committee are the Cities
of Monash, Knox and Whitehorse, Vic
Roads (easement of the Scoresby Freeway),
Melbourne Water (Winton Wetlands north
of Boronia Road and the Police Road
Retarding Basin between Wellington Road
and Stud Road), Boral Bricks (which man-
ages a quarry and a farmland) and Kingston
Links Golf Course. The study site includes
four golf courses: Kingston Links,
Tirhatuan, Waverley Municipal and

269

Research Reports

Dandenong
4 Creek
Boronia Rd Jc inten Wetlands
Vermont d
Morack Golf (/
Gauss Koomba Park
Burwood Hwy. C Mountain Hwy
Bushy Park Wantirna South
Wetlands ie
Glen Waverley
High Street Rd

Shepherds Bushy"

Waverley Golf
Course ~

Proposed
Scoresby
Freeway

Waverley Road

Corhanwarrabul
Ferntree Gully Rd

Corhanwarrabul

Creek Section * Kingston Links

~ Golf Course

Wellington Rd

Rowville

Dandenong
Creek

Scale (km)

Fig. 1. Dandenong Creek Valley forms the bor-
der between the municipalities of Whitehourse
and Monash to the west and Knox to the east.

Morack. Two waste transfer stations are
present and large blocks of Parks Victoria,
Vic Roads and Melbourne Water land are
used for horse agistment, cattle grazing and
fruit and flower growing. We believe such
co-operation between all landholders in fox
control in a semi-urban setting is unique
and a necessary condition if successful
management of foxes is to be achieved.

The study area is fringed on all sides by
residential and factory development and
some open land. Dandenong Creek has been
severely modified over time and is prone to
heavy flooding up to three times per year.

Study design and methods
The project began in 1998. In the first

two years the aims were to:

+ describe the diet of foxes in the area to
determine the effects on wildlife;

- describe the potential for foxes to spread
Blackberry Rubus fruticosus spp. agg. via

270

seed germination from their scats;

e survey for sites of weed harbour suitable
as refuge for foxes;

* survey the site for foxes and dens;

+ review methods suitable for urban envi-
ronments for the control of foxes.

Fox seats were collected weekly from
January 1998 till January 1999 from
Koomba Park and Boral Bricks sites. As
well, less frequent, opportunistic collecting
occurred over this period elsewhere, with
monthly collections from transects estab-
lished in the Police Road Retarding Basin,
Corhanwarrabul Section, Jells Park,
Shepherds Bush, Bushy Park Wetlands,
Morack Golf Course, Kingston Links Golf
Course and Drummies Bridge Reserve (Fig.
1). Scats were identified by size and shape
(Triggs 1996), collected in manila
envelopes, sterilised, washed, sorted and the
food items identified and categorised using
techniques described in Brunner and Wallis
(1986) and Wilson and Wolrige (2000).
Feathers found in the scats (and in den litter)
were identified against specimens held at the
Bird Observers Club of Australia premises;
mammal bones were identified using speci-
mens at Deakin University.

Fox scats containing Blackberry seeds were
collected over a 6-week period during
February and March 1998. They were
washed in water with a 10% solution of com-
mercial fungicide to inhibit fungal growth,
Blackberry seeds were placed onto filter
paper in petrie dishes and kept in a germina-
tion cabinet at 18°C over a 9-month period,
where they were kept moist and monitored
for seedling development. Some of these
seeds were kept at 25°C for a period, Another
group of seeds was first treated at 0°C and
4"C for a 3-week period before being placed
into the germination cabinet at 18"C.

Approximately half of the seeds were
tested for viability using tetrazolium chlo-
ride. Tetrazolium salts are used to indicate
the presence of dehydrogenase enzymes,
which provide reduction processes in liv-
ing tissue. Dehydrogenase enzymes reduce
tetrazolium salts to red or blue substances
which appear as stains on viable seeds.
Dead tissues remain unstained as there are
no active enzymes present (MacKay
1972). The Blackberry seeds were placed
in a 0.1% tetrazolium chloride and buffer
solution at neutral pH and the number of

The Victorian Naturalist

stained seeds counted to estimate viability.
In December 1998 the remaining seeds
(those not used in viability testing) were
placed into potting mix in four pots which
were exposed to normal sunlight and rain.
There was no additional watering.

In April 1998 six scats containing
Blackberry seeds were planted directly into
potting mix. Fifty seeds from four of these
were washed in a 10% fungicide solution
and individually placed in the potting mix.
The other two scats were not treated, but
were planted whole into the pot. The pots
were then placed in a glasshouse where they
received regular watering and monitoring.

The public was kept informed about the
project through frequent articles in the
local press and talks to local resident
groups. As well, 1200 flyers were distrib-
uted to households around the study site
which sought reports from residents seeing
foxes - people were invited to phone a
"fox hotline’ at the university.

Each management section was systemati-
cally searched for den sites, Dens were
classified as active if foxes were actually
observed using them, if there were obvious
signs of fox prints in freshly raked sand
placed near the entrances or if there was
den litter nearby (Coman 1972). The pres-
ence of scats and flattened grass around a
den also provide good evidence of activity
(Macdonald 1987), Dens and weed har-
bour patches were mapped using a GPS;
volunteers, Green Corps personnel, student
classes and Parks Victoria staff all assisted
in these tasks.

The recommendation of appropriate con-
trol measures for each of the eight organi-
sations was an aim of this research. These
recommendations took into account cur-
rent legislation, community concerns and
the safety of domestic animals, Fumigation
of den sites using Denco-fume carbon
monoxide cartridges was selected as the
most humane and species-specific method
of control in an urban environment,

Results and Discussion
Diet studies

A total of 719 scats was collected from
seven areas. Bones were the most frequent-
ly detected food item (56% of all fox scats
examined contained bones). These includ-
ed bones of mammalian and avian prey as

Vol. 119 (6) 2002

Research Reports

a
Table 1. Occurrence of specific mammals in
219 fox scats containing hair collected over one
year trom seven major locations in the
Dandenong Valley Parklands and surrounding,
sites. Percentages in parentheses, * denotes
introduced species. Fox grooming hairs ignored.
Hrs ris e aae Ik ia See id indc aaa ea ach Ld d

Species Number of
occurrences
European Rabbit/Brown Hare 83 (29)

Oryctolagus cuniculus:
Lepus eurapaeus*
Common Ringtail Possum
Pseudocheirus peregrinus
Black Rat Rattus rattus*
Common Brushtail Possum
Trichosurus vulpecula
House Mouse Mus domesticus*
Brown Rat Ratrus norvegicus*
Sheep Ovis aries*
Sugar Glider Petaurus breviceps
Cattle Bos taurus
Cat Felis eatus*
Total seats with hair 29

71 (24)

56 (19)
40 (14)

— 4

-—— [304 cC i

well as those which probably came from
garbage. Hair was found in 40% of scats.
Other frequently found items included
invertebrate (mainly insect) remains
(55%), plant material (non-seed) (5394),
seeds (41%) and feathers (28%). Of the
298 scats with seeds, 168 had Blackberry,
all of which were found in summer/early
autumn. Most invertebrate remains detect-
ed in scats were collected in spring, sum-
mer and autumn. On the other hand, feath-
ers occurred in roughly similar percentage
frequencies in all seasons.

Table | shows the European Rabbit
Oryctolagus cuniculus as the preferred food
item followed by the Common Ringtail
Possum Pseudocheirus peregrinus. Three
scats contained the remains of the locally
rare Sugar Glider Petaurus breviceps.
Remains of birds were found in 204 scats
and 17 species of birds were identified
from their feathers (Table 2). Four of these
species were also found as den litter,

Foxes in the Dandenong Creek Valley
were shown to consume a wide variety of
prey. Mammals, birds and invertebrates
(chiefly insects) are taken together with
seeds such as Blackberry and fruit. Whilst
rabbits, rats and mice appear frequently in
fox seats, three native mammal species are
eaten. The appearance of Sugar Glider
remains in fox seats, whilst rare, is of

271

Research Reports

Table 2. Bird species taken as prey by foxes in
the study area. Feathers found in fox scats were
compared with those from specimens. * indi-
cates species also found as “den litter’ around a
natal den in Jells Park between September and
November 1998. * denotes an introduced
species.

Common name

Australian White
Ibis
Purple Swamphen
Australian Wood
Duck
Pacific Black Duck
Silver Gull
Buff-banded Rail
Sulphur-crested
Cockatoo
Platycercus eximius" Eastern Rosella
Trichoglossus haematodus Rainbow Lorikeet
Accipiter fasciatus Brown Goshawk
Grallina cyanoleuca Magpie-lark
Gymnorhina tibicen* Australian Magpie
Corvus mellori Little Raven
Streptopelia chinensis* Spotted Turtle-Dove
Acridotheres tristis * Common Myna
Sturnus vulgaris* Common Starling
Turdus merula** Common Blackbird

Scientific name

Threskiornis molucca

Porphyrio porphyrio”
Chenonetta jubata

Anas superciliosa
Larus novaehollandiae
Gallirallus philippensis
Cacatua galerita

concern. This species is highly vulnerable
to predation by cats and foxes and its dis-
tribution in urban areas is rapidly declining
(Brunner et al. 1991). Dandenong Valley
Parklands is one of the closest sites to
Melbourne inhabited by Sugar Gliders.

Scats collected from different sites in the
study area had different proportions of prey.
For example, hair from Sugar Gliders was
only found in scats collected from Koomba
Park, while scats from Boral Quarry had
higher numbers of feathers and bones than
other areas. Although these feathers were
not identified, both Boral Quarry and the
proximate Knox Tip supported wetland bird
populations. A large population of scav-
enger birds including Silver Gull Larus
novaehollandiae and Australian White Ibis
Threskiornis molucca was present at the
Tip. Many of the bone fragments were from
domestic rubbish accessible to foxes at the
open landfill site of Knox Tip.

Rabbits and Brown Hares Lepus europaeus
were common in fox scats collected at Boral
and the Police Road Retarding Basin. Our
data are similar to those reported in Wallis ef
al. (1996) for fox scats collected in 1989 and
1990, although the proportion of scats

272

containing the remains of rabbits has
increased significantly.

As well, foxes are likely to prey on frogs
and lizards — one Blue-tongued Lizard
Tiliqua sp. was found with its skin
removed beside the entrance of a fox natal
den. Wallis et al. 1990 reported that
Dandenong Valley Parklands represents a
linear reserve of modified bushland in a
‘sea’ of suburbia providing largely unsuit-
able habitat for sustainable wildlife com-
munities. They compared the diet of foxes,
dogs and cats in the Dandenong Valley
Parklands and from the evidence found
that predation by these three introduced
species threatened the continued survival
of some native vertebrates in the
Parklands.

Germination trials

Those seeds tested for viability with tetra-
zolium chloride showed a positive response
with 120 of 391 seeds (or 31%) showing
viability. However, there was no germina-
tion from seeds whilst in the germination
cabinet. In December 1998 the remaining
seeds were planted into potting mix. In
August 1999 these seeds germinated with
166 seedlings growing from a total of 660
seeds, a germination rate of 25%, 18 months
after being deposited by the foxes (Table 3).

In comparison, the scats that were planted
directly into potting mix in March 1998
germinated in August 1998 just five
months after deposit. The seats planted
whole germinated at a much faster rate than
the individual seeds. Of 50 individual seeds
planted into each of four pots only 29 ger-
minated — a maximum of 15% germination.
Growth was uneven with one pot yielding
32%, but the others only 12%, although
seedling growth in all four was strong and
vigorous. In the other two pots planted with
whole scats, many more seedlings germi-
nated — 32 and >50 — but as the number of
seeds planted were not counted no germi-
nation rate could be calculated. Many of
these seedlings were still present in 1999
although growth was limited by the lack of
resources in the pot. Once these seedlings
were removed more seedlings germinated
in August 2000. One whole scat planting
had 43 new seedlings present.

We have demonstrated that Blackberry
seeds in fox scats are viable and capable of

The Victorian Naturalist

Research Reports

Table 3. Treatments of various groups of Blackberry seeds collected in fox scats during

February/March 1998. GC, germination cabinet; R/F, refrigerator/f

Group A

Date collected — Feb/Mar ‘98
Ist treatment GC at 18°C
2nd treatment — -

Group B
Feb/Mar *98
GC at 18°C
Tested for

viability

Jul *98
Planted Dec ‘98 Dec *98
Germination Aug ‘99 Aug *99

ezer.

Group C Group D Group E

Feb/Mar ‘98 Feb/Mar ‘98 Mar ‘98

GC at 18°C R/F 3 weeks =

GC at 25°C GC at 18°C -
Aug/Sep *98

Dee *98 Dec '98 Mar *98

Aug *99 Aug ‘99 Aug “98,

Aug ‘00

germination, Fox scats may contain more
than 50 seeds, which remain viable for at
least two years in storage and pots. Since
foxes may travel up to 10 km per night
(Saunders et al. 1995), there is potential
for widespread dispersal of Blackberry
throughout this area. Blackberries may
have a laxative effect on foxes as there are
many more scats collected during the
Blackberry fruiting season than at other
times of the year. Blackberries provide an
easy resource for foxes and require mini-
mal energy expenditure to harvest them.

Blackberry appears to benefit from dis-
persal by foxes. The whole scats planted in
potting mix without other treatment pro-
duced higher germination rates than for
seeds that were washed in fungicide. Seeds
from the germination cabinet showed no
germination until they were planted into
potting mix, while none of the seeds, from
the germination cabinet or direct sowing,
germinated before late August. This may
indicate that the length of daylight pro-
vides the trigger for growth. The nutrient
package provided by the fox may also
improve the number of seeds germinating.

Brunner et al. (1976) collected Blackberry
seeds from scats of foxes and found germi-
nation did not commence until early Spring,
some six months after they had been col-
lected. Overall they found germination rates
of between 22% and 35%. Our conclusions
concur with those of Brunner et al. (1976)
and Amor and Richardson (1980) who con-
cluded that foxes are a major dispersal agent
for Blackberry in Victoria, particularly in
sites which are prone to flooding after
heavy rainfall.

Weed mapping

The presence of Blackberry, Gorse Ulex
europeaus and Wandering Tradescantia
Tradescantia fluminensis was mapped and
the maps used by land managers to control

Vol. 119 (6) 2002

infestations. Other weeds were also
mapped if it was thought they were in such
high densities that they might provide
cover for foxes. Many organisations
involved have since carried out weed con-
trol and in some places have followed with
landscaping and revegetation programs.
'These and other weeds found are declared
noxious weeds and pest plants in the study
area (Knox City Council and Maroondah
City Council 1999).

Fox Hotline

Some 46 people telephoned the ‘lox
Hotline’ reporting sightings, although
some were for foxes seen outside the study
area. Interestingly, a number expressed
concern for the safety of their own pets,
including aviary birds, from fox predation.
Others were concerned with the noise
made by foxes.

Density estimate

Marks and Bloomfield (1999) used natal
den counts to estimate family group density
in Melbourne's suburbs. They found foxes
living, and in many cases breeding, in all
parks larger than 20 ha within 10 km of the
GPO. Foxes were especially common in
parkland associated with creeks and rivers.
Our study site was most similar to their
Box Hill/Camberwell site in terms of hous-
ing, parkland and presence of thick vegeta-
tion and weed infestations used by foxes
for diurnal shelter. An average of 1.13
independent active natal dens/km with a
mean distance to the nearest neighbour of
1.09 km was found in that area, The natal
dens in our study area were distributed in a
similar fashion. Furthermore, Marks and
Bloomfield (1999) found a mean of 4.36
cubs produced per den, a figure which is
comparable to urban den litter sizes record-
ed for foxes elsewhere (Harris and Smith
1987). Typically in Australia the predomi-

273

Research Reports

nant social group consists of an adult
breeding pair and cubs with perhaps one or
two non-breeding vixens (often young of
previous litters). However, in sites that
have high resource availability (food and
shelter), workers have observed polyga-
mous social groups (Macdonald 1979) or
tolerance of the presence of non-breeding
males (both itinerant and non-dispersing)
(Lovari et al. 1994).

Using these numbers and an area of 16.5
km' it is reasonable to estimate that there
may be approximately 80 foxes living in
the study area. Given the observations
mentioned in overseas studies this may be
à conservative estimate with numbers vari-
able across the year.

Fox control

There are few options available in urban
areas for fox control. The poison 1080
(monosodium fluoroacetate) is one of the
most selective and safe means of fox control
and has been used successfully in Western
Australia where native species have a higher
tolerance to the poison (Mellroy 1986; King
and Kinnear 1991). Fox poisoning has been
especially successful when baits have been
buried in bait stations (Saunders ef al. 1995),
Poisons like 1080 and cyanide cannot be
used in urban sites because of the risk to
non-target species at bait stations or cache
sites. Foxes are known to cache food and
might carry a bait up to 800 m before
caching it. Domestic dogs could be at risk if
a fox were to cache a bait in a residential
area (Kay et al. 1997).

Similarly, shooting was ruled out because
of the dangers involved in using firearms
in built-up areas. Trapping with soft-jawed
traps is an inefficient means of fox control
(Saunders er al. 1995; Kay et al. 2000),
The only practicable method for fox con-
trol in the Dandenong Valley Parklands
and surrounding sites was for natal den
fumigation and destruction and removal of
weed harbour used as cover by foxes.

To date, 33 fox dens have been located,
although not all were being used in the
study period. Five were used as natal dens,
whilst six others showed signs of recent
use. Family groups tended to use multiple
dens and den sites were regularly dispersed
throughout the study site. Many dens were
found under the cover of weeds such as
Blackberry, Gorse and Wandering

274

Tradescantia. Other dens were found in
mounds in the open with introduced grass
cover, under trees with introduced or nat-
ural grass cover, in drains or dug into dam
walls (with or without overstorey cover).
The most common factor for den sites was
the use of a mound or earth wall which
provided easy digging for the fox.

A professional pest controller was
engaged to supervise den fumigation and a
total of 29 of the 33 possible sites was
checked by his dogs. Nine sites were
selected for fumigation from the amount of
interest shown in the den by the dogs. Two
other sites were unable to be fumigated
due to the complete Blackberry cover.

Two weeks later, three fumigated sites
were opened by a backhoe to assess the
success of the control exercise. Two sites
had been reopened by foxes since the
fumigation, one of which was still in use as
a natal den. One cub was dispatched at this
site. Six other sites were unable to be back-
hoed due to the nature of the surrounding
vegetation or terrain. Three of these were
in sensitive remnant vegetation or were
dug into dam walls. There was no access
for the backhoe at the final three sites.

Fumigation is neither as cost-effective
nor as efficient as 1080 baiting. Locating
fox dens is labour-intensive and time-con-
suming and fumigation must be carried out
within the first eight weeks after the birth
of the cubs. After thís time both vixen and
cubs leave the den. The dog fox and any
sub-dominant females do not use the den
but may support the vixen and cubs with
the supplv of food (Macdonald 1979). At
best, fumigation will remove only one gen-
eration of foxes, whilst leaving other adult
foxes to continue the attack on native
mammals and birds.

This study has shown that foxes will
reopen den sites that have been closed for
fumigation if the site is not levelled with a
backhoe. Unfortunately, not all sites are
accessible to a backhoe as they were locat-
ed in important remnant vegetation. These
sites are then available for reuse by foxes.

Although studies indicate that fumigation
may reduce predation by 92% (Hewson
1986) this would assume that no new foxes
could inhabit the area. Marks and
Bloomfield (1999) have estimated that
there may be as many as 16 foxes/km? in
residential areas. Once an area becomes

The Victorian Naturalist

vacant due the effects of a fox control pro-
gram there will be immigration from areas
of greater fox density. For these reasons
fumigation cannot be considered by itself
as a successful method of fox control.

The manipulation of habitat might
become the means of keeping fox numbers
to a minimum in the Dandenong Creek
Valley. This would necessitate the removal
of Blackberry and Gorse, which provide
food and harbour resources for foxes, and
some of their favoured prey species, rab-
bits and rodents. Blackberry also provides
harbour for these prey; the removal of
Blackberry and other weed species would
support a program of fox control and aid in
the removal of rabbits and rodents. A con-
tinuation of this study will consider what
effect habitat manipulation will have on
the success rate of a control program.

The next stage

Follow-up fox surveys will be carried out
in the next phase of the project after weed
harbour and den destruction. As well, we
will continue to collect and process fox
scats to monitor change in diet over the
seasons. The Co-ordinated Fox Control
Committee which oversees the project was
recently successful in receiving funding
from the Natural Heritage Trust to com-
pare fox behaviour, use of daytime habitat
and home range in sites where weed har-
bour has been removed with a control site.
We hope this will provide insights into
how fox activity and movements change in
response to habitat modification.

Acknowledgements

We thank members of the Dandenong Creek
Valley Co-ordinated Fox Control Committee for
their support and advice, the National Feral
Animal Control Program (NFACP) for funding,
Hans Brunner for advice and help with hair identi-
fication and Clive Marks for expert advice. We
also thank those who helped with weed mapping
and den location: Mick Van de Vreede, Craig
Lupton, làn Morrisson and Gary Lavlor (Parks
Victoria), the Dandenong Valley Parklands Green
Corps group and students from Deakin University.

References

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275

Naturalist in the Mountains

Flying Spiders and Crawling Flies

December has come and the new year is
fast approaching; with summer settling in
it tends to be a quiet time in the mountains
for the bigger wildlife. The birds have set-
tled down, most of the migratory birds
having arrived in the mountains for the
summer in September or October; breeding
is past the raucous courtship phase and
parents are busy feeding their young. Even
the Crescent Honeyeaters Phylidonyris
pyrrhoptera have quietened down.

The small mammals, Dusky and Agile
Antechinus Antechinus swainsonii and A.
agilis, and Mountain Pygmy Possum
Burramys parvus, are carrying pouch
young. The female antechinuses are hard
to cateh, while the males are long gone,
and at higher altitudes the young are diffi-
cult to catch until some time into February.
The Bogong Moths Agrotis infusa are in
their aestivation sites and not going any-
where in a hurry. At this time the flower-
ing plants have it all their own way with
the tourists, while the more obvious insects
in your field of view tend to be the pollina-
tors: butterflies, wasps and bees.

But summer days drift on by to the sounds
of everyone's least favourite, fies: March
Flies, bushtlies, blowflies, mosquitoes and
the knuckle-biting black flies — these are the
ones that grab all the attention. Out of the
limelight of the tourist-packed Main Range
with its multitude of flowering plants and
butterflies, other animals are equally busy.

Everywhere spiders and flies are creeping,
Creeping flies? The Soldier Fly Boreoides
subulatus (Stratiomyidae) is one of the
insects most commonly brought to entomol-
ogists for identification and one which peo-
ple only have to begin describing at this time
of year for me immediately to know what
they are talking about. This strange looking
fly, in which the wingless female is many
times the size of the male, looks vaguely
threatening, Not so vaguely to some, who
think they have been attacked by one when
it tumbles out of a tree on top of them. At
the lower altitudes of Canberra, there is a
spring and an autumn brood, whereas in the
snow-country Boreoides has just one breed-
ing season per year. The female emerges
already gravid and is mated by more than
one of the winged males. Afier mating, the
females climb — up a post, a tree, a tent pole,
it doesn't matter — looking for a hole in

276

which to lay their eggs. They can be found
with ovipositor buried in a hole carved out
by beetle or moth larvae. The female does
no digging herself but in some holes half of
her body might disappear from view. Any
hole will do. I have had them lay eggs in a
fence post at home or in the apex of my tent
way back when | had an A-frame tent. The
female seems to lay all her eggs in the one
hole; certainly she lays a lot. The young will
later descend to live in the soil.

Also creeping at this time (and the reason
that years ago I went off the A-frame tent
without the floor), are male Funnelwebs,
one of the two still-undescribed species of
Hadronyche (Hexathelidae) found in the
mountains, It is at about this time that the
males go in search of the females, who
remain in their holes or beneath an exfoliat-
ing slab of granite, For most people, the evi-
dence won't be there until January when
bodies of male Funnelwebs litter the alpine
grassland. But at night the grasslands
abound in them, to the extent that going out
of the tent in bare feet in December can be a
nerve-racking activity. For me it was at
night in my floorless tent, with a Funnelweb
glistening by candlelight on my pillow
beside me as I read my book. A torch-lit
search outside revealed another 13 males
within a couple of metres of the tent. A tent
with sewn-in floor was purchased soon
after! The density is astounding, and the
holiday season is the time when a number
of bites (none yet fatal) occur.

I have recorded flies among a horde of
insects, moths, and wasps migrating across
Dead Horse Gap (a Mecca for anyone inter-
ested in migration of birds or insects; see
Green 1982), but spiders also migrate. On
summer days the gossamer threads, 2-3 m
long, of spiders floating through the air can
be seen against the sun. Young spiders use
this technique, called ballooning, for disper-
sal from nest sites. When ready for migra-
tion, the abdomen is raised and silk extrud-
ed, drying and being caught by the wind
which finally lifts the spiderling from the
ground, sometimes to heights in excess of
4000 m. The lines of silk on the ground are
often the results of failed take-off attempts.
Malaise traps (normally set to catch flying
insects) set in the alpine zone between the
top of the Thredbo chairlifts and Mt
Kosciuszko during the “Biodiversity Blitz

The Victorian Naturalist

in January this year (Green 2002) collected
over 100 such dispersing juvenile spiders.
About 90% of the specimens collected in
the malaise traps were thomisids
(Thomisidae), comprising what appears to
be two species of Diaea. Despite all this
gossamer, aerial web building spiders are
uncommon in the mountains in comparison
to lower altitudes. The short summer sea-
son, strong winds and snow probably make
the capture of aerial food with a web a little
problematie, and most spiders rely on ter-
restrial hunting. But spiders have no diffi-
culty getting to the mountains or even being
some of the earliest colonisers of new
mountains or newly exposed parts of old
mountains, as snow and glaciers retreat.
Ballooning is their major mode of transport
and they are able to exist even above the
limit of plant growth. This is what is called
the aeolian zone, where life forms exist on
the material brought in by the wind (a fairly
obvious phenomenon from the Snowy
Mountains to Canberra in October this year
when a fair bit of the cropland out west
came visiting). At times, on high moun-
tains, spiders may be the only life form pre-
sent (except man). Leslie Stephen wrote in
1874 ‘The only creature besides myself that
could fairly be called living was a small
black spider, which had been led by an
apparently misguided spirit of enterprise to
seek for prey in this loftiest zone of organic
existence' (4 Bye Day in the Alps).

Of the nearly 70 families of spiders
found in Australia, only 27 have been
recorded above the winter snowline (1500
m) in the Snowy Mountains, and 20 of
those are only known from one or two
species (Green in press). The best known
are the Lycosidae (Wolf Spiders) where
half of the eight known species have been
described. In the herbfields and grasslands,
burrows of Wolf Spiders are common, and
you can sometimes find a burrow every
one or two square metres. In December,
the female Wolf Spiders sit at the entrance
to their web-covered burrow, holding on to
their egg case — scuttling back inside when
danger arrives, Later in the summer they
can be seen with young on their backs.
Sometimes when you disturb them on a
sunny day you can see hundreds of spider-
lings scuttling back to their natal burrow
on your approach.

Vol. 119 (6) 2002

Naturalist in the Mountains

One thing that constantly keeps me on my
toes in the mountains is the appearance of
the unusual among the mundane. Bogong
Moths are well known for their gregarious
aestivation in cracks among the rock tors
and blockstreams. During my studies of the
moths and associated arsenic last year
(Green er al. 2001) I found the relative
humidity in à bogong moth site over sum-
mer averaged around 75% with the average
air temperature below 8°C. These cool
damp conditions also attract blowtlies, ever-
present in the snow-country — even in win-
ter — and small flies. These join the giant
springtails and other detritivores crawling
around on the Bogong Moth debris on the
floors of caves. All this (apart from the
arsenic) was expected, but as | checked a
cave on one hot day 1 found a particularly
fuzzy rock. On closer examination it was
covered in hundreds, maybe thousands, of
small flies, lined up side by side, end to end,

just like Bogong Moths. These turned out to

be Fungus Gnats Exechia sp. (Myceto-
philidae) which are often associated with
damp places, including caves, as are their
relatives the glow worms. Were these
Fungus Gnats just taking advantage of the
cool moist environment to avoid the dessi-
cating effects of altitude and alpine sun? Or.
like the two species of parasitic nematodes
Amphimermis bogongae and Hexamermis
cavicola (Mermithidae) that do not depart
with their host moths but await their return
each year, do the Fungus Gnats have an
organic connection with the caves? Are they
also dependent upon the annual cycle of
Bogong Moth migration, death, decay and
fungal growth? These are the sorts of ques-
tions that face the naturalist in our moun-
tains, and yet another good reason for me to
spend the summer sticking my head into
any cave big enough to admit me.

References

Green K (1982) Migration of an ichneumonid wasp over
a subalpine pass. The Victorian Naturalist 99, 260,

Green K (2002a) The biodiversity blitz. The Victorian
Naturalist 119, 36-37.

Green K (ed) (2002) Biodiversity in the Snowy Mountains,
(Australian Institute of Alpine Studies: Jindabyne)

Green K, Broome L, Heinze D and Johnston S (2001)
Long distance transport of arsenie by migrating
Bogong Moths trom agricultural lowlands to mountain
ecosystems. The Victorian Naturalist 118, 112-116.

Ken Green

National Parks and Wildlife Service
PO Box 2228, Jindabyne, NSW 2627
Email ken.greeninpws.nsw.gov.auu

277

Book Reviews

Grassed Up: Guidelines for Revegetating with
Australian Native Grasses

by Cathy Waters, Wal Whalley and Charles Huxtable

Publisher: NSW Agriculture, 2001. 72 pages, black and white illustrations.
RRP $33.00

The status of grasslands has been largely
overshadowed in recent times by the envi-
ronmental focus on planting trees, and
more recently the inclusion of shrubs,
Native grasses have generally been forgot-
ten in the revegetation process, primarily
because information on harvesting, dor-
mancy and establishment has been scarce.
Over the last few years, however, there has
been a growing interest in native grasses in
terms of pasture management and their
environmental values, and consequently
there has been a shift in emphasis from
trees and shrubs to native vegetation as a
whole. Landowners and land managers are
now becoming aware of the role native
grasses can play in rehabilitation of sites
and are desperate for information on har-
vesting and establishment techniques for
native grasses. Grassed Up: Guidelines for
Revegetating with Australian Native
Grasses is the first of such information, It
provides practical advice with step by step
instructions on native grass collection and
establishment.

The Guidelines is an A4, 72 page spiral-
bound booklet which the authors have
divided into two easy-to-read sections, The
first section provides general information
on native grasses (chapter 1) and up to date
techniques on collection (chapter 2), stor-
age (chapter 4) and establishment (chapter
5). Whilst individual species’ responses to
fertilisers are covered generally in section
two under individual species information, |
would have liked to see the authors discuss
soil fertility in more detail and the likeli-
hood of increased weed competition under
higher soil fertility.

Throughout the Guidlelines the authors
have included a number of black and white
photos to complement the text. The photos
of the current machinery being used and
diagrams of how they work are invaluable.

The second section contains details on
fourteen native grass species considered to
have a high priority for commercial devel-
opment or those that are already being
used in revegetation, Each information
sheet is complete with photos and a
description with information on morpholo-
gy, distribution, harvesting, sowing and
management. Whilst a lot of information is
still unknown for many species, it would
have been worthwhile for the authors to
include information sheets on other native
grasses such as Austrostipa and Poa
species, making it more applicable to sites
outside of NSW.

Included in the back of the book is a
series of guidelines produced by
FloraBank which outlines the protocols for
seed collection and storage. There is also a
useful list of publications on grass identifi-
cation, harvesting and germination.

Grassed Up: Guidelines for Revegetating
with Australian Native Grasses is a well-
written account of native grass revegeta-
tion techniques, It is aimed at landowners,
land managers or anyone interested in the
practical application of the establishment
of native grasses in revegetation, regenera-
tion or the rehabilitation of degraded sites.

Cheryl O’ Dwyer

Institute of Land and Food Resources
The University of Melbourne

Dookie College, Victoria 3647

For assistance in preparing this issue, thanks to Kate Smith (desktop publishing), Ann

Williamson (label printing) and Dorothy Mahler (administrative assistance).

278

The Victorian Naturalist

Book Reviews

Platypus: the Extraordinary Story of How a Curious Creature
Baffled the World

by Ann Moyal

Publisher: Allen & Unwin, 2001. 2

Today, scientists confidently classify the
platypus and echidnas as monotremes:
egg-laying, milk-producing, fur-clad mam-
mals. Based mainly on fossil evidence, it is
believed that the many attributes which
distinguish monotremes from marsupials
and placental mammals — including differ-
ences in the structure of the ear, backbone,
limb girdles, brain and sex chromosomes —
reflect the fact that monotremes have been
evolving independently of other mammals
for 100 million years or more.

Ann Moyal has produced a fascinating
and meticulously researched account of
how the platypus’s place in nature has come
to be defined, and how that process was
interwoven with the rise of evolutionary
theory. The story begins in 1798 when John
Hunter, governor of the penal colony at
Botany Bay, watched ‘a Small Amphibious
Animal of the mole kind" being speared by
an aboriginal man near the Hawkesbury
River. The animals skin was preserved in a
keg of spirits and sent (along with a wom-
bat) to the Literary and Philosophical
Society of Newcastle-upon-Tyne, which
had recently honoured Hunter with a corre-
sponding membership. Upon its arrival, the
keg was collected by a woman servant, who
was carrying the container on her head
when the bottom gave way and soaked her
with alcohol. Looking down, her distaste
grew with the realisation that she had been
carrying the remains of ‘a strange creature,
half bird, half beast, lying at her feet’.

The bemused reaction of George Shaw,
the first professional zoologist to examine
a platypus (a dried specimen arriving in
England in 1799) was fundamentally the
same as that of the servant. ‘Of all the
Mammalia yet known, it seems the most
extraordinary in its conformation, exhibit-
ing the perfect resemblance of the beak of
a Duck engrafted on the head of a
quadruped.’ Suspecting that the specimen
was a hoax, he carefully probed at the line
where the bill joins the head, but could

Vol. 119 (6) 2002

26 pages, hardback. RRP $29.95

find no stitching or other evidence of
forgery. The mystery only deepened when
a British anatomist, Everard Home, applied
himself to the task of dissecting preserved
platypus organs. Home’s studies revealed
that the structure of the platypus bill was
actually quite unlike that of a duck's beak.
However, after examining the reproductive
systems of males and females, he conclud-
ed that they differed in many respects from
the accepted mammalian pattern — and
most closely resembled the organs of
lizards which hatch eggs internally.

Scientific consensus about whether or not
the female platypus can produce milk was
not reached for another three decades.
Heated debate about the platypus’s mode
of reproduction ~ live-bearing or egg-lay-
ing — would not be resolved until 1884, As
a research biologist, | particularly appreci-
ated Moyal's description of the obstacles
facing the most brilliant academic minds
of the 19" century in discovering the truth
about the platypus, including their appar-
ently universal reluctance to countenance
first-hand observations made by colonials.

Moyal devotes relatively few pages to
platypus-related achievements in the 20"
century, though some interesting material
is presented, including a description of the
mission undertaken by David Fleay to sat-
isfy Winston Churchill’s desire to see a
live platypus midway through the Second
World War.

The text is complemented handsomely by
illustrations of early platypus paintings and
portraits of the zoologists and natural his-
torians who contributed to our knowledge
of the species. | commend the book to any-
one with an interest either in Australian
fauna or the argumentative and all-too-
human process by which scientists reach
agreement on how nature functions.

Melody Serena
Australian Platypus Conservancy

PO Box 84
Whittlesea, Victoria 3757

279

Book Reviews

Working on Country:
Contemporary Indigenous Management of
Australia's Land and Coastal Regions

Edited by Richard Baker, Jocelyn Davies and Elspeth Young

Publisher: Ox/ord University Press, 2001. xxiv * 351 pages, 27 figures, 5 tables.
ISBN 0195512170. RRP $49.95

One of the more significant changes over
the past 20 years or so, in a range of field
disciplines that focus on natural resources,

has been the increasing involvement of

Aboriginal people. It has taken too long to
recognise the rights of indigenous peoples
or to give respect to their knowledge sys-
tems, so that meaningful collaborations
could occur. As this book illustrates, how-
ever, indigenous people now not only have
input, but are directing projects in all parts
of Australia. Indeed, the major part of this
volume consists of reports of close collab-
orations between indigenous and non-
indigenous people, working in situations
that give accord to indigenous knowledge
and experience. Working on Country is
both a statement for the cause of eross cul-
tural collaboration and a testament to its
enormous value,

Of course, collaborations and joint pro-
jects of the kind reported in this volume do
not happen easily or without attendant chal-
lenges. Aboriginal understandings and uses
of local land and sea resources are often at
odds with those of non-indigenous man-
agers, administrators, and academics; issues
relating to ownership of resources and
appropriate access also need to be resolved.
And as the editors of this book point out,
even the concepts of “work? and ‘country’
can have different meanings for different
groups. The resolution of these issues is
often a matter of protracted discussion but,
if nothing else, such resolution and the
resulling collaborations can be seen as
points of reconciliation at a grass roots
level,

This book ts divided into four parts. In the
first, headed ‘Contemporary indigenous
management’, two summary chapters pro-
vide an overview of the prime issues within
the field, These essays are followed by a
series of case studies grouped in three the-
matic sections: * Approaches to managing

280

country’, ‘Sharing knowledge: tools and
communication’, and ‘Negotiating manage-
ment’. The first paper in each of these three
parts, written by the editors, introduces the
relevant issues,

The studies that are detailed in this book
were undertaken in different contexts and
for a variety of purposes, including the man-
agement of land and sea resource, biological
surveys, and heritage studies. The chapters
focus on locations in almost every state of
Australia, the exception being Victoria,
which barely receives a mention in the entire
book. However, this omission should not be
taken as an indication that Kooris and
Gubbas (non-Indigenes) are not working
together in Victoria. Although not represent-
ed in this volume, collaborations of not dis-
similar kinds are certainly taking place here.

Working on Country works well as an
introduction to the ways in which seeming-
ly opposed perspectives can be reconciled.
The book's breadth of coverage and the
emphasis on non-technical language are
aimed in the direction of a wide audience.
Although they work in academic environ-
ments, the editors have maintained a style
that is both easy to read and informative.
An interesting aspect of the presentation is
the use of ‘boxes’ providing detail on a
wide range of related subjects. These are
inset in many of the chapters, and could
usefully be read on their own.

This book is an important contribution to
the subject of resource management. It
provides interesting insights into the nature
of connections between indigenous popu-
lation and land and should also help to dis-
pel the myth of a monocultural indigenous
population in Australia. Por these reasons
alone, it should be recommended reading.

Gary Presland

40 William Street
Box Hill, Victoria 3128

The Victorian Naturalist

Book Reviews

Gardening With Australian Rainforest Plants

by Ralph Bailey and Julie Lake

Publisher: B/oomings Books, Melbourne. 2001. 144 pp, paperback.
RRP 832.95

There is considerable interest in
Australian rainforest plants for their con-
servation as well as cultivation and they
are certainly worthy of our attention.

Rainforests contain some of the world's
richest biological systems. It is vital that
we appreciate these natural habitats and do
all we can to ensure the preservation of the
rainforest areas we still have in Australia,
and, indeed, in other areas of the world.

Ralph Bailey and Julie Lake, in Gardening
With Australian Rainforest Plants, remind
us that a staggering 95% of Australia's rain-
forests have been felled in the last 200 years.

We sometimes think of rainforest plants
as being applicable only to tropical regions
but rainforests extend right down the east-
ern coast of Australia to Tasmania, with
some pockets of rainforest also in Western
Australia and the Northern Territory,

The introductory chapter covers the vari-
ous types of Australian rainforests includ-
ing Monsoon rainforest; Lowland and
Upland tropical rainforests; Lowland and
Upland subtropical rainforests; Littoral
rainforest, Warm temperate rainforest,
Cool temperate rainforest and to make sure
that absolutely nothing is missed there is a
further heading of Other Rainforests.

Many rainforest species are very adaptable
to cultivation and quite a number of plants
from north-eastern Australia are adaptable
to cultivation in cool temperate zones.

The book proceeds to dispel some of the
common myths about rainforests and rain-
forest plants. These include the incorrect
assumptions that rainforest plants are diffi-
cult to grow and don't have attractive flow-
ers and that all rainforest trees are tall. The
book also points out that rainforest plants do
not necessarily need a lot of water.

Five chapters give detailed advice on
planning and planting a rainforest garden
and its subsequent care. Small gardens,
courtyards and container cultivation are
covered in separate sections.

Vol. 119 (6) 2002

Rainforests and rainforest gardens can be
wildlife havens and the chapter on Wildlife
in the Rainforest Garden covers this aspect
in considerable detail. Birds, butterflies
(including caterpillars), frogs, other pond
life and lizards are all included together
with fungi, lichens and mosses which are
all an intrinsic part of the total ecology.
Even unwanted creatures receive a men-
tion and some helpful advice.

Edible rainforest plants and a section on
propagating rainforest plants are included
prior to the final chapter which gives
detailed descriptions of one hundred
favourite rainforest plants.

Gardening With Australian Rainforest
Plants is illustrated throughout with clear
colour photographs and there are also a
number of helpful line drawings to com-
plement the text.

There is a Bibliography and comprehen-
sive Index to complete its usefulness.

Ralph Bailey is a landscape architect who
has been involved with Australian plants
for more than 30 years. He has written
extensively on rainforest plants, bush tuck-
er and ecologically sensitive design.

Julie Lake is a horticultural writer and
consultant with a particular interest in
Australian rainforest plants. An active
member of the Society for Growing
Australian Plants and the Horticultural
Media Association, she is a regular contrib-
utor to several Australian magazines and
lectures on rainforest plant identification,

Gardening With Australian Rainforest
Plants is à very easy-to-use book which
will show readers how to turn a garden
into a cool, green haven, and to transform
the landscape into something distinctively
Australian.

Gwen and Rodger Elliot
PO Box 655
Heathmont, Victoria 3135

281

Guidelines for Authors — The Victorian Naturalist

Submission of all Manuscripts

Authors may submit material in the form of
research reports, contributions, naturalist notes,
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Research Report is a succinct and original sci-
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including abstract, introduction, methods, results
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ural history. The scope of a contribution is broad
and little defined to encourage material on a
wide range of topics and in a range of styles.
This allows inclusion of material that makes a
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Cite references used for taxonomic names,
References used by The Victorian Naturalist are
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The following abbreviations should be used in
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ul; pers. obs.; unpubl. data; and pers. comm.
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pers. comm. 3 May). Use ‘subsp.’ for subspecies.

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Authors are advised to note the layout of head-
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issues of the Journal, Single spaces are used
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In all papers, at the first reference of a species,
please use both the common name and binomial.
However, where many species are mentioned, a
list (an appendix at the end), with both common
and binomial names, may be preferred. Lists
must be in taxonomic order using the order in

which they appear in the references recommend-
ed below,

The journal uses capitalised common names
for species followed by the binomial in italies
without brackets, e.g. Kangaroo Grass Themeda
triandra.

References

References in the text should cite author and
year, e.g. Brown (1990), (Brown 1990), (Brown
1990, 1991), (Brown 1995 unpubl.), (Brown and
Green 1990), (Brown and Green 1990; Blue
1990; Red 1990). If there are more than two
authors for a paper use Brown ef al. (1990),
These should be included under References, in
alphabetical order, at the end of the text (see
below). The use of unpublished data is only
accepted if the data is available on request for
viewing. Pers. obs. and pers. comm. should not
be included in the list of references. Journal
titles should be quoted in full.

Leigh J, Boden R and Briggs J (1984) Extiner
and Endangered Plants of Australia.
(Maemillan: South Melbourne)

Lunney D (1995) Bush Rat. In The Mammals of

Australia, pp 651-653, Ed R Strahan.
(Australian Museum/Reed New Holland:
Sydney)

Phillips A and Watson R (1991) Xanthorrhoea:
consequences of ‘horticultural fashion’, 7he
Vietorian Naturalist 108, 130-133.

Smith AB (1995) Flowering plants in north-
eastern Victoria, (Unpublished PhD thesis,
University of Melbourne)

Wolf L and Chippendale GM (1981) The natural
distribution of Eucalyptus in Australia.
Australian National Parks and Wildlife
Service, Special Publications No 6, Canberra.

Other methods of referencing may be acceptable
in manuscripts other than research reports, and
the editors should be consulted. For those using
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guide for The Victorian Naturalist is available
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Government Publishing Service: Canberra).

Manuscript Corrections

Authors ean verify the final copy of their man-
uscript before it goes to the printer, A copy of
their article as ‘ready for the printer’ will be sent
and only minor changes may be made at this
stage.

Complimentary Copies

After publication of an article in the journal,
five complimentary copies of that issue are sent
to the author(s) for each paper. Authors of
Naturalist Notes and Book Reviews will receive
two complimentary copies of the journal.

Additional copies of The Victorian Naturalist:
25 copies, $50.00 (4 postage); 50 copies, $90.00
(+ postage), including GST,

Checking species names is the responsibility of authors. The books we would like used as references
for articles in The Victorian Naturalist are listed below. Authors should refer to the source used for
species names in their manuscripts. In every case, the latest edition of the book should be used.

Mammals Menkhorst PW (ed) (1995)
Mammals of Victoria; Distribution, Ecology
and Conservation, (Oxford University Press:
South Melbourne)

Reptiles and Amphibians — Cogger H (2000)
Reptiles and Amphibians of Australia, 6 ed.
(Reed Books: Chatswood, NSW)

Insects — CSIRO (1991) The Insects of Australia:
a textbook for students and research workers.
Vol I and Il. (MUP: Melbourne)

Birds — Christidis L and Boles W (1994) The
Taxonomy and Species of Birds of Australia
and its Territories. Royal Australian
Ornithologists Union Monograph 2. (RAOU:
Melbourne)

Plants — Ross JH (ed) (2000) A Census of the
Vascular Plants of Victoria, 6 ed. (Royal
Botanic Gardens of Victoria: Melbourne)

Please submit manuscripts and enquiries to:

The Editor

The Victorian Naturalist
Locked Bag 3, P.O.
Blackburn, Victoria 3130

Phone/Fax (03) 9877 9860. Email fnev@vienet.net.au

Web address: http://www.vicnet.net.au/~fnev/vienat.him

The Field Naturalists Club of Victoria Inc.

Reg No A0033611X

Established 1880
In which is incorporated the Microscopical Society of Victoria

OBJECTIVES: To stimulate interest in natural history
and to preserve and protect Australian flora and fauna.

Membership is open to any person interested in natural history and includes
beginners as well as experienced naturalists.
Registered Office: ENCV, I Gardenia Street, Blackburn, Victoria 3130, Australia.
Postal Address: FNCV, Locked Bag 3, PO Blackburn, Victoria 3130, Australia.
Phone/Fax (03) 9877 9860; International Phone/Fax 61 3 9877 9860.

Patron
John Landy, Ac, MBE, The Governor of Victoria

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Printed by Brown Prior Anderson, 5 Evans Street, Burwood, Victoria 3125.

The Victorian
Naturalist

Index to

.— Volume 118, 2001

Compiled by KN Bell

Authors

Adams R, Simmons D and Lewis C, 16

Ahern L, 285

Archbold NW, 66 (Tribute), 178, 234

Ashton DIT, 321

Barker R, 61 (book review)

Bell KN, 101 (book review)

Bock P, 256

Boyd S, Coventry AJ, Dixon J, Gomon
M, O'Loughlin M, Poore G, Walker K
and Yen AL, 242

Broadberry J, 64 (book review), 65 (book
review)

Broome L, Heinze D, Johnston S and
Green K, 112

Burgess DR and Wright MF, 117

Butcher BW, 226

Carkeek M, 151

Clemann N, 30 (book review)

Conole L and Mac Nally R, 56

Coventry AJ, Dixon J, Gomon M,
O'Loughlin M, Poore G, Walker K,
Yen AL and Boyd S, 242

Dann P, Jessop R and Healy M, 76

Darragh TA, 160

Di Stefano J, 46

Dixon J, Gomon M, O'Loughlin M,
Poore G, Walker K, Yen AL, Boyd S
and Coventry AJ, 242

Earp C, 82

Editors, Vic. Nat., 107, 159, 233

Fleming A, 210

Fletcher RJ, 27 (book review)

Gerdtz WR, 231

Gibson M, 102 (book review)

Gillbank L, 297

Gomon M, O'Loughlin M, Poore G,
Walker K, Yen AL, Boyd S, Coventry
AJ and Dixon J, 242

Green K, Broome L, Heinze D and
Johnston S, 112

Grey E, 74, 100

Grey P, 104 (software review)

Healy M, Dann P and Jessop R, 76

Heinze D, Johnston S, Green K and
Broome L, 112

Hell AJ, 294

Holland GJ, 123

Houghton S, 314

Jessop R, Dann P and Healy M, 76

Johnston F and Pickering CM, 21

Johnston S, Green K, Broome L and
Heinze D, 112

Kasel S, 127

Leary EJ, Parris KM, McDonnell MJ and
Williams NSG, 4

Lebel T, 38

Lewis C, Simmons D and Adams R, 16

McCann D, 146, 148, 165, 309, 319

Mace B, 287

McDonnell MJ, Williams NSG, Leary EJ

and Parris KM, 4

McLean A and Meagher D, 92

Mac Nally R and Conole LE, 56

Maroske S, 305

May T, 44

Meagher D and McLean A, 92

Milne J and Thies AW, 89

Morton A, 62, (book review), 142 (book
review)

O’Brien M, 60 (book review)

O’ Loughlin M, Poore G, Walker K, Yen
AL, Boyd S, Coventry AJ, Dixon J and
Gomon M, 242

Parkin A, 2

Parris KM, McDonnell MJ, Williams
NSG and Leary EJ, 4

Pascoe G, 193

Pickering CM and Johnston F, 21

Pickering CM and Scherrer P, 93

Pierson R, 219

Poore G, Walker K, Yen AL, Boyd S,
Coventry AJ, Dixon J, Gomon M and
O'Loughlin M, 242

Rasmussen C, 200

Sault T, 106 (Tribute)

Scherrer P and Pickering CM, 93

Schleiger N, 29 (Tribute), 266

Seebeck J and Warneke, RM 277

Simmons D, Lewis C and Adams R, 16

Sinclair B, 143 (book review)

Slattery D, 25 (book review)

Stefano J Di, 46

Thies AW and Milne J, 89

Traill A, 58, 59

Walker K, Yen AL, Boyd S, Coventry
AJ, Dixon J, Gomon M, O’ Loughlin
M, and Poore G, 242

Warne MT, 283

Warneke RM and Seebeck J, 277

Weste G, 140

Wilkinson I, 186

Williams NSG, Parris KM, McDonnell
MJ and Leary EJ, 4

Wright MF and Burgess DR, 117

Yen AL, Boyd S, Coventry AJ, Dixon J,
Gomon M, O'Loughlin M, Walker K
and Poore G, 242

Birds

Eudyptula minor, at sea, Western Port, 76

Little penguins, at sea, Western Port, 76

McCoy-Gould correspondence, 210

Red-chested Button-quail, identity, ecol-
ogy, status, 56

Turnix pyrrhothorax, identity, ecology,
status, 56

Book Reviews

Birds of French Island Wetlands, D
Quinn and G Lacey (M O’Brien), 60

Celebrating our Parks: Proceedings of
the First Australian Symposium on
Parks History, ed E Hamilton-Smith
(D Slattery), 25

Common Australian Fungi: A
Bushwalkers Guide, T Young (R
Barker), 61

Field Guide to the Orchids of NSW and
Victoria, 2 ed, T Bishop (J
Broadberry ), 64

Flora of Australia, v 48: Ferns,
Gymnosperms and Allied Groups, (M
Gibson), 102

Guide to Squid, Cuttlefish and Octopuses
of Australasia, M Norman and A Reid
(K Bell), 101

Kosciuszko Alpine Flora, 2 ed, AB
Costin, M Gray, GJ Totterdell and DJ
Wimbush (RJ Fletcher), 27

Native Orchids of Southern Australia: A
Field Guide, D and B Jones (J
Broadberry), 65

Nature Photography, K Griffiths (A
Morton), 142

Pythons of Australia; A Natural History,
G Torr (N Clemann), 30

Wildflowers of the Brisbane Ranges, C
and M Trigg (A Morton), 62

Wildflowers of Victoria, M Corrick and B
Fuhrer (B Sinclair), 143

Botany

Achillea millefolium, weed threat,
Australian Alps, 21

Allocasuarina luehmannii, galls and
mistletoes on, 117

Alpine vegetation, grazing, tourism, cli-
mate effects, 93

Australian Edelweiss at Mount Hotham, 2

Buloke, galls and mistletoes on, 117

Eucalyptus erenulata, Buxton Silvergum
Reserve, 16

Ewartia nubigena at Mount Hotham, 2

Dieback disease, impact, Mount
Stapylton, 46

FUNGIMAP, target species notes, 44

Leafy liverwort, Yarra Ranges NP, 92

Mistletoes, galls on Buloke, 117

Moss collections, Lord Howe Island,
addendum, 89

Native grasslands, potential freeway
impact, 4

Pedinophyllum monoicum in Yarra
Ranges, NP, 92

Phytophthora cinnamomi, Mount
Stapylton, 46

System Garden, University of
Melbourne, 193

Truffles, native Australian, 38

Yarrow, weed threat, Australian Alps, 21

Entomology
Agrotis infusa, arsenic carriers, 112
Ant behaviour, 100
Bogong moths, arsenic carriers, 112
Galls and mistletoes on Buloke, 117
Leptopius sp. c.f. L. gravis, Simpson

Desert, 74

Weevil, large, Simpson Desert, 74

Geology
Black coal McCoy-Clarke dispute, 219
Cranbourne meteorites, fate of, 305
Geological time scale, 150
Tanjilian (Early Devonian) fossils,
Dungaree Creek, 82

Honours
David Ashton OAM (Gretna Weste), 140
Lyle Courtney OAM (The Editors), 107
Gwen Elliot AM (FNCV), 141
Roger Elliot AM (FNCV), 141

Localities

Australian Alps, Yarrow weed threat, 21

Buxton Silver Gum Reserve, historic
changes, 16

Dungaree, Creek, Central Victoria,
Tanjilian fossils, 82

Hume Freeway-F2 link, impact on grass-
lands, 4

Kosciuszko NP, grazing, tourism, climate
effects on, 93

Lord Howe Island, moss collection in
MEL, addendum, 89

Mount Hotham, Australian Edelweiss, 2

Mount Macedon, McCoy's property at, 319

Mount Stapylton, Grampian NP, dieback
at, 46

Northern Victoria, Red-chested Button-
quail, 56

Snowy Mountains, Bogong moths as
arsenic carriers, 112

Victorian Alps, Bogong moths as arsenic
carriers, 112

Western Port, Little penguins at, 76

Wimmera, galls and mistletoes on
Buloke, 117

Yarra Ranges NP, leafy liverwort in, 92

Yellingbo Nature Conservation Reserve,
human and natural impacts on, 127

MeCoy, Frederick
Animal acclimatization society, 297
Antievolutionism, 226
Birds, Gould correspondence, 210
Black coal dispute with Clarke, 219
Brachiopoda, 178
Bryozoa, 256
Cranbourne meteorites, fate of, 305
The late F McCoy (extract Vic. Nat.

1899), 276
FNCV, 314
Graptolites, 266
Grave, McCoy’s, 304
Ichthyosaur, 294
Irish years, 160
McCoy society, 321
Mammals, McCoy’s, 277
Name, spelling of, 234
National Museum of Victoria, 200
Naturalist tradition, 309
Ostracoda, 283
Overview of his life, 151
Prodromus of Zoology of Victoria, 242
Property at Mount Macedon, 319
Snake, Western Brown, 285
Stratigraphic palaeontology, 165
System Garden, University of
Melbourne, 193

Tasmanian devil, 231
Thylacoleo, 287
Timeline of his life, 148
University of Melbourne, 186

Mammals

McCoy's mammals, 277

Petaurus norfolcensis, opportunistic ver-
tebrate predation by, 123

Squirrel glider, opportunistic vertebrate
predation by, 123

Tasmanian devil, 231

Thylacaleo, 287

Miscellaneous
100 vears ago, 15, 105, 140
Animal acelimatization society, 297
Antievolutionism, MeCoy, 226
Australian Ornithological Conference, 43
Flora and Fauna Guarantee Act 1988, 91
ENCV and McCoy, 314

FUNGIMAP National Conference, 45

Geological timescale, 150

Human and natural impacts, Yellingbo
Nature Conservation Reserve, 127

International Ornithological Conference,
126

International Year of Mountains 116

Introduction, McCoy Special Issue, 146

Irish years, McCoy, 160

McCoy grave, 304

McCoy society, 321

Name, spelling of McCoy, 234

National Museum (extract Vic. Nat.), 209

National Museum of Victoria, 200

Naturalist tradition, 309

Overview of McCoy's life, 151

Prodromus of Zoology of Victoria, 242

McCoy's property at Mount Macedon, 319

Timeline of MeCoy's life, 148

University of Melbourne and McCoy, 186

VENCA Annual Camp, 141

Palaentology

Brachiopoda, 178

Bryozoa, 256

The late F McCoy (extract ie. Nat.
1899), 276

Graptolites, 266

Ichthyosaur, 294

Ostracoda, 283

Stratigraphic palaeontology, 165

Tanjilian (Early Devonian) fossils,
Dungaree Creek, 82

Tasmanian devil, 231

Thylacoleo, 287

Reptiles
Black snake feeding, 59
Lampropholis guichenoti, mating behay-
iour, 58
Pseudechis porphyriacus, feeding, 59
Skink mating behaviour, 58

Software Review
Compendium of FUNGIMAP Target Species,
V. 1.0 [CD-ROM] (P Grey), 104

Tributes
Ilma Dunn, 110
Stefanie Rennick (T Sault), 106
John Paul Stewart (N Schleiger), 29
George Anthony Thomas (NW
Archbold), 66

Issues 5 and 6 form the
McCoy Special Issue