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Volume 99 Part 1
June 2016

CONTENTS

PREFACE i

Mammal assemblages in Boonanarring Nature Reserve,

Dandaragan Plateau, Western Australia from 1986 and 2012

T L Moore, A H Burbridge, T Sonneman & B A Wilson 1

Environmental characteristics of ephemeral rock pools
explain local abundances of the clam shrimp Paralimnadia
badia (Brachiopoda Spinicaudata: Limnadiidae)

A Calabrese, C McCullough, B Knott (deceased) & S C Weeks 9

Chronostratigraphic context for artefact-bearing palaeosols
in late Pleistocene Tamala Limestone, Rottnest Island,

Western Australia

I Ward, T J Pietsch, E J Rhodes, G H Miller, J Hellstrom

& C E Dortch 17

Male sterility in Corymbia calophylla (marri)

J McComb 27

The Royal Society of Western Australia

To promote and foster science in Western Australia

Patron

Her Majesty the Queen

Vice-Patrons

Her Excellency the Honourable Kerry Sanderson AO
Governor of Western Australia

Professor Lyn Beazley AO FTSE MA PhD

COUNCIL 2015-2017

President Kate Triajstic
Immediate Past President Hugo Bekle
Vice-President Bill Loneragan
Vice-President Chris Florides
Hon Treasurer Greg Benjamin
Hon Secretary Kelly Freeman
Hon Editor Patrick Armstrong

Councillors

Elizabeth Re
Charlotte Oskam
David Haig
Brian Devine

EDITORIAL BOARD 2015-2016

Editor-in-chief

Patrick Armstrong, University of Western Australia

Editorial Assistant

Desiree Moon

Board Members

Hugo Bekle, Enviro Systems

Alex Bevan, Western Australian Museum

Rob Davis, Edith Cowan University
Chris Florides, Murdoch University
David Haig, University of Western Australia
Philip Ladd, Murdoch University
Kate Trinajstic, Curtin University
Mike Van Keulen, Murdoch University

The Royal Society of Western Australia was founded in 1914. The Society promotes exchange among scientists from all fields in Western Australia
through the publication of a journal, monthly meetings where interesting talks are presented by local or visiting scientists, and occasional symposia
or excursions on topics of current importance. Members and guests are encouraged to attend meetings on the third Monday of every month
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Events, or online.

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The Journal of the Royal Society of Western Australia was first published in 1914 and circulates widely within Australia and throughout the world.
The journal has a large number of personal subscribers, many of whom are scientists working in Western Australia.

Cover design: The images symbolise the diversity of sciences embraced by the Royal Society of Western Australia. Counter-clockwise from the top
they are: Wolfe Creek Meteorite Crater; the world-famous stromatolites at Shark Bay; the numbat (Myrmecobius fasciatus), Mangles' kangaroo paw
(Anigozanthos manglesii) and Gogo fish (Mcnamaraspis kaprios), which are the faunal, floral and fossil emblems of Western Australia, respectively;
a zircon grain (Western Australian rocks have yielded the oldest zircon dates in the world, up to 4.37 Ga); and the black swan ( Cygnus atratus ) that
appears on the logos of the Royal Society and the coat of arms of the Government of Western Australia.

Blank Page Digitally Inserted

Journal of the Royal Society of Western Australia, 99(1): i, 2016

PREFACE

The Journal of the Royal Society of Western Australia is proud to include four important papers
in issue 99/1, the first issue to be published entirely online.

Ecologists are frequently enjoined to utilise the ecosystem concept for segments of nature of any
magnitude, from the entire biosphere down to the tiniest natural entity, and in this issue we see a
careful study of clam shrimp populations in relation to their environment in small ephemeral rock
pools on granite outcrops in WA's Wheatbelt.

Scientists are also encouraged to take 'a long-term view' and to study the nature of changes in
the environment by repeating observations at relatively long intervals. It is thus a pleasure to be
able to present a study undertaken on mammal assemblages in a Western Australian Nature
Reserve in 1986 and 2012.

Rottnest Island has served the purpose of an 'outdoor laboratory' for Western Australian schools
and universities for over a century and this issue includes a significant contribution to the geology
of that island. The return of earth science papers to the journal is particularly welcome and serves
to emphasise the interdisciplinary nature of the journal.

The marri is often taken as an organism for special study in the field and in the laboratory in the
State; the final paper reveals that there is much to be found out about a most familiar organism.

P H Armstrong

Editor-in-Chief

Nedlands
June 2016

Journal of the Royal Society of Western Australia, 99(1): 1-8, 2016

Mammal assemblages in Boonanarring Nature Reserve, Dandaragan
Plateau, Western Australia from 1986 and 2012

T L MOORE 15 *, A H BURBIDGE 3 , T SONNEMAN 2 , B A WILSON 14

1 Department of Parks and Wildlife, Swan Region, Corner ofHackett and Australia II Drive, Crawley, Western Australia, 6009

2 Department of Parks and Wildlife, West Kimberley District, 111 Herbert Street, Broome, 6725

3 Department of Parks and Wildlife, Woodvale Research Centre, Locked Bag 104, Bentley Delivery Centre, Western Australia, 6983

4 School of Life and Environmental Sciences, Deakin University, 221 Burwood Highway, Burwood, Victoria, 3125, Australia.

5 School of Veterinary and Life Science, Murdoch University, 90 South St, Murdoch, Western Australia, 6150
* Corresponding author: t.moore@murdoch.edu.au

ABSTRACT

Although long term monitoring can provide land managers and researchers with insights into
faunal changes there are few such programmes in Australia, and many conservation actions are
implemented without assessment of their biodiversity benefits or costs. This study investigated
the current status of native small mammals in the Boonanarring Nature Reserve (BNR), Western
Australia, aiming to compare the contemporary distribution of mammals to those recorded 26 years
ago.

Of particular importance is the evidence that no small mammal species have been lost from the
reserve in the last 26 years; Pseudomys albocinereus, Tarsipes rostratus and Sminthopsis (sp)p. were
recorded in both years' surveys. Records of P. albocinereus in this study are significant as they
confirm the persistence of the species on the Dandaragan Plateau, whereas on the adjacent Swan
Coastal Plain the species has not been recorded since 1987.

Overall, the persistence of small mammals in this reserve, unlike the nearby reserves on the Swan
Coastal Plain (SCP), could be attributed to the mix of vegetation types within the reserve and the
larger size of Boonanarring Nature Reserve. However, deficiencies in the monitoring programme
were identified. There is a need to improve the long-term monitoring of small mammals within the
BNR with long-term, repeat-measures, analysis and reporting.

KEYWORDS: Swan Coastal Plain, Pseudomys albocinereus, Tarsipes rostratus, Sminthopsis, ash
grey mouse, monitoring

INTRODUCTION

Long term studies can provide researchers and land
managers with insights into faunal changes over time,
often in association with abiotic and biotic variables of
change. However, biodiversity monitoring in Australia
is limited and poorly coordinated (Natural Resource
Management Ministerial Council 2010, Lindenmayer &
Gibbons 2012, Lindenmayer et al. 2012). Managers and
researchers who have worked in conservation biology
in Australia are often disappointed by the absence of
effective biodiversity monitoring that allows evaluation
of how well Australia's natural heritage is managed
(Garkaklis 2014). There are few long-term programmes,
and many conservation actions are implemented
without assessment of their biodiversity benefits or costs.
Ultimately, a management decision must be made, but a
manager needs to feel confident in making management
intervention decisions (Varcoe 2012). There is a need
to improve this with long-term, repeat-measures data,
followed by its analysis and reporting. This is particularly
relevant given Australia's significant extinction rate since
European settlement (Woinarski et al. 2015).

Mammal extinctions and declines have occurred
across the continent (McKenzie et al. 2007). Australian
mammals, including those in the biodiversity hotspot
of south-west Western Australia, have seen some
of the highest extinction rates and include 50%
of the world's extinction events since European
settlement (Short & Smith 1994, Woinarski et al. 2015).
Many factors have been implicated in these declines
including climate change and a lack of rainfall,
introduced predators, introduced herbivores and habitat
modification (Burbidge & McKenzie 1989, McKenzie
et al. 2007, Wilson et al. 2012, Woinarski et al. 2015). To
conserve mammals, quantifying population trends
over time in relation to management practices such as
prescribed burning, predator control and abiotic changes
such as habitat and climate change factors are pivotal in
predicting populations in current and future conditions.
This study employed sampling at two time points, 1986
and 2012, in Boonanarring Nature Reserve (BNR) in the
south-west of Western Australia to examine changes in
the species richness and abundances of mammals over
recent decades.

Faunal declines in small isolated reserves are not
unexpected (Fahrig 2002, Henle et al. 2004, Fischer &
Lindenmayer 2006) but fauna populations in larger
reserves may also be at risk. Boonanarring Nature

Journal of the Royal Society of Western Australia, 99(1), 2016

GINGIN
PERI H

BUNBURY

Legend

Boonanarring NR Boundary

GERAlLDTON

Figure 1. Location of Boonanarring Nature Reserve on the Swan Coastal Plain, Western Australia.

Reserve is located on the Dandaragan Plateau, which
borders the Swan Coastal Plain (SCP) and Darling Scarp,
near Gingin, Western Australia. Much of the Dandaragan
Plateau has been extensively cleared and the dominant
land use is dry-land agriculture (92.6%). Only 6.8% is
remnant vegetation under conservation management and
much of this is present in isolated patches (Department
of Conservation and Environment 1983). Despite its
isolation BNR has special significance, not only because
it is one of the larger nature reserves covering >9000
ha with a level of connectivity to areas of surrounding
natural vegetation totalling approximately 10, 000 ha. It
contains the highest quality and most extensive example
of conserved Banksia woodlands on Dandaragan soils
with unique flora and fauna (Burbidge et al. 1996). It was
surveyed in 1986 for vegetation and fauna (Burbidge et al.
1996) and broad scale vegetation mapping was conducted
by Beard (1979) and Heddle et al. (1980), but no other
biological surveys have been conducted in the reserve.
Fauna surveys were thus carried out in 2012 to determine
the current status of vertebrate fauna and mammal
populations in the reserve.

Burbidge et al. (1996) captured three significant native
small mammal species ( Pseudomys albocinereus (ash grey
mouse), Tarsipes rostratus (honey possum) and Sminthopsis
griseoventer (grey bellied dunnart) in BNR, and identified
fire and introduced predators as likely threats to native
mammals. Within conservation lands, fire management
by the Department of Parks and Wildlife aims to protect

human life and property, and to a lesser degree biological
diversity by reducing fuel loads that could lead to
large wild fires. Prescribed burning occurs in BNR but
planning for such burns is currently not focused on
biological diversity or conservation of native species.
Feral predators and weed invasion are other management
issues across reserves in south-western Australia but
currently no feral predator or weed control occurs
in BNR. There are additional, potentially deleterious
activities occurring within and surrounding the reserve,
including gravel extraction and exploration for oil and
gas (Coffey Natural Systems Pty Ltd. 2008; unpublished
data).

The objectives of this paper were thus to (i) assess
the current status of small mammals (abundance and
species richness) in BNR, (ii) compare the current
status of mammals to that described 26 years ago, and
(iii) evaluate any problems associated with long term
monitoring.

METHODS

Site description

Boonanarring Nature Reserve is a 'C' class reserve
of 9250 ha, 15 km north of Gingin, with connections to
10,000 ha of protected remnant vegetation in Moore
River Nature Reserve. It is situated to the southern

T. Moore et al.: Mammal assemblages in Boonanarring Nature Reserve

Table 1. Fire history and description of each site from the 2012 surveys at Boonanarring Nature Reserve, AB indicates
that the site associated with the grid has PVC pit traps. FERA is a fire reference exclusion area. Vegetation data originate
from Burbidge et al., (1986).

Site Number

Fire age (years)

Vegetation type

Corymbia calophylla over heath

2 and AB1

Banksia woodland

3 and AB2

Eucalyptus wandoo woodland

Eucalyptus marginata and Corymbia calophylla woodland

5 and AB3

Banksia woodland

Eucalyptus marginata and Corymbia calophylla woodland

14 (FERA)

Heathland

14 (FERA)

Heathland

Breakaway- low vegetation

Unusual tall heath on slope with Eucalyptus sp. woodland

end of the Dandaragan Plateau and is bordered by
the SCP to the west and the Darling Scarp to the east
(Figure 1). The biological significance of the area was
first assessed in 1971 (N. McKenzie pers. comm.) and
on the basis of this report it was later recommended
that the then unallocated Crown land be combined with
smaller reserves and an area of 400 ha of private land
along Gingin Brook to form the BNR (Department of
Conservation and Environment 1983). The reserve was
gazetted as a 'C' class reserve in 1991, and vested in the
National Parks and Nature Conservation Authority.
Boonanarring Nature Reserve has a range of soil types
(mostly lateritic, but also with white, grey or yellow
sands) supporting over 570 plant species, 13% of which
are recorded as being of special interest. The reserve
supports a mix of vegetation types including banksia
(.Banksia grandis, B. attenuata and B. menziesii ), Corymbia
calophylla (marri). Eucalyptus marginata (jarrah), and E.
wandoo (wandoo) woodlands (Table 1). The reserve is
significant as it is rich in flora and vegetation types not
present together on any other conservation reserves and
being large enough to provide some protection from
degradation owing to edge effects. It has a Mediterranean
climate of wet winters and dry, hot summers (Burbidge et
al. 1996, Coffey Natural Systems Pty Ltd. 2008).

At the time of the 2012 survey, the majority of the
fauna survey sites had been burnt within the last 14
years (Table 1). The reserve contains a centrally located
Fire Exclusion Reference Area (FERA) of 845 hectares for
breeding and resting sites of the endangered Carnaby's
black-cockatoo ( Calyptorhynchus latirostris ). Prescribed
burns are planned for this reserve but a lack of optimal
burning conditions had restricted implementation in the
previous six years.

Monitoring surveys

1986 survey

Seven (2 ha) sites were trapped from the 17 th to the
23 rd of March 1986 (Table 2). Pitfall traps (12.5 x 60 cm)
were employed at sites one to four and seven, and Elliot
traps at sites five and six. Sites one to four had three lines
of pitfall traps, sites five and six had three lines of Elliot
traps and site seven had two lines of pitfall traps. Trap
nights equated to 600 nights (Burbidge et al. 1996).

2012 survey

Trapping sites were established in the reserve in
autumn and spring 2012. The sites were located in
similar positions as those in the 1986 surveys in order to
document any changes in mammal faunal assemblages.
An additional five sites (7-11) were located in different
vegetation types (heathland, breakaway and unusually
tall heath) and/or fuel age in spring 2012 in an attempt
to survey as many different habitats as possible. The trap
efforts for each year are detailed in Table 2.

During two survey periods in 2012, in autumn (15 th -
23 rd April) and spring (15 th to the 22 nd November), pitfall,
Elliott and funnel traps were employed. Autumn surveys
monitored six pitfall sites and in spring an additional five
sites were surveyed. In autumn each pitfall site had 10
buckets connected by drift fences in a Y formation with
two funnel traps at the end of each arm and five Elliott
traps distributed within approximately 30m of the pitfall
buckets. In spring the same set up was used, except that
funnel and Elliot traps were not used. Three additional
sites (AB 1, 2 and 3), closely connected to the other
trapping locations (details in Table 2), consisting of only
pitfall traps (12.5 x 60 cm) were established for the spring
survey with a total of six traps per site as it was suspected
that Pseudomys albocinereus could jump out of the 20L
buckets. All pitfall and funnel traps were opened for
eight nights and Elliot traps were open for six nights (for
capture of mammals, amphibians and reptiles; however
we only report on the mammals in this paper). In autumn
and spring, trapping was conducted over 948 and 1084
trap nights respectively.

Table 2. Trap nights and survey effort of the 2012 and
1986 surveys in Boonanarring Nature Reserve.

Survey Method

Trap nights
in 1986

Trap nights in 2012
Autumn Spring

Pitfall traps (20L buckets)

480

880

Funnel traps

288

PVC pitfall traps

300

144

Elliot traps

300

180

Total trap nights

600

948

1084

Journal of the Royal Society of Western Australia, 99(1), 2016

ANALYSIS

As the trapping effort differed between years the mean
mammal abundance, species richness and individual
mammal species' abundances (as defined by mammal
captures) were standardised to ten trap nights using
data from all pitfall, Elliot and funnel traps (e.g. mean
mammal abundance per ten trap nights at each site).
Individual species were only analysed if their captures
were above 20 individuals. In 2012 three individuals of
Sminthopsis were captured but not identified to species.
Captures of Sminthopsis were added irrespective of the
initial species identification, as identification of animals
from this genus in the field can be problematic (Kemper
et al. 2011) and these individuals are herein termed
Sminthopsis sp(p).

A direct comparison of 1986 and 2012 may be
influenced by the season in which the survey was
completed. Surveys were conducted in autumn in 1986
and 2012, and in spring 2012. One-way ANOVAs (Inc
2007) were thus performed at the site level to determine:
1) the relationship between mean mammal abundance,
mammal species richness and individual mammal
species' abundance and the season of trapping (autumn
and spring) in 2012, and 2) relationships between
mammal abundance, mammal species richness and
individual mammal species' abundance and the year
(1986 and 2012) and 3) relationships between mammal
abundance, mammal species richness and individual
mammal species' abundance in 1986 and 2012 using only
autumn data.

Capture data were compiled for four species, T.
rostratus, M. musculus, P. albocinereus and Sminthopsis
sp(p). ( Sminthopsis griseoventer in 1986), including capture
numbers, sex ratio and body weights. Weight data for
2012 included mean, standard deviation and range for
each species, whereas the weight data for 1986 were less
detailed and only included the weight range for each
species.

RESULTS

1986 survey

Four mammal species including one introduced
species, Mus musculus, and three native species.

Pseudomys albocinereus, Sminthopsis griseoventer and
Tarsipes rostratus, were captured in 1986 (Burbidge et al.
1996).

2012 survey

Four small mammal species were captured during
the 2012 surveys including one introduced species,
M. musculus and three native species, P. albocinereus,
Sminthopsis sp(p). and T. rostratus.

Mus musculus abundance was significantly different
between the autumn and spring 2012 surveys with more
M. musculus captured in autumn than spring (Table 3 and
Figure 2). Total mammal abundance, species richness
and abundances of the other individual mammals did
not differ between the autumn and spring 2012 surveys
(Table 3).

Capture data for individual mammal species

Body weights for both sexes and sex ratios between
2012 and 1986 for all species were similar (Table 4; no
statistical analysis has been performed). There were some
differences in the numbers of P albocinereus of each sex
captured. Male P albocinereus captures were higher in
spring 2012 (10), than females (5). In autumn 1986 and
2012 similar numbers of each sex were caught (1986: 10
males and 13 females; 2012 one male and female; Table 4).

Sites

Figure 2. Mus musculus abundance in autumn and spring
2012. Only sites one to six were trapped in autumn and
all 11 sites were trapped in spring 2012.

Table 3. Relationships between total mammal abundance, mammal species richness and individual mammal species'
abundance, and the season of trapping (autumn and spring) in 2012, the year (1986 and 2012) and in autumn only in 1986
and 2012 (excluding the spring 2012 survey data) using a one way ANOVA (F and P values in bold are significant).

Autumn and

1986 and

Autumn only

Spring 2012

2012

2012 and 1986

Dependant Variables

F and P values

Total mammal abundance

2.74, 0.11

0.70, 0.41

3.61, 0.08

Mammal species richness

0.436, 0.51

9.733, 0.007

1.103, 0.316

Mus musculus abundance

10.05, 0.006

4.78, 0.04

3.49, 0.08

Pseudomys albocinereus abundance

2.13, 0.16

11.81, 0.003

8.645, 0.01

Sminthopsis sp(p). abundance

1.951, 0.183

0.025, 0.877

0.923, 0.357

Tarsipes rostratus abundance

4.01, 0.06

4.58, 0.04

4.04, 0.06

T. Moore et ah: Mammal assemblages in Boonanarring Nature Reserve

Comparison of 1986 and 2012

Mammal species richness in 1986 was statistically
higher than in 2012 (Table 3; Figure 3a). The abundance
of Mus musculus was significantly higher in 2012 than
1986 (Table 3; Figure 3b). The abundance of Pseudomys
albocinereus and Tarsipes rostratus were significantly lower
in 2012 than 1986 (Table 3; Figure 3c and d). Mammal
abundance and Sminthopsis sp(p). abundance were
unchanged between the two survey years (Table 3).

Sites

Figure 3. Mammal species richness (a), and the abundances
rostratus (d) at each site in 1986 and 2012.

When examining the differences between the autumn
surveys in 1986 and 2012 (excluding spring 2012 surveys
to account for the effect of seasonality on trapping
results) there was a significant difference between the
years for P. albocinereus (Table 3). There were less P.
albocinereus in autumn 2012, similar to the results using
both 2012 surveys (autumn and spring; Figure 4). Mean
mammal abundance, mammal species richness and M.
musculus, Sminthopsis sp(p). and T. rostratus abundances
were unchanged between the two surveys (Table 3).

Sites

of Mus musculus (b), Pseudomys albocinereus (c), and Tarsipes

Table 4. Capture data for Tarsipes rostratus, Mus musculus, Pseudomys albocinereus and Sminthopsis sp(p)., including the
number of captures, sex ratio and body weights of males and females in 1986 (range) and 2012 ( ± SD, range); 1986 data
taken from Burbidge et al. (1996).

Species

Captures
in 1986

Captures
in 2012

Sex ratio
(M:F) 1986

Sex ratio
(M:F) 2012

Body weight
males 1986

Body weight
males 2012

Body weight
females 1986

Body weight
females 2012

Tarsipes rostratus

14:16

12:10

5.3-7.0

6.78 ± 4.19
(3.1-17.5)

6.0-11.3

7.5 ± 3.92
(2-12)

Mus musculus

234

9:5

67:97

8-17.5

11 ±4.14
(4-32)

8.5-14.5

9.54 ± 2.62
(3-19)

Pseudomys albocinereus 23

10:13

11:6

16.2-32

17.93 ± 6.45
(10-28)

15.5-28

20.4 ± 6.78
(10.5-25.5)

Sminthopsis sp(p).

5:2

14:12 (3
unknown
sex)

9.5-11.5

8.25 ± 4.43
(5-19)

9.5-13.5

10.4 ±4.87
(4.1-18)

Journal of the Royal Society of Western Australia, 99(1), 2016

Sites

Figure 4. Pseudomys albocinereus abundance in 1986 and
autumn 2012 at each of the six sites.

DISCUSSION

Current status of small mammals in Boonanarring
Nature Reserve

Ecosystems on the Dandaragan Plateau and northern
SCP, including BNR, have been severely impacted by
clearing, fragmentation, and decreasing rainfall over
the last 40 years but there is little knowledge of the
significant impacts on mammal fauna and their habitats
(Wilson et al. 2012, Woinarski et al. 2015). This study has
contributed important information to the knowledge of
the distribution and abundance of native small mammal
species in BNR, a significant and large reserve on the
southern Dandaragan Plateau (Burbidge et al. 1996).

Of particular importance is the information obtained
on P. albocinereus , for which records have been infrequent
over the last few decades. This native rodent has a broad
distribution over coastal and inland areas of Western
Australia, ranging from Bernier Island in the north¬
west to Israelite Bay in the south-east (Van Dyck &
Strahan 2008). While it was trapped at a range of sites
in woodland and heath habitats on the northern SCP
by Kitchener et al. (1978) it has not been recorded there
since 1978. The records in this study are significant as
they confirm the presence of the species at least on the
Dandaragan plateau, some 26 years after previous studies
(Bamford 1986; Burbidge et al. 1996).

Pseudomys albocinereus is classified as Least Concern
on the IUCN Red List (Morris et al. 2008), and not listed
under the Western Australian Wildlife Conservation
Act 1950 (Department of Parks and Wildlife 2014), even
though its former range has been reduced by clearing for
agriculture and infrastructure (Bleby et al. 2009). Recent
work on P. albocinereus showed a preference for relatively
dense vegetation cover at ground level compared to
available microhabitat and suggested that the species
may compete with Mus in some habitats. Home range
size was estimated to be ca. 1.7 ha and most shelters
were in burrows or grass trees (Xanthorrhoea sp.) (Smith
2015). Up until this study and Smith's (2015) there was
a lack of knowledge about the habitat selection and
behaviour of P albocincereus (Bamford 1986). However,
more data and information are required to develop

a detailed understanding of the relationship of P
albocinereus with habitat complexity and fire age, as well
as nesting and habitat use behaviour, in order to facilitate
informed management. This is particularly important as
ongoing habitat fragmentation, decreasing rainfall and
inappropriate fire regimes endanger the current range
of this species, therefore putting local populations at risk
(Bleby et al. 2009, Wilson et al. 2014, Woinarski et al. 2014).

One interesting piece of information on P. albocinereus'
ecology that was noted, was the slightly higher number
of captures of males in spring. In the 2012 trapping,
twice as many males were caught as females. In 2014
trapping of the same sites, again more males were
captured than females (Smith 2015). Yet in autumn 1986
and 2012 surveys similar numbers of each sex were
caught. This suggests that males are more mobile in
spring, presumably searching for mates, but during
non-breeding seasons (autumn) do not disperse as
far. However, more research is required to test this
hypothesis.

Captures of T. rostratus at eight sites scattered
across the reserve in 2012 were indicative of long term
persistence of the species in BNR some 26 years after the
Burbidge et al. (1996) study. Tarsipes rostratus is endemic
to the south-west of Western Australia. Although
the species is not considered to be threatened, it is
noteworthy taxonomically as it is the only species in the
Lamily Tarsipedidae. The species is restricted to coastal
sandplain heaths and low open woodlands with healthy
understorey (Wooller et al. 2004, Bradshaw et al. 2007).
Most studies of the species have been conducted in a
cool climate in continuous habitats across the south-coast
of Western Australia and may not be representative of
much of the range of this species, which is distributed
from Shark Bay to the edge of the Nullarbor Plain
(Garavanta et al. 2000, Wooller et al. 2004, Bradshaw et al.
2007, Dundas et al. 2013). It is also commonly captured
in studies on the northern SCP, where the habitat is
more open and lacks the connectivity of the south-
coast habitat, often being surrounded and fragmented
by pine plantations (Clancy 2011, Wilson et al. 2012).
Tarsipes rostratus is evidently an adaptable species found
in a range of vegetation types, from those with a dense,
flowering understorey to open woodlands, both of
which are found in Boonanarring Nature Reserve on the
Dandaragan Plateau.

In 1986 S. griseoventer was captured at only two sites
in BNR, whereas in the 2012 survey, individuals of
Sminthopsis were captured at all sites except for two.
Only one individual (considered likely to have been S.
griseoventer ) has been recorded recently on the SCP in
2007-08 (Wilson et al. 2012) indicating that at least one
Sminthopsis taxon is still extant, albeit in small numbers.
The confirmation of the presence of a considerable
Sminthopsis sp(p) population in BNR is important as there
is evidence that other vegetation remnants on the SCP are
largely devoid of the species.

Status of mammals in 2012 compared to 26 years
previously

Species richness (overall number of species) of small
mammals in BNR was maintained between the two
survey years (1986 and 2012) with four mammal species
recorded: M. musculus, P. albocinereus , Sminthopsis sp(p)

T. Moore et al.: Mammal assemblages in Boonanarring Nature Reserve

(S. griseoventer in 1986) and T. rostratus. The significant
difference in the mean species richness between autumn
2012 and 1986 surveys is most likely a reflection of
differences in captures at the site level. There is therefore
no evidence that any small mammal species has been lost
from the reserve in the last 26 years.

Overall, the trapping revealed more M. musculus
captures in 2012 than 1986, with the majority of the 2012
captures being in autumn. The results are consistent with
the findings of Bamford's (1986) study at a nearby reserve
where M. musculus was captured more commonly in
autumn than any other time of year. Autumn provides
optimal resources for breeding and survival for this
introduced rodent (Brown & Singleton 1999). Increased
presence of M. musculus is particularly concerning as
the species may compete with native rodents for nesting
and shelter sites as well as food resources (Smith & Quin
1996).

In contrast to Mus, the abundance of P. albocinereus and
T. rostratus appeared to be lower in 2012 than 1986. There
is a need to confirm if these differences are the result
of a declining population trend, by undertaking more
repeat measure monitoring. If these differences reflect a
decline, they could be a result of many factors including
inappropriate fire regimes, changes in water availability,
introduced weeds and predators (Torre & Diaz 2004,
Bleby et al. 2009, Craig et al. 2010, Wilson et al. 2014).
Several studies in the northern SCP and the south-coast
of Western Australia demonstrate that higher abundances
of T. rostratus are found in complex habitat with a fire
age of 15-20 years (Bamford 1992, Garavanta et al. 2000,
Friend & Wayne 2003, Clancy 2011). Future studies
should measure habitat complexity (as well as fuel age)
to determine the habitat requirements of T. rostratus and
P. albocinereus present in the reserve. Water is required for
plants and shrubs to provide food (flowers or seeds) for
T. rostratus and/or P. albocinereus (Wilson et al. 2012); with
declining rainfall in this area over multiple decades, such
factors may be playing a role in the declining abundance
of these small mammals. Lastly, although our traps were
not appropriate for capturing introduced predators,
passive sampling methods did reveal the presence of
Felis catus and Vulpes vulpes in both 1986 and 2012, and
Canis familiaris in 1986 (Burbidge et al. 1996; unpublished
data), all of which are predators of small native mammals
(Doherty et al. 2015). Future work should expand on the
knowledge of T. rostratus and P. albocinereus in BNR and
their habitat requirements.

Evaluation of problems associated with long term
monitoring in this project

For many long term studies, trap design, location and
effort are difficult to replicate. In this study there were
minor differences in the 2012 surveys compared to the
1986 surveys. In 2012 it was difficult to determine the
exact location of the traps from the 1986 survey using
only GPS coordinates, so the traps were located as close
as possible. Positions of the trap arrays may have differed
by tens of metres between 1986 and autumn 2012, and it
is likely that different microhabitats were sampled, and
the differences in trap array design (lines vs Y-shapes)
would have meant that, for animals with small home
ranges, different numbers of home ranges would have
been intersected, and different proportions of dispersing

animals might encounter a trap (Friend et al. 1989). The
number of survey nights also varied, with six nights
in autumn 1986 and eight nights each in autumn and
spring 2012 (Table 2). The number and types of traps
used also differed between the two surveys (Table 2).
With these caveats in mind we chose to compare trapping
rates as captures/trap nights. Taking these caveats into
consideration is also important when assessing any
statistical differences identified (Friend et al. 1989, Rolfe
& McKenzie 2000, Garden et al. 2007, Environmental
Protection Authority & Department of Environment and
Conservation 2010).

CONCLUSION

Boonanarring Nature Reserve to date has provided a
sustainable habitat for small native mammals including P.
albocinereus, Sminthopsis sp(p). and T. rostratus, most likely
due to its large area of remanent vegetation encompassing
many habitat types. The continued presence of these
species requires active management actions such as
appropriate burning regimes and predator control,
followed by evaluation of population trends.

ACKNOWLEDGEMENTS

Many thanks to all those who assisted in the field.
Special thanks go to Karen Bettink and Alice Reaveley
who were involved in site selection, trap installation and
survey work in 2012. Thank you also to Nicole Godfrey,
Ben Kreplins, Craig Oljenik and conservation employee
crew members from Swan Coastal District. This work was
completed with approval from the Department of Parks
and Wildlife Animal Ethics Committee (DPAW 2011 and
2013/31). Thanks also go to Natalie Warburton for her
editorial comments.

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Journal of the Royal Society of Western Australia, 99(1): 9-15, 2016

Environmental characteristics of ephemeral rock pools explain local
abundances of the clam shrimp, Paralimnadia badia (Branchiopoda:
Spinicaudata: Limnadiidae)

ALISSA CALABRESE 1 , CHERIE MCCULLOUGH 2 , BRENTON KNOTT 34
& STEPHEN C. WEEKS 1

1 Program in Integrated Bioscience, Department of Biology, University of Akron, Akron, OH 44325-3908, USA.

2 School of Natural Sciences, Edith Cowan University, 270 Joondalup Drive, Joondalup, WA 6027, Australia.

3 School of Animal Biology, University of Western Australia, 35 Stirling Highway, Crawley, WA 6009, Australia

ABSTRACT

The conditions of ephemeral freshwater pools are highly variable through time, and their
inhabitants must be able to tolerate these changing conditions to survive. Although much research
has focused on large branchiopod hatching requirements, there is comparatively little information
available on the environmental conditions endured by adult clam shrimp populations. A suite of
physical and chemical characteristics, especially pool depth, influence the presence or absence of
clam shrimp populations in rock pools on granite outcrops in the Wheatbelt region of Western
Australia. Here we examine multiple environmental variables of temporary rock pools and how
they may affect adult populations of the clam shrimp, Paralimnadia badia.

KEYWORDS: habitat, physiochemical, temporary pool, pool morphology, hydroperiod,
environmental variability

INTRODUCTION

Clam shrimp are branchiopod crustaceans
(Branchiopoda: Spinicaudata) that are obligate dwellers
of temporary waters (Dumont & Negrea 2002).
Populations survive dry periods in the form of resting
eggs produced by mature females or hermaphrodites.
Their eggs lie dormant until appropriate conditions are
met for hatching (Brendonck 1996, Brendonck & Riddoch
2000, Brendonck & De Meester 2003). Upon inundation,
a fraction of resting eggs hatch, leaving behind a bank
of dormant eggs in the sediment (Brendonck 1996,
Brendonck & De Meester 2003). Occasionally, eggs
will continue to hatch even after the initial inundation,
resulting in multiple generations of shrimp in a single
pool (Benvenuto et al. 2009). Often, rain will fill the pool
and eggs will hatch but not reach sexual maturity before
the basin dries. In these instances, the egg bank is vital to
ensure long-term population persistence (Brendonck &
De Meester 2003). In this way shrimp 'hedge their bets'
against the unpredictability of their environment which
has led to speculation as to which factors might promote
this pattern of hatching (Mossin 1986, Brendonck et al.
1996, Kuller & Gasith 1996, Simovich & Hathaway 1997,
Beladjal et al. 2007).

Many physical environmental parameters, including
dissolved oxygen, pH, osmotic pressure, light, and
temperature, have been individually implicated in

4 Deceased March 2013

regulating hatching in various large branchiopod
species (Bishop 1967, Belk 1977, Scott & Grigarick 1979,
Mitchell 1990, Schonbrunner & Eder 2006). However,
initial hatching conditions are not sole predictors of
clam shrimp presence, as temporary pool environmental
conditions change throughout the duration of their
inundation and drying regimes (Jocque et al. 2007a).
Here we aim to determine environmental factors that are
associated with maintenance of adult populations of clam
shrimp.

For a hatched population to be sustained, the
conditions of the pool must meet survival requirements
after initial hatching. Long term maintenance of
populations of several species of Daphnia have been
assessed in relation to habitat characteristics and each
species appeared to thrive under different environmental
conditions, occupying different pools as a result (Pajunen
& Pajunen 2007). Both the size and permanence (length
of hydroperiod) of temporary pools have been found
to play important roles in predicting species richness
(Kiflawi et al. 2003). Additionally, since larger pools will
resist evaporation for longer periods of time, pool size
has been shown as an important factor structuring local
communities (Jocque et al. 2007b)

Our study sought to answer how temporary pool
environments relate to adult clam shrimp populations.
Here we determine which parameters best explain
abundances of adult populations of the clam shrimp
Paralimnadia badia (Wolf 1911; the Australian species
Limnadia have recently been moved to a new endemic
genus Paralimnadia, Rogers et al. 2012) in a variety of
temporary rock pools on granite outcrops in the semi-
arid, Wheatbelt region of Western Australia.

Journal of the Royal Society of Western Australia, 99(1), 2016

METHODS

Study Sites

Although the Wheatbelt region of southwestern. Western
Australia is generally flat, occasional large granite
outcrops (or 'inselbergs' sensu stricto, Withers 2000) can
be found in the area (York Main 1997, Withers 2000). The
rainy season in this part of Australia lasts approximately
from May to October and is the milder time of the year,
with mean maximum air temperatures ranging from
20.3°C to 23.7°C (Commonwealth of Australia, Bureau of
Meteorology, 26 Sept 2011).

Three outcrops received adequate rainfall for
sustained hydration: Holland Rock (Shire of Kent,
33°21.259'S; 118°44.639'E; 13 pools sampled). Dingo Rock
(Shire of Lake Grace, 33°0.558’S; 118°36.321’E; 8 pools
sampled), and Rockhole Rock (Shire of Bruce Rock,
31°55.970'S; 117°45.209'E; 11 pools sampled). Data from
Bruce Rock were collected at a later date than Holland
and Dingo (May 2009) and were only used to construct
Fig. 3.

Field Sampling

Study pools were selected for a range of sizes and depths,
and were sampled for physiochemical parameters and
quantities of clam shrimp and other macroinvertebrates
in late April to late May 2007. Water quality [temperature,
pH, dissolved oxygen (DO), and electrical conductivity
(EC)] of each of the three sites was monitored for five
consecutive days at four times per day (12:00, 14:00,16:00,
and 18:00) to assess diurnal change, however for most
analysis only the average of the mid-afternoon readings
(14:00 and 16:00) were used as a result of incomplete
sampling across the other two time periods. Pools on
Holland and Dingo Rocks were sampled with a YSI 556
multi-probe (YSI, USA). Pools on Rockhole Rock were
sampled with a Hydrolab Quanta multi-parameter meter
(Hydrolab, USA). Pool dimensions measured included
maximum length (1), maximum width (w) and mean
depth (d) (each averaged from three measurements). Pool
volumes were estimated from these dimensions assuming
the shape to be half of an ellipsoid (Formula: 4/37ilwd,
Baron et al. 1998).

Each pool was sampled across the entire volume
for three, 3-min periods with sweep nets of mesh sizes
2.0 mm and then 0.5 mm. Paralimnadia badia abundance
approximately halved each sampling event over the three
successive rounds of sampling. Consequently, after three
rounds in each pool approximately 90% of clam shrimp
had been captured. Paralimnadia badia were sorted from
other macroinvertebrates in white plastic collecting trays,
species were identified, counted, and then returned to the
pool alive.

Analysis of habitat variables and species distributions

Volume and EC values were log 10 transformed to achieve
normality. All other variables had normally distributed
residuals (using Shapiro-Wilk test for normality). One¬
way ANOVAs compared the means of environmental
variables between pools with and without P. badia.

All parametric analyses were made with JMP Pro 10
(SAS Institute Inc. 2012). Ordinations of environmental

data were produced by Principal Components Analysis
(PCA) in PRIMER multivariate software (PRIMER-E
2006). Although not suitable for analysis of biotic
community structure, the implicit underlying Euclidean
Distance matrix of this method makes it suitable for
environmental data, with an advantage over nMDS
ordination in that ordination axes are interpretable
(Clarke & Warwick 2001). Correlations between pairs
of environmental variables were then examined with
draftsmen plots. Combinations of highly correlated
variables (Spearman rank higher than 95%) were reduced
to a single, representative variable.

The PRIMER BEST procedure (Clarke & Ainsworth
1993) identified the combination of mean water quality
and habitat variables for each pool, which best rank-
correlated with clam shrimp abundances. These best
correlating environmental variables are likely to be
those that are most important in defining clam shrimp
abundance. Prior to BEST analysis, environmental
variables were normalized to the maximum value
encountered (Olsgard et al. 1997, Clarke & Warwick 2001)
and draftsman plots were created to determine which
variables were highly correlated with each other (i.e.,
95% or greater. Bob Clarke, Plymouth Marine Laboratory
UK pers. comm.). Data were then log 10 transformed to
enhance a linear relationship between variables and
finally standardized to account for different variable
scales (Clarke & Warwick 2001).

A canonical discriminate analysis (CDA) was then
used (JMP Pro 10, SAS Institute Inc.) to determine which
environmental variables best described the variation
between groups (presence / absence of P. badia; see
below). Bivariate regressions were then made to describe
relationships between P. badia abundance, estimated
fitness and habitat pH. Fitness was estimated from the
size of the carapace, which is directly related to the
number of resting eggs per clutch in female clam shrimp
(Weeks et al. 1997).

RESULTS

Of the 21 pools studied on Holland and Dingo Rocks,
13 contained clam shrimp populations. Pools with and
without P badia had different abiotic conditions. Pool
depth was related to pool volume and surface area
(shown as factor loadings clustering on the loading plot;
Fig. 1), which is expected because of the common formula
used to derive pool volume. Examined individually, pH
was the only abiotic parameter that differed significantly
between the two groups (Table 1). Pools containing P
badia typically had lower pH than those without clam
shrimp. Pools containing P badia had overlapping ranges
of all variables with pools lacking P badia. The water
temperature of the pools without P badia was also higher,
but not significantly so (p=0.06). Pool volume, surface
area, conductivity, and TDS were higher in the pools that
had P badia populations, albeit not significantly.

Mean depth was the most important sole
environmental variable explaining clam shrimp
abundance according to the BEST analysis (Spearman
Rank correlation: rho = 0.375). When pH was added to
this model, this water quality variable further helped to
explain clam shrimp distribution at rho = 0.413.

Temperature
Dissolved Oxygen
pH
Depth

Surface Area
Conductivity
Volume

Principal Component 1
-0.35*

-0.07

- 0.21

0.49*

0.52*

0.02

0.56*

Principal Component 2
0.46*

-0.49*

0.55*

0.26

0.10

-0.38*

0.12

Figure 1. PCA of pool
environmental variables. Length
of vector indicates strength of
correlation between variable
and component axes. Asterisks
indicate significant correlation of
the component with the original
variable (*p<0.05). Pools with P.
badia present are filled circles. Pools
without P. badia are empty circles.

Table 1. Habitat characteristics of 21 total pools with P badia present (n=13) and absent (n=8). Bold type indicates
statistically significant difference (p<0.05) between pool environmental variable with and without P badia.

Pools without P. badia Pools with P. badia

Mean

Max

Min

Mean

Max

Min

F-ratio

p-value

Temperature (°C)

21.2

0.41

26.1

17.3

20.3

0.25

25.8

17.2

3.95

0.061

Conductivity (mS/cm)

0.24

0.03

0.47

0.06

0.27

0.04

0.93

0.04

0.07

0.799

Dissolved oxygen (%)

113

2.75

138

97.3

109

1.60

141

88.9

1.46

0.243

7.09

0.17

9.03

5.91

6.64

0.13

8.57

5.53

4.39

0.049

Depth (mm)

30.7

5.36

70.7

4.33

39.0

2.48

68.3

17.7

2.56

0.126

Volume (L)

136

41.3

420

1.71

276

59.9

930

24.9

2.83

0.109

Surface area (m 2 )

7.32

1.41

14.2

0.75

12.2

2.14

28.5

1.98

2.75

0.114

Journal of the Royal Society of Western Australia, 99(1), 2016

18.0-

‘5

16.0-

14.0-

12 . 0 -

10 . 0 -

28 29

Canonical 1

p-value

Temperature

-0.48

0.006*

Dissolved Oxygen

-0.18

0.096

3.08

0.004*

Depth

0.05

0.027*

Surface Area

-0.03

0.022*

Conductivity

-0.94

0.721

Volume

-0.91

0.020*

Figure 2. Results of a canonical discriminate analysis (CDA) on all seven dependent variables using data from Holland
and Dingo outcrops only. Pools without P. badia present are empty circles; pools with P badia present are filled circles.
Canonical correlation 1 explains -100% of the differences between present/absent pools. Ellipses show the means of the
centroids of P badia -present pools and P badia-absent pools. Asterisks indicate significant correlations of the canonical
scores with each of the original variables (*p<0.05).

Figure 3. Regression of CC1 on the
quantity of P badia present in all pools.
Canonical 1 (consisting largely of
volume, depth, and surface area) is
negatively correlated with the quantity
of P badia (p=0.0052).

Figure 4. Diurnal variation in pH levels in pools on Dingo and Holland Rocks.

The CDA correctly classified 17 of 21 pools in its
assignment of pools into groups that either had P. badia or
not, or 81% of the total pools in this comparison (Fig. 2).
The first canonical correlation (CC1) explained -100% of
the important variation that differed between pools with
and without P badia. Five original dependent variables
were correlated strongly with CC1: temperature, depth,
and surface area were negatively correlated with CC1,
while volume and pH were significantly positively
correlated with CC1 (Fig. 2). Volume, depth, and surface
area are all measures describing pool size. Temperature
and pH are variable that are closely associated and
influenced by pool size, which suggests that CC1 is
mainly describing the size of the pools. This 'pool size'
variable is negatively correlated with the quantity of P.
badia (Fig. 3). Figure 4 displays the diurnal variability of
pH in pools on Dingo and Holland rocks.

DISCUSSION

Our study confirms that temporary rock pools are highly
variable environments (Bayly 1982, Williams 2001). To
survive this environment, pool inhabitants must cope
with extended periods of drought between wet phases
as well as the widely variable conditions throughout the
inundation phase.

There is a wealth of literature available that quantifies
the habitat requirements for large branchiopods to
break dormancy (Moore 1963, Belk 1977, Brendonck
1996, Schonbrunner & Eder 2006). The information
available regarding the maintenance of populations of
clam shrimp mainly suggests that the size of the pools
is of key importance (Kiflawi et al. 2003 & Jocque et
al. 2007b). Frequently, clam shrimp nauplii will hatch
from resting eggs, but then the population will crash
before they have reached maturity (Jocque et al. 2007b).
Much of the available information regarding hatching
or sustaining populations is focused on the range of a
single parameter (i.e., only temperature or pH; Belk 1977,
Scott & Grigarick 1979, Mitchell 1990, Schonbrunner &

Eder 2006). However, by combining all parameters into
a multivariate BEST analysis, we showed that pool depth
combined with pH best explained P. badia abundance
in temporary outcrop pool habitats. A canonical
discriminate analysis confirmed pool depth and pH as
the most vital explanatory variables, while additionally
implicating volume, surface area, and temperature as
important habitat variables. Temperature was negatively
correlated with volume, depth, and surface area, as might
be expected: as pool sizes increase, maximum pool water
temperatures decrease because larger pools can be more
resistant to temperature change. The sampling times
used in this analysis were from the warmest parts of the
diurnal cycle, and thus the larger volume pools should
have been more resistant to warming, explaining the
negative correlation.

The environmental variables that described pool
morphology were all positively correlated with one
another and with CC1. Canonical correlation 1 correctly
predicted which pools contained P. badia 81% of the
time, illustrating that pool morphology was a key habitat
characteristic for P badia abundance. This may well be
because pools that are on the more ephemeral end of the
continuum evaporate before clam shrimp populations
have reached sexual maturity.

The apparent association between pH and the
presence of P badia (Table 1) is likely an artifact of the
sampling regime (i.e., pH values used in this analysis
were only from the middle of the day and thus do
not take into account the significant diurnal change
observed; Fig. 4). It is possible that the variability of pH,
like temperature is associated with the size of the pool
and this is why it appears significant here. Otherwise it
is not obvious why such a subtle difference in mean pH
between the two groups (0.45) would have an impact on
P badia populations. More information is necessary to
determine the significance of this observation.

Clam shrimp species previously reported from these
pools were Eulimnadia dahli Sars, 1896 and Paralimnadia
badia (Timms 2006, Weeks et al. 2006). Whether these

Journal of the Royal Society of Western Australia, 99(1), 2016

genera never coexist within the same pool or whether
one succeeds the other in the same pools is still unknown.
Preliminary evidence suggests that P. badia may
predominate in pools in the winter (rainy) season, while
E. dahli may be more common in the summer (Timms
2006).

These data represent the results of an observational
and not a manipulative study and so can only suggest
a potential relationship between the above variables,
not necessarily a direct causal relationship. There
may also be other factors beyond pool morphology
that could affect the distribution of this species. It is
possible that clam shrimp had not yet dispersed to the
non-P. badia pools, although this scenario is unlikely
considering the proximity of the pools to each other on
each outcrop. Overflow and wind-mediated propagule
dispersal have been shown to be extremely effective in
temporary pool metacommunities (Brendonck & Riddoch
1999, Vanschoenwinkel et al. 2008a, Vanshoenwinkel
et al. 2008b). Another possibility is that the presence
of a predator excluded P badia from some pools and
not others, although this seems unlikely. Some clam
shrimp were consumed by predacious water beetle
larvae (Dytiscidae) during sampling, but this was a rare
occurrence in our observations. Furthermore, in every
pool in which these predacious larvae were present, P
badia was also present. Additionally, 81% of pools that
harboured anuran larvae also had P badia. Our sampling
was not exhaustive enough to prove that no predator
had an effect on P badia populations, but simply suggests
that we did not observe such a phenomenon during our
sampling period.

Generally, there is thought to be a continuum of
factors structuring communities in habitats ranging from
permanent freshwater bodies to temporary freshwater
environments (Wellborn et al. 1996). Some work has
suggested that predation is the dominant structuring
force in the most permanent communities while
competition is more powerful in structuring pools that
are less permanent (Wellborn et al. 1996, Wilbur 1987).
Indeed, priority effects may play a role in early colonizers
monopolizing particular pools (De Meester et al. 2002,
Jocque et al. 2010).

Another factor that may affect the habitat quality of
P badia is dryland salinity. Extensive land clearing in
the region has resulted in increased salinities in many
aquatic habitats (Anon 1996, National Land and Water
Resources Audit 2001). Dust and saline solutes from salt
flats around the rock outcrops are likely to be both more
alkaline and also more buffered. Some outcrop pools
are also showing higher alkalinity through increased
bicarbonate concentrations than would be expected from
rainfall interaction with acidic granite rock (Pinder et al.
2005).

Pool morphology appears to influence the presence of
P badia in temporary rock pools. Temperature may also
be an influential factor, especially as it is correlated with
the size of the pools. Many of these outcrops are already
functioning as islands in an ecological desert that is the
mono-agricultural Wheatbelt landscape. Consequently,
loss of these island habitats to clam shrimp at a local scale
may lead to a significant threat to sustained existence for
clam shrimp at a more regional scale.

ACKNOWLEDGEMENTS

We would like to dedicate this paper to our colleague
Brenton Knott, who passed away March 2013. His
friendship, advice, and humour are dearly missed. We
are very grateful to Chiara Benvenuto, Wally Gibb, Debra
Judge, Kerry Knott, Sadie Reed Stimmell, Danny Tang,
Magdalena Zofkova and the School of Animal Biology at
the University of Western Australia for their instrumental
support in collecting this data. We thank Tony Cockbain,
Patrick Armstrong, Brian Timms and one anonymous
reviewer for their valuable advice. This project was
financed by the Department of Biology at the University
of Akron and the Kevin E. Kelleher Memorial Fund.

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Journal of the Royal Society of Western Australia, 99(1): 17-26, 2016

Chronostratigraphic context for artefact-bearing palaeosols in late
Pleistocene Tamala Limestone, Rottnest Island, Western Australia

I WARD T J PIETSCH 2 , E J RHODES 3 , G H MILLER \ J HELLSTROM 5 & C E DORTCH 6

1 School of Social Sciences, University of Western Australia WA 6009, Australia

2 Australian Rivers Institute, Griffith University, Qld 4111, Australia

3 Department of Earth, Planetary and Space Sciences, University of California, CA 90095, USA
4 INSTAAR and Dept. Geological Sciences, University of Colorado, Boulder, CO 80309 USA

5 School of Earth Sciences, University of Melbourne, Vic 3010, Australia

6 24 Howard Street, Fremantle, WA 6160 Australia

* Corresponding author ilSSI Ingrid.ward@uwa.edu.au

Abstract

This paper presents absolute dates for artefact-bearing palaeosols intercalated with Tamala
Limestone aeolianite successions on Rottnest Island (Wadjemup), Western Australia. The absolute
chronology for the sub-aerial part of the island's constituent Tamala Limestone is based on 20
optically stimulated luminescence (OSL), two thermoluminescence (TL) age estimates, one U-series
assay and three radiocarbon dates. The oldest of these estimates is an OSL age of 140 ± 14 ka at
Fairbridge Bluff for the aeoliniate beneath the Rottnest Limestone member - a marine member of
the Tamala Limestone succession. The palaeosols range in age from 49 to 10 ka. These age estimates
provide, for the first time, a chronostratigraphic context for the isolated archaeological finds on
the island as well as contributing to the timing and nature of late Quaternary sequences within the
Tamala Limestone of the Perth Basin.

KEYWORDS: Rottnest Island, late Quaternary, Tamala Limestone, palaeosols,
chronostratigraphy, luminescence dating

INTRODUCTION

Rottnest (or Wadjemup) is the largest of a chain of offshore
islands and reefs composed of Tamala Limestone, the
constituent rock of Quaternary age throughout the Perth
Basin (Playford 1983, 1988, 1997) (Figure 1). Separating
the sandy beaches around the island are aeolian
calcarenite cliffs and headlands that formed when wide
areas of the continental shelf was exposed and carbonate
productivity was high - this is the Tamala Limestone
(Playford 1988, 1997). These aeolianite successions
comprise moderately cemented, fine to coarse bioclastic
sands and shelly deposits, mainly in large scale cross
bedded and planar bedforms (Hearty 2003). Intercalated
within the aeolianite successions are moderately
cemented calcretes and palaeosols (the "protosols"
of Vacher & Hearty 1989), characterised by higher
proportions of quartz and clay, calcified roots (rhizoliths),
fossil land snails (mainly the gastropod Austrosuccinea
sp.) and fossil weevil ( Leptopious sp.) pupal cases (see also
Playford 1997; Hesp et al. 1999).

It is within these palaeosol units that a number of
isolated Eocene fossiliferous chert and calcrete artefacts
are recorded at Bathurst Point, Little Armstrong Bay,
Charlotte Point, Fish Hook Bay (Figure 1) along with
seven other surface finds. Although few in number, the
stone artefacts identified in situ within Tamala Limestone
paleosols on Rottnest Is. (Dortch & Hesp 1994), along
with a further nine found on Garden Island (Dortch &

Morse 1984; Dortch 1991; Dortch & Hesp 1994; Dortch
& Dortch 2012) indicate human presence on this part of
the emergent continental shelf during the late Pleistocene
and early Holocene. Minimal ages for these artefacts have
been estimated from the time of the island's separation
from the mainland around c. 6500 years ago (Churchill
1959; Playford 1997). This paper provides a more
complete stratigraphic context and absolute chronology
for the artefact-bearing palaeosols.

In situ artefact find sites and previous dating

The Tamala Limestone provides a register of Quaternary
sea-level events (Teichert 1950; Fairbridge 1954, 1961;
Playford 1988, 1997; Kenrick & Wyrwoll 1991; O'Leary
et al. 2013; Brooke et al. 2014) and is also important from
a regional archaeological perspective (see also Dortch
& Dortch 2012). During the Last Interglacial (Marine
Isotope Stage (MIS) 5), Rottnest Island existed as shallow
submerged shoals and reefs overlying Tamala Limestone,
as evident from the Rottnest Limestone member at
Fairbridge Bluff (Figure 2A). The reef/beach sequence
here has been dated between 132 - 121 ka (MIS 5e) (Szabo
1979; Stirling et al. 1995, 1998; Price et al. 2001). Mean sea
level (MSL) was about 3 - 4 m above present for much
of this period, increasing to at least 8 m above present
around 118 ka (O'Leary et al. 2013). This reef/beach
sequence is capped by a reddish calcrete and terra rossa
palaeosol with deep root casts filled with ferruginous,
well-rounded, well-sorted quartz-rich dune sands (Figure
2A), indicating sub-areal weathering of the reef towards
the end of MIS 5e (or possibly MIS < 5d, Hearty 2003;
Hearty & O'Leary 2008).

Journal of the Royal Society of Western Australia, 99(1), 2016

Little

Armstrong
Bay

City of York
Bay

Fish Hook
Bay

Thompson
Bay

2 km

Figure 1. Rottnest Island map showing location of artefact-bearing palaeosols, including Bathurst Point, Little Armstrong
Bay, Charlotte Point, City of York Bay and Fish Hook Bay. Inset A shows location of Rottnest Is. in relation to the Perth
Metropolitan Region, central Swan Coastal Plain and Swan River/estuary. Inset B shows sea level curve showing major
marine isotope stages (sourced from Waelbroeck et al. 2002).

Additional absolute ages relate to MIS 4 and are
provided from Bathurst Point (Figure 1). Here four
prominent stratigraphic units are observed - the first
is a basal aeolianite unit 3 - 7 m ASL to below MSL,
extending along the c. 500 m cliff length, dated by
thermoluminescence (TL) at 67 ± 9 ka (Price et al. 2001)
and by Optically-Stimulated Luminescence (OSL) dating
to 77 ± 12 ka (Playford et al. 2013; Brooke et al. 2014).
This unit is capped (trunctated?) by a massive calcrete/
breccia calcrete unit (Figure 2B), defined by Hearty
(2003) as a well-developed brecciated soil (rendzina) with
abundant limestone clasts. Along parts of the eastern
side of Bathurst Point, this calcrete/breccia unit can been

seen to be overlain by a dark brown to nearly black silty
sand (Figure 3). It is within this brecciated calcrete that a
fossiliferous chert core was found (Dortch & Dortch 2012:
their Figure 7), cemented and overlain by a thin layer (0.5
- 1 cm) of carbonate cement (see also Hearty 2003).

Overlying this calcrete/breccia unit is a younger
aeolianite succession, exposed mainly on the western
side of Bathurst Point, and dated by OSL to 27 ± 4.5 ka
(Brooke et al. 2014) and 20 ± 2 ka by TL (Price et al. 2001)
or MIS 2. This aeolinite unit is not present on the east
side of Bathurst Point, at the find site of the chert artefact.
Here the calcrete/breccia unit is immediately capped by
a light brown (7.5 YR 5/3) rhyzolithic-rich palaeosol that

I. Ward et al .: Dating of palaeosols on Rottnest

Fairbridge Bluff

10 r

Complex dune
cross-bedding

Terra rossa palaesol,
with deep root casts filled with
well-rounded, red quartz sand

Rottnest limestone reef
-MLW

47 3+4.5 ka

105+17 kyr
(W2631)

132+181
(W2630)
140 + 14

TgI sand
^ aeolianite
jfn palaeosol
calcrete
$ rhizolith
- shell debris
® shell (Turbo sp.)
O eggshell
© charcoal
0 OSL
© artefact
(g coral

Bathurst Point

Unconsolidated brown
(10 yr 8/1) carbonate sands

Strongly cemented, brown
(7.5 YR 5/3) rhyzolithic
palaeosol with Leptopius sp

ka (R12/K0035) 10

- MLW

East

kVesf

YA =
OA =

carbonate lens

f7± 1.7 ka (BP2.2)

Brecciated calcrete
42,0+2.7 ka
(R11/K0034)

Low angle cross-beds

Little Armstrong Bay

East

Calcarenite

boulders

35.9+2.1 ka ..
(R3/K0020)

(R2/K0019)

Moderately cemented, mod sorted,
med-coarse, brown (10YR 4/4),_
quartz carbonate sand with
Austrosuccinea sp., Grading into
light brown (10YR 8/4) med sand

(GU S.1)

(GU 8.2)

(GU 8.3!

(GU 8.4)

0 ~ MLW

Charlotte Point

0 ~ MLW

12.7+1.25 ka
8.5)

12.1 + 1.01 ka
(GU 8.6)

Unconsolidated light brown (10 YR 8/1) W-jP*

"7 carbonate sand with abundant EjBt!

Bothriembryon sp. shells

Oelcrete (1 - ? cm) JWH

Aeolianite (horizonal beddina) HOB!

ZzEEsC^'xStrongly cemented, med-coarse, brown (7.5 YR 5/3)

^ 00 .

quartz carbonate sand with Austrosuccinea sp. shells
and Leptopius sp. pupal cases grading into lighter

brown (7.SYR 8/2) moderately cemented sand

Complex dune cross bedding

—-

Figure 2. Schematic stratigraphic sections of Rottnest Island Eocene fossiliferous artefact find sites, including A.
Fairbridge Bluff, B. Bathurst Point, C. Little Armstrong B and D. Charlotte Point. Inset petrographic photos show the
quartz-rich (with inherited ferruginous rinds) terra rossa palaeosol at Fairbridge Bluff (A), the contrasting carbonate-rich
palaeosol (with post-depositional ferriginisation) at Little Armstrong Bay (C) and Charlotte Point (D), and the sharp
boundary created by the calcrete overlying the aeolianite unit at Bathurst Point (B) (scale bar is 500 pm).

Journal of the Royal Society of Western Australia, 99(1), 2016

Figure 3. Brecciated calcrete unit on
the eastern side of Bathurst Point
overlain by a dark brown silty sand
and capped by a very thin carbonate
lens (photo by IW).

can be traced through the largely vegetated dunes across
to the western side of Bathurst Point where it overlies the
younger aeolianite (Figure 2B). The succession is overlain
by unconsolidated to lightly cemented Holocene dune
sands containing abundant Bothriembryon and introduced
land snail taxa (Hearty 2003).

A further two sites. Little Armstrong Bay and Charlotte
Point (Figure 1), have each yielded a single in situ Eocene
fossiliferous chert artefact from a Tamala Limestone
palaeosol unit, previously described by Dortch and Hesp
(1994). In contrast to the terra rossa palaesol at Fairbridge
Bluff, these palaeosols are mainly composed of shelly
carbonate, and the minor (5 - 10%) quartz that is present
is more angular (Figure 2C and 2D) indicating a less
distant source. The artefacts include a stone tool from the
foot of sea-cliff on the western side of Little Armstrong
Bay (LAB west) and a retouched tool, found half
protruding from a prominent palaeosol on the eastern
side of the bay (LAB east). The latter was associated
with a tiny fragment of emu eggshell ( Dromaius
novaehollandiae) - the sole vertebrate fossil specimen
known from the island. An earlier series of Amino Acid
Racemisation (AAR) measurements, using the land snail
Austrosuccinea sp. found in situ in palaeosols intercalated
within aeolianite successions at three Rottnest Island sites
- including Little Armstrong Bay, City of York and Fish
Hook Bay - gave a provisional ages of > 50 ka (Hesp et
al. 1999). However, these dates have been questioned by
Brooke et al. (2014), and Hesp et al. (1999: 11) themselves
regard their AAR dates as provisional due to "the paucity
of amino acid data concerning the racemisation kinetics
in [the genus] Austrosuccinea".

A further two fossiliferous chert artefacts were found
eroded from the summit of the cliffs at Fish Hook Bay,
on the island's south-west, and could have been derived
from the deflation of the uppermost palaeosol and
aeolianite units at the top of the cliff. The following
presents new dates for Fish Hook Bay, City of York

Bay and the artefact find sites at Bathurst Point, Little
Armstrong Bay and Charlotte Point.

DATING METHODS

OPTICALLY STIMULATED LUMINESCENCE (OSL)

Aeolianite (7) and palaeosol (5) units were sampled (by
EJR) in 2003 - 2004 at City of York, Fairbridge Bluff,
Bathurst Point and Little Armstrong Bay. A series of
small aliquot Optically Stimulated Luminescence (OSL)
measurements were undertaken at the Australian
National University (ANU), using a conventional
SAR approach for small aliquots, and standard Riso
equipment as described by Rhodes et al. (2010).
High OSL sensitivity and favourable luminescence
characteristics were observed, although some samples
displayed relatively high over-dispersion (statistical
scatter) between aliquots (up to 12%), possibly related to
spatial variability in beta dose rate within the carbonate
and quartz-rich aeolianite (Table 1). Dose rates were
determined using NAA-measured radionuclide contents,
along with in-field gamma spectrometry for gamma dose
rate calculation.

The above ANU dates are complemented by single¬
grain OSL dating of palaeosols and aeolianites at Little
Armstrong Bay and Charlotte Point (Table 1). These
OSL measurements, undertaken at Griffith University,
augment published OSL dating of samples collected at
Bathurst Point by the same laboratory (Table 1; Brooke
et al. 2014; Playford et al. 2013). The modified single
aliquot regenerative dose protocol of Olley et al. (2004)
was used on standard Riso equipment, as described in
detail elsewhere (Olley et al. 2004; Pietsch et al. 2013;
Brooke et al. 2014). Given over-dispersion values of 18
- 36% (Table 1), burial ages have been calculated using
the minimum age model. Radionuclide contents for each
GU sample were determined using Neutron Activation

I. Ward et ah: Dating of palaeosols on Rottnest

Analysis (NAA - Becquerel Laboratories, Mississauga,
Ontario, Canada), with lithogenic dose rates calculated
using the conversion factors of Stokes et al. (2003); with
(3-attenuation factors taken from Mejdahl (1979) and
cosmic dose rates calculated from Prescott & Hutton
(1994) using our best estimate of a time weighted long
term burial depth (Table 1) based on examination of the
field stratigraphy.

Uranium-series (U-series)

Six small sub-samples of between 20 and 150 mg were
cut using a hand-held dental drill from an 8 x 8 cm block
of carbonate cement containing the fossiliferous chert
core at Bathurst Point. The most indurated available
material was targeted, with a preference for the lightest
coloured. Analyses of six sub-samples followed the
procedure of Hellstrom (2003) and Drysdale et al. (2012).
Following Hellstrom (2006) an initial 230 Th/ 232 Th ratio
of 3.51 ± 0.87 was found to bring the Th-corrected ages
of all six analyses into agreement with respect to their
uncertainties (i.e. Mean square weighted deviation
(MSWD) = 1), meaning it is unlikely that the sample has
been affected by uranium mobility since its deposition
(Table 1).

Radiocarbon

Radiocarbon (Accelerator Mass Spectrometry (AMS))
radiocarbon dating of the emu eggshell fragment
(Dromaius novaehollandiae) (CURL-16668) recovered from
the palaeosol on the eastern side of Little Armstrong
Bay was undertaken at the University of Colorado. In
addition in situ charcoal fragments (Wk-37948) were
collected from a palaeosol 150 m south east of the
fossiliferous chert artefact find location and closely
adjacent to OSL sample positions at Charlotte Point.
The collective fragments were submitted to Waikato
Radiocarbon Laboratory for AMS dating.

RESULTS OF THE DATING PROGRAM

FAIRBRIDGE BLUFF/CITY OF YORK

Two new OSL dates, not associated with artefacts, were
obtained from Fairbridge Bluff. The oldest date of 140 ±
14 ka for sample R7/K0024 (Table 1, Figure 2A), produced
using small aliquot OSL methods, is from an aeolianite
sequence immediately overlying and broadly correlating
with the Rottnest Limestone member (Figure 4). A
second, younger age of 47 ± 4.5 ka (R6/K0023; Table 1),
also obtained using small aliquot OSL methods, is from
the palaeosol unit overlying the aeolianite.

Three other small aliquot OSL dates, not associated
with artefacts, were obtained from City of York and
include a palaeosol unit sandwiched between two
aeolianite sequences (Figure 4). The palaeosol dated at
49 ± 3.3 ka (R5/K0022) and underlying aeolianite giving
an equivalent age within uncertainties of 46 ± 2.9 ka
(R6/K0023) (Table 1, Figure 4). Together these provide
a weighted mean age of 48.1 ±3.1 ka. The overlying
aeolianite yielded an age of 36 ± 2.3 ka (R4/K0021).

Bathurst Point

The massive/breccia calcrete unit immediately underlying
a cemented fossiliferous chert artefact is dated by small
aliquot OSL to 42.0 ± 2.7 ka (R11/K0034), and gives a
maximum age for the artefact. The U-series sample from
the carbonate lens overlying and cementing the calcrete/
breccia unit yielded a weighted average age of 17.1 ±
1.2 ka (Table 1). A second block sample cut into the top
of rhyzolithic palaeosol unit is dated by small aliquot
OSL to 32.9 ± 3 ka (R12/K0035) (Figure 2B). This date is
discordant with both the U-series age estimate and the c.
20 ka inferred maximum age for the rhyzolithic palaeosol
unit near the top of the section 60 m to the west (Figure
2B). Here, the rhyzolithic palaeosol unit rests directly on
the underlying younger aeolianite dated by single aliquot

Figure 4. Tamala Limestone
stratigraphical succession at City
of York Bay (cf. Figure 1; photo by
EJR).

Table 1. Age estimates for City of York Bay (COY), Fairbridge Bluff (FB), Charlotte Point (CP), Bathurst Point (BP) and Little Armstrong Bay (LAB) from Single-grain (SG)
OSL estimates, single aliquot (SA) OSL age estimates, isotopic (U-series) age estimates (*2 sigma uncertainty) and radiocarbon dating ( 14 C). Water contents were in the range
of 3 - 10%. Burial depth calculated as time weighted mean depth, i.e. the time-weighted average distance between the sampling point and the surface, based on our best
estimate of the aggradation history at each site. Note, over-dispersion values for small aliquot (SA) OSL is expected to be significantly lower than for single grains (SG),
owing to signal averaging effects. De is the dose (in Gy) of laboratory b irradiation equivalent to the dose received in the field from all sources (a, b, g, cosmic). s d is the
population overdispersion, it represents the degree of spread in the data beyond that which can be explained by known sources of uncertainty (i.e. measurement uncertainty
on each individual single grain or single aliquot De). Non-zero s d values are almost universally found for single grain dose distributions. The greatest component of this is
traditionally attributed to partial bleaching (e.g. Olley et al. 2004) however there are other important contributors, most notably b-dose heterogeneity (Nathan et al. 2003) and
variations in instrument uncertainty (Jacobs et al. 2006; Pietsch 2009). Radiocarbon dates are reported at 95.4% probability and calibrated using the SHCall3 curve (Hogg et
al. 2013) in OxCal v.4.2.3 (Ramsey 2013).

Site

Lab code

Unit

Method

Depth

(m)

De (Gy)

s d

U (ppm)

Th (ppm)

K (%)

Dose rate
(mGya 1 )

Age

(ka)

COY

R4/K0021

Upper aeolianite

SA OSL

4.0

8.82±0.24

4.2

0.740±0.002

0.440±0.022

0.081±0.001

0.28±0.02

36.3±2.3

COY

R5/K0022

Palaeosol

SAOSL

5.5

28.07±1.13

0.560±0.003

1.840±0.092

0.463±0.002

0.66±0.03

49.1±3.3

COY

R6/K0023

Lower aeolianite

SA OSL

7.5

23.72±0.84

9.6

0.610±0.003

1.420±0.071

0.410±0.002

0.60±0.03

46.2±2.9

R8/K0025

Palaeosol

SAOSL

0.3

42.97±1.39

8.2

0.460±0.025

3.430±0.172

0.571±0.029

1.048±0.09

47.3±4.5

R7/K0024

Aaeolianite

SAOSL

1.0

65.31±2.39

9.9

0.520±0.026

1.330±0.067

0.219±0.011

0.506±0.05

140.0±14.0

GU8.5

Palaeosol

SG OSL

3.0

8.28±0.46

0.8±0.04

2.1±0.11

0.164±0.082

0.63±0.06

12.7±1.25

GU8.6

Palaeosol

SG OSL

3.5

8.72±0.16

1.0±0.05

1.8±0.09

0.199±0.099

0.68±0.06

12.1±1.01

Wk 37948

Palaeosol

AMS 14 C

2.0

10.3±0.06

BP (West)

R9/K0026

Aeolianite

SAOSL

3.0

6.31± 0.152

4.2

0.700 ± 0.001

0.450 ± 0.023

0.072 ± 0.001

0.294 ± 0.037

24.9 ± 3.2

BP (West)

R10/K0027

Aeolianite

SAOSL

4.5

6.2 ± 0.157

4.4

0.640 ± 0.001

0.380 ±0.019

0.115 ±0.001

0.324 ± 0.027

22.2 ±1.9

BP (East)

R12/K0035

Palaeosol

SAOSL

3.5

7.71 ± 0.256

7.7

0.320 ± 0.016

0.690 ± 0.035

0.058 ± 0.003

0.270 ± 0.027

32.9 ±3.0

BP (East)

R11/K0034

Palaeosol

SAOSL

4.0

11.36±0.271

2.9

0.420 ± 0.021

0.770 ± 0.039

0.066 ± 0.003

0.321 ± 0.037

42.0 ±2.7

BP (East)

BP2.2

Calcrete

U-series

3.9

17.1 ± 1.7*

LAB (East)

GU8.1

Aeolianite

SG OSL

24.0

4.02±0.07

1.1±0.06

0.5±0.03

0.069±0.003

0.39±0.04

10.3±1.04

LAB (East)

GU8.2

Aeolianite

SG OSL

24.0

3.49±0.12

0.8±0.04

0.5±0.03

0.050±0.003

0.30±0.03

11.6±1.27

LAB (East)

GU8.3

Palaeosol

SG OSL

26.5

7.58±0.19

1.1±0.06

2.0±0.10

0.211±0.011

0.60±0.05

12.6±1.10

LAB (East)

GU8.4

Palaeosol

SG OSL

27.5

8.07±0.33

0.9±0.05

1.0±0.05

0.178±0.009

0.48±0.04

16.8±1.59

LAB (East)

CURL-16668

Palaeosol

AMS 14 C

27.5

28.2±0.18

LAB (West)

R1/K0018

Upper aeolianite

SAOSL

4.0

5.02±0.159

8.3

0.730±0.001

0.570±0.029

0.050±0.050

0.267±0.04

22.2±3.3

LAB (West)

R2/K0019

Palaeosol

SAOSL

4.5

8.08±0.221

1.4

0.850±0.003

1.630±0.082

0.194±0.001

0.475±0.02

19.7±1.1

LAB (West)

R3/K0020

Lower aeolianite

SAOSL

6.5

9.51±0.337

11.6

0.640±0.001

0.550±0.028

0.133±0.001

0.308±0.01

35.9±2.1

Journal of the Royal Society of Western Australia, 99(1), 2016

I. Ward et ah: Dating of palaeosols on Rottnest

OSL to between 25 - 22 ka (Table 1). This age discrepancy
may be explained by the presence, within the upper
surface of the rhyzolithic palaeosol unit at Bathurst Point
east (i.e. artefact find site), of large clasts of what may be
older calcarenite. Hence the true age of the rhyzolithic
palaeosol unit is considered to be < 20 ka.

Little Armstrong Bay

At Little Armstrong Bay (LAB) east, two single-grain
OSL dates 12.6 ± 1.1 ka (GU8.3) and 16.8 ± 1.6 ka (GU8.4)
pertain to samples taken just above, and at the same
depth in this palaeosol, as the Eocene fossiliferous
chert artefact and the emu eggshell fragment (Table
1). Significantly older than these age estimates is the
calibrated radiocarbon date for the eggshell fragment at
28.2 ± 0.18 ka (CURL-16668). The discrepancy between it
and the younger OSL dates from the same position in this
palaeosol indicates that the eggshell fragment is probably
re-worked from an older deposit - or recrystallised.

Two single-grain OSL dates of 10.3 ± 1.0 ka (GU8.1)
and 11.6 ± 1.3 ka (GU8.2) were obtained from the
aeolianite immediately overlying the fossiliferous chert
artefact and emu eggshell-bearing palaeosol at LAB
east (Figure 2C). These age estimates are much younger,
but in chronological sequence, with the single-grain
OSL date of 16.8 ± 1.6 ka for the palaeosol itself at LAB
east and the single aliquot date of 19.7 ± 1.1 ka from the
same palaeosol at LAB west (Table 1). The single aliquot
OSL age estimate of 22.2 ± 3.3 ka, for the aeolianite
overlying the palaeosol at LAB west (Table 1), is out of
chronological sequence, indicating possible mixing of
older grains in this sample. The single aliquot OSL age
estimate of 35.9 ± 2.1 ka (R3/K0020) from the aeolianite
below the palaeosol at LAB west provides a likely age
for the corresponding aeolianite underlying the artefact¬
bearing palaeosol at LAB east (Figure 2C).

Charlotte Point

Two single-grain OSL dates (GU 8.5 and GU 8.6) from
the artefact-bearing palaeosol at Charlotte Point give an
absolute age of 12.7 - 12.1 ka (Table 1, Figure 2D). The
palaeosol unit comprises a strongly cemented, brown
(7.5 YR 5/3) medium-coarse grained quartz carbonate
sand grading into a lighter brown (7.5 YR 8/2) and
more loosely cemented quartz carbonate sand with
Austrosuccinea sp. shells (Figure 2D). Charcoal from a
pedogenically similar palaeosol 150 m south-east of
the chert artefact find location, has yielded a slightly
younger radiocarbon age of 10.3 ka (Wk 37948, Table 1).
This date is however fully acceptable; together with the
two OSL dates, it records the period through which the
original sediments were deposited and developed into
a soil.

DISCUSSION

DATED TAMALA LIMESTONE HISTORY FOR
ROTTNEST ISLAND

A summary of the dated aeolianite sequences on Rottnest
Is. is given in Figure 5. Eighteen previously unpublished
OSL dates (Table 1) augment the absolute chronology
for Rottnest Island's Tamala Limestone succession given

by Price et ah (2001); Playford et ah (2013) and Brooke
et ah (2014). The latter two papers note three major
phases of dune deposition in the Tamala Limestone of
Rottnest Island. A possible fourth or earlier phase of dune
deposition is dated by single aliquot OSL to 140 ± 14 ka
(MIS 6/5e), confirming Playford's (1997:725) suggestion
that 'part of the Tamala Limestone near Fairbridge Bluff
underlies the [Rottnest Limestone] reef, and must have
formed a little earlier perhaps at about 135 - 140 ka'.
Hence the major shoaling of Rottnest began during, or
just after 125,000 yr ago (see also Hearty 2003).

A second major phase of dune deposition dates from
around 77 ka (MIS 4) (Playford et ah 2013; Brooke et ah
2014). A third phase of dune deposition is dated to 49 - 36
ka (MIS 3) at Bathurst Point find site. City of York (Figure
4) and Little Armstrong Bay (Figure 2C). The final phase
of aeolianite deposition represented at Bathurst Point and
Fish Hook Bay broadly correlates with MIS 2 (27 - 23 ka),
approximately the initial stages of the LGM (Andrews
2013; Clark et ah 2009). Thus the main periods of
carbonate aeolianite deposition show a general younging
seaward trend, dating to MIS 5e (-140 ka), MIS 5a (-80
ka), MIS 3 (~ 46 - 36 ka) and MIS 2 (~ 25 ka) on Rottnest
Island and MIS 5e (-115 ka) and later on the mainland
(Figure 5). Hearty (2003) adds a further two aeolinite
stages on Rottnest Is. at MIS 5c (-100 ka) and MIS 1 (-15
ka). However, these latter dates are based on contestable
amino acid racemisation (AAR) ages of 'whole-rock'
biogenic samples, that likely date reworked calcareous
marine invertebrates rather than the depositional age of
the dunes (Dortch & Hesp 1994; Brooke et ah 2014). We
would concur. Alongside Brooke et ah (2014), our OSL
chronology indicate that coastal carbonate barriers and
dune fields form very rapidly and under a wide range of
climatic conditions and sea levels (c.f. Hearty 2003). Given
the high potential for reworking of this shelf and coastal
sediment prior to its final deposition and cementation,
the data also show that OSL, especially single-grain
OSL, is an effective and reliable method for dating these
deposits (see also Playford et ah 2013).

Dated palaeosol history for Rottnest Island

According to Semeniuk (1986), fossil soils and calcretes
in this region mark periods of local interruption in
dune building during more humid periods of the
Pleistocene, albeit with variable rates of lithification.
Like the aeolianite sequences, the sometimes very
thick (~ 1 - 2 m) palaeosol sequences on Rottnest Is. are
laterally discontinuous and, from the chronology, appear
to have formed in relative quick succession within the
aeolianites but under a wide range of climatic conditions.
The latter may well result in quite different palaeosol
characteristics, as implied by the very preliminary
analyses undertaken in this study.

A summary of the dated palaeosol sequences on
Rottnest Is. is given in Figure 5, which indicate palaeosol
units corresponding to MIS 3 (~ 49 - 33 ka) and MIS 2 (~
17-10 ka) on Rottnest Island. The earliest of these date
to 49 ± 3.3 ka and 47.3 ± 4.5 ka (MIS 3) at City of York Bay
and Fairbridge Bluff respectively. Alongside the palaeosol
units at Little Armstrong Bay and Charlotte Point, these
two palaeosol units also contain significantly higher
proportions of carbonate material than the older (MIS
5e/5d), almost pure quartz terra rossa palaeosol recorded

Journal of the Royal Society of Western Australia, 99(1), 2016

Point Reran
7& ± 5 ka

Ins&t key;

Bassendean Sand

■ Tamala Limestone

Pol&nlial submarine ■anient of Tamala Limestone
Epproximaled by the -70 m AHD bathymplric cgnlgyr

Garden Is

2 km

12 ka

I0±1.0ka

13 + 1.1 ka
17 ± 16 ka

22 ± 3 3 ka
36 ±2.1 ka

36 ± 2.3 ka

49 1 3.3 ka

46 + 2.9 ka

Little
Armslrong

City of York

Bay

Thompson
Bay

17 +16 ft*

22 ± 1.9 ka
27 ± 4.5 Ka 1
42±2.7 ka
77 + 12 ka 1.

{Fish Hook
Bay

Figure 5. Summary of TL/OSL age estimates for aeolianite and palaeosol (in bold italics) sequences on Rottnest Is. and the
adjacent mainland (inset figure), including previously published dates of 1. Brooke et al. (2014) and 2. Price et al. (2001).
Inset figure also shows potential offshore extent of Tamala Limestone (after Smith et al. 2011).

at Fairbridge Bluff (Figure 2A), and also at Minim Cove
and Cottesloe (Brooke et al. 2014). These contrasting
carbonate-rich and quartz-rich palaeosols likely reflect
the variable movement of mainland (westward) and
exposed shelf (eastward) sediments across the shelf in
association with changing sea level (Hearty & O'Leary
2008, 2010).

Other palaeosols, such as those at Charlotte Bay and
Fish Hook Bay (Dortch and Hesp 1994), also contain
variable amounts of charcoal (and fossil plant material)

but this does not appear to be age-related. Likewise,
although the palaeosols at the City of York Bay and
Little Armstrong Bay and pedogenetically very similar,
and their stratigraphic positions and heights above sea
level are also much the same (Dotch and Hesp 1994:
26), nevertheless they yield different depositional ages
(Figure 5). In contrast, the exposed palaeosol units on
the east and west side of Little Armstrong Bay do show
a similar age and pedogenic characteristics and hence
likely belong to a single depositional unit. The palaeosol

I. Ward et ah: Dating of palaeosols on Rottnest

horizon at Charlotte Point also share similar sediment
characteristics (albeit with inclusion of charcoal) and
age to the Little Armstrong Bay. However, as indicated
by Dotch and Hesp (1994), the palaeosols often cannot
be traced laterally very far and caution should be taken
before assuming any correlation between these.

The artefact-bearing palaeosols mainly date to the late
(17-10 ka) stages of MIS 2 and possibly to MIS 3 if the
oldest date of 42 ka for Bathurst Point is assumed. This
brecciated calcrete/palaeosol unit forms partly as a result
of mechanical fracturing by roots (see Arakel 1982), which
are clearly evident in the overlying rhyzolithic palaeosol
dated to around 17 ka (Figure 2B). Dissolution and re¬
cementation are common in such calcretes, as evident
from the thin calcrete unit that caps the brecciated
calcrete (Figure 3) and also in thin section (Figure 2B).
Subsequent erosion of the rhyzolithic palaeosol, possibly
as sea-level encroached Rottnest Is., may have resulted in
exposure of the embedded artefact (see also Ward et al.
2016). Indurated palaeosols and duricrusts intercalated
within Tamala Limestone aeolianite successions occur
along many parts of the West Australian coastline
(Playford et al. 2013, p.102) hence the possibility of other
embedded artefacts being similarly exposed.

Palaeosols, occasionally featuring rhizotubules and
Leptopious sp. pupal cases are revealed in many places
along the littoral zone. At Little Armstrong Bay a massive,
brown sandy palaeosol unit continues c. 200 m within
the intertidal zone. Playford (1997) also describes root
pipes in the Tamala Limestone extending below sea level
at many localities around the coast (e.g. at Fairbridge
Bluff). The luminescence dating of the 'older aeolianite'
at Bathurst Point (Figure 2B) implies that some, perhaps
even all of the palaeosol units exposed in the littoral
are older than c. 77 - 66 ka (MIS 4). Correspondingly
Smith et al. (2011) indicate the Tamala Limestone (and by
implication intercalated palaeosol units) may extend as
far as the -70 m bathymetric contour (Figure 5). However,
no artefacts have been identified in any of these older or
partially submerged palaeosol units exposed on Rottnest
Island.

CONCLUSION

With an absolute chronology spanning MIS 6 to MIS 2 (~
140 ka - 10 ka), Rottnest Island is today one of Australia's
best dated Late Quaternary localities and one of Western
Australia's most iconic geoheritage sites (Ward 2013). This
secure chronology, particularly of the palaeosol units, is
of great importance to further investigation of prehistoric
archaeological sites as well as to other Quaternary field
studies on the island. The artefact-bearing palaesol units
mainly date to MIS 2 (17 - 10 ka). However, further
dating and pedogenic characterisation of palaesol units
around the island is needed before considering the
possibility of a widespread distribution of one or more
archaeologically significant palaeosol units (see Dortch
and Hesp 1994). Similarly more detailed chronological
studies - such as this one - are needed to further define
dune-building and calcrete-forming episodes in the
Tamala Limestone (Playford 1997). With continued
evaluation of the Rottnest Island absolute chronology,
the island will become an increasingly valuable site for
Quaternary investigations.

ACKNOWLEDGEMENTS

I.W. and C.E.D. sincerely thank Perth Nyungar
community elders who endorsed the archaeological
investigations that resulted in the chronology reported
here. They also thank staff of Cultural Heritage
Services, Rottnest Island Authority (RIA) for the
issuance of numerous ferry travel vouchers, and for
contributing significantly to the project in many other
ways. Many thanks to Phil Playford OA, who advised
on Tamala Limestone geology and kindly arranged
with WA Geological Survey for the sponsoring of
charcoal 14C date (Wk 37948) reported here. The
authors also thank the reviewers, Karl-Heinz Wyrwoll
and David Haig. Lastly, thanks to Joe Dortch (UWA
Archaeology) for his welcome comments on early
versions of the text.

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Journal of the Royal Society of Western Australia, 99(1): 27-30, 2016

Male sterility in Corymbia calophylla (marri)

JEN MCCOMB

School of Veterinary and Life Sciences, Murdoch University, Perth, Western Australia 6150
; H jmccomb@murdoch.edu.au

Most Corymbia calophylla trees have inflorescences of hermaphrodite flowers with a variable
number of male flowers. Some trees have been identified that have either only female flowers, or a
seasonally variable production of female flowers at the beginning of the flowering season, changing
to production of hermaphrodite flowers after the first 2-3 weeks of flowering. In the flowers of the
female trees, the anthers develop normally until the stage of meiosis when premature disintegration
of the tapetum is followed by degeneration of the pollen mother cells. Female trees appear to be rare
but widespread.

KEYWORDS: Female flower; Pollen sterility.

INTRODUCTION

Early illustrations of buds and flowers of marri
(Eucalyptus calophylla (Lindl.) KD Hill & LAS Johnson)
such as those by Baker and Smith (1920, Fig. 2 in Carr
et al. 1971) showed flower buds of two shapes - some
pear- shaped and others more spherical or top-shaped.
This remained unremarked by botanists until Carr
et al. (1971) reported that the pear-shaped buds were
hermaphrodite, while the spherical ones were male, i.e.
female sterile. Earlier observers may have thought the
degenerated ovary contents in spherical buds were a
result of fungal or insect damage, but this is not the case
with most corymbs having some male flowers amongst
the hermaphrodite ones. The proportion of male flowers
varies and some trees have only male flowers (Carr et al.
1971). The variety maideniana Hochr. from the Darling
Range, was shown to be based on a type specimen with
only male flowers (Carr & Carr 1972).

A further variation of floral structure is reported here.
During collection of pollen for hand pollinations, some
marri trees were identified that had only female flowers.

METHODS

An attempt to collect pollen from newly opened buds
of marri in the Kalamunda hills and foothills resulted in
the identification of a number of trees that produced no
pollen. Anther development was examined to determine
the cause of the male sterility. Hermaphrodite and male
sterile buds at different stages of development were
collected (Table 1), fixed in formalin acetic alcohol,
processed for sectioning in paraffin wax, cut at 8 pm and
stained in 0.5% toluidine blue . Buds were fixed over the
period 14th December 2013 to 4 th April 2014.

Observations were made over several years to
determine the consistency of expression of male sterility.

Herbarium specimens were examined at the Perth
herbarium and for those without pollen obvious on

the anthers, anthers of flowers or unopened buds were
crushed in aceto-orcein and examined for the presence of
pollen.

RESULTS

In a number of Kalamunda trees initially only female
flowers (i.e. male sterile flowers), were present but after
one to two weeks the trees produced hermaphrodite
ones. The initial occurrence of female flowers varied with
season. Four trees were identified that produced only
female flowers over the entire flowering season, at least
to the height of 5 m. This was consistently the case for
two of the trees that were observed over the flowering
periods of 2013 to 2016 which included years of abundant
flowering as well as sparse years. A third female tree did
not flower in 2015, and the fourth tree was burned in 2015
and did not flower in 2016. The female trees set the usual
crop of capsules and seeds. Trees with female flowers also
had some female sterile flowers in their corymbs making
these flowers totally sterile.

The female flowers on living trees and herbarium
specimens were generally smaller than the
hermaphrodite ones (Figure 1) and this size difference
is the best indication that a tree may be female. Anthers
of female flowers after anthesis, in living specimens and
herbarium specimens are pale and small, but otherwise
similar to those of male or hermaphrodite flowers from
which the pollen had been shed and collected by insects.
Thus to identify a female flower it is necessary to examine
the anthers of mature unopened buds. It is possible that
the herbarium collections referred to as 'female' could be
from trees that later in the season produce hermaphrodite
flowers, but the flower size would suggest they are from
female trees.

The anthers from hermaphrodite buds had darkly
stained premeiotic pollen mother cells with a well
defined tapetum. During meiosis (Figure 2A) and
through to the stage of tetrads (Figure 2B) and
uninucleate pollen, the tapetum was still in place
although becoming vacuolated. The mature anthers
showed the normal development of the fibrous layer and

Journal of the Royal Society of Western Australia, 99(1), 2016

Figure 1. Corymbia calophylla trees
from Kalamunda. A. Female flower,
B. Hermaphrodite flower. C. Left
female. Right Hermaphrodite
flowers.

Table 1. Corymbia calophylla trees: locations and flower fertility

Location/herbarium number

Sex of flowers

Histology

Remnant tree on road verge at No 20 James Rd. Kalamunda.

Hermaphrodite and male, 2013-16

Sections of hermaphrodite
flowers

Remnant tree on road verge of 29 Betti Rd (between James
and Grace corners) Kalamunda

Hermaphrodite and male 2013-2016.
Initially female in first weeks of
flowering, 2013, 2016.

Sections of hermaphrodite
flowers

Remnant tree on road verge SW corner of Robbins Rd
and Grace Rd. Kalamunda.

Female, 2013-14,16

Sections of female flowers

Remnant tree on road verge at No 5 Betti Rd. Kalamunda.
(PERTH 08733619)

Female 2013-16

Sections of female flowers

Remnant tree on road verge at No 18 James Rd Kalamunda
(PERTH 08733627)

Female 2015,16.

Tree in a remnant patch of trees at corner of Crystal Brook
Road and Welshpool Road, Wattle Grove

Female 2015.

Herbarium specimen Between Dunsborough and Cape
Naturaliste (PERTH 01319256)

Female 1982

Herbarium specimen Marangup Reserve off Toodyay Rd
(PERTH 08029164)

Female 2007

tricolpate pollen with dark stained deposits at each pore
(Figure 2C).

In the anthers of the two female, (male sterile) trees
sectioned, there were no major differences from male
fertile anthers until the onset of meiosis. At meiosis in
the female trees, the pollen mother cells rounded up
and the tapetal cells were very faintly stained and more
vacuolated than in the hermaphrodite trees. The most

advanced meiotic stages were pachytene or diplotene
(Figure 2D). Degeneration of the sporogenous tissues
followed quickly (Figure 2E) with pollen mother cells
becoming vacuolated then cell contents of each locule
degenerating into a small deeply stained area. This
reduced the size of the anther and distorted its shape,
but the subepidermal fibrous layer developed normally
with the empty locules splitting open at anther maturity

J. McComb: Male sterility in the marri

Figure 2. Corymbia calophylla anther development. Trees from Kalamunda A-C. Fertile anthers from hermaphrodite
flowers. A, anthers at early prophase of meiosis, B, tetrad stage, C. Pollen in mature anthers. D-F. Sterile anthers from
female flowers. D. anthers at early prophase, note vacuolate tapetum, E. degeneration of pollen mother cells. F. anthers
with empty locules but mature walls with fibrous layer developed, t, tapetum, mpmc meiotic pollen mother cells, te
tetrads, dpmc degenerating pollen mother cells, f fibrous layer, o oil cells, p pollen.

(Figure 2F). The development of the oil cells in the
filament was similar to the fertile anthers.

The female trees set crops of capsules comparable to
those on nearby hermaphrodite trees.

The two herbarium specimens observed to have only
female flowers had short pale anthers and although there

were a few small pollen grains on the surface of open
flowers of the Dunsborough specimen, an unopened
bud showed no pollen in the anthers and this tree is
considered a putative female (Figure 3). Similarly no
pollen was present in anthers from unopened flowers of
the Toodyay specimen.

Journal of the Royal Society of Western Australia, 99(1), 2016

DISCUSSION

Flowers of marri are usually hermaphrodite or male,
but female flowers and completely sterile flowers also
occur. The production of female flowers may be transient
at the beginning of the flowering season or a tree may
produce only female flowers. In the female flowers the
pollen mother cells and tapetum degenerate at the onset
of meiosis, but the anther wall matures as in the fertile
anthers. Tapetal abnormalities are frequently related
to male sterility though mostly reported in herbaceous
plants (Kaul 2012).

If the first flowers that open on a tree are female they
must be outcrossed to set fruit, while in female trees all
flowers will be outcrossed. Hermaphrodite flowers may
be geitonogamously pollinated or outcrossed. Eucalypts,
as with other forest trees, show inbreeding depression
associated with self pollination (Sedgley and Griffin
1989).

The mix of male flowers, or sterile flowers in the
corymb probably adds to its attraction for insects without
being a sink for resources following seed set. The stigma
of the hermaphrodite flowers does not become receptive
until 7-9 days after anthesis (J. McComb data not shown)
and this long period means that each corymb has a long
lasting display of flowers.

Pryor (1976) reported that some trees of E. pulverulenta
Sims had pollen with no protoplasmic content, and a
detailed study by Peters et al. (1990) showed a wide
range of pollen sterility in both a natural and cultivated
population, with number of trees in the cultivated stand
having no fertile pollen. Pryor (1976) also reported an
unusual type of male sterility from E. grandis. In this case
the pollen grain wall released no protein on to a stigma or
a sucrose or agar gel and the grains failed to germinate.

While male flowers have previously been reported
from C. calophylla and closely related species, as well as
from other genera of Myrtaceae (Bentham, 1869), there
are few reports of female flowers. Melaleuca cornucopia
Byrnes is recorded as having female flowers and can be
monoecious or gynoecious (Byrnes 1985).

REFERENCES

Barker R T & Smith H G 1920. Research on the Eucalypts, especially
in regard to their essential oils. 2 nd .ed. Sydney.

Bentham G 1869 Notes on Myrtaceae. Botanical Journal of the
Linnaean Society 10,101-66.

Byrnes N T 1985. A revision of Melaleuca L. (Myrtaceae) in
northern and eastern Australia. Baileyana 2,131-146.

Carr SGM & Carr D J 1972. Eucalyptus calophylla var. Maideniana
Hochr.: A male tree. Proceedings of the Royal Society of Victoria
85, 49-51.

Carr SGM, Carr D J & Ross F L 1971. Male flowers in eucalypts.
Australian Journal of Botany 19, 73-83.

Kaul M L H 2012 Male Sterility in Higher Plants. Springer Verlag,
Berlin.

Peters G B, Lonie J S & Moran G F 1990. The breeding system,
genetic diversity and pollen sterility in Eucalyptus pulverenta,
a rare species with small disjunct populations. Australian
Journal of Botany 38, 559-70.

Pyror L D 1976 Biology of Eucalypts. Edward Arnold, London.

Sedgley M & Griffin A R 1989. Sexual Reproduction of Tree Crops.
Academic Press, London.

ACKNOWLEDGEMENTS

I thank Patsy Stasikowski for helping collect buds in 2013

and Gordon Thompson for assistance with the histology.

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