The Status of the Greater Glider "Petauroides Volans" in the Illawarra Region Kevin S

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2007 The status of the greater glider "Petauroides volans" in the Illawarra region Kevin S. Maloney University of Wollongong

Recommended Citation Maloney, Kevin S, The status of the Greater glider "Petauroides volans" in the Illawarra region, MSc-Res thesis, School of Biological Sciences, University of Wollongong, 2007. http://ro.uow.edu.au/theses/59

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The Status of the Greater Glider Petauroides volans in the Illawarra Region

A thesis submitted in partial fulfillment of the requirements for the award of the degree

Masters of Science-Research

from

University of Wollongong

by

Kevin Shane Maloney BenvSc (Hons) Grad. Dip. Ed

School of Biological Sciences

2007

i

CERTIFICATION

I, Kevin S. Maloney, declare that this thesis, submitted in partial fulfillment of the requirements of the award of Masters of Science-Research, in the School of Biological

Sciences, University of Wollongong, is wholly my own work unless otherwise referenced or acknowledged. The document has not been submitted for qualifications at any other academic institution.

Kevin S. Maloney

18 November 2007

ii Abstract

Anecdotal evidence suggested that the Greater glider Petauroides volans Kerr 1792 had been eliminated from Royal National Park by wildfires in 1994. This thesis is concerned with the distribution of the Greater glider in the Illawarra region, the reasons why it appears unable to recolonise an area in which it was formerly found, and the potential for re-establishing the former population. The specific aims of this study were to (1) clarify the taxonomy of this species; (2) review the distribution and abundance of

Greater gliders in the Illawarra area and the current threats to populations; (3) conduct a detailed field study in the region; and (4) develop a translocation proposal for reintroduction of Greater gliders to Royal National Park.

Early accounts of the Greater glider Petauroides volans (Marsupialia:

Pseudocheiridae), started with Arthur Phillips’ 1789 account in The Voyage of

Governor Phillip to Botany Bay, but, since then, the species has had a quite peripatetic and confusing taxonomic history. It has been listed as a member of 10 genera, with about 23 different binomial names. The taxonomy and early descriptions of the species’ morphology, dentition, behaviour, distribution and abundance are complex, and P. volans has frequently been confused with a number of other gliding possums, particularly the Yellow-bellied glider, Petaurus australis. Early descriptions of the morphology of P. volans were given only in broad general terms. More value can be placed on the early behavioural observations, and on the earliest records of their occurrence.

i Recent distribution records of the Greater glider in the study area show that its range and numbers have declined over a 35 year period. Many factors may have contributed to this decline including; removal of habitat and den trees, predators, and the timing and frequency of fire events. The population of Greater gliders that was present at Royal

National Park prior to 1994 was depleted by the shooting of 21 individuals between

1978 to 1980. A number of barriers in the landscape will limit the ability of the Greater glider to disperse from adjacent areas back to Royal National Park.

My detailed field study, using spotlighting, at 19 sites confirmed that it is no longer present in many areas in which it was once observed. It is present in areas that are conserved as part of Sydney catchment Authority, and is indeed absent from Royal

National Park. This information suggests that a reintroduction of the Greater glider to

Royal National Park would be worthwhile, particularly as the Greater glider was formerly abundant at Royal National Park.

The biological and ecological factors required by the New South Wales National Parks and Wildlife Service policy document for the translocation of fauna are considered for the translocation proposal of the Greater glider to Royal National Park. These factors include; the reintroduction of 18 individuals using a sex ratio of 1:2 (6 males and 12 females), at two sites (Jersey Springs and Bola Creek) with three lots of F-M-F at each site (3 males and 6 females). Monitoring of the translocated individuals would use radio collars, reflective tags and spotlighting to determine initial success of the reintroduction.

The low population numbers of the Greater glider in the Illawarra region call for sourcing the individuals from other areas. The taxonomy and the current and former range of the Greater glider are reviewed.

ii

The Greater glider lives for 10 – 15 years with adult females having one young per year, with greater success in forests with higher nutrient levels. The species is solitary and virtually silent, with populations ranging from 0.01 to 5 individuals per hectare. It is a hollow dependent, nocturnal folivorous marsupial which feeds high in the canopy and consumes some 33 eucalypt and ten non eucalypt species across its distribution. The home range size for the Greater glider is 1.4 to 2.6 ha for males and 0.8 to 2.5 ha for females. Threats to this species include habitat clearing, logging, fire and predation with ten predators reported. The only disease reported is Chlamydia which appears not to have any effect, while three ectoparasites and eight endoparasites were also recorded for this species. There have been no previous translocation programs undertaken for the

Greater glider; other analogous species which have been translocated include the Sugar glider, Koala and Common Brushtail possum.

The source population for the translocation of the Greater glider to Royal National Park, should come from an area other than the Illawarra as these populations are in low numbers and are of local conservation priority.

iii Preface

The work presented in this thesis is predominantly my own work; publications and contributions from other people are detailed below:

The work presented in chapter two was undertaken in collaboration with Jamie M.

Harris and resulted in the following manuscript:

MALONEY, K.S. AND HARRIS, J.M. (2008). Early natural history of the greater glider Petauroides volans (Kerr, 1792). Proceedings of the Linnean Society of New

South Wales 129, 39-55.

iv Acknowledgements

There are many people to thank when undertaking a research thesis.

Many thanks to Marieann, Declan and Ruby (the three amigos) who have been helpful and understanding in many ways, such as listening to the rants and raves of a fanatic and also putting up with the mess around the house.

The NSW NPWS personnel who helped with data provision; Peter Ewin and Kylie

Madden. Field work; Helen Jessup. Advice; Debbie Andrew and Geoff Ross.

The staff of the Sydney Catchment Authority for access to Authority lands

Thanks to the following for help with field work (DD’s provided); Matt Clancy, Declan

Maloney, Jamie Harris, Michelle Haynes and Damon Maloney (the Demon).

Big thanks to the library staff at the Australian Museum (AM) Sydney; Natural History

Museum, London; and the University of Wollongong. For translating French articles, I am grateful to Betty Hassen, and for translating German, Peter Simmons. I also thank

Sandy Ingleby for access to the mammal collection in the AM and Cécile Callou for correspondence and photograph of a specimen from the MNHN.

Finally, I owe a debt of gratitude to Rob Whelan as supervisor for helpful discussion, encouragement and comments on this work.

v Table of Contents

Abstract i

Preface iv

Acknowledgements v

Table of Contents vi

List of Tables xi

List of Figures xii

Chapter 1 General Introduction 1

1.1 The landscape context for habitat 1

1.2 Habitat fragmentation components 2

1.3 Responses of mammals within fragmented landscapes 3

1.4 Declining mammalian fauna 6

1.5 Arboreal mammals 7

1.6 Status of Australian arboreal marsupials 7

1.7 Greater glider 12

1.8 Aims 18

Chapter 2 Historical accounts and discovery of the greater glider 20

2.1 Introduction 20

2.2 Taxonomy and nomenclature 20

2.3 Morphology 30

2.4 Dentition 33

vi 2.5 Habitat and diet 35

2.6 Distribution and abundance 37

2.7 Behaviour 39

2.8 Conclusion 42

Chapter 3 Review of the past knowledge of distribution and abundance of the greater glider in the study area, and threats to populations 45

3.1 Introduction 45

3.2 Recent records (1908-1980) 45

3.2.1 Records of Norm Robinson 45

3.2.1.1 Summary 48

3.2.2 Specimens held in institutions 48

3.2.2.1 Summary 50

3.3 More recent records (1980-2002) 50

3.3.1 Illawarra escarpment survey 51

3.3.1.1 Summary 53

3.3.2 Records from the NSW NPWS data base 53

3.3.2.1 Summary 53

3.4 Threats to populations of the greater glider 54

3.4.1 Introduction 54

3.4.2 Removal of den trees 55

3.4.3 Predators 55

3.4.3.1 Powerful owl 56

3.4.3.2 Sooty owl 57

vii 3.4.3.3. Spotted-tailed quoll 57

3.4.3.4 Foxes 58

3.4.3.5 Dog, Cat and Dingo 58

3.4.3.6 Lace monitor and Carpet python 58

3.4.3.7 Wedge-tailed eagle 59

3.4.3.8 Summary 59

3.4.4 Fire 59

3.4.5 Habitat removal 60

3,5 Barriers to recolonising Royal National Park 61

Chapter 4 Detailed field study 63

4.1 Introduction 63

4.2 Aims 63

4.3 Methods 64

4.3.1 Site selection 64

4.3.2 Detection of greater gliders 66

4.4 Results 67

4.4.1 Greater glider absence 67

4.4.2 Greater glider presence 67

4.5 Conclusion 68

Chapter 5 Translocation 70

5.1 Introduction 70

viii 5.2 Host site population parameters 71

5.2.1 Sex ratio 71

5.2.2 Number of Individuals 71

5.2.3 Spacing 74

5.2.4 Monitoring 74

5.2.5 Source population 75

5.3 Taxonomy of the greater glider 75

5.4 Distribution of the greater glider in NSW and across its range ,

including current and historical range 77

5.5 The relevant ecological requirements of the greater glider 80

5.5.1 Life cycle 80

5.5.2 Reproductive biology 80

5.5.3 Population dynamics 81

5.5.4 Social behaviour 82

5.5.5. Group composition 82

5.5.6 Home range size 83

5.5.7 Shelter and food requirements 84

5.5.7.1 Shelter 84

5.5.7.2 Food requirements 84

5.5.8 Foraging behaviour 85

5.5.9 Predators 87

5.5.10 Diseases 88

5.6 Known and potential threats 88

5.7 The success of previous translocation programs of the same or

analogous species 90

ix 5.8 Conclusion 91

References cited 93

x List of Tables

Table 1. The possum and glider species listed in Australian state and

federal conservation legislation. 8

Table 2.1. Dental formulas provided in the early natural history literature for

Petauroides volans. 35

Table 2.2. New synonymy based on the current review 44

Table 3.1. Records of the Greater glider from Norm Robinson 46

Table 3.2. Specimens of Petauroides volans held in the Australian Museum

(AM) and the CSIRO from the study area. 49

Table 3.3. The ten predators of the Greater glider obtained from a literature

review. 56

Table 4.1. Sites used in this study for spotlight presence or absence of the

Greater glider. 64

Table 4.2. The presence or absence of the greater glider from the

chosen 19 sites for this study. 68

Table 5.1. Taxonomy of the Greater glider. 76

Table. 5.2. The results of six studies on the home range size (estimates)

of the Greater glider. 83

Table 5.3. Reported food tree species of the Greater glider across its range. 86

Table 5.4. The ten predators of the Greater glider obtained from literature

reviews. 87

Table 5.5. The ectoparasites and endoparasites collected from the Greater glider.

88

xi List of Figures

Figure 1. The distribution of the Greater glider (Petauroides volans) in

Australia. 13

Figure 2.1: Black flying opossum (=Petauroides volans) drawn

by P. Mazell and published in Phillip (1789). 21

Figure 2.2: Long-tailed opossum Didelphis macroura

(=Petauroides volans) from Zoology of New Holland

(1794) by George Shaw 23

Figure 2.3: A print from Waterhouse (1841) that is clearly Petauroides

volans because of the length of the tail and the hairy ears 27

Figure 2.4: Petaurista taguanoides from Gould (1863) (= P. volans). 32

Figure 3.1. The recent records of the Greater glider from the records

of Norm Robinson (1977, 1985, 1988) in the study area.. 47

Figure 3.2. Records and habitat model of the Greater glider provided by

the NPWS (2002) for the Commission of Enquiry into the

Illawarra escarpment. 52

Figure 3.3. Distributional records of the Greater glider in the Illawarra. 54

Figure 3.4. Satellite image of the northern half of study area. 62

Figure 4.1. Location of the 19 sites surveyed in the field study. 65

Figure 5.1. Proposed translocation sites for the Greater glider 73

Figure 5.2. Distribution of the Greater glider in Australia. 77

Figure 5.3. Distributional records of the Greater glider in NSW. 78

Figure 5.4. Distribution and predictive habitat model of the Greater

glider in Sydney bioregion. 79

xii

Chapter 1

General Introduction

1.1 The landscape context for habitat

The distribution of natural vegetation is dependent on the geology, soil patterns and climate within a landscape (Benson et al. 1996). Landscapes consist of many habitat types at a regional scale, which are nested within the much larger continental landscape.

The composition and arrangement of the surrounding landscape can have a strong effect on the presence, abundance and activities of local populations. Most organisms move, eat, reproduce, live and die in spatially heterogeneous landscapes; their habitat.

Habitat has been defined as being specific to a particular species and is the sum of resources needed by the species (Franklin et al. 2002). The greatest alterations to ecosystems have occurred in landscapes where natural habitats have been completely removed and replaced by different land-uses (Dobson et al. 1997, van der Ree and

McCarthy 2005). The removal of native vegetation in Australia has continued since the establishment of the penal colony in Sydney in 1788 (Benson 1999). As a result, the indigenous mammals in Australia have declined in abundance and many have become extinct (Strahan, 1995). One of the earliest mentions of the removal of native vegetation and the effect on the fauna was by Partington (1837), who stated…

“The description of characters that have been sent from Britain to New

Holland, to labour in the forests as a sort of penalty for crimes committed

in the mother country, are exactly such as we might expect to destroy

1 every living creature that comes in their way, without paying the least

attention to its manners. In consequence of this, the native mammalia

have been exterminated from all the cleared lands of the colonies, and as

it does not appear that any of them, with the exception perhaps of the

great kangaroo, have the slightest tendency to be social, there is little

chance of their again returning, even when ornamental plantings shall

have in part covered and sheltered those grounds which have been so

indiscriminately denuded by the desolating zeal of the axe”.

Since 1788, the removal of native vegetation and consequent changes in habitat type and quality, have been caused by anthropogenic disturbances in the landscape, these disturbances include:- agricultural practices, stock grazing, housing developments, electricity transmission lines, roads, forestry (including the cutting down of forests and introduction of plantations) and the introduction of herbivores (see Keith and Pellow

2005) and predators. Natural disturbances such as fire, flood and drought, have been causing habitat change in Australian landscapes for a greater period (Benson 1999).

1.2 Habitat fragmentation components

Dodson (1998) argued that natural communities can be viewed as “islands in a sea of development”, illustrating the process of habitat fragmentation (Fahrig 2003,

Lindenmayer and Fischer 2006). Fragmentation of habitat can be considered to comprise both the reduction in the total amount of a habitat type available in a landscape, and the apportionment of this remaining habitat into smaller, more isolated patches (Ewers and Didham, 2006).

2 The size, shape and spatial arrangement of habitats affect the extinction rate, population size and dispersal patterns of individuals among local and regional populations (Fahrig and Merriam 1994, Gibbs 2001, Swihart et al. 2006).

Three components of habitat fragmentation that contribute to an increased likelihood of population extinction include:-

(1) The overall loss of habitat in a landscape - which creates discontinuity in the spatial

distribution of resources and conditions present in an area, affecting the occupancy,

reproduction and survival of a particular species (Franklin et al. 2002).

(2) An increase in separation between remaining patches – resulting in the decline and

viability of a population, as there is not sufficient continuity between patches with

suitable habitat (Hanski 2005).

(3) A decrease in area per patch of habitat – which tends to sustain smaller populations,

causing susceptibility to genetic drift (Byers et al. 2005), inbreeding and stochastic

demographic challenges (Burgman et al. 1993). The maintenance of genetic

variation is considered essential for the long-term survival of a species, since

genetic diversity provides the raw material for evolution (Frankel and Soulé 1981,

Gibbs 2001).

1.3 Responses of Mammals within Fragmented landscapes.

Many mammals require a range of resources, which are naturally patchily distributed across a landscape, so they need to move around between resource sites. Habitat fragmentation can limit the mobility of species through the matrix (Laurance 1997,

3 Selonen and Hanski 2003). For example, roads both remove individuals from populations through collisions with vehicles (Taylor and Goldingay 2004, Ramp et al.

2006), and also form barriers to movement for resources and mates (Richardson et al.

1997). Culverts and exclusion fencing have been used to mitigate road mortalities and aid animal movement (Taylor and Goldingay 2003, McDonald and Cassady St Clair

2004).

The fragmented areas are also surrounded by a modified landscape ‘the matrix’ (Ewers and Didham 2006). The matrix can be hostile or suitable to a species, depending on its particular requirements and characteristics (Saunders et al. 1991) and the biology of the species. Matrix tolerance has been suggested as the key predictor of vulnerability for mammals in fragmented rainforest habitats in tropical Queensland (Qld) (Laurance

1990, 1997). Species that use or exploit the matrix remained stable or increased in fragments, while those that avoided the modified habitat declined or disappeared

(Laurance 1997). Species that avoid the matrix are unable to disperse between fragments, making recolonisation of fragments following a local extinction event quite rare (Laurance 1990, 1997; Fahrig 2003, Ewers and Didham 2006).

The home-range of a species can be affected by habitat fragmentation. Species that have a high home-range affinity, show an increase in mortality rates after habitat removal

(Tynedale Biscoe and Smith 1969a, Newell 1999). For example, Lumholtzs’ tree- kangaroos (Dendrolagus lumholtzi) were found to be using marginal home-ranges in woody debris left behind after habitat removal, and subsequently died a short time later from stress and predation (Newell 1999). Wilson et al. (2007) reported that the loss of canopy connectivity through the maintenance of power line corridors and roads had a

4 negative impact on the movement and home-range of the Lemuroid ringtail possum

(Hemibelideus lemuroides).

A human-altered disturbance regime, such as urban development and expansion, influences the species-level response to habitat fragmentation (Koprowski 2005, Ewers and Didham, 2006). Tigas et al. (2002) reported that bobcats (Felis rufus) and coyotes

(Canis latrans), which persisted in an urban environment, showed behavioural adjustments to habitat fragmentation and human activities. Both species reduced their daytime activity and showed avoidance of developed areas during times of high human activity (Tigas et al. 2002). Lopes and Ferrari (2000) found that the abundance and biomass of game species primates in eastern Amazonia had decreased with an increase in hunting pressure, and that the non-game species had increased in abundance with increasing forest disturbance (see also Lehman et al. 2006).

The change in size of a habitat can alter vegetation structure, with the edge of the remaining habitat being exposed to greater physical changes (Saunders et al. 1991,

Laurance 1997). These edge effects can influence the feeding and time budgets of species (Umapathy and Kumar 2000), with more fruiting trees available (Lopes and

Ferrari 2000, Lehman et al. 2006), resulting in a higher mortality rate (Ewers and

Didham 2006) through easier detection by predators (May and Norton 1996, Newell

1999).

Climatic changes over time can lead to altered distribution patterns of flora and the dependent fauna. Oshida et al. (2005) found differences in the genetic structure of

Russian flying squirrels (Pteromys volans) that indicated isolation (or fragmentation

5 into refugia) during the Pleistocene. The Mountain Pygmy-possum (Burramys parvus) is the only marsupial restricted to alpine regions in Australia (Kerle 1994; Mansergh

1994), and the fossil record indicates that its distribution was much wider during the

Pleistocene (Broom 1895; Wakefield 1960).

Individual species have varied responses to habitat fragmentation, from resilience and survival, to being sensitive and therefore unable to persist in the fragment. Particular characteristics such as mobility, home-range, susceptibility to barrier, edge effects and the matrix are all important parameters for a species response.

1.4 Declining mammalian fauna

Many animal species have been declining and disappearing worldwide (Soulé et al.

1979). Australia has been credited with approximately one-third of the global mammal extinctions since European settlement in 1788 (Maxwell et al 1996, Burgman and

Lindenmayer 1998). The Australian federal governments’ Environment Protection and

Biodiversity Act 1999 (EPBC Act, Australian Government 2007) lists 27 mammals that have become extinct since 1788. For the extant species, the EPBC Act lists three that are critically endangered, 33 that are endangered, 54 that are vulnerable, and one that is conservation dependent. In New South Wales (NSW) approximately 25% of the native mammalian species that were present in 1788 are now extinct (NPWS 2003).

6 1.5 Arboreal mammals

Arboreal mammals around the world have been reported as being particularly sensitive to fragmentation (Laurance 1990, Lopes and Ferrari 2000, Umapathy and Kumar 2000), mainly because even only partly cleared forest is a most effective barrier to movement

(Braithwaite 1996, Lindenmayer 1996a, Laurance 1997, Pausas and Austin 1998,

Goldingay 2000). The conservation of arboreal mammals in forests is vital as they perform many important ecosystem functions, including pollination, seed dispersal, and many are the primary prey of large forest owls (Kavanagh 1988, Goldingay 2000).

1.6 Status of Australian arboreal marsupials

Since European settlement, no Australian arboreal marsupial has become extinct

(Lindenmayer 2002a, Goldingay and Jackson 2004). They are reliant on the forests of the continent (Strahan 1995). The areas of forest where the arboreal fauna are located are characterised by fertile soils (Braithwaite 1996), these are the same areas which are sought after by the forest and agricultural industries (Braithwaite 1984). For example, only approximately 1% of the Sydney blue gum (Eucalyptus saligna) forests remain after two centuries of settlement, when compared to the relatively intact communities

(heath and woodland) found on the poorer soils (Benson and Howell 1990).

The possum and gliding arboreal marsupials have been one of the most studied suite of mammal taxa in Australia (Archer 1984), due to the effects of habitat removal and fragmentation on the biology and ecology of individual species (Smith and Hume 1984,

7 Goldingay and Jackson 2004). Among the possum and gliders, twelve species are listed as either rare, vulnerable or endangered in federal or state legislation (Australian government 2007, Carthew 2004, Eyre 2004, Jones 2004, Kavanagh 2004a, van der Ree et al. 2004, Winter et al. 2004) (see Table 1).

Table 1. The possum and glider species listed in Australian state and federal conservation legislation.

Species Status/Listing Habitat Range Acrobates pygmaeus Endangered in South Forests South East Australia (S.A) Australia (SEA) Gymnobelideus leadbeateri Endangered EPBC and Vic. Forests Endemic Vic. Petaurus norfolcensis Endangered in S.A and Vic, Forests/Woodlands SEA Vulnerable in NSW Petaurus gracilis Endangered EPBC and Qld. Woodlands Endemic Petaurus australis Endangered in S.A, Forests SEA 1 = Wet tropics population Vulnerable EPBC (1), NSW and Qld (1). Pseudocheirus occidentalis Vulnerable EPBC Forests/Woodlands sw Western Australia Phalanger intercastellanus Rare in Qld. Rainforest Cape York peninsula and New Guinea (NG) Spilocuscus maculatus Rare in Qld. Rainforest, Cape York woodland and peninsula and NG mangroves Pseudochirops archeri Rare in Qld. Upland to highland Endemic to wet rainforest tropics Pseudochirulus cinereus Rare in Qld. Upland to highland Endemic to wet rainforest tropics Pseudochirulus herbetensis Rare in Qld. Upland to highland Endemic to wet rainforest tropics Hemibelideus lemuroides Rare in Qld. Upland to highland Endemic to wet rainforest tropics

Of these, nine are from the eastern states, one from Western Australia and one from

South Australia. The four endemic wet tropics species; Green ringtail possum

(Pseudochirops archeri), Daintree River ringtail possum (Pseudochirulus cinereus),

Herbert River ringtail possum (Pseudochirulus herbetensis) and Lemuroid ringtail

8 possum (Hemibelideus lemuroides) are listed as rare on the basis of restricted geographical distribution (Winter et al. 2004). They have been reported to be vulnerable to habitat fragmentation and isolation (Laurance 1997). Winter et al. (2004) suggested that their status should be changed from rare to vulnerable, listing global warming as the main threat to these four species. The Southern common cuscus (Phalanger intercastellanus) and Common spotted cuscus (Spilocuscus maculatus) are restricted to

Cape York Peninsula in Australia and are both listed as rare. They have a wider geographic range and are found in New Guinea and neighbouring islands (Flannery

1994), climate change and habitat destruction is not a major threat to these two species

(Winter et al. 2004). The decline of the Western ringtail possum (Pseudocheirus occidentalis) has been attributed to removal of up to 80% of its habitat, fuel reduction burning, and fox predation (Jones 2004).

The endangered Leadbeater’s possum (Gymnobelideus leadbeateri) illustrates the conflict between natural resource exploitation (i.e. timber harvesting) and species conservation (Lindenmayer 1996a). This endemic arboreal species is found in a restricted area of the central highlands of Victoria (Vic.), where some of the most economically valuable forest areas are also found (van der Ree et al. 2004).

Lindenmayer (1996) reported ‘forestry operations must change if they are to become ecologically sustainable and are to preserve leadbeater’s possum and other species in the long term’.

Four gliding marsupials are listed in Table 1, the Feather-tailed glider (Acrobates pygmaeus), Squirrel glider (Petaurus norfolcensis), Mahogany glider (Petaurus gracilis) and the Yellow-bellied glider (Petaurus australis).

9

The Feather-tailed glider is rare in South Australia, the western edge of its range

(Carthew 2004). It is only known from two locations, where suitable habitat is limited.

Surveys have reported the species as uncommon in the state, with low numbers captured

(Carthew 2004).

The Squirrel glider has been affected by woodland clearing for agriculture, and is now primarily restricted to woodland remnants or linear networks (roadside vegetation) set within cleared farmland in south-east Australia (van der Ree 2000, van der Ree et al.

2003). A study on the population ecology of the Squirrel glider within linear habitats found that their density was equal to or greater than studies undertaken in continuous forest elsewhere (van der Ree 2002). The responses of Squirrel gliders to habitat fragmentation vary, showing population loss and declines throughout its former range

(Lindenmayer 2002a) and population increases in linear habits (van der Ree 2002).

For over 100 years the Mahogany glider was generally referred to as either a subspecies or synonym of P. norfolcensis (Jackson 2000), until it was given full species status (Van

Dyck 1993). It is endemic and restricted to an area of north Qld. around Cardwell

(Flannery 1994, Strahan 1995). Lindenmayer (2002a) reported that the Mahogany glider is ‘highly endangered with a high risk of extinction in the coming decades due to poor land management practices such as land clearing for agriculture’. The Mahogany gliders’ original habitat, has been reduced to 20% of its former size (Winter et al. 2004).

At present only a small amount of the remaining habitat is conserved in protected areas

(Goldingay and Jackson 2004).

10 The Yellow-bellied glider was found to be lost from forest fragments (some >120 ha) that were embedded in pine plantations in the Tumut area of NSW (Lindenmayer

2002a). This species has a very large home range (up to 60 ha), forages extensively for preferred feeding trees, and is hollow dependent (Goldingay and Jackson 2004). The reduction in frequency and intensity of fires have been reported as causing loss of yellow-bellied glider (wet tropics population) habitat due to rainforest species capturing forested areas in north Qld (Winter et al. 2004), which may affect its ability to use food resources (Goldingay and Quin 2004).

All the other possum and glider species are protected and, while many are not listed in conservation legislation, they have shown declines in areas of their range. The Common ringtail possum (Pseudocheirus peregrinus) and the Common brushtail possum

(Trichosurus vulpecula) have been reported to be declining in areas where natural habitat has been removed (Kerle 2004, Paull and Kerle 2004). These two species have become common in urban areas (Maloney and Harris 2006a) and T. vulpecula has become a pest after introduction to New Zealand (Efford and Cowan 2004).

The conservation legislation lists the threats which are the cause of the decline in possums and glider species. These include:-

• Loss and fragmentation of habitat through clearing and the activities associated

with clearing;

• Logging of old growth elements removes the number of hollow bearing trees

available for nesting;

• Inappropriate fire regimes reduces the availability of food resources and isolates

populations making them vulnerable to regionally catastrophic events;

11 • Predation by introduced animals (foxes and cats).

1.7 Greater Glider

The focus of this study is the Greater Glider Petauroides volans Kerr 1792, which is a nocturnal, hollow dependent, folivorous arboreal marsupial (McKay 1995). The Greater

Glider is the most common species found in the taller and wetter forests on the east coast of Australia, but not in Tasmania (Archer 1984, McKay 1995, Lindenmayer

2002a, Menkhorst 2004) (Figure 1).

Although the Greater Glider is not yet considered vulnerable or endangered as a species, it has many characteristics that make other Australian mammals sensitive to fragmentation:

(i) Low mobility and small home-range;

(ii) Dependence on particular eucalypt species for food;

(iii) Reliance on old growth forest for dens.

12

Figure 1. The distribution of the Greater Glider (Petauroides volans) in Australia. (Darker shading on the eastern states). (Source Wildlife Preservation Society of Queensland 2005).

As a result of these characteristics, a fragmented population of Greater Gliders in the

Eurobodalla Local Government Area is listed as an endangered population under the

NSW Threatened Species Conservation Act 1995 (Department of Environment and

Climate Change NSW 2007) because of fragmentation of habitat by roads and farms.

The population has been deemed to be disjunct from other occurrences of the species due to these barriers restricting their dispersal ability, and continuing loss of habitat will further isolate the population.

13 Fleay (1947) stated, for the Greater Glider:

‘the fact that the animals have an inseperable attachment for dense forested lands, and do no economic damage whatever, is not likely to lead to any molestation by man’.

During the 1960s, a study on the effects of clearing of native forest for a Pinus radiata plantation at Tumut (NSW) found that 90% of the displaced Greater Gliders were never recaptured in adjoining forest (Tyndale-Biscoe and Smith 1969b). Tyndale-Biscoe and

Smith (1969b) concluded ‘the displaced gliders die in situ rather than emigrate to occupied forest and die there through failure to become established’. Greater Gliders recaptured in the same season as initial capture, had lost up to 25% of body weight and many breeding females had lost pouch young, due to the stress of habitat loss (Tyndale-

Biscoe and Smith 1969b).

Lindenmayer (2002a) stated that ‘extensive field surveys conducted in radiata pine plantations over the past three decades have failed to find gliders inhabiting these areas’.

Stands of exotic softwoods lack the resources needed by Greater Gliders, such as appropriate hollows and food (Lindenmayer et al. 2000, Pope et al. 2004). This also suggests that Greater gliders do not use this sort of matrix, thus making them vulnerable to population declines or extinction within fragmented habitat (Laurance 1997).

Remnant patch sizes of <3 ha (within the radiate pine plantation) were reported to be of limited value as a habitat for the species (McCarthy and Lindenmayer 1999a).

Nevertheless, many larger patches (e.g. >5 ha) of eucalypt remnants which have been surrounded by these pine forests for 65 years supported Greater Glider populations

(Lindenmayer 2002b).

14 Surveys for aboreal marsupials in Australian forests have reported that the Geater Glider is rare or absent in areas that were logged (Lunney 1987, Kavanagh et al. 1995,

Goldingay and Daly 1997, McCarthy and Lindenmayer 1999a). This species is reliant on old-growth forest, which supports a higher number of individuals than regenerating forest (McFarlane 1988, Stockwell et al. 1990, Comport et al. 1996). Recently the

Greater Glider has been reported as having declined or disappeared over a 25 year period in uncleared woodland sites in central Queensland. (Woinarski et al. 2006), and to have a probability of 50-94% of remaining extant in the outer local government areas of Melbourne (van der Ree and McCarthy 2005).

The home-range of the Greater Glider varies between 1.4 ha to 2.6 ha for males and 0.8 ha to 2.5 ha for females across its range, with overlap only between males and females

(Henry 1984, Kehl & Borsboom 1984, Norton 1988, Comport et al. 1996, Kavanagh and Wheeler 2004, Pope et al. 2004). Pope et al. (2004) reported that home-range decreases with decreasing patch size, with small patch areas having a larger number of

Greater Gliders per unit area. This species shows a high affinity to its home-range, remaining in situ when the habitat is cleared, subsequently becoming prey to a range of predators (Tynedale-Biscoe and Smith 1969b, Maloney and Harris 2006b). Kavanagh and Wheeler (2004) reported that Greater Gliders restricted their home-ranges to the unlogged part of their range after intensive logging (see also Tynedale-Biscoe and

Smith 1969b). Although this species is the largest glider, it does not require a large home-range as suitable food resources are abundant and continuous (Lindenmayer

2002b).

15 The Greater Glider has been reported to use the linear strips of unlogged forest or

‘wildlife corridors’ set aside for wildlife management in timber production forests

(Lindenmayer 1996a). Downes (1997) reported that they were not found in corridors

<32 m in width in the Strathbogie Ranges Vic. The Greater Glider was also recorded in corridors ranging in width between 40-160m in the Mountain Ash forests in Vic.

(Lindenmayer 1996a, 1997). Menkhorst (1984) reported that they used nest boxes that had an opening of 8 cm, set at a height of 8 m with a north orientation, other studies have reported that orientation had no significance (Norton 1988). The den and nest trees are tall and of large diameter, the entrance is through a spout or hollow branch 40m high

(Lindenmayer 1997). The number of hollows is reported to be a limiting factor in the occurrence of the Greater Glider (Incoll 2001). Tree hollows used as nest and den sites are the key habitat requirement for virtually all species of Australian arboreal marsupial

(Lindenmayer et al. 2000).

Fire is a natural disturbance in Australian ecosystems; and is used as a management tool in forests that are close to urban areas (Benson 1999). The use of fire as a management tool causes changes in the occurrence and frequency of fire at a site. The timing, intensity and frequency of a fire event can have a detrimental affect on the life cycle attributes of a species during and after a fire event (Whelan et al. 2002). Kavanagh et al

(1995) suggested the Greater Glider was found in forests with a sparse understorey, which is probably maintained by frequent fire.

The effects of fire on the Greater Glider was reported by Fleay (1947) who stated:

‘Bushfires are the scourge and bane of this species. The animals

are utterly hopeless, and I have watched them helplessly twirling

16 in agonized bewilderment down into the onrush of consuming

flames’. …. ‘In the holocaust of early 1939 tens of thousands of

these volplaning creatures perished in the Victorian ranges, and

even many that survived in odd sheltered gullies died

subsequently of starvation, because leaves of their food trees,

where not consumed, were blasted and withered’

This report illustrates that, during a fire event, Greater Gliders are killed due to lack of mobility during the event, and if they survive in parts of the landscape that is spared by fire, they are likely to die subsequently from a lack of food resources. The Greater

Gliders’ dependence on tree hollows makes them likely to be susceptible to death by smoke inhalation during a fire event.

Lunney (1987) reported that gullies that are spared by natural calamities, such as fire and logging practices, serve as refuges for this arboreal marsupial. van der Ree and

Loyn (2002) found that the Greater Glider was more abundant in 1939 than 1983 regrowth, due to the post fire salvage logging after 1983, which reduced the number of hollow bearing trees in the latter (see also Lindenmayer and Ough 2004). Macfarlane

(1988) agreed with this interpretation and added ‘they are able to recolonise regrowth after a wildfire from the untouched gullies as the habitat becomes suitable, in around 40 years’. The interactions between fire and logging processes have been suggested to have an impact on the Greater Glider because, in some areas, the amount of old-growth forest is small after logging and could be destroyed by fire (Lindenmayer 2002a).

Wildfires reduce the number of available tree hollows in the landscape for arboreal marsupials (Gibbons and Lindenmayer 1997).

17

1.8 Aims

The Greater Glider is distributed in fire-prone forests in south-east Australia, many of which are being increasingly fragmented by infrastructure (road and rail) as well as clearing for agriculture, forestry and urban development. Since European settlement, the continuity of the habitats of many species in Australia has been compromised.

Anecdotal evidence suggested that the Greater Glider, Petauroides volans Kerr 1792, was eliminated from Royal National Park as a result of large-scale and intense bushfires in January 1994 (Andrew 2001). This species had been readily seen in some parts of the park prior to this fire but large-scale fauna surveys conducted by the National Parks and

Wildlife Service (NPWS) in 1996 and 1997 indicated that they had disappeared, even though the vegetation had regenerated (Andrew 2001).

This thesis is concerned with the distribution of the Greater Glider in the Illawarra region, and the reasons why it is unable to recolonise an area in which it was formerly found. On the face of it, this species would seem to be an ideal candidate for a translocation program at Royal National Park because it was formerly abundant, the remaining habitat is isolated, it is excluded by fire, and it is now unlikely to recolonise by itself. The ultimate aim of this study was to prepare such a plan (Chapter 5).

In order to design an effective management strategy for this species at Royal National

Park, including a translocation program, a thorough understanding of its distribution in the regional landscape is required. Further, preliminary work suggested that the

18 taxonomy of the Greater Glider was not entirely clear – potentially leading to erroneous location records for the species. Therefore, the proximate aims of this study were to:

(1) clarify the taxonomy of the species and interpret historical accounts

(Chapter 2),

(2) review current knowledge of the distribution and abundance of Greater Gliders in

the study region, and review the current threats to populations (Chapter 3),

and

(3) conduct a detailed field study in the region, to determine current location of

populations (Chapter 4).

19 Chapter 2

Historical Accounts and Discovery of the Greater Glider

2.1 Introduction

The Greater Glider, Petauroides volans, is relatively conspicuous and was quickly noticed by the early colonists (Phillip 1789). Subsequently, descriptions of this species were included in many of the earliest zoological accounts of the Australian fauna.

However, few modern zoologists are aware of the historical significance and value of this old literature as it relates to this and other species (see also Harris 2006). Whilst some of this literature on P. volans has been reviewed by McKay (1982), this was limited to aspects of the nomenclature of the genus name Petauroides (and also

Petaurus). This chapter provides a comprehensive survey of the early natural history literature pertaining to P. volans, including information on discovery, taxonomy, dentition, morphology, distribution, abundance, diet and behaviour. This chapter was designed to clarify the taxonomy of the Greater Glider, one of the requirements of a translocation program.

2.2 Taxonomy and nomenclature

Governor Arthur Phillip reported ‘black flying opossum’ from NSW (Phillip 1789). A male specimen owned by Henry Constantine Nowell was illustrated (Figure 2.1), although no details on the precise collection locality were published. Presumably it was found in the vicinity of Port Jackson. Phillip (1789, 1790) recognised that it represented

20

Figure 2.1: Black flying opossum (=Petauroides volans) drawn by P. Mazell and published in Phillip (1789). Note the opposable clawless hallux and syndactylous digits on each of the hind feet.

21 a new species and suggested taxonomic affinity with American Didelphis, although a specific name was not offered. A few years later, Kerr (1792) named Phillip’s specimen

Didelphis volans, derived from the Latin word ‘volare’ meaning ‘to fly’ (Strahan 1981).

A year later, Meyer (1793), named Phillip’s specimen D. voluccella, and a year later still, Shaw (1794) proposed the name D. macroura. In Shaw’s work, The Zoology of

New Holland, a juvenile specimen drawn by James Sowerby was illustrated (Figure

2.2). Shaw (1800) explained that it was sent to him by John White, who was the first

Surgeon-General for the colony of NSW.

Cuvier (1798) followed use of the name D. volans (Kerr 1792), but questioned the affiliation with the genus Didelphis. Nevertheless, Shaw (1800) continued the use of D. macroura. Bechstein (1800) elevated the name Voluccella, used by Meyer (1793), to generic level and proposed V. nigra for the subject species, but he evidently confused the Greater Glider and the Yellow-bellied Glider Petaurus australis in synonymy. His proposed V. nigra incorporated D. voluccella Meyer, 1793 (=Petauroides volans) and

“Hepoona Roo” White, 1790 (=Petaurus australis). It is understood that Hepoona Roo is P. australis and not Petauroides volans (McKay 1982, 1988). Bechstein (1800) also advanced V. macroura as a separate species that incorporated D. volans Kerr, 1792 and

D. macroura Shaw, 1794. Thus, V. nigra and V. macroura are both synonyms of P. volans. Voluccella Bechstein, 1800 was discontinued for the subject species because this genus name had already been advanced by Fabricius (1794) for a species of fly

(Diptera: Bombyliidae) (Thomas 1888; McKay 1988; Evenhuis 1991). Hence,

Voluccella Bechstein, 1800 is a junior generic synonym for Petauroides but not

Voluccella Fabricius, 1794.

22

Figure 2.2: Long-tailed opossum Didelphis macroura (=Petauroides volans) from Zoology of New Holland (1794) by George Shaw. The figure was drawn by James Sowerby. This illustration was also reproduced in Shaw (1800) and Desmarest (1820).

23 Phalanger volans was used by Lacépède (1801), whilst Desmarest (1803) and

Tiedemann (1808) placed it under Phalangista (see also Schinz 1821; Thomas 1888).

Turton (1806) mistakenly thought that the descriptions by Kerr (1792; D. volans) and

Shaw (1794; D. macroura) represented two separate species. Oken (1816) made a similar mistake, but also erroneously included the Petaurus australis in the synonymy for one of his proposed species. This was Petaurus niger, and the epithet was a gender change of Bechstein’s (1800) nigra (see also Iredale and Troughton 1934). Oken’s

(1816) second species was Petaurus macroura.

Desmarest (1817) listed three species (Petaurus macrourus, P. peronii and P. taguanoides). Petaurus macrourus included a slight change in the epithet to standardise the gender of the binomial. Desmarest’s explanation that the membrane of P. peronii

“terminates at the elbow” is good evidence that this specimen was also P. volans. For P. taguanoides, however, the synonymy was confused with the Yellow-bellied glider [i.e.

Didelphis petaurus of Shaw (1791) and “Hepoona Roo” of White (1790)] and the descriptions about the patagium ending at the wrist suggested to us that this specimen was not the Greater Glider. However, according to the publications of the Muséum

National d'Histoire Naturelle (MNHN) the type specimen of P. taguanoides as described by Desmarest is indeed P. volans (de Beaufort 1966; Julien-Laferrière 1994).

To confirm this identification I contacted the MNHN directly, and obtained a photograph of the specimen (number CG1990-408) and although no patagium was evident in the photograph, it looks like a Greater glider because of its long tail and hairy ears. The arrangement of Desmarest’s (1817) terminology was later followed by Cuvier

(1826), Lesson (1827, 1828, 1830, 1838), and Fischer (1829). Bennett (1837) also used

24 Desmarest’s (1817) terminology, although he appears to have used P. peronii in reference to the sugar glider Petaurus breviceps.

Desmarest (1820) applied Petaurista to supersede Petaurus, and maintained Petaurista taguanoides, P. macroura and P. peronii as separate species (later followed by Cuvier

1827, 1829). However, this was flawed, because Petaurista had already been advanced for the giant flying squirrels (Rodentia) by Link (1795) (see also Fischer 1814; Thomas

1888; Sherborn 1902; Palmer 1904). Waterhouse (1838b), Gloger (1842), Gould (1863) and Thomas (1885), persisted with this invalid generic name for the Greater Glider.

Frédéric Cuvier (1825) mentioned Petaurus didelphoides Geoffroy, an apparent new name for the subject species (Thomas 1888; Iredale and Troughton 1934; de Beaufort

1966). However, later works by F. Cuvier and also his brother Georges, made no reference to P. didelphoides (Cuvier 1826, 1827, 1829). de Beaufort (1966) noted that

Cuvier (1825) offered no specific descriptions, and stated that he was unable to find any reference to Geoffroy as the authority for the name. It is uncertain whether Cuvier intended this name for the Greater Glider. Iredale and Troughton (1934) considered it a vernacular name.

Lesson (1828, 1830, 1838) listed the “Black Flying Opossum” of Phillip (1789) (=the

Greater Glider) as a junior synonym of Petaurus taguanoides. This was subsequently repeated by Fischer (1829), Wagner (1843), Schinz (1844) and Giebel (1859).

Waterhouse (1838a) then stated that two specimens of Petaurista taguanoides were held in the Museum of the Zoological Society of London (ZSL), one of which was a ‘white variety’. Waterhouse (1841) included an illustration of a Greater Glider (Figure 3.2),

25 and stated that “Specimens which are totally white, and others which are white and irregularly variegated with grey, are not rare”. Waterhouse (1841) was wrong when he suggested that P. macrourus is P. flaviventer (=P. australis) (see also Wagner 1855;

Giebel 1859; Gould 1863). Descriptions of taguanoides specimens in many 19th century publications subsequent to Waterhouse (1841) also appear to be wrong, taxonomically, and actually represent the Greater Glider (e.g. Owen 1841, 1845; Gloger 1842;

Waterhouse 1846; Gervais 1855; Gerrard 1862; Brehms 1880; Flower 1884; Forbes-

Leith and Lucas 1884; Krefft 1864; Haswell 1886; Jentink 1886; Lucas 1890).

Major T.L. Mitchell collected a presumed new species of glider “from the banks of the

Murray”, named it Petaurus leucogaster and “deposited [it] in the Australian Museum

(AM)” (Bennett 1837; Mitchell 1838). Gray (1841) suggested that it “may only be a variety of [the] P. taguanoides” of Waterhouse (=the Greater Glider) (see also mention of P. leucogaster in Gray 1842, 1843; Krefft 1864). Several authors considered leucogaster to be synonymous with P. volans (Gould 1863; Thomas 1888; Iredale and

Troughton 1934; McKay 1982). However, McKay (1988) stated that P. leucogaster was

Incertae sedis (of uncertain position) because the specimen could no longer be found at the AM. He suggested that the locality for Mitchell’s specimen was outside the range of

P. volans and may have been Petaurus norfolcensis.

M.R. Oldfield Thomas, of the British Museum of Natural History (BMNH), revised the taxonomy of the subject species several times during the period 1879-1923. Thomas

(1879) noted that the specific name volans Kerr antedated taguanoides Desmarest, and maintained that the correct binomial was Petaurus volans. A few years later, however, he listed it as Petaurista volans (Thomas 1885). After finding that Petaurista was unavailable, Thomas (1888) advanced Petauroides to replace the previous generic

26 names. He listed two sub species: Petauroides volans typicus as the southern form; and

P. v. minor as the northern form (following Collett 1887). Later, Thomas (1923) received further examples from Qld and considered that there were two additional subspecies: P. v. incanus and P. v. armillatus.

Figure 2.3: A print from Waterhouse (1841) that is clearly Petauroides volans because of the length of the tail and the hairy ears. This image was also reproduced in Waterhouse (1843) and Lydekker (1896).

27 Thomas (1923) mentioned that Ogilby (1892) referred to “Dr Ramsay’s P. cinereus” and that it “seems never to have been described”. However, Ramsay (1890) did indeed publish a description of a supposed new species, which he named Petaurides cinereus.

This was based on two specimens obtained from the Bellinden-Ker Range, north-east

Qld. The name Petaurides is a definite misspelling of Petauroides Thomas 1888 (see

Ramsay 1890). It is also noted that these specimens had earlier been exhibited at a meeting of The Linnean Society of NSW under the name of Belideus cinereus (Anon

1890).

The next taxonomic contribution was by Iredale and Troughton (1934). They argued that the generic name Schoinobates Lesson 1842 had been published before Petauroides

Thomas 1888, and advanced the name S. volans with four subspecies: S. v. volans; S. v. incanus, S. v. armillatus and S. v. minor. Subsequently, S. volans was in use for around

50 years (Fleay 1947, 1968; Tate 1945; Anon 1946; Troughton 1935, 1941; Marlow

1958, 1962; de Beaufort 1966; Ride 1970; Strahan 1980, 1981). However, the nomenclatural change by Iredale and Troughton (1934) was groundless. McKay (1982) pointed out that Schinobates was first used by Lesson (1842) to supersede Petaurista leucogenys Temminck, 1838 (=Pteromys leucogenys; the Japanese flying squirrel). In fact, this was an error on Lesson’s part because there are no marsupials in Japan (Palmer

1904). Nevertheless, it was highly irregular for Iredale and Troughton to amend the type locality of P. leucogenys from “Japan” to “Sydney”. Probably, Iredale and Troughton

(1934) did not view the original account and illustration of P. leucogenys in Fauna

Japonica (Temminck 1838), which clearly depicts a sciurid. Schoinobates Lesson,

1842, is therefore properly placed as a junior synonym of Petaurista Link, 1795. Thus,

28 McKay’s (1982) assessment that the name Schoinobates was unavailable and that the name Petauroides must therefore stand was justified.

Iredale and Troughton (1934) also nominated Petaurus maximus as a synonym for the subject species, listing Partington (1837) as the authority. This was accepted by McKay

(1982) and Flannery (1994). However, McKay later attempted unsuccessfully to uncover the original reference and stated that the relevant page in the book he examined

“contains no reference to this or any other mammal” (McKay 1988). McKay (1988) misread Iredale and Troughton’s (1934) reference to Partington (1837: 424), because P. maximus is indeed described in The British Cyclopædia of Natural History, but not in the The British Cyclopædia of Arts and Sciences, which was read by McKay (1988).

After reading Partington (1837) with its reference to some “almost white” specimens, I accept P. maximus can be accepted as synonymous with the Greater Glider (following

Iredale and Troughton 1934). The preceeding literature review of taxonomy of the

Greater Glider is presented in Table 2.1.

Common names for the subject species have included ‘black flying opossum’ (Phillip 1789), ‘flying opossum’ (Kerr 1792; Turton 1806; Waterhouse 1841), ‘long-tailed opossum’ (Shaw 1794, 1800; Turton 1806; Waterhouse 1841), ‘large-tailed Petaurista’, ‘Peron’s Petaurista’ (Cuvier 1827), ‘white-bellied flying squirrel’ (Bennett 1837), ‘grey flying squirrel’ (Bennett 1837; Waterhouse 1841) ‘large-tailed flying squirrel’ (Bennett 1837), ‘taguan flying opossum’ (Waterhouse 1838b), ‘taguan flying phalanger’ (Waterhouse 1846; Thomas 1888, 1923; Fleay 1933), ‘greater flying phalanger’ (Gould 1863; LeSouef and Burrell 1926; Fleay 1933), ‘the brill’ (De Vis 1886), ‘flying phalanger’ (Haswell 1886), ‘great flying oposssum’, ‘flying squirrel’ (Lucas 1890),

29

Table 2.2: Proposed new synonymy based on the current review.

Petauroides Thomas, 1888 Petaurus Shaw, 1791 Didelphis Kerr, 1792 Voluccella Bechstein, 1800 Phalanger Lacepede, 1801 Phalangista Desmarest, 1803 Petaurista Desmarest, 1820 Petauroides Thomas, 1888 Petaurides Ramsay, 1890 Belideus Anon, 1890 Schoinobates Iredale and Troughton, 1934 Petauroides volans (Kerr, 1792) Petauroides volans volans (Kerr, 1792) Didelphis volans Kerr, 1792 Didelphis voluccella Meyer, 1793 Didelphis macroura Shaw, 1794 Voluccella nigra Bechstein, 1800 Voluccella macroura Bechstein, 1800 Phalanger volans Lacepede, 1801 Phalangista volans Desmarest, 1803 Petaurus macroura Oken, 1816 Petaurus niger Oken, 1816 Petaurus taguanoides Desmarest, 1817 Petaurus macrourus Desmarest, 1817 Petaurus peronii Desmarest, 1817 Petaurista taguanoides Desmarest, 1820 Petaurista macroura Desmarest, 1820 Petaurista peronii Desmarest, 1820 Phalangista macroura Schinz, 1821 Petaurus didelphoides Cuvier, 1825 Petaurus maximus Partington, 1837 Petaurus volans Thomas 1879 Petaurista volans Thomas 1885 Petauroides volans typicus Thomas, 1888 Petauroides volans incanus Thomas, 1923 Petauroides volans armillatus Thomas, 1923 Schoinobates volans volans Iredale and Troughton 1934 Schoinobates volans incanus Iredale and Troughton 1934 Schoinobates volans armillatus Iredale and Troughton 1934 Petauroides volans minor (Collett, 1887) Petaurista volans minor Collett, 1887 Belideus cinereus Anon, 1890 Petaurides cinereus Ramsay, 1890 Schoinobates volans minor Iredale and Troughton 1934

30 ‘dusky glider’ (Fleay 1933; Ride 1970), ‘Greater glider-possum’ (Iredale and Troughton

1934; Anon 1946), and ‘Greater glider’ (Marlow 1958). Stability in the vernacular name was achieved in 1980 when a committee of the Australian Mammal Society formalised it as the ‘Greater glider’ (Strahan 1980).

2.3 Morphology

The morphology was first described by Phillip (1789). He stated that the “tip of the nose to root of tail [was] 20 inches [=508 mm], tail 22 inches [=559 mm], loins 16 inches

[=406 mm].” The ears were described as “large and erect”, the fur “glossy black” on top, “mixed with grey”, and “the under parts...white”. It was noted that the fur

“continued to the claws”, and that the membrane “expanded on each side of the body”.

Phillip (1789) also described and illustrated the foot (Figure 2.1). He observed that the

“fore legs have five toes on each foot, with a claw on each; the hinder ones four toes, with claws, (the three outside ones without any separation) and a thumb without a claw”. Following Phillip (1789), similar descriptions were also published by subsequent authors based on his original account and from the illustration provided (i.e. Kerr 1792;

Meyer 1793; Bechstein 1800). Shaw (1794) provided morphological descriptions based on the illustration reproduced in Figure 2.2.

One diagnostic feature of P. volans is the flying membrane which runs from the elbow to the knee, and this was noted by several early zoologists (Kerr 1792; Turton 1806;

Desmarest 1817; Waterhouse 1841, 1846). Thomas (1888) added that the membrane is

“very narrow along the sides of the forearm and lower leg”. Ramsay (1890) stated that the “parachute” or “wing membrane” commences a little in front of the elbow-joint, and

31 extends to about half-way below the knee-joint. Numerous early authors also noted the syndactylous hind feet (Kerr 1792; Shaw 1794, 1800; Bechstein 1800; Lacepede 1801;

Tiedemann 1808; Desmarest 1820; Partington 1837).

Some authors have compared the size of this species to animals known from Europe.

For example, it has been suggested to be about the size of a “black rat” (Shaw 1800),

“flying squirrel” (Desmarest 1803; Tiedemann 1808), “surmulot” (Cuvier 1817;

Desmarest 1820; Lesson 1827), “squirrel of Europe” (Desmarest 1820; Lesson 1827), and “brown rat’ (Partington 1837). More recently, it has been suggested to be about the size of a domestic cat (Flannery 1994).

Colouration was also frequently commented on. For example, Cuvier (1817) reported that the fur exists in different tones of brown; with many varieties, and others are whitish. Fully white specimens were also noted (Lesson 1827; Waterhouse 1841; Krefft

1864; Le Souef and Burrell 1926). Gould (1863) stated that “it is subject to very great variety in the colouring of its fur, some specimens being entirely blackish brown [see

Figure 2.4], grey to cream and others quite white”. Krefft (1871) reported that the species “varies much from creamy-white to spotted black and white and perfect black, beneath the fur is always white.” Le Souef and Burrell (1926) stated that “as a rule [the] colour [is] darker in winter than in summer.” They also stated that “animals from

Gippsland (Victoria) [were] dead black above and on tail; pure white on undersides”; whereas Qld and NSW specimens were “usually smoky grey” and “white specimens

[were reportedly] common.”

32

Figure 2.4: Petaurista taguanoides from Gould (1863) (= P. volans). Note: The front arms of the background glider are shown in the wrong position as P. volans tucks them under the chin when gliding (Fleay 1933; Grzimek 1967; McKay 1989).

33 Other notable morphological features described in the early literature include the ears, tail and size differences in the sexes. Waterhouse (1838a, 1841, 1846) stated that “the ears are entirely covered externally with long and dense fur, flesh-coloured and almost bare within” (see also Krefft 1864, Thomas 1888, Ramsay 1890). The tail was reported as not being prehensile (Lacepede 1801; Tiedemann 1808; Partington 1837), and longer than the body (Shaw 1800; Turton 1806; Cuvier 1817). Thomas (1888) described and illustrated the naked tip of the tail. Gould (1863) stated the “sexes offer no external difference, except that the female is somewhat smaller than the male” (see Flannery

1994, as this is erroneous). Various other aspects of the morphology of this species are discussed in the literature, but lack of space precludes a detailed discussion here. These aspects include skull structure (Waterhouse 1846; Collett 1887; Thomas 1888) and myology (Haswell 1886).

2.4 Dentition

Phillip (1789) stated that in “the upper jaw forwards are four small cutting teeth, then two canine ones, and backwards five grinders: the under jaw has two long large cutting teeth, five grinders, with no intermediate canine ones, the space being quite vacant”.

Similarly worded descriptions were provided by Kerr (1792) and Turton (1806).

A dental formula for the species was first provided by Desmarest (1820) (see Table 2.2).

He counted six upper and two lower incisors, but was uncertain about the number of canines and premolars. This uncertainty led him to indicate a total of 32 or 34 teeth.

Cuvier (1825) and Lesson (1827) counted a total of 38 teeth. Cuvier (1825) reported that the space between the incisors and molars is occupied by two rudimentary teeth.

34 Waterhouse (1838b, 1841) and Owen (1841, 1845) mentioned they had never observed any of these diminutive teeth in the specimens they had examined. Waterhouse (1841) suggested that Cuvier (1825) may have inadvertently described the dentition of

Phalangista cookii (=Pseudochirus peregrinus; common ringtail possum). These two species do have great similarity in their dental characteristics, as noted by early zoologists (Owen 1841, 1845; Giebel 1853, 1855; Thomas 1885; Collett 1887) and more modern authors (Tate 1945; Triggs 1996). Waterhouse (1838b) provided a dental formula indicating a total of 34 teeth. Subsequent authors concurred with this observation (Waterhouse 1841, 1846; Wagner 1843; Collett 1887; Ramsay 1890). Early illustrations of the dentition in Cuvier (1825, 1827), Waterhouse (1846) and Giebel

(1853, 1855), support the dental formula of Waterhouse (1838b).

Krefft’s (1871) dental formula (Table 2.2) was for a total of 40 teeth (see also Collett

1887). Thomas’ (1885) assessment was that the number of teeth varied from 34 to 40, dependent on the presence or absence of a small canine and two premolars in the lower jaw. Thomas (1888) attempted to improve his earlier dental formula by changing the position of the lower canine to the incisor position (Table 2.2), and remarked that the

“presence or absence of the minute teeth is not of any systematic importance”. Thomas

(1888) provided illustrations of the upper and lower jaw of P. v. volans and P. v. minor, although these are not consistent with his dental formula. Later reviewers have alluded to a socket in the lower jaw where a small incisor would be present (i.e. Archer 1984;

Triggs 1996). Twenty-one P. volans specimens in the AM were recently examined, and four (19%) were noted to have minute teeth between the incisors and pre-molars.

35

Table 2.2: Dental formulas provided in the early natural history literature for Petauroides volans. Abbreviations: I = Incisors; C = Canines; M = Molars; P = Premolars. For Thomas (1885, 1888) an asterisk indicates that the tooth is sometimes or commonly absent.

Source Dental formula

Desmarest 6 1 1 1 1 6 6 7 7 I, ;C, or ;M or = 32 or 34 1820 2 0 0 2 2 6 6 6 6 Cuvier 1825, 6 0 0 8 8 I ;C, ; M , = 38 1826 2 0 0 6 8 Lesson 1827, 6 0 0 8 8 I ;C, ; M , = 38 1830, 1838; 2 0 0 7 7 Fischer 1829

Waterhouse 3 3 1 1 3 3 4 4 I ;C, ; P ; M , = 34 1838b 1 1 0 0 1 1 4 4

Krefft 1871; 3 3 1 1 3 3 4 4 I ;C, ; P ; M , = 40 Collett 1887 1 1 1 1 3 3 4 4 Thomas 1 2 3 1 1 2 3 1 2 3 4 I C P M x2 = 34 or 40 1885 1 0 0 1* 1* 2 * 3 1 2 3 4 Thomas 1 2 3 1 1 0 3 4 1 2 3 4 I C, P, (or1) M , = 17 + (at most) 3 (or 16 + 4) × 2 = 40 1888 1 2 * 0 0 1* 0 3 * 4 1 2 3 4

2.5 Habitat and diet

Some information on the habitat and diet of P. volans is available in the early literature.

Gould (1863) stated that the species seeks “blossoms of the Eucalypti…together with the tender buds and shoots of the same trees”. Similarly, Le Souef and Burrell (1926) stated that the “food consists of the leaves and buds of eucalyptus-trees”. They also added that:

‘careful examination of the contents of several stomachs of animals taken from the

forests has not revealed anything else, but in the Myall Lakes district [NSW]… we

36 have observed this species on the casuarina-trees; in one such case the contents of

the stomach, although much masticated, seemed to be the casuarina-leaves. Mr.

Ralph C. Blackett, forest ranger at Queanbeyan [NSW]…, states that they chiefly

feed on E. regnans, and to a lesser extent on E. viminalis, E. fastigata, E.

australasiana, and other narrow-leaved peppermints.’

In captivity, P. volans has been observed to eat E. sieberiana readily, “being especially fond of the flowers, and preferring the bark of the branches to the leaves” (Le Souef and

Burrell 1926). Fleay (1933) stated “one of the chief difficulties in captivity is the maintenance of an abundant supply of the tender leaves of acceptable species of eucalypts” and reported on collecting trips to obtain sufficient amounts of leaf from E. elaeophora and E. australiana. He also reported that “captive specimens could be persuaded to acquire an additional taste for bread and milk spread with a sweet jam, but only as an adjunct to the diet of eucalypt leaves.” Grzimek (1967) stated “because [P. volans] are exclusive in their diet, like koalas, no specimen has ever reached a European zoo alive.” Menkhorst and Knight (2004) stated that it “eats only eucalypt leaves and buds.” However, Maloney and Harris (2006b) reported feeding observations from several non-eucalypts.

In terms of habitat, Gould (1863) wrote that it “is strictly an inhabitant of the extensive brushes which stretch along the south-eastern and eastern portions of New South

Wales”. It has also been reported to occur in Eucalyptus forests (Le Souef and Burrell

1926, Anon 1946). Fleay (1933) stated that the species was found “favouring the taller timber areas and generally inhabiting dead trees in the gullies of mountainous country”.

Marlow (1958) reported that P. volans was more abundant in dry than wet sclerophyll

37 forests and less common in open woodland. Ride (1970) stated that “the habitat is sclerophyll forest and tall woodland”.

2.6 Distribution and abundance

The earliest statements on the distribution of the subject species was that it inhabits

NSW (Phillip 1789; Kerr 1792) or “New Holland” (=Australia) (Meyer 1793; Shaw

1794; Cuvier 1798; Bechstein 1800). The earliest specific localities mentioned were for places in NSW, such as Botany Bay, Port Jackson, Sydney, Blue Mountains, Port

Macquarie, Bathurst, Maitland, Clarence River and Goulburn Plains (Cuvier 1826,

1827; Lesson 1830; Bennett 1837; Waterhouse 1841; Gray 1841; Krefft 1864). Other early distributional records for NSW include Sutherland (1908, AM M2003),

Helensburgh (1909, AM M2051), Bowral (1918, AM M2724), Myall Lakes (1922, AM

M33762), Gerringong and Milton (Troughton 1935, 1941), Geehi Gorge (Mt

Kosciuszko area) (Anon 1946), Armidale and Tidbinbilla Nature Reserve (1974), (see

Maloney and Harris 2006).

Early literature records from Queensland are north of the Herbert River (de Vis 1886),

Herbert Vale, Coomooboolaroo, Calliungal (Collett 1887), Bellenden-ker Range

(Ramsay 1890), Eidsvold, Gin-Gin (Thomas 1923), Atherton tablelands, Evelyn

Station, Dimboola and Mount Spurgeon Stations (Tate 1945).

In Victoria, the species has been reported from Templestowe around 1865; east and north-east of Melbourne (Lucas 1890), and also from the south, south-west and questionably north-west areas of the State (Forbes-Leith and Lucas 1884; Lucas 1897).

38 Other Victorian distributional records include Allambee East, Newham, Bullengarook

(1905), Dandenong (1923), Mitta Mitta (1931), Upper Beaconsfield, Traralgon,

Daylesford, Bendoc (1933); Buchan (1960), Matlock (1961), Healesville, Yellingbo,

Powelltown (1963), Woori Yallock, Darlimurla (1966), Upper Thompson Valley

(1968), Marysville (1969), Porepunkah, Mount Buffalo and Upper Lerderderg Valley

(1970) ( Maloney and Harris 2006b).

Gould (1863) believed that its range was from “Port Phillip to Moreton Bay”. Krefft’s

(1864) assessment was that it occurred in the “mountainous coast districts of the

Australian continent”, from Victoria to Qld; also that it was “not found upon the plains of the interior”. Thomas (1888) and Lydekker (1896) reported that its range was from

Qld to Victoria. Fleay (1933) stated that the range “extends down the highlands of eastern Australia from southern Qld. to Victoria”, and that he had “never observed the species further west than the Ballarat-Daylesford forest” in Victoria. Marlow (1958) found that the western limits of its distribution in NSW were Barraba, Orange and

Tumut. Ride (1970) reported the distribution to be from the Dandenong Ranges

(Victoria) to Rockhampton, Qld.

In terms of abundance, the species has been described as the “most abundant of the arboreal marsupials in the forests to the east and north-east of Melbourne” (Lucas

1890), “very plentiful in the heavy eucalypt forests” of eastern Australia (Le Souef and

Burrell 1926); and “among the most numerous of arboreal marsupials” in East

Gippsland (Fleay 1933). Marlow (1958) reported that P. volans was “abundant” in

NSW (see also Calaby 1966; Flannery 1994; McKay 1995). Currently, P. volans is not listed as threatened in the three states that it occurs, and recent distribution maps are

39 provided by Eyre (2004) and Winter et al (2004) for Qld; Kavanagh (2004) for NSW and van der Ree (2004) for Victoria.

2.7 Behaviour

The gliding ability of P. volans was first reported by Phillip (1789) and then by Shaw

(1794, 1800), Cuvier (1798) and Turton (1806). Later authors remarked that it moves with a gliding motion, but this was not true flying (Desmarest 1817; Lesson 1827;

Owen 1841, 1845; Lydekker 1896). Le Souef and Burrell (1926) record a “flight by one of these animals from the top of one tall eucalypt to the base of another was 80 yards

[=73 m]; another flight, of 55 feet [=17 m], occupied 1 ½ seconds.” Troughton (1935,

1941) stated that it is “the record glider of the possum world” and reported that one individual was observed at Milton NSW, covering a distance of 590 yards [=540 m] in six successive glides. Two of these glides were 120 yards [=110 m], and one of 70 yards

[=64 m] from a tree 100 feet [=30 m] high. Wakefield (1970) stated “that some long glides, attributed in the literature to P. volans, belong in fact to Petaurus australis”. He discussed the report by Troughton (1935, 1941) and stated:

‘The 70 yard [=64 m] glide from a 100-foot [=30 m] tree indicates an angle

of descent of 26 degrees to the horizontal, and, even allowing for sloping

ground and a margin of error in the measurements, this performance, though

well within the capabilities of Petaurus, is quite outside that of Petauroides.

Also, for the 120-yard [=110 m] glides P. volans would require for its 40

degree descent, a take-off point approximately 300 feet [=90 m] high, while

Petaurus would need a 200-foot [=60 m] tree. Other features of the Milton

resident’s report – that during the performance the animal “lost no time in

ascending three more trees” and that “it uttered its peculiar squealing call” –

40 leave no doubt that the “record glider” was, in fact, Petaurus australis and

not Petauroides volans.’

The voice and gliding accomplishments of Petaurus australis have sometimes been credited erroneously to P. volans, which is in fact, a sedentary, slow-moving, silent animal of minor gliding ability (Wakefield 1970; McKay 1989). Many authors have mistakenly accredited P. volans with the vocalisations of P. australis, for instance

Lydekker (1896) was the first to erroneously report “when disturbed, or in flight, they utter a loud piercing scream, audible for a long distance” (see also Le Souef and Burrell

1926; Troughton 1935, 1941; Fleay 1933, 1947; Calaby 1966 for similar reports).

It was also recognised quite early that this species was nocturnally active and utilised tree hollows as den sites during the day (Oken 1816; Desmarest 1817; Lesson 1827;

Partington 1837; Waterhouse 1846; Thomas 1885; Collett 1887; Aflalo 1896). Gould

(1863) stated that “on the approach of evening [it] emerges from its retreat.” Lydekker

(1896) reported that they “spend the day in some hollow branch or the stem itself, whence they issue forth for their nocturnal flight”.

Le Souef and Burrell (1926) suggested that the only predators of P. volans are the powerful owl Ninox strenua and the introduced fox Vulpes vulpes; ‘the latter occasionally catches them on the ground’ (see also Fleay 1933, 1947, 1968). However,

Maloney and Harris (2006b) reported P. volans falling prey to a range of other predators, such as the cat (Felis catus), dog (Canis familiaris), fox (V. vulpes), wedge- tailed eagle (Aquila audax), quoll (Dasyurus maculatus) and Sooty owl (Tyto tenebricosa). Other recorded predators of the Greater Glider include the dingo (C. f.

41 dingo; Robertshaw and Harden 1985), lace monitor (Varanus varius; Weavers 1989) and carpet python (Morelia spilota; Lindenmayer 2002b).

Fleay (1933) reported:

‘Wandering under the trees on a still night, when the dusky gliders [P. volans]

are feeding overhead, rarely leads to their discovery without resort to intent

listening. Perhaps the faint sound of a leaf being pulled from a stalk, or a

sudden rustle as the animal plunges its weight from one slender limb to

another, betrays its position to a searching torch beam held so that the

observer’s eyes look straight along the path of light. Then the blazing orbs of

the animal, certainly the most brilliant light reflectors that I know of among

the marsupial family, regard the intruder with some curiosity’.

In terms of its reproduction and breeding behaviour, Desmarest (1817) reported

“females have a pouch under the belly, where the young spend the first part of their existence”. Fleay (1933) made the following observations on captive specimens “only two mammae are found in the pouch” and “only one embryo is reared at a time.” He also reported as follows:

‘In Vic. this minute naked creature seems to appear usually in July or August,

and it is difficult to realize that such a mite, no larger than the head of a

drawing-pin, may indulge some day in graceful aerial “flights”. Gradually as

the youngster increases in bulk, it is noted that the limbs and tail are

extraordinarily long, the loose volplaning membrane from fore limb to hind

limbs is plainly visible, and the colour of the furless embryo is pink with very

dark ears. The little fellow becomes free of its inseparable attachment to the

mamma when some six weeks of age. Later the eyes open and a covering of

short fur indicates plainly the contrast between the black and white of the

upper and lower surfaces respectively. It then spends the daylight hours out of

42 the pouch, and by night is carried around as a large bulge in it. At four months

it has become too bulky to be contained in the pouch any longer. Between the

growing of fur and the forsaking of the mother’s “pocket nursery” the young

Taguan Phalanger [P. volans] is one of the most curious and pathetic babes

that one can imagine with its lanky legs, very long tail and thin weekly body.

Having outgrown the pouch, though still being nourished from it, the little

phalanger clings to its mother’s back during her nocturnal wanderings, though

perhaps the gliding leaps are out of the question unless the youngster remains

in the home tree or sleeping hollow’.

2.8 Conclusion

Petauroides volans has had a long and sometimes confusing taxonomic history. It has been listed as a member of 10 genera (Belideus, Didelphis, Petaurista, Petaurides,

Petauroides, Petaurus, Phalanger, Phalangista, Schinobates, and Voluccella) and there have been at least 23 different binomial names used for it since its discovery. This geographically widespread species was sent to different museums throughout Europe by collectors, and given different designations by 19th Century zoologists. These early zoologists were often rivals, each of whom was more anxious to discover and name species, than to find out the habits of the species already known (Partington 1937).

Consequently errors were made, and some of these have persisted into the modern literature. For example, Flannery (1994) mistakenly listed Hepoona Roo (= Petaurus australis) as synonymous with P. volans.

Early descriptions of the morphology of P. volans such as its colouration, size and the presence of a gliding membrane, are given in broad general terms but nevertheless they do have value from a historical viewpoint. Dental descriptions in the early literature

43 vary, and some confusion with the similarly structured dentition of Pseudocheirus peregrinus is evident. Early behavioural observations include the ability to glide, and that it is nocturnally active using tree hollows as den sites. The earliest records of occurrence were centered about the Sydney district. As the colony expanded so did its recorded range. Gould (1863) reported that its distribution was from Port Phillip

(Victoria) to Moreton Bay (Qld), and this is reasonably accurate when compared to our understanding of its current range.

44

Chapter 3

Review of the past knowledge of the distribution and abundance of the Greater

Glider in the study area, and threats to populations

3.1 Introduction

This chapter includes a review of the more recent and current knowledge of the distribution of the Greater Glider in the study region, and the current threats to populations. Information concerning the distribution of a species is available from sources such as naturalists, government agencies and museum collections. The last section of this chapter looks at the barriers to the recolonisatsion of Royal National Park by the Greater Glider.

3.2 Recent records (1908-1980)

Recent records for the Greater Glider in Royal National Park and surrounding areas were obtained by consulting available sources of information: including the records of

Norm Robinson, an Illawarra Naturalist (1945-1970), the Australian Museum (AM)

(1908-1980) and the CSIRO wildlife collection.

3.2.1 Records of Norm Robinson

The naturalist Norm Robinson (1977, 1988a, and 1988b) undertook a detailed account of records of mammals in the Illawarra. He conducted trapping, spotlighting and also interviewed people with a long history in the area. The earliest date for records of the 45 Greater Glider, based on interviews, was from the Coledale and Helensburgh areas in

1945. Other records provided by the interviews include Darkes Forest (1967) and

Coledale to Scarborough (1969) (Table 3.1). The records provided by the surveys of

Robinson (1977, 1988a, and 1988b), indicate the Greater Glider was present in low numbers in areas of the Sydney Catchment Authority (Fire road 6B and Fire road 8).

Compared to the observations at Lilyvale (4) and Lady Carrington Drive (8). Both of these sites are located along the Hacking River Catchment, one within Royal National

Park and the other is in close proximity.

Table 3.1 Records of the Greater Glider from Norm Robinson

Location Date Numbers / observer

Coledale 1945 Unknown / L. Carrick

Helensburgh 1945 Unknown / W. Clark

1st hill north of Mt. Ousley 1966 Road victim / N. Robinson

Darkes Forest 1967 Unknown / F. Short

Fire road 6B 1968 1 / N. Robinson

Fire road 8 1969 1 / N. Robinson

Coledale to Scarborough 1969 Unknown / A. Cheatham

Lilyvale 1970 4 / N. Robinson

Lady Carrington Drive 1970 8 / N. Robinson

The records of Norm Robinson shown in Figure 3.1 are from the surveys conducted between 1966 and 1970 (Robinson 1977, 1988a, and 1988b). Robinson (1977) had suggested that a wildlife corridor was required to link the Illawarra escarpment park and water catchments to areas north such as Royal National Park. Robinson (1977) reported

46

Figure 3.1. The recent distributional records of the Greater glider in the study area from Robinson (1977, 1988a, 1988b). The blue diamonds are indicative of records for the Greater glider during this time. The blue arrows indicate records from, (top) Royal National Park, (middle) Lilyvale area, (bottom) Thirroul/Coalcliff area (see table 3.1). The green arrow indicates the corridor position suggested by Robinson (1977).

47 that a number of mammal species inhabiting reserves within the Illawarra were declining, due to altering patterns of land usage and wide road construction. The Greater

Glider was one of the species that would benefit by the inclusion of a corridor at this site, facilitating movement of the species between areas (Robinson 1977).

3.2.1.1 Summary

The records of the Greater Glider provided by Robinson (1977, 1988a, and 1988b), indicate that this species was present at Royal National Park and Lilyvale in 1970. It was found above the Illawarra escarpment at Mt. Ousley, Darkes Forest and fire roads 6 and 8 between 1966 to 1969, and below the escarpment between Coalcliff and

Scarborough in 1970. Earlier records indicate that this species was present at Coaldale and Helensburgh in 1945.

3.2.2 Specimens held in institutions (AM and CSIRO)

A total of 23 records of the Greater Glider were provided by the Australian Museum

(AM) and the CSIRO wildlife collection for the study area (Table 3.2). The Australian

Museum provided 22 records held in their collection, two specimens were from

Sutherland and Helensburgh collected in the early 1900s. Of the other 20 records in the

AM collection, ten were collected in 1978 between February and April, the other ten were collected in February 1980, all 20 from the same locality, Lady Carrington Drive in Royal National Park. The CSIRO provided one record from this last location, also collected in 1978. Which included fourteen females (weight range 1200-1650 g) and six males (weight range 725-1400g), two of these males were juveniles with weights of

725g and 800g respectively (Table 3.2).

48

Table 3.2. Specimens of Petauroides volans held in the Australian Museum (AM) and the CSIRO from the study area.

Source/Reg Date Location Co-ord Collector Sex weight Form No collected AM-M2003 13-08-1908 Sutherland 340200S- Zool Soc n/a n/a Skin & skull 1510400E NSW AM-M2051 28-07-1909 Helensburgh 341100S- Frank n/a n/a Skin & skull 1505900E Farnell AM-M29518 15-02-1978 Royal NP 340800S- G. McKay F 1600g Skull (1) 1510400E AM-M29527 15-02-1978 Royal NP 340800S- G. McKay F 1600g Skull & Skin in 1510400E spirit AM-M29524 28-03-1978 Royal NP 340800S- G. McKay F 1450g Skull & Skin in 1510400E spirit AM-M29526 28-03-1978 Royal NP 340800S- G. McKay M 1270g Skull & Skin in 1510400E spirit AM-M29440 14-04-1978 Royal NP 340800S- G. McKay F 1420g Skull & Skin in 1510400E (2) spirit AM-M29520 17-04-1978 Royal NP 340800S- G. McKay F 1280g Skull & Skin in 1510400E spirit AM-M29521 17-04-1978 Royal NP 340800S- G. McKay F 1650g Skull & Skin in 1510400E (2) spirit AM-M29522 24-04-1978 Royal NP 340800S- G. Mckay F 1400g Skull & Skin in 1510400E spirit AM-M29523 24-04-1978 Royal NP 340800S- G. McKay M 1370g Skull & Skin in 1510400E spirit AM-M35345 24-04-1978 From S end 340400S- G. McKay F 1200g Skull of (1) - RNP 1510300E CSIRO- 24-04-1978 6km from 340400S- n/a n/a n/a n/a M16188 Audley (1) 1510400E AM-M29460 6-02-1980 Royal NP 340800S- Leon F 1400g Skull & Skin in 1510400E McQuade spirit AM-M29461 6-02-1980 Royal NP (1) 340800S- Leon M 1350g Skull 1510400E McQuade AM-M29464 7-02-1980 Royal NP (1) 340800S- Leon F 1400g Skull 1510400E McQuade AM-M29481 7-02-1980 Royal NP (1) 340800S- Leon F 1550g Skull & Skin in 1510400E McQuade spirit AM-M29474 12-02-1980 Royal NP 340800S- Leon M 725g Skull 1510400E McQuade AM-M29470 12-02-1980 Royal NP 340800S- Leon M 800g Skull 1510400E McQuade AM-M29516 13-02-1980 Royal NP (1) 340800S- Leon M 1400g Skull 1510400E McQuade AM-M29462 14-02-1980 Royal NP (1) 340800S- Leon F 1500g Skull 1510400E McQuade AM-M29469 14-02-1980 Royal NP (1) 340800S- Leon F 1350g Skull 1510400E McQuade AM-M29479 14-02-1980 Royal NP (1) 340800S- Leon F 1350g Skull 1510400E McQuade

49 Over a two-year period (1978-1980); these specimens were removed from the Royal

National Park population by shooting (L. McQuade pers. comm. L. McQuade collected the specimens from Royal National Park as a post graduate student at Macquarie

University). There are several consequences with collecting gliders from a population with this technique. The difficulty in distinguishing between the sexes with a spotlight, as this species resides high in the canopy, leads to the possibility that the remaining gliders may all be male. As a result, no new gliders would be born into the population, and the population would die out naturally. The impact of predation could eliminate a population with low numbers, for example Powerful owls (Ninox strenua) have been reported to reduce a population of Greater gliders by 90% (Kavanagh 1988).

3.2.2.1 Summary

The records provided by the Australian Museum and the CSIRO indicate that a large sized population (>21) of Greater Gliders was present in Royal National Park between

1978 and 1980, and it was significantly reduced by shooting to collect specimens.

Urbanisation has claimed the habitat for this species in the Helensburgh and Sutherland area (100 odd years since collection).

3.3 More Recent Records (1980-2002)

The recent records of the Greater Glider were obtained from the Sydney Catchment

Authority and NSW National Parks and Wildlife Service, including surveys undertaken by National Parks and Wildlife Service for the Illawarra escarpment (NSW NPWS

2002).

50 3.3.1 Illawarra Escarpment Survey

The NSW state government, in response to the 1999 Commission of Inquiry into planning and management of the Illawarra escarpment, conducted a bioregional assessment (NSW NPWS 2002). This assessment was based on a survey of the fauna of the Illawarra region sampling many different habitat types. A set of priority species was selected from those listed in state or federal conservation legislation, or via nomination by local fauna experts. The Greater Glider is not listed in state or federal conservation legislation, but it is nevertheless protected under the National Parks and Wildlife Act,

1974, and was included as a priority species in the assessment. The reasons for including the Greater Glider were that it had a restricted, declining population in the region and that core habitat was present in the area (NPWS 2002).

The assessment produced a total of 22 Greater Glider records from the tall escarpment forests on the plateau. These included records from 1990 for the Illawarra area and

Royal National Park (see Figure 3.2). No records of the Greater Glider were obtained in the forests in the northern part of the district north of Thirroul below the escarpment, where the species had been found previously (Robinson 1988a). A habitat model provided by the NPWS, indicated that abundance should increase with eucalypt height

(NPWS 2002 p.191). The habitat area for the Greater Glider was mapped from this model (Figure 3.2), and it includes the area below the escarpment from Bulli north towards Royal National Park (two records from RNP were used in the data). The

Illawarra region has a large area of suitable habitat (10,384 ha) for the Greater Glider, the majority of this habitat (63.7%) is conserved in the Sydney Catchment Authority lands (NPWS 2002).

51

Figure 3.2. Records and habitat model (red shading) of the Greater Glider provided by NPWS (2002) for the commission of enquiry into the Illawarra escarpment.

52 3.3.1.1 Summary

The records of the Greater Glider provided by the bioregional assessment of the

Illawarra Escarpment indicate that this species is no longer found below the escarpment north from Bulli, but is found in suitable habitat within Sydney Catchment Authority lands above the escarpment.

3.3.2 Records from the NPWS fauna database.

The distributional records provided by the NSW NPWS for the Greater Glider in the

Illawarra region (Wollongong Local Government Area) are restricted to areas above and west of the escarpment, where the tall eucalypt forests are found (Figure 3.3). The 35 records from 1990 till 2006 are from areas in and adjacent to Sydney Catchment

Authority lands, where the habitat for this species is protected (NPWS 2002). Incoll et al. (2001) found similar results for the Greater Glider in the Melbourne Water

Catchments. There are no recent records (>1990) from north of the Bulli area, where this species was recorded thirty five years ago (Robinson 1988a).

3.3.2.1 Summary

The records provided by the NSW NPWS data base indicate that the Greater Glider is found above the escarpment, but not below the escarpment.

53

Figure 3.3. Distributional records of the Greater Glider in the Illawarra, from the NSW NPWS Fauna Atlas.

3.4 Threats to populations of the Greater Glider

3.4.1 Introduction

A range of factors threaten the survival of the Greater Glider in the forested ecosystems of eastern Australia (Lindenmayer 2002a). These include habitat loss and land clearing; through the processes of logging, plantation establishment, road construction and urbanisation. Threatening processes do not act alone and species such as the Greater

Glider must contend with the interaction of the above factors. The following section assesses the effects of these threatening processes at the habitat level for the Greater

Glider.

54 3.4.2 Removal of den trees

The Greater Glider is dependent on tree hollows for its denning requirements (Gibbons and Lindenmayer 1997). The habitat of the Greater Glider in areas of its range coincides with high production forest which is sought after by wood production companies

(Lindenmayer 2002a). The removal of den trees from the habitat of the Greater Glider would have a negative effect on the survivorship and abundance of a population.

Kavanagh et al. (1995) reported that the Greater Glider can survive in areas that are selectively logged, whereas it is vulnerable to intense logging (Lunney 1987, Kavanagh

2000). Some studies have reported that the distribution of the Greater Glider is closely related to hollow availability (Milledge et al. 1991). This species utilizes many dens in its home range, individual Greater Gliders have been observed using from four to eighteen dens (Kehl and Borsboom 1984). Andrew (2001) reported that after the 1994 fires at Royal National Park, up to 120 large hollow bearing trees were felled along

Lady Carrington Drive by NPWS for reasons of public safety. These trees would have contained many suitable hollows for this species.

The reduction in the availability of tree hollows, through land-management practices, logging and wildfire, threatens the viability of populations of hollow-dependent fauna

(Gibbons and Lindenmayer 1997).

3.4.3 Predators

A literature search provided a list of ten predators of the Greater Glider (see Table 3.3 for list and sources). Three of these, the Powerful owl (Ninox strenua), Sooty owl (Tyto

55 tenebricosa) and Spotted-tailed quoll (Dasyurus maculatus), are listed as threatened under NSW conservation legislation. Two, the Lace monitor (Varanus varius) and

Carpet python (Morelia spilota) are reptiles. The Wedge-tailed eagle (Aquila audax), is a diurnal predator and the Dingo (Canis familiaris dingo) is a predator of mainly terrestrial species. The Cat (Felis catus), Dog (Canis familiaris) and Fox (Vulpes vulpes) have been introduced into Australian ecosystems for various reasons.

Table 3.3. The ten predators of the Greater Glider obtained from a literature review.

Scientific Name Common Name Source Ninox strenua Powerful Owl 1,2,3,4,5,6,7,8,9,10,11,12,13,14,15,16,17 Tyto tenebricosa Sooty Owl 14,15,17 Canis familiaris Dog 17,18,19,20 Vulpes vulpes Fox 1,2,17,18,19,20,21,22, 23, 24 Felis catus Cat 17,25 Varanus varius Lace Monitor 26, 27 Dasyurus maculatus Spotted-tailed Quoll 17,28,29 Canis familiaris dingo Dingo 30 Aquila audax Wedge-tailed Eagle 17,31 Morelia spilota Carpet Snake 27

Sources 1. Le Souef and Burrell (1926), 2. Fleay (1947), 3. Fleay (1968), 4. James (1980), 5. Kavanagh (1988), 6. Pavey (1992), 7. Chafer (1992), 8. Kavanagh (1992), 9. Pavey (1994), 10. Pavey et al (1994), 11. Lavazanian et al (1994), 12. Pavey (1995), 13. Schultz (1997), 14. Kavanagh (2004), 15. Bilney et al (2006), 16. Cooke et al (2006), 17. Maloney and Harris (2006b), 18. Lunney et al (1990), 19. McFarlane (1988), 20. Mitchell and Banks (2005), 21. Wallis and Brunner (1986), 22. Coman (1973), 23. Brunner et al (1975), 24. Roberts et al (2006), 25.Coman and Brunner (1972), 26. Weavers (1989), 27. Lindenmayer (2002). 28. Glen and Dickman (2006), 29. Belcher and Darrant (2006), 30. Robertshaw and Harden (1985), 31. Tyndale-Biscoe and Smith (1969b).

3.4.3.1 Powerful Owl

The large number of sources (17, see Table 3.3) over an 80 year timeframe, which reported the powerful owl as a predator of the Greater Glider, indicates that the Greater

Glider is an important prey source for this species, across the distribution of the two species. Kavanagh (1988) reported that a pair of powerful owls had depleted a population of Greater Gliders, by 90 % over a 46 month period. The Greater Glider was

56 the preferred prey for a pair of Powerful owls in an unlogged and undisturbed area of forest in south-eastern NSW (Kavanagh 1988, see also Pavey 1992 for comment and

Kavanagh 1992 for reply).

3.4.3.2 Sooty Owl

In the Sydney region, Kavanagh (2004) calculated that the Greater Glider comprised

10% of the prey biomass for the Sooty owl, based on seven owl territories. Since

European settlement in east Gippsland, Victoria, the number of terrestrial mammals in the diet of the Sooty owl has decreased. To compensate, the Sooty owl has increased the consumption of arboreal prey from 55% to 81% of their diet, including the Greater

Glider (Bilney et al. 2006). These authors attributed this change to the predation on terrestrial mammals by introduced predators, such as the fox.

3.4.3.3 Spotted-tailed Quoll

The Greater Glider has been reported as making up 35% of the diet of the spotted-tailed quoll in northern NSW (Glen and Dickman 2006). In Victoria, cave deposits of bones of the Greater Glider of Holocene-Late Pleistocene age have been attributed to predation by quolls (Wakeford 1960). The Spotted-tailed quoll has been observed removing

Greater Gliders from their dens (Belcher and Darrant 2006). This information suggests that Greater Gliders have been a prey item of the quoll for a very long time and, as such, the Greater Glider should be recognized as a priority species for conservation management, as it is a preferred prey for a threatened species (Glen and Dickman 2006).

57 3.4.3.4 Foxes

The fox has been reported as a predator of the Greater Glider for 80 years in Australia

(see Table 3.2 for sources). Many reports of Greater Gliders in the diet of foxes have come from the analysis of scats during biodiversity surveys (see table 3.2 for sources).

It was presumed that foxes scavenged on the remains of Greater Gliders found on the ground, left by powerful owls (Kavanagh 1988, Lindenmayer 2002a) or when they walked along the ground (Fleay 1933, 1947, 1968). Foxes are very capable climbers

(personal observation) and Le souef and Burrell (1926) observed them in daylight hours with Greater Gliders in their mouths. When startled dropping the victim, which was very much alive. This report suggests that foxes remove Greater Gliders from their dens.

3.4.2.5 Dog, Cat and Dingo

The Dog, Cat and Dingo are all opportunistic predators and it is highly unlikely that these species would have removed a Greater Glider from its den through climbing

(Lindenmayer 2002a). More than likely, they would have scavenged on the dropped remains of Greater Gliders. Powerful owls have been observed snipping the tails and heads from Greater Gliders and letting them drop to the ground, where these three predators would have likely consumed the carrion (Kavanagh 1988, Lindenmayer

2002a).

3.4.3.6 Lace Monitors and Carpet Pythons

Lace monitors and Carpet pythons are opportunistic predators and good climbers, both species have been reported preying on Greater Gliders if they occur in their range

(Weavers 1989, Lindenmayer 2002a).

58

3.4.3.7 Wedge-tailed Eagles

Wedge-tailed eagles are diurnal predators and would only be able to prey on Greater

Gliders in the day time. The report of predation by a Wedge-tailed eagle on the Greater

Glider by Tyndale-Biscoe and Smith (1969b) was observed as a forest was logged and they glided to the ground, becoming active during the day as they searched for new shelter sites (see also Maloney and Harris 2006b).

3.4.4 Summary

Eight of the ten species which have been reported here as being predators of the Greater

Glider have been recorded in Royal National Park. These are the Wedge-tailed eagle,

Powerful owl, Sooty owl, Lace monitor, Carpet snake (Diamond python), Dog, Fox and

Cat (Andrew 2001). The Spotted-tailed quoll is thought to be locally extinct and of uncertain status within Royal National Park (Andrew 2001).

3.4.4 Fire

The NSW Threatened Species Conservation Act 1995 lists high frequency fire as a key threatening process in Schedule 3 of the Act. ‘High frequency fire resulting in the disruption of life cycle processes in plants and animals and loss of vegetation structure and composition’. The loss of vegetation after a fire event has been reported to cause the death of the folivorous Greater Glider (Fleay 1947). Greater Gliders have been observed dying as the flames consume those which attempt to escape the fire (Fleay 1947). They are able to survive fire events in unburnt refuges (Lunney 1987), and can recolonise burnt areas when the vegetation recovers (Mcfarlane 1988).

59

High intensity wildfires, burn over a broad area and burn in conditions that leave few unburnt areas/patches. The adverse impacts of such fires include:-

• Forest habitat burnt from ground to canopy;

• Widespread removal of food resources (post fire recovery);

• Large numbers of senescent, hollow bearing habitat trees are removed;

• Improved access by introduced predators;

• Loss of vegetation in gullies.

There have been a number of fire events in the study area, which have burnt extensive areas above and below the Illawarra escarpment. They occurred in September 1939,

October 1968, November 1980, January 1994, December 1997, December 2002.

3.5.5 Habitat Removal

The removal of habitat is a major threat for the survival of the Greater Glider. Surveys conducted which compared the species in logged and unlogged forest, reported they were rare to absent in the logged areas (Lunney 1987, Kavanagh et al. 1995, Goldingay and Daly 1997, McCarthy and Lindenmayer 1999a). The establishment of pine plantations has had a negative effect on populations of the Greater Glider (Lindenmayer

2002a). The clearance of native forests is the cause of stress and death among individual

Greater Gliders (Tynedale-Biscoe and Smith 1969b) and isolates surviving populations

(Lindenmayer 2002b). There has not been any research conducted on the effects of native hardwood plantations on glider populations, but they would be expected to have the same effects as the pine plantations (Lindenmayer 2002a).

60 The use of barbed wire fencing around native forests to keep stock from entering has been reported to be lethal for Greater Gliders (Fleay 1933, 1947, Maloney and Harris

2006b). Fleay (1933) reported that the Greater Gliders catch their membrane on the fence and suffer a long and agonizing death (see also Lindenmayer 2002a).

3.5 Barriers to recolonising Royal National Park

The three areas from which the movement of Greater Gliders would be able to recolonise Royal National Park are shown in Figure 3.4. The top arrow shows the movement from Heathcote National Park, the middle from Garrawarra State Recreation

Area and the bottom from the Illawarra Escapment. The barriers to movement from these sites are as follows.

(1). Heathcote National Park

i) The Princess highway, which has four lanes of traffic;

ii) The urban fringe where vegetation is managed for fire retardation;

iii) The absence of Greater Gliders from the area due to no core habitat.

(2). Garrawarra state recreation area

i) The old Princes highway;

ii) The F6 expressway;

iii) No habitat for dispersal routes.

(3). Illawarra Escarpment

i) Urban development along the escarpment at Stanwell Park, Stanwell Tops

and Helensburgh;

61 ii) Lack of corridors connecting the escarpment park with contiguous core

habitat;

iii) Roads connecting southern suburbs with Royal National Park and

Helensburgh.

Powerline corridors also represent barriers, because they are maintained with vegetation management strategies, thus making these cleared strips unsuitable for volplaning across a surrounding vegetation. The barriers that stop the Greater Glider from recolonising Royal National Park are mostly anthropogenic.

Figure 3.4. Satellite image of the northern half of study area. Circle is the known former habitat of the Greater Glider in Royal National Park, and arrows indicate the possible movement of Greater Gliders to recolonise RNP. (Top = Heathcote, Middle = Garrawarra, Bottom = Illawarra escarpment). (Source Google Earth).

62 Chapter 4

Detailed Field Study

4.1 Introduction

The Greater Glider is considered to be one of the most abundant arboreal marsupial species found in the forested lands of the eastern coast of Australia (Kavanagh 2004).

This species was readily seen along Lady Carrington Drive in the Royal National Park, south of Sydney prior to a wildfire in 1994. Fauna surveys conducted after the 1994 fire event in Royal National Park suggested that the Greater Glider had disappeared from this former stronghold (Andrew 2001, fire and not shooting for specimen collection was considered in Andrew 2001 assessment). This chapter reports on a detailed field study undertaken on the presence and absence of the Greater Glider in Royal National Park and the surrounding Illawarra district.

4.2 Aim

The aims of the detailed field study were; (i) to establish the presence or absence of

Greater Gliders at Royal National Park; (ii) to determine the location of the nearest population to Royal National Park for the possibility of translocation of this species, and

(iii) to test the suitability of a habitat model provided by the NSW NPWS (Figure 3.2).

63 4.3 Methods

4.3.1 Site selection

To ascertain whether the Greater Glider is present or absent in Royal National Park and the surrounding areas, a total of nineteen (19) sites were chosen. The criteria for a chosen site were (i) that the Greater Glider had been recorded previously from the site, and (ii) that the site was located within the habitat model indicated by the NSW NPWS

(Figure 3.2). The names and number of each of the nineteen (19) chosen sites are listed in Table 4.1; the locations of these sites are shown in Figure 4.1.

Table 4.1. Sites used in this study for spotlight presence or absence of the Greater Glider. # = Sydney Catchment Authority Lands; * = Cordeaux area; ● = Cataract area; ■ = Loddon area

Site Number Name of Site 1 Lady Carrington Drive RNP 2 Lady Wakehurst drive RNP 3 Helensburgh/Stanwell Tops 4 Coalcliff 5 Coledale 6 Thirroul 7 Bulli Pass 8# Fire road 7I ■ 9# Fire road 7K ● 10# Fire road 7B ● 11# Fire road 7 ● 12# Fire road 7D ● 13# Fire road 7C ● 14# Fire road 8 ■ 15 Mt. Keira 16# Cordeaux road * 17# Morans Road * 18# Fire road 6E * 19 Mt. Kembla

64

1

2

3

4

5

8 6 10 9

11 7 12 14 13

12

16 15 18 17

19

Figure 4.1. Location of the 19 sites surveyed in the detailed field study (see table 4.1).

65

4.3.2 Detection of Greater gliders

Spotlighting was chosen as the method to determine the presence or absence of the

Greater Glider at a site. The Greater Glider is a readily detectable species from spotlighting (Lindenmayer et al. 2001); the eyeshine from this species is a white reflected light, and its colouration and size is easily discernable from other species.

Unlike Petaurus spp. the Greater glider is silent and cannot be detected by vocalizations

(Wakeford 1970). Wintle (2003) reported that negative observations resulting from periodic absences from a given site within its home-range are likely to be quite low for this species, due to its ease of detection.

Spotlighting was undertaken along a 1 km transect at each site. Between 2003 and 2006

I surveyed each site on 12 occasions. The average duration of time undertaken per transect was 1.5 hours. Spotlighting was undertaken at a walking pace with Oldham mining cap lamps (4 volt battery and 6 volt quartz globe). Searches were confined to relatively clear nights to avoid extreme weather conditions, which restrict observation of the Greater Glider. The survey was undertaken on both sides of the transect. The presence or absence of the Greater Glider at a site was recorded during each visit.

66 4.4 Results

4.4.1 Greater Glider absence

The Greater Glider was not recorded from eight of the 19 sites (1-7 and 15) over a four year period during this study (Table 4.2). Sites 1-3 are located within or close to Royal

National Park. Sites 4-7 north of Bulli, are located below the Illawarra escarpment, and site 15 was located at Mt. Keira. Greater Gliders had been recorded previously at or near these sites (Robinson 1988a, Andrew 2001). The availability of hollows and tall trees did not appear to be a limiting factor at these sites.

4.4.2 Greater glider presence

Greater Gliders were recorded from 11 of the sites (8-14 and 16-19) (Table 4.2), which were above and west of the escarpment, in or close proximity to SCA protected lands.

Few Greater Gliders were observed at these sites (Table 4.2). The maximum number observed at any one time was three gliders in a 1 km transect at site 17, Morans Road.

The forests in which the Greater Glider was observed were tall and old growth, and many hollows were present. On one occasion, at site 17 (Morans Road), a white phase glider was observed exiting an old senesced stump at a height of 10 metres. At site 16,

Cordeaux Road, the gliders were recorded on nine of twelve visits (Table 4.2).

67 Table 4.2. The presence (numbers observed)) or absence (blank) of the Greater glider from the chosen 19 sites (Table 4.1 and Figure 4.1) for this study.

Site 2003 2003 2003 2004 2004 2004 2005 2005 2005 2006 2006 2006 Total 1 2 3 4 5 6 7 8 1 2 1 2 1 1 2 1 1 1 1 1 15 9 2 1 1 2 1 1 2 1 1 1 1 1 15 10 1 2 1 1 2 1 2 1 2 1 1 2 17 11 1 1 1 1 1 2 2 1 1 1 1 1 14 12 2 2 1 1 2 1 1 1 1 1 1 1 15 13 1 1 1 1 1 2 1 1 1 1 1 1 13 14 1 1 1 1 1 1 1 1 1 1 2 2 14 15 16 1 2 1 1 2 1 1 1 1 11 17 3 2 3 2 1 2 2 3 1 2 1 2 24 18 2 1 2 1 1 2 1 1 2 1 1 1 17 19 1 2 2 1 1 1 1 1 1 1 2 1 15 Total 16 17 15 14 15 15 16 13 13 11 12 13

4.5 Conclusion

The Greater Glider was present at sites above the escarpment. These areas are conserved

within the Sydney Catchment Authority lands. They are present in low numbers at these

sites (8-14 and 16-19). The Greater glider was absent from sites (1-7) from Bulli to

Royal National Park, which were below the escarpment. It was also absent from site 15

at Mt. Keira.

Lindenmayer et al. (2001) reported that spotlighting underestimates the actual

population size of the Greater Glider. This study was undertaken to determine the

presence or absence of the Greater Glider at nineteen (19) chosen sites. Wintle (2003)

reported that approximately five (5) visits to a site were required to be 90% sure that

68 this species was truly absent. The amount of survey effort undertaken for this study would have maximized the likelihood of detection, if the Greater Glider was present at a site. This study indicates that the Greater Glider is absent from sites where it was previously present (Robinson 1988a). The habitat model (Figure 3.2) was true for areas above the escarpment and not true for areas below the escarpment north of Bulli to

Royal National Park.

The Greater Glider has been reported to disperse distances of 1 to 7km, through radiata pine (Pinus radiata) plantations surrounding remnant eucalypt patches at Tumut NSW,

(Pope et al. 2004, Taylor et al. 2007). Cunningham et al. (2004) suggested that these distances were achieved by utilising available food resources in the P. radiata plantations. The nearest population to Royal National Park was observed at site 8 (Fire road 7I) approximately 15km away. This is a much larger distance to disperse than has been reported for the species, as well as the many barriers and habitat discontinuity they would face during dispersal.

The Greater Glider is indeed absent from Royal National Park, as reported in Chapter 3 there have been many processes which have attributed to this absence. In the study area the Greater Glider is present in low numbers, and further population decline is possible from wildfires and predation. To recolonise Royal National Park, Greater Gliders would have to disperse a very large distance and overcome many barriers. The preceding factors all contribute to an unlikely event that the Greater Glider in the study area will have the ability to recolonise Royal National Park. This information strongly suggests that Royal National Park would make a good site for a translocation program.

69 Chapter 5

Translocation

5.1 Introduction

This chapter integrates the information presented in the previous chapters into a proposal for translocation, based on the requirement for a reintroduction set by the

Department of the Environment and Climate Change (DECC) (NPWS NSW 2001). The previous chapters;

(i) clarified the taxonomy of the species and interpreted historical accounts

(Chapter 2),

(ii) reviewed the current knowledge of the distribution and abundance of Greater

Gliders in the study region, and current threats to populations (Chapter 3),

(iii) described the results of detailed field studies in the region conducted to

determine the current location of populations (Chapter 4).

Translocation refers to the movement of living organisms from one area with free release to another (NPWS NSW 2001). The objectives of a translocation are to conserve organisms in areas where they have declined or disappeared (IUCN 1998). The types of translocation include introduction, reintroduction and supplementation (NPWS NSW

2001). Reintroduction is defined as

“an attempt to establish a species in an area which was once part of its historical range, but from which it has become extirpated or become extinct”

70 Serena and Williams (1994) argued that an ideal candidate species for reintroduction is one that is known to have declined in the wild, but is not yet threatened. The information gained from such a reintroduction may be used to reverse the decline of that species and assist more endangered species (Serena and Williams 1994). The Greater

Glider satisfies these criteria; it is not a threatened species and has been missing from

Royal National Park since 1994 (around 13 years) (Andrew 2001), thus making it a worthwhile candidate species for reintroduction into a former stronghold.

While the Greater Glider is not listed as a threatened species in New South Wales conservation legislation, a population in the Eurobodalla Local Government Area has been determined as endangered. The objective of a translocation of the Greater Glider to

Australia’s oldest National Park would be an iconic reintroduction to re-establish a recently extinct population. There were many processes outlined previously (see

Chapter 3) that were possibly involved in the decline of this species at Royal National

Park. I consider that the most significant was the removal of 21 animals by shooting between 1978 and 1980. The remaining animals survived till the 1994 major fire event

(Andrew 2001), which was the likely ultimate demise of this population.

For translocation to achieve a successful reintroduction, information concerning the background biology and ecology of the species must be considered (Lindenmayer

1994). This information is to help sustain a free-ranging, self-sustaining viable population in an area of suitable habitat, with minimum management input

(Lindenmayer 1994). The aim of this chapter is to develop a translocation proposal for the Greater Glider in order to justify and guide a reintroduction into Royal National

Park.

71 The following sections of this chapter outline the factors considered for the background biology and ecology of the Greater Glider as outlined in the NPWS NSW guidelines for a translocation proposal (NPWS NSW 2001).

5.2 Host site population parameters

5.2.1 Sex ratio

The sex ratio of the Greater Glider has been reported to vary from 1:1 to 1:4, depending on the patch size utilised (Pope et al. 2004). Norton (1988) suggested that the nutrient quality of the habitat was a determining factor for the sex ratio of a site. Nevertheless, the ratio of males to females removed from Royal national Park from 1978 to 1980 was

1:2.3 (6 males and 14 females). As the actual sex ratio for this site is unknown, it is deemed most appropriate to use a sex ratio of 1:2, which is within the range reported by

Pope et al. (2004).

5.2.2 Number of individuals

To achieve a sex ratio of 1:2, the total number of individuals to be translocated would be 18 (6 males and 12 females). This number of individuals is based on Pope et al.

(2004); who reported that after a 35 year period a fragmented patch of habitat with an area of 6 ha supported a population of 10 Greater Gliders. A greater number of individuals would involve more resources for capture, transport and maintenance. It is proposed to utilise two sites along Lady Carrington Drive (Bola Creek and Jersey

Springs approximately 4 km apart see Figure 5.1). The number of individuals would be

72 the same at both sites (3 males and 6 females). This would enable the comparison of success of translocation at two sites, expansion from each site and the possibility of dispersal between sites.

Figure 5.1. The proposed translocation sites for the Greater Glider reintroduction to Royal National Park along Lady Carrington Drive (yellow line). Blue arrow = Bola Creek and Green arrow = Jersey Springs. (Scale 1 cm = 500 m) (Source CMA map Royal National Park, photo of section by S. Maloney)

73

5.2.3 Spacing

The Greater Glider has been reported to be evenly spaced in the forest they inhabit

(Tynedale-Biscoe and Smith 1969). Males overlap the home ranges of females, and have either a monogamous, polygamous or polygynous mating system depending on the density of the population (Norton 1988, Lindenmayer 2002a). The spacing of individuals in a translocation program can also influence the mating system of individuals. The spacing to be utilised would be dependent on the release of individuals.

At the two sites a 400 meter length of Lady Carrington Drive would be utilised. The set up would be F-M-F every 100 metres giving the male access to two females 50 meters either side of the midway point (3 lots at each site).

F-M-F – 50 m space - F-M-F – 50 m space - F-M-F over the 400 meter area.

5.2.4 Monitoring

The monitoring of translocated individuals is an important part of the program, as the success of the program and the fates of individuals can be ascertained (Norton 1994).

The collaring of individual Greater Gliders for radio tracking has been undertaken successfully for home range studies (Henry 1984, Kehl & Borsboom 1984, Comport et al 1996, Kavanagh and Wheeler 2004 and Pope et al, 2004). The same gliders had reflective tape and ear tags attached to identify individuals if and when the batteries failed. The monitoring of individuals will assess changes in population size at the host site, and use of resources (i.e. den use, breeding success). Monitoring of the source population will determine any detrimental effects on these populations, if required.

74 5.2.5 Source population

The source population for a translocation of the Greater Glider to Royal National Park as has been detailed in Chapter 4 should not come from the nearby populations of this species in the Illawarra region, because of the low number of individuals in fire prone habitat, these populations have been reported to be of high local conservation priority

(DECC 2007b). The possible sources are from State Forest areas that have been targeted to be cleared for wood production or replaced by plantations. Individuals from a source of this type would not have any detrimental effects on the targeted population, and eliminate monitoring of the source population.

5.3 Taxonomy of the species.

The Greater Glider, Petauroides volans (Marsupialia: Pseudocheiridae), is the largest gliding marsupial and is endemic to eastern mainland Australia (McKay 1995).

Currently, there are two recognised sub-species: P. volans volans, which occurs in south eastern Australia (from Victoria in the south, through mainly coastal New South Wales

(NSW) to the Rockhampton district in north-east Queensland (Qld)), and P. v. minor, which occurs in very far north-east Qld (from the Dawson River to the Barron River)

(Flannery 1994). It is around the size of a domestic cat, with females being larger than males (Flannery 1994; Kavanagh and Wheeler 2004). Most individuals are jet black on the dorsum and creamy white on the ventrum, but pure white forms are not uncommon and intermediate colours are also found (Flannery 1994; McKay 1995; Lindenmayer

2002a). This species is nocturnal, arboreal and folivorous and is dependent on tree hollows for its nesting requirements.

75 The following table of taxonomy for the Greater Glider has been taken from the literature review conducted for Chapter 2 of this thesis. The information provided is an up-to-date review of the previously accepted (McKay 1988) taxonomy of the Greater

Glider Petauroides volans Kerr 1792.

Table 5.1: Taxonomy of the Greater Glider.

Petauroides Thomas, 1888 Petaurus Shaw, 1791 Didelphis Kerr, 1792 Voluccella Bechstein, 1800 Phalanger Lacepede, 1801 Phalangista Desmarest, 1803 Petaurista Desmarest, 1820 Petauroides Thomas, 1888 Petaurides Ramsay, 1890 Belideus Anon, 1890 Schoinobates Iredale and Troughton, 1934 Petauroides volans (Kerr, 1792) Petauroides volans volans (Kerr, 1792) Didelphis volans Kerr, 1792 Didelphis voluccella Meyer, 1793 Didelphis macroura Shaw, 1794 Voluccella nigra Bechstein, 1800 Voluccella macroura Bechstein, 1800 Phalanger volans Lacepede, 1801 Phalangista volans Desmarest, 1803 Petaurus macroura Oken, 1816 Petaurus niger Oken, 1816 Petaurus taguanoides Desmarest, 1817 Petaurus macrourus Desmarest, 1817 Petaurus peronii Desmarest, 1817 Petaurista taguanoides Desmarest, 1820 Petaurista macroura Desmarest, 1820 Petaurista peronii Desmarest, 1820 Phalangista macroura Schinz, 1821 Petaurus didelphoides Cuvier, 1825 Petaurus maximus Partington, 1837 Petaurus volans Thomas 1879 Petaurista volans Thomas 1885 Petauroides volans typicus Thomas, 1888 Petauroides volans incanus Thomas, 1923 Petauroides volans armillatus Thomas, 1923 Schoinobates volans volans Iredale and Troughton 1934 Schoinobates volans incanus Iredale and Troughton 1934 Schoinobates volans armillatus Iredale and Troughton 1934 Petauroides volans minor (Collett, 1887) Petaurista volans minor Collett, 1887 Belideus cinereus Anon, 1890 Petaurides cinereus Ramsay, 1890 Schoinobates volans minor Iredale and Troughton 1934

76 5.4 Distribution of the species in NSW and across its range, including current and

historical range.

The current distribution of the Greater Glider ranges from north Queensland to Victoria, restricted predominantly to the wet sclerophyll forests and rarely in the dry sclerophyll forests of eastern Australia (Tynedale-Biscoe 1973, Flannery 1994, Strahan 1995, see

Figure 5.1). The distribution and abundance of the Greater Glider from the early natural history literature is reviewed in Chapter 2 (Section 2.6).

Figure 5.2. Distribution of the Greater Glider in Australia. (Source- Wildlife Preservation Society of Queensland 2005)

77 In New South Wales the Greater Glider is found in the wet tall forests of the ranges along the eastern coast (Figure 5.2). Marlow (1958) undertook a review of the mammals of NSW, and reported that the Greater Glider was abundant in the taller forested areas.

The distributional records provided by NPWS (see Figure 5.2) confirm that the greater glider is widespread in NSW.

Figure 5.3. Distributional records of the Greater Glider in NSW. (Source DECC Fauna Atlas 2006)

The recent distributional records of the Greater Glider from the southern half of the

Sydney Bioregion (Figure 5.3) indicate that there are more records in the western parts of this bioregion than the eastern part. Much of the natural habitat has not been cleared in this region compared to the intensely cleared eastern part (Benson and Howell 1990).

The predicted occurrence of the Greater Glider in suitable habitat is shown in Figure

5.3. The western half of the bioregion is predicted to have more Greater Gliders than the eastern half, because more suitable habitat is available in the western area. 78

Figure 5.4. Distribution and predictive habitat model of the Greater Glider in the Southern Sydney bioregion. (Source Department of Environment and Climate Change 2007b).

79 5.5 The relevant ecological requirements of the species.

5.5.1 Life-cycle

Tyndale-Biscoe (1973), with limited information, reported that the female Greater

Glider had a life expectancy of four years: More recently, life expectancy has been reported as being 10 - 15 years (Lindenmayer and Lacy 1995, Lindenmayer 1997,

Lindenmayer 2002a). Animals living in the wild have an expected shorter life span than captive animals. The Greater Glider has been reported to have a 20% juvenile mortality rate which is male biased (Tyndale-Biscoe 1973, Kerle 2001). Agonistic behaviour by adult males towards young males is thought to be the cause of these deaths (Kerle 2001,

Lindenmayer 2002a), as well as predation as juveniles disperse (Norton 1988). The body weight of an adult Greater Glider ranges from 1000 to 1700g.

5.5.2 Reproductive biology

Mating occurs between February and May (Norton 1988), and the Greater Glider gives birth to a single young per year between April and June (Kerle 2001, Lindenmayer

2002a). The pouch contains two teats and the weaning time is 7.5 months (Lindenmayer

2002a).The reproductive rate is only 60-75 % per adult female per year, with a greater success in forests with higher nutrients (Norton 1988, Kerle 2001). Females are able to reproduce in their second year; any young lost are not replaced (Tyndale-Biscoe and

Smith 1969a). Coinciding with this is the fact that males produce sperm only during the breeding season, with the testes regressing after May (Kerle 2001). The mating system of the Greater Glider can be monogamous, polygamous or polygynous, depending on

80 the density of the population (Norton 1988, Lindenmayer 2002a). Males are solitary and occupy a home range that is inclusive of females (Henry 1984, Kehl & Borsboom 1984,

Comport et al. 1996, Kavanagh and Wheeler 2004). Greater Gliders, due to their larger size, have a lower reproductive potential than the other gliding species because they have a maximum of one offspring per year (Kerle 2001, Lindenmayer 2002a).

Lindenmayer and Lacy (1995) calculated a λ (lambda) value of 1.047 and a generation time of 5.78 years for this species, within old growth Montane Ash forest in Victoria..

5.5.3 Population dynamics

The size and distribution of a population of Greater Gliders is dependent on the amount of hollows and nutrient levels in the forest they inhabit (Kehl and Borsboom 1984,

Norton 1988, Braithwaite 1996, Kavanagh 2004). The two sexes have been reported to be evenly spaced in the forest, with very little overlap (Tyndale-Biscoe and Smith

1969a). The males and females only spend time together during the breeding season, at other times being solitary (Lindenmayer 2002a). Exceptions being the time the young spend with the female, till they disperse (Fleay 1933). The population density of Greater

Gliders has been reported as ranging between 0.01 to 5 animals per hectare in the differing forest types across its range (Norton 1988, Gibbons and Lindenmayer 1997,

Lindenmayer 2002a, Pope et al. 2004). Norton (1988) found that population fecundity was lower in forests with low concentrations of nutrients (nitrogen, phosphorous and potassium) in the foliage.

81 5.5.4 Social behaviour

The Greater Glider has been reported as being virtually silent (Wakefield 1970) and solitary, with exclusive home ranges that overlap with the home range of one or more females (Henry 1984, Pope et al. 2004). The co-occupancy of den trees is rare except between matched pairs in the breeding season (Lindenmayer 2002a). They communicate through odoriferous secretions from specialised glandular areas (Kerle

2001). With a limited vocal ability, it has been suggested that these scents promote the spacing of individuals throughout the forest (Lindenmayer 2002a). Agonistic behaviour between females has been observed, when one animal moves through or into the other animal’s territory (Henry 1984). The size of the home-range and population density is related to the levels of nutrient foliage available (Norton 1988); that is, low foliage nutrient levels relate to larger home-ranges and smaller populations, in contrast to areas with higher nutrient foliage levels where smaller home-ranges and larger populations are found (Norton 1988, Lindenmayer 2002a).

5.5.5 Group composition

The Greater Glider is a solitary species and has only been reported to share a den with another Greater Glider during the mating season (Fleay 1933, Henry 1984, Norton

1988, Comport et al. 1996, Lindenmayer 2002a). The young stays with the female till they disperse (Fleay 1933).

82 5.5.6 Home range size

There have been five published studies (Henry 1984, Kehl & Borsboom 1984, Comport et al. 1996, Kavanagh and Wheeler 2004 and Pope et al. 2004) and one unpublished study (Norton 1988) on the home range of the Greater Glider. These studies were undertaken across the distribution of the species, and results varied between 1.4 ha to

2.6 ha for males and 0.8 ha to 2.5 ha for females.

Table. 5.2. The results of six studies on the home range size (estimates) of the Greater Glider. P = polyynous males, M = monogamous males, # = mean home- range area (ha).

Area Study No. of Males# Females# All# Animals Southern Henry 24 (P) 2.1 1.3 1.5 Victoria (1984) (M)1.4 South-east Kehl & Queensland Borsboom 10 2.6 2.5 2.6 (1984) North Comport et 11 2.2 1.0 1.6 Queensland al. (1996) South- Kavanagh eastern and 9 2.0 0.8 1.4 NSW Wheeler (2004) South- Pope et al. 23 2.6 2.0 2.3 eastern (2004) NSW South- Norton 10 1.4 1.4 1.4 eastern (1988) NSW

The difference in the home-range size of the Greater Glider has been attributed to the nutrient levels and hollow availability of the forest type that it inhabits (Norton 1988,

Kavanagh 2004). Although this species is the largest glider, it does not require a large home-range as suitable food resources are substantial and continuous in suitable forest types (Lindenmayer 2002a).

83 5.5.7 Shelter and food requirements

5.5.7.1 Shelter

The Greater Glider is dependent on tree hollows with an entrance of >5cm for its nesting requirements (McKay 1995, Gibbons and Lindenmayer 1997). Milledge et al.

(1991) suggested that the distribution and abundance of the Greater Glider is closely related to hollow availability. In the central highlands of Victoria, the number of trees with hollows was found to be a significant factor in models describing habitat requirements of the Greater Glider (Lindenmayer et al. 1990). This species utilizes many dens in its home range, individual Greater Gliders have been observed using from four to eighteen dens (Kehl and Borsboom 1984). The den and nest trees are tall and of large diameter, the entrance is through a spout or hollow branch 40m high

(Lindenmayer 1997).

5.5.7.2 Food requirements

The Greater Glider has been reported as being folivorous, eating the buds, leaves and bark (Marples 1973) from only a few selected eucalypt species in its range (McKay

1995), favouring the young leaves when seasonally available (Kavanagh and Lambert

1990). Maloney and Harris (2006b) reported that the Greater Glider had been observed feeding upon other species than eucalypts (see Table 5.2) and also being predatory on kookaburra eggs. Other species reported as being eaten by the Greater Glider include

Acacia spp. and the mistletoes that occur on eucalypts; Amyema spp. (Lindenmayer

2002a) (see also Table 5.2). Lindenmayer (2002a) suggested that the Greater Glider

84 consumes more mistletoe than has been recognised, as mistletoes are comparatively high in nitrogen and phosphorous with fewer toxic chemicals than the eucalypts they grow on. Le Souef and Burrell (1926) reported that they had observed Greater Gliders in Casuarina forests in the Myall Lakes area, and examination of stomach contents revealed Casuarina spp. had been eaten. A review of the preferred food tree species eaten by the Greater Glider revealed that 33 eucalypt species and ten non eucalypt species are consumed across its distributional range (Table 5.2). Hume (1999) reported that this species is close to the minimum size requirement to sustain a strictly folivorous diet. The Greater Glider utilises hindgut microbial fermentation to maximise the energy it gains from a folivorous diet (Foley 1987, Foley and Hume 1987).

5.5.8 Foraging behaviour

The foraging behaviour of the Greater Glider is dependent on where the preferred food and den tree species occur in its home-range (Lindenmayer 2002a). The gliders emerge from the den 35-100 minutes after dusk dependent on the season (Lindenmayer1997,

Kavanagh and Wheeler 2004); and take a known course to the preferred feed tree, either by a succession of glides or through a connected canopy (Fleay 1947, Norton 1988,

Kerle 2001). Leaves are eaten all year round; eucalypt buds are eaten in winter to spring, and they feed high up on the edge of the canopy in the largest trees (McFarlane

1988, Kavanagh and Lambert 1990). The approximate time outside the den is spent resting (40%), feeding (30%), moving (12%), grooming (10%) and other activities (e.g. social interactions, 8%) (Kehl and Borsboom 1984, Norton 1988, Kavanagh and

Lambert 1990, Comport et al. 1996).

85

Table 5.3. Reported food tree species of the Greater glider across its range. Area and source of report in bold. # = non eucalypt, ** non native Victoria, Fleay 1947 South-east NSW, Norton 1988 Eucalyptus radiata Eucalyptus radiata E. ovata E. viminalis E. viminalis E. dalrympleana E. stuartiana E. fastigata E. rostrate E. piperata E. goniocalyx E. gummifera E.sieberi E. pellita E. consideniana Tumut NSW, Marples 1973 North Queensland P. volans minor Eucalyptus robertsoni (radiata) Comport, Ward and Foley 1996 E. viminalis Eucalyptus acmenoides E. pauciflora E. intermedia E. dalrympleana E. citriodora E. tereticornis E. creba Allocasuarina torulosa # Lophostemon suaveolens # Captive fed specimens South-east NSW, Kavanagh and Lambert Hume, Foley and Chilcott 1984 1990 Eucalyptus radiata Eucalyptus obliqua E. fastigata E. viminalis E. ovata E. radiata E. cypellocarpa Southern Victoria, Henry 1984 Records from the Victorian Naturalist Eucalyptus globoidea 1884 – 2005. Maloney and Harris 2006 E. sideroxylon Eucalyptus cypellocarpa E. bridgesiana E. dives E. cypellocarpa E. globulus E. obliqua E. goniocalyx E. globulus E. macrorhyncha E. nortonii E. obliqua E. ovata E. radiata E. viminalis Acacia melanoxylon # Amyema pendula # Mullerina eucalyptoides # Pandores pandorana # Parsonsia straminea # Pomaderris aspera # South-east Queensland Tumut NSW Cunningham et al, 2004 Kehl and Borsboom 1984 Eucalyptus viminalis/E. dalrympleana Eucalyptus umbra E. radiata E. intermedia E. camphora E. drepanophylla Pinus radiata ** E. trachyphloia E. macrorhyncha Melaleuca quinquernervia # E. dives Acacia melanoxylon # E. pauciflora

86 5.5.9 Predators

There have been ten reported predators of the Greater Glider in the literature (see Table

5.3). Three of them are listed as threatened in NSW. The Greater Glider is an important prey source for these three species, and as such should be a priority species for conservation management (see section 3.4.2 for individual species appraisal). Eight of the ten species which have been reported here as being a predator of the Greater Glider have been recorded in Royal National Park. These are the wedge-tailed eagle, powerful owl, sooty owl, lace monitor, carpet snake (diamond python), dog, fox and cat (Andrew

2001). The spotted-tailed quoll is thought to be locally extinct and of uncertain status within Royal National Park (Andrew 2001).

Table 5.4. The ten predators of the Greater Glider obtained from literature reviews. (# = species listed on the NSW Threatened Species Conservation Act 1995).

Scientific Name Common Name Source Ninox strenua# Powerful Owl 1,2,3,4,5,6,7,8,9,10,11,12,13,14,15,16,17 Tyto tenebricosa# Sooty Owl 14,15,17 Canis familiaris Dog 17,18,19,20 Vulpes vulpes Fox 1,2,17,18,19,20,21,22, 23, 24 Felis catus Cat 17,25 Varanus varius Lace Monitor 26, 27 Dasyurus maculatus# Spotted-tailed Quoll 17,28,29 Canis familiaris dingo Dingo 30 Aquila audax Wedge-tailed Eagle 17,31 Morelia spilota Carpet Snake 27

Sources 1. Le Souef and Burrell (1926), 2. Fleay (1947), 3. Fleay (1968), 4. James (1980), 5. Kavanagh (1988), 6. Pavey (1992), 7. Chafer (1992), 8. Kavanagh (1992), 9. Pavey (1994), 10. Pavey et al (1994), 11. Lavazanian et al (1994), 12. Pavey (1995), 13. Schultz (1997), 14. Kavanagh (2004), 15. Bilney et al (2006), 16. Cooke et al (2006), 17. Maloney and Harris (2006), 18. Lunney et al (1990), 19. McFarlane (1988), 20. Mitchell and Banks (2005), 21. Wallis and Brunner (1986), 22. Coman (1973), 23. Brunner et al (1975), 24. Roberts et al (2006), 25.Coman and Brunner (1972), 26. Weavers (1989), 27. Lindenmayer (2002a). 28. Glen and Dickman (2006), 29. Belcher and Darrant (2006), 30. Robertshaw and Harden (1985), 31. Tyndale-Biscoe and Smith (1969b).

87 5.5.10 Diseases

The only report of a disease for the Greater Glider is that some populations have contracted Chlamydia (Lindenmayer 2002a). Lindenmayer (2002a) suggested that it

‘does not appear to compromise breeding success in the species as it has done in some populations of the koala’. Fleay (1933) reported that one of his captive Greater Gliders had a mite which affected the animal around the eyes, becoming so bad that the animal eventually died. Nematodes, recorded in their millions from the caecum of the Greater

Glider, have been suggested to aid in the digestion of their folivorous diet (Lindenmayer

2002a). While only one disease has been reported for the Greater Glider, many ecto and endoparasites have been reported (see Table 5.5).

Table 5.5. The ectoparasites (= #) and endoparasites (= *) collected from the Greater Glider. Source Lindenmayer (2002a) and Spratt (2002).

Fleas # Ticks # Cestoda * Nematoda * Choristopsylla ochi Ixodes fecialis Bertiella mawsonae Paraustrostrongylus spp. Ixodes tasmani Bertiella petaurina Marsupostrongylus minesi Austroxyuris finalysoni Paraustroxyuris parvus Breinlia spp. Cercopithifilaria johnstoni

5.6 Known and potential threats

While the Greater Glider is not listed as a threatened species, it is susceptible to the threatening processes listed for possum and glider species in state and federal conservation legislation. These threatening processes include the following;

• Loss and fragmentation of habitat through clearing and the activities associated

with clearing

88 • Logging of old growth elements removing the number of hollow bearing trees

available for nesting

• Inappropriate fire regimes reducing the availability of food resources and

isolating populations making them vulnerable to regionally catastrophic events

• Predation by introduced animals (foxes and cats)

Threatening processes do not act in isolation; typically they act in a cumulative way

(Lindenmayer 2002a). For example, the loss of habitat removes hollow bearing trees and causes fragmentation of the remaining habitat. This can be detrimental for dispersing gliders, as the distance between the fragmented patches cannot be traversed by gliding, bringing them to the ground where they are susceptible to predation. The

Greater Glider has been recorded as the prey for ten species (see table 5.3); eight of these predators have been recorded in Royal National Park (Andrew 2001), the introduced dog, cat and fox included.

A fragmented population of Greater Gliders in the Eurobodalla Local Government Area, has had a determination made as an endangered population under the NSW Threatened

Species Conservation Act 1995 (Department of Environment and Climate Change NSW

2007). The population has been deemed to be disjunct from other occurrences of the species due to barriers restricting their dispersal ability, and continuing loss of habitat will further isolate the population. See also a review of the threats to populations of the

Greater Glider in Chapter 3 (section 3.4) of this study.

89 5.7 The success of previous translocation programs of the same or analogous

species

There have been no published reports of any previous translocations of the Greater

Glider in NSW. Maloney and Harris (2006b) reported that two Greater Gliders were liberated at Wilsons Promotory (Vic.) one in 1929 the other in 1934, their fate was uncertain. Male Greater Gliders that were released into woodland adjoining the State

Forests offices in west Pennant Hills survived for several years in this area (Rod

Kavanagh pers. comm.).

The Greater Glider is a folivorous, arboreal gliding marsupial that is hollow-dependent

(Flannery 1994, McKay 1995). There is no other analogous species that has these same requirements; nevertheless there are some similar species which have been translocated.

The Koala (Phascolarctos cinereus) is a folivorous arboreal marsupial, but it is not hollow dependent (Strahan 1995). In Victoria, populations of the Koala have been translocated since the 1920s from French and Phillip islands (Martin 1989). This species has been translocated into many parts of its former geographic range; however the fate of individuals and population dynamics of some of these translocated individuals is poorly known, whereas some have become pests (Norton 1994).

The Sugar glider (Petaurus breviceps) is an arboreal glider that utilizes smaller hollows than the Greater Glider, and is not folivorous (Strahan 1995, Lindenmayer 2002a). The

Sugar glider was successfully introduced into Tasmania in 1935 (Gunn 1851), and is now found in suitable habitat across that state (Lindenmayer 2002a). Suckling and

90 Goldstraw (1989) successfully used nest boxes to help establish a population of Sugar gliders in Box-Ironbark forests in northern Victoria.

The Common brushtail possum (Trichosurus vulpecula) is a nocturnal arboreal herbivore, which uses hollows when available (Pietsch 1984, Strahan 1995). The

Common brushtail possum was successfully introduced into New Zealand, and is now a pest in that country (Efford and Cowan 2004). Other studies have reported that translocated individuals from urban areas to native bushland, either died from predation or dispersed to other areas (Pietsch 1994). Cowan (2001) reported that translocated possums in New Zealand suffered the same fate as in Australia.

The successful translocation of a species is dependent on the receiving area being vacant of the same species, eliminating competition between translocated and resident animals (Pietsch 1994). The monitoring of translocated individuals is important to understand the fate of individuals and the success of the programme (Norton 1994,

Serena and Williams 1994).

5.8 Conclusion

The Greater Glider is not a threatened species and was commonly observed in Royal

National Park, but it has been missing since the 1994 fire event, thus making it a worthwhile candidate for a translocation project, as outlined by Serena and Williams

(1994).

The translocation programme would be a reintroduction of this species to Royal

National Park. The individuals required would need to be sourced from an area other

91 than the Illawarra. The results from a detailed field study (Chapter 4) found that while the Greater Glider is found in the Illawarra they are in low numbers. Populations of the

Greater Glider in this area have declined and are a local conservation priority (DECC

2007b). If they were sourced from this area, unnecessary pressures would be placed on these populations, with possible extinction resulting. Other possible sources include the western part of the study area shown in Figure 5.4.

The reason for translocating the Greater Glider to Royal National Park (RNP) is that it is missing from RNP and also reported to be declining in some areas (DECC 2007b, van der Ree and McCarthy 2005 and Woinarsky et al 2006). Translocation is an option for conservation action for this species.

92

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