Factors Influencing the Distribution of Bush Rats

Home , Acrotriche divaricata, Bush rat, Epacris microphylla, Grevillea sphacelata, Kunzea ambigua, Leucopogon amplexicaulis

Factors influencing the distribution of Bush rats

Rattus fuscipes

Bush rats (photo Wendy Gleen)

Bush rat habitat (photo Wendy Kinsella)

A thesis submitted for Master of Philosophy at the University of NSW

2014

Wendy Kinsella

The School of Biological, Earth and Environmental Science

The University of New South Wales Sydney NSW, Australia

Table of Contents Student Declaration ...... v

Acknowledgement ...... vi

List of Figures ...... vii

List of Tables ...... ix

Abstract ...... xi

Chapter 1 ...... 1

General Introduction ...... 1 1.1 Habitat requirements for a small mammal ...... 1

1.2 Australia’s Urban Landscape ...... 2

1.3 Relationship between habitat and species...... 3

1.4 Research objectives ...... 4

Chapter 2 - Habitat characteristics associated with bush rat populations in Eastern Australia...... 5

2.1. Introduction ...... 5 2.1.1 Habitat and Mammal association ...... 5

2.1.2 Impacts of land clearing on native vegetation ...... 6

2.1.3 Mammal translocation and reintroduction ...... 7

2.1.4 Habitat Characteristics ...... 8

2.2 Methods ...... 11

2.2.1 Study area ...... 11

2.2.2 Study sites ...... 11

2.2.3 Method of Survey site selection ...... 17

2.2.4 Field methods ...... 18

Vegetation layers ...... 20

2.2.5 Vegetation Community Profiles ...... 21

2.2.6 Data analysis ...... 23

2.3 Results ...... 24

2.4 Discussion...... 28

2. 4.1 Bush rat Habitat ...... 28

2.4.2 Ground cover density ...... 29

2.4.3 Other structural variables ...... 31

2.4.4 Floristic variables ...... 32

2.4.5 Environmental variables ...... 32

2.4.6 Implications for the reintroduction of bush rats ...... 35

2.5 Conclusion ...... 36

Chapter 3 – Competition interactions with black rats limit bush rat distribution within its native range ...... 40

3.1 Introduction ...... 40

3.1.1 Understanding establishment of Communities and Competitors ...... 40

3.1.2 Reintroduction and impacts from competitor species ...... 40

3.2 Methods ...... 43

3.2.1 Coexistence of Bush rats and Black rats ...... 43

3.2.2 Data collection ...... 44

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3.3 Analysis ...... 45

3.4 Results ...... 46

3.5 Discussion...... 47

3.5.1 Small Mammal Communities ...... 47

3.5.2 Community assembly and coexistence with invasive species ...... 48

3.5.3 Coexisting and available resources ...... 49

3.5.4 Community structure and competition ...... 49

3.6 Conclusion ...... 50

Chapter 4...... 52

General Discussion ...... 52

4.1 Why habitat matters ...... 52

4.2 Why determine factors that influence Rattus fuscipes (bush rat) distribution...... 53

4.3 What I found in relation to high density of bush rat populations ...... 53

4.4 Variables suitable for bush rat establishment in reintroduction sites ……………………….55

References ...... 57

Appendix ...... 65

Student Declaration

ORIGINALITY STATEMENT

'I hereby declare that this submission is my own work and to the best of my knowledge it contains no materials previously published or written by another person, or substantial proportions of material which have been accepted for the award of any other degree or diploma at UNSW or any other educational institution, except where due acknowledgement is made in the thesis. Any contribution made to the research by others, with whom I have worked at UNSW or elsewhere, is explicitly acknowledged in the thesis. I also declare that the intellectual content of this thesis is the product of my own work, except to the extent that assistance from others in the project's design and conception or in style, presentation and linguistic expression is acknowledged.'

Signed

Date

Acknowledgement I’ve been so privileged to have an excellent supervisor in Dr Stephen Bonser to guide encourage and tease me in this journey over the last 4 or is it 5 years? I remember dearly the time he told me he and Angela where expecting their first child Sam. I must have looked surprised as he quickly said “OH it won’t affect your studies” and what surprised me was that he took the trouble to inform me as it was excellent news. I really appreciated the warmth care and understanding I received working in the Bonser lab.

My one regret was not having more to do with Dr Peter Banks just time did not allow me to visit Sydney uni as frequently.

Then there is Dr Grainné Cleary who was the inspiration of why I got involved in this project.

Grainné was always incredibly helpful when I could not work my computer or deal with some issue she would solved every problem to help me find my way. Bill Cleary was also a treasure a computer whiz at fixing a graph or putting Mrs Doyle (Father Ted (Crilly’s) house keeper) as my screen saver.

A big thankyou also goes to Wendy Gleen who believed that I could reach the end of this report thanks Wendy. I love you for your dedication to education us about the beauty and uniqueness of Australian fauna and your passion for their protection in the wild. Also wish to thank the wonderful Rebecca Spindler and Duncan Bourne they are two of the most generous people I know. Duncan Bourne and Emily Smith, both fantastic field assistants it was a pleasure to ID plants with you both.

Big thanks also goes to my 2 sisters Jane and Judith who were so understanding with my preoccupation with this project and a special thanks to Jane who spent many hours helping me with words thoughts and challenges.

Finally I’d like to acknowledge the 3 bush rats, the two Wendy’s and Hattie who lived for 2 years 10 months during this project and travelled extensively in Taronga’s Zoo Mobile.

List of Figures

Fig. 2.1 Map of the central east coast of New South Wales showing the location of Sydney with Ku-ring-gai National Park to the north and The Royal National Park to the South within the Sydney Basin highlighted in red on map of Australia.

Fig. 2.2 Map of the 9 survey areas at Ku-ring-gai Chase National Park with the 9 sample sites marked with black dots. The red triangles represent records of Rattus fuscipes bush rats from

Atlas of NSW Wildlife records 2012.

Fig. 2.3 Map of the 13 survey areas at The Royal National Park marked with black dots. Red triangles represent records of Rattus fuscipes bush rats as held in Atlas of NSW Wildlife records 2012.

Fig. 2.4 Logistic regressions demonstrating a significant (P = 0.025) relationship between bush rat density and percent fallen timber cover across sample plots (a), and a marginally significant (P = 0.05) relationship between bush rat density and percent leaf litter across sample plots (b). Symbol size is presented relative to the number of overlapping data points for a given value of percent fallen timber or leaf litter (smallest points represent a single sample plot and largest point’s represent four overlapping sample plots).

Fig. 3.1 Mapping occurrence records of Bush rats Rattus fuscipes in Australia. Darker colour indicates higher density of bush rats. Source The Atlas of Living Australia (www.ala.org.au).

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Fig. 3.2 Mapping occurrence records of Black rats Rattus rattus in Australia. Darker colour indicates higher density of black rats. Source The Atlas of Living Australia (www.ala.org.au).

Fig. 3.3 Map of the Sydney Metropolitan Area showing black rats (Rattus rattus) in urban areas and their impact into surrounding areas of bush rat (Rattus fuscipes) populations along the east coast of New South Wales. (Source Atlas of NSW accessed October 2009).

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List of Tables

Table 2.1 The Royal National Park landscape and vegetation characteristics based on

SMCMA region (2009 DECC) Vegetation Community Profiles.

Table 2.2 Ku-ring-gai National Park landscape and vegetation characteristics based on

SMCMA region (200(DECC) Vegetation Community Profiles

Table 2.3 Vegetation Survey Field Sheet

Table 2.4 Summary of all records collected from the field at Ku-ring-gai National Park

Table 2.5 Summary of all records collected from the field at The Royal National Park &

Heathcote National Park

Table 2.6 Multiple Logistic Regression results of bush rat population density (variable in the equation percent fallen timber > 5cm) for B=-0.236, S.E=0.105, Wald=5.013, d.f. =1, Sig.

=0.025, Exp (B) =0.790 results significant.

Table 2.7 Multiple Logistic Regression results of bush rat populations (variables not in the equation) results show variable that do not significantly affect the predictive power of the model but list variable in order of importance to bush rat habitat.

Table 2.8 Summary Plant List: Ku-ring-gai Chase National Park.

Table 2.9 Summary Plant List: The Royal & Heathcote National Park.

Table 3.1 Binomial Tests for coexistence between Bush rats and Black rats.

Table 3.2 List of sites selected in Sydney Basin where each site had numerous transects.

Source: The Living Atlas of Australia.

Table 3.3 Case Study Wyong (South & West) New South Wales.

Abstract Translocation and reintroduction of species back to their former range is an excellent goal but reintroductions can only be achieved successfully with extensive knowledge of the different factors controlling species distributions. This study investigates the factors controlling the distribution of the bush rat (Rattus fuscipes). The Sydney Basin Region of Australia contains core habitats that support bush rat populations but bush rats tend not to be found in habitats heavily modified by humans. The absence of bush rats in these habitats could be due to habitat modification, or due to the increased presence of an invasive competitor, the black rat

(Rattus rattus). The first goal of this research was to identify habitat characteristics associated with high density bush rat populations. The second goal was to test if there is a negative association between the distribution of bush rats and a potential competitor, the black rat. To test for habitats associated with high density bush rat populations, habitats were surveyed over 22 sites in national parks surrounding Sydney. High and low densities of bush rat data were identified using the Wildlife Atlas. Field records of surveys covered 21 habitat variables. I found that the only significant habitat variable predicting the presence of bush rats was the presence of percent timber greater than 5cm in the ground cover. To test negative associations between bush rats and black rats, I collected records from Living Atlas of

Australia within the Sydney Basin Region. Records covered 39 areas where longitude and latitude bearings grouped to give individual trap lines over 30 years. I found that bush rats and black rats have broadly overlapping distributions. However, bush rats and black rats tend not to coexist in the same place at the same time. My results suggest that competition with black rats can limit the distribution of bush rats. My studies demonstrate that habitats with complex understory cover (fallen timber) and the absence of black rat competitors both promote the presence of bush rats in a habitat. Both habitat quality and invasive competitors will impact the success of reintroductions of bush rats within urban habitat fragments.

Chapter 1

General Introduction

1.1 Habitat requirements for a small mammal

The goals of this thesis are to understand the habitat characteristics of a small mammal Rattus fuscipes (bush rat) and to test for co-existence with an urban invader Rattus rattus (black rat).

Describing ecological niche or habitat of any species requires detailed knowledge of that organism and its environment which is very difficult to obtain (Villee et al. 1985) but this knowledge is critical in understanding patterns of biological diversity, conservation and in efforts to reintroduce species into former habitats.

Worldwide biodiversity is declining due to habitat loss from vegetation clearing habitat fragmentation and environmental degradation due to population growth (Cardillo et al. 2004;

Sodhi et al. 2004). In Australia around 22 terrestrial mammal species are estimated extinct

(Firth 2010). No single factor is thought to explain the decline in Australia but one theory is some mammal populations are thought to have passed a threshold where they are no longer capable of withstanding ongoing threats of habitat change, predators and disease from introduced exotic animals (Woinarski et al. 2011). Australian mammal extinctions are thought to have occurred mainly from 1890-1950 with European settlement and when little research into the complexity of Australian wildlife was carried out (Woinarski et al. 2011).

Research into factors of habitat characteristics and competition from invaders are especially relevant today since fragmentation of remnant vegetation and on ground works of restoring remnant vegetation will determine if a given species will have core habitat areas within restored vegetation. Habitat characteristics help us understand wildlife and help protect both

1 the land and the species that depend on it. Australia’s native vegetation is unique with 85% of our plant species found nowhere else (State of the Environment Committee 2011). In turn, these vegetation communities support a unique diversity of animal species. The paradox of the bush suggests that while we superficially recognise the outback and the bush as part of our essence, the level of recognition is skin deep and we don’t seem to recognise when the bush is falling apart (McDonald & Woinarski 2011).

1.2 Australia’s Urban Landscape

Over eighty percent of the population of Australia live within 50km of the coast with seven of the largest cities located on the coast with few large cities located inland. Almost two thirds

(64%) live in the eight capital cities while the proportion of Australians living in urban areas is 87% (State of the Environment Committee 2011). Urban expansion in Australia is continuing, and is having a particularly strong impact in areas near the coasts (Gurran et al.

2007; Bradshaw 2012).

Over the decade from 2001 to 2010 in Australia an average of 1 million hectares was cleared annually (State of the Environment Committee 2011). Fragmentation of ecosystems by land clearing in Australia is concentrated in a number of regions and coastal areas. Land clearing has stabilised recently but the clearing of woody vegetation has fluctuated for the past 20 years in New South Wales. The impact on habitats is shown by the decline of species with still limited understanding of complexity of Australian vegetation communities and assemblages. Habitat fragmentation initiates the decline of other factors of habitat such as size, shape and isolation (Salek et al. 2010). These factors of habitat are crucial for any organism, as fragment size can influence many aspects of population demography (Holland

& Bennett 2010) as habitat is the physical space that an organism uses to gain resources essential for growth, survival and reproduction.

1.3 Relationship between habitat and species

The bush rat (Rattus fuscipes) is a common small native mammal in coastal areas of

Australia. It occurs as four subspecies and ranges from south-west Western Australia to northern Queensland (Taylor & Calaby 1988). This thesis focuses on the bush rats found in the bushland around Sydney NSW. Bush rats can occupy a range of habitats (Dickman et al.

2000) but are susceptible to logging, fire and habitat fragmentation (Lindenmayer et al.

2005). Urbanisation has confined the bush rat’s habitat to areas in National Parks North and

South of Sydney (Fig 2.1). Bush rats are found to the west of Sydney but only in very small numbers due to extensive land clearing (T.Leary, personal communication). The challenges then are what factors influence the distribution of the bush rat in coastal forests and shrublands. The bush rat is commonly trapped in the field and has been a target of research in a variety of areas including habitat interspecific association, succession, fire, genetics, metapopulations, diet, social and forestry (Steward & Barnett 1983; Carron et al. 1990; White et al. 1996; Downes et al. 1997; Banks 1999; Hinten et al. 1999; Tasker et al. 1999; Maltz &

Dickman 2001; Lindenmayer et al. 2005; Peakall et al. 2006; Fox et al. 2007; Frazer et al.

2007; Garden et al. 2007; Holland & Bennett 2007; Parrott et al. 2007; Claridge et al. 2008;

Macqueen et al. 2008; Stokes et al. 2009a; Vernes & Dunn 2009).

This research was initially started under the arm of a larger project to reintroduce bush rats to suitable urban remnants in Sydney Harbour National Park. The aim of the reintroduction project was to release and monitor bush rats known to exhibit interspecific competition with black rats Rattus rattus in the wild (Stokes et al. 2009b). Black rats are considered a threat to native small mammal species (Downes et al. 1997; Cox et al. 2000; Harper et al. 2005;

Cassaing et al. 2007) with increasing presence and instrumental in spreading disease to native species (Woinarski 2011). The aim of my research is to understand fundamental habitat features of the bush rats and the presence of a competitive species so as to assist reintroducing bush rats back into suitable urban areas. Sampling ideal habitat will contribute or defining a set of observation needed before undertaking future reintroduction. Another researcher with the project also focused their master’s thesis on habitat of the bush rat but looked at marginal features rather than fundamental features along the urban edge (Gleen

2014). Previous research suggest that bush rats favour Eucalyptus woodland (Breed & Ford

2007) and tend to be found in habitats with dense ground cover which usually occurs in gullies where low shrubs, grasses, leaf litter and fallen trees provide shelter (Downes et al.

1997; Maitz et al. 2001; Holland et al. 2007; Claridge et al. 2008). However, we know little about the specific habitat requirements of bush rats. This species was ideal to identify habitat associations and interactions with invasive rat species that support or limit bush rat populations and find habitat features that would assist reintroduction of the bush rat back into suitable urban matrix.

1.4 Research objectives

1. To identify habitat characteristics associated with high density bush rat populations.

2. To test if there is a negative association between populations of bush rats and a

potential competitor, the black rat.

Chapter 2 - Habitat characteristics associated with bush rat populations in Eastern Australia

2.1. Introduction

2.1.1 Habitat and Mammal association

Habitat conditions play a major role in the distribution of mammal species, with many species found in a relatively narrow range of habitat types (Pagel et al. 1991). The most common habitat preferences of terrestrial mammals are forests with shrubs and to lesser degree grasses (IUCN 2014). Forests are species rich ecosystems (Lindenmayer 1999) with ground dwelling mammal’s abundant and distribution varying due to forest structure and habitat complexity (Catling et al. 2001). Habitats are complex as they have many features such as cover, food availability and distance to water that could influence the persistence of a given mammal species. Research into small mammal assemblages in fragmented forest has shown habitat structure to be important in determining mammal assemblages (Garden et al.

2007). Dense vegetation and habitat structure provides ground dwelling animals protection from predation (Lindenmayer et al. 1999a). Further studies have focused on succession of small mammals assemblages after disturbance by fire; these studies found successional changes in vegetation drove the changes in small mammal communities (Monamy and Fox

2000). I found records of bush rats in a range of habitats from rainforest, woodland, forest, scrub and heath. This is consistent with previous studies where Rattus species are known to have a relatively broad range of habitats compared to other small mammals (Dickman et al.

2000). These findings are similar to other studies where mammal species composition is determined by the complexity of the understorey (Catling & Burt 1994; Maitz & Dickman

2001: Gardner et al. 2007; Fox & Monamy 2007). Fragmentation of the forest cover must impact the population viability and structure of forest dwellers by changing connectivity of populations (Dept of Ag. 2013). Today fifty percent of Australia’s forest has been completely

5 cleared or modified (Bradshaw 2012). Bush rats still remain a common species due to their broad diet and generalist habitat range where species with specialist habitat and diet are more likely to be threatened though no species can be considered entirely free of threats (Dickman et al. 2000).

Reintroduction of native fauna is a major goal in conservation and restoration as habitats face increasing human induced ecosystem changes (Seddon 2010). The ability to re-establish populations of endangered or threatened species has potential to be a powerful conservation tool (Fisher & Lindenmayer 2000). Though understanding the root causes of native animal extinctions is critical for implementing successful reintroduction or translocation programs

(Dodd & Seigel 1991). Causes may be by one or all of a combination of factors such as the reduction in habitat area, changes to the structure of remaining habitat fragments and/or the introduction of competitive non-native animals (Gardner et al. 2010). By examining the habitat that supports native fauna we should understand how to replicate these habitats and manage them so that reintroduction and translocation programs have a greater success rate.

2.1.2 Impacts of land clearing on native vegetation

Urbanisation and human induced changes in land use have precipitated the decline and local extinction of native fauna and flora (Tilman et al. 1994). Habitat fragmentation along the coastal areas of Australia presents ecological, economic and conservational challenges

(Gardner et al. 2007; Fleming 2013). These challenges have developed as size and composition of human populations have expanded in coastal areas with human induced changes placing extra pressure on land sensitive to changes (Gurran et al. 2007). Agriculture was once the main cause of land clearing in Australia, now urbanisation is modifying coastal habitat (State of the Environment Committee 2011). Urban areas occupy a small part of the

6 country but have a pervasive influence on the natural environment. Expanding settlements and infrastructure will continue to impact on the environment with a great deal of uncertainty about how species and ecological systems will cope with cumulative effects of development along coastal strips. Economic growth places demands on natural resources with a major impact on biodiversity which has declining since European settlement. The legacy of land clearing effects is said to last for years or even decades (State of the Environment Committee

2011; Hahs & McDonnell 2014). Current research recently shows a rapid decline in small mammal species in northern Australia with the impact of changed fire regimes and invasive species (McDonald & Woinarski 2011). Fragmentation of remnant vegetation and habitat in

Australia has impacted on fauna populations altering the in situ structure, composition and spatial pattering within the landscape (Garden et al. 2007).

2.1.3 Mammal translocation and reintroduction

Previous studies of translocation and reintroduction of species found success and failure varied dramatically between taxa in different trophic levels and life cycle stages when relocated (Griffith et al. 1989; Burke 1991; Dobb & Seigel 1991; Sheean et al. 2011). These reports show that despite decades of research, even defining what is successful in a reintroduction is not clear. Seddon (1999) questions if we really can define successful reintroduction or should we even try. There is no general agreement on the definition of success for species relocation, though the aim of the International and Australian programs is to establish a viable, self-sustaining population (IUCN 2012; AWMS 2014). Then how do you define standards or criteria to measure these factors over long term monitoring or population increase?

Reintroduction success is influenced by effective management and organisation of the program, removal of competitors and type of release (Sheean et al. 2011) while failure can be

7 due to lack of finance, resources and unpredictable environmental factors (e.g. drought). In an

11 year translocation program of hackings (chick translocation) on islands off the Iberian

Peninsula researchers found it was more important to consider evolutionary life history and detailed information on habitat quality (Oro et al. 2010). This study and others showed that translocation programs were very likely to fail if lower quality habitat was targeted (Sheean et al. 2011) especially for highly dispersing nomadic animals (Oro et al. 2011).

2.1.4 Habitat Characteristics

Habitat characteristics are one of the main factors influencing the success of animal reintroductions (Griffith et al. 1989; Sheean et al. 2007; Rantanen et al. 2010). I will test habitat requirements of a common mammal species Rattus fuscipes (the bush rat) in its natural environment. Successful reintroductions require an understanding of how individual species respond to changes in the vegetation community (Kearney et al. 2007). Landscape composition and configuration to patch size and local scale are important to habitat variables for small mammal assemblages and reptile populations (Garden et al. 2010). Mammal distribution and association with home ranges on a landscape can be limited by what resources are available in that landscape (Mitchell & Powell 2004).

The bush rat habitat in eastern Australia is diverse (Lunney, 1991) while variables that define bush rat habitat can differ. Some populations have been found to have a positive correlation with leaf litter, vegetation density and period since the last fire (Fox & Monamy 2007) while others found habitat variables of fallen logs and mid story vegetation density (Spencer et al.

2005). Generally there appears to be a habitat association with dense ground cover where bush rat populations can reach densities of around 10 individuals per hectare (Maitz &

Dickman 2001). This previous research suggests that there are likely to be key habitat

8 characteristics that support bush rat populations. While no research so far has evaluated the importance of a wide range of habitat characteristics that identify suitable habitat requirements for successful reintroductions. Good quality habitat or optimal habitat preferences (Fox & Monamy 2007) will assist translocation of species for they depend on these factors for survival, reproduction and importantly persistence in a habitat.

Establishment of an animal in a region rely in physical factors such as temperature, light, water and biotic factors that provide food supply (Villee et al. 1985). Cover is important especially for protection from predators where space can be important for reproduction and growth. Habitat selection is a specialised process and so understanding the optimal habitat preferences of a common mammal species the bush rat may inform us how to adapt for the particular combination of features found in that habitat. While shelter, food, and water are basic requirements, wildlife survival knowledge of these requirements can be used for translocation of species especially in fragmented habitats.

I test the predictions that bush rat population density will be those associated with 1) ground cover 2) land elements, 3) light levels, 4) diversity of plant communities and 5) structure of the canopy. Past research indicates that dense ground cover is important for small mammal populations (Goertz 1964: Fox & Monamy 2007; Gardner et al. 2007). Testing what variables of the ground cover or vegetation community are significant will assist persistence of the species. Previous research of ground cover density was successfully tested by the use of light readings. Measurements were taken above vegetation and at ground level to assess density.

The assumption was that leaves and stems contributed to vegetation density and that lower layer was important for small mammal habitat (Fox & Monamy 2007; Kearney et al. 2007).

Land elements of creek distance, gully, lower, mid, upper slope and ridge were tested because there was little evidence that these factors had been tested in the past for bush rat population density. Vegetation is known to vary over these different elements. Aspect was tested as it is known to vary from fieldwork where vegetation changes occur with different aspects. Early research showed floristic as well as structural habitat requirements determine small mammal habitat (Braithwaite & Gullan 1978) though (Wilson et al. 1986) found none of the species of small mammals showed an overall preference for any floristic or structural vegetation types.

Though conflicting information regarding floristic had been found, testing this information may clarify vegetation community and be a factor in defining available food and cover for why bush rats persist in certain habitats. Determining which of these habitat components are important to bush rat presence and absence presented the need to collect as much field data as possible covering all these areas.

Understanding the key characteristics of species that are adaptable, such as the bush rat which is capable of living within or close to city boundaries (Breed & Ford 2007) we can cater to the needs of these species and not just provide green spaces that fail to support their continued existence.

2.2 Methods

2.2.1 Study area

The study was carried out in the Sydney Basin Bioregion (insert Fig 2.1) on the central east coast of NSW (NSW NPW&S 2003). The Sydney Basin is one of the most species diverse regions in Australia (NSW NP&WS 2003) but it also has one of the greatest threats, urbanisation. The study area is also part of the main distribution area of bush rats along the east coast of NSW.

2.2.2 Study sites

Three study areas were selected within the Sydney Basin, Ku-ring-gai Chase National Park,

The Royal National Park and Heathcote National Park each on the edge of the urban areas of

Sydney. These areas were selected because they represent large areas of relatively undisturbed vegetation reflecting Sydney’s distinct bush land character which is disappearing because of urbanization. Royal National Park, Heathcote and Ku-ring-gai National Park were selected as bush rat data from previous trappings for fox bating showed high numbers of bush rats, M. Hall (personal communications 2009 Ranger Ku-ring-gai Chase National Park). The

NSW Wildlife Atlas also showed the presence of bush rats. Data were also collected from

National Park survey records over the last 10 years.

Fig.2.1 Map of the central east coast of New South Wales showing the location of Sydney with Ku-ring-gai National Park to the north and The Royal and Heathcote National Park to the South within the Sydney Basin highlighted in red on the map of Australia.

Ku-ring-gai Chase National Park (33⁰39’3.6”S, 151⁰12’3.6”E) is located within the

Sydney metropolitan area and surrounded by medium density and semi-rural properties

(Fig.2.2). The parks boundaries are defined by the F3 Sydney-Newcastle Freeway to the western and the Hawkesbury River and tributaries to the north with Pittwater to the east and

Mona Vale Rd to the south. The Park covers 14,709 ha and was created in 1894 and was once the traditional lands of the Guringai Aboriginal People.

Fig 2.2 Map of the survey area with the 9 sample sites marked with black dots. The red triangles represent records of Rattus fuscipes bush rats from Atlas of NSW Wildlife records

2012. Red line marks the boundary of Ku-ring-gai Chase National Park.

Urbanisation is placing increased pressure on Ku-ring-gai Chase National Park due to the growth of Sydney’s metropolitan area and the estimated 6,500 surrounding neighbours (NSW

NPWS 2002a). The area is part of a major structural unit of Permian and Triassic age which consists almost entirely of horizontally bedded sedimentary rocks (NSW NPWS 2002).

Hawkesbury Sandstone is the most extensive rock type in the park and forms the plateaus and hill slopes of the national park. Plant communities consist of over 1,000 plant species with

Ku-ring-gai listed on the National Heritage register for its natural values and important from a conservation perspective as it contains relatively undisturbed vegetation that is significant in a regional and local content (Benson & Howell 1994). Urban development is taking its toll on the Park with 36 animal species and 27 plant species in the region declared threatened or

13 endangered under the Threatened Species Act 1995. Impact on the bushland are from nutrient enriched run-off, sewage overflows, high flow stormwater, soil disturbance, vegetation clearing, dumping of fill and garden waste and garden escape plants. Other major impacts are from a high demand for hazard reduction burning for surrounding property protection.

Mammal trapping has been carried out in Ku-ring-gai to capture the northern limit of the population of southern brown bandicoot. This trapping program has also captured other common small mammals. For the years 2009-2011, bush rats have been the most common small mammal found from the annual trapping program (personal observation). I have been involved in the trapping over this period providing evidence of habitat sites to survey.

Suitable sites where observed when captured bush rats were released after overnight trapping and scampered to nearby hollows in the ground. Trapping carried out in Ku-ring-gai National

Park during the period 2009 till 2011 consisted of one Elliott aluminium trap and one wire cage possum trap at each census station. Batting in each trap consisted of peanut butter. Trap period consisted of 5 consecutive nights over 2 weeks. Captured bush rats were recorded but no details of weight, sex, juveniles or reproductive conditions were obtained before releasing.

The Royal National Park (34⁰ 7’21” S, 151⁰ 3’50” E) is adjacent to the southern limits of the Sydney metropolitan area bounded by the suburbs of Cronulla, Sutherland, Engadine and

Heathcote (Fig.2.3). The park is located approximately 30 km south of Sydney covering an area of just over 15,000 ha with its eastern and northern boundaries defined by the salt water of the Pacific Ocean and Port Hacking respectively with Princess Highway to the west and

Garawarra State Recreation Area to the south.

Fig. 2.3 Map of the survey area with 13 sampling sites marked with black dots. Red triangles represent records of Rattus fuscipes bush rats as held in Atlas of NSW Wildlife records 2012.

Red line marks the boundary of Royal National Park and Heathcote National Park, surrounded by the towns of Bundeena, Helensburgh and Heathcote.

Heathcote National Park (34⁰ 07’49”S, 150⁰ 58’17⁰ E) covers 2,251 ha and lies to the west of the Royal National Park bounded by Holsworthy Army Range and Sydney Catchment

Authority land on its eastern side (Fig 2.3). The Royal and Heathcote are part of the

Woronora Plateau, which lies east and south of the Cumberland Plain (NSW NP&WS

2002b). Steep slopes, ridges, rocky outcrops and streams punctuated by small waterfalls and pools are a feature of the landscape (Fairley 1995). The geology is predominantly

Hawkesbury Sandstone while in the upper Hacking River valley and the Southern Coastal escarpment of the Park areas of Narrabeen Shale are exposed. These sheltered sections and

15 coastal valleys of rich, shale-derived soils support pockets of closed moist forest and rainforest (NSW NPWS 2002b).

The climate statistic for the survey sites was taken from Sydney as climate variation between sites was minimal. Generally average maximum temperature 21.7⁰C and average minimum is

13.8⁰C (Australian Bureau of Meteorology 2013). Mean annual rainfall is 1214.1mm/yr.

These figures have been estimated over the last 154 years (Australian Bureau of Meteorology

2013).

Major plant associations in The Royal and Heathcote National Parks include wet and dry heathlands shrublands, woodlands, open forests, tall open forest, warm temperate, subtropical and littoral rainforests estuarine and freshwater wetlands and coastal dunes (Andrews 2001).

The Royal has 1000 plant species recorded, including 26 species listed nationally as rare or threatened. Many of the plant species in The Royal and Heathcote are fire sensitive and rely on seed stock stores in the soil or need to recolonise from surrounding areas after fire. The opportunity for recolonising is now drastically reduced because of surrounding urbanisation

(NSW NP&WS 2002).

A survey carried out by NPWS (Andrews 2001) found 247 species of native vertebrates, comprising of 16 frogs, 37 reptiles, 164 birds and 30 mammals showing a decrease in species to the previous survey which showed 358 species with 20 frogs, 40 reptiles 268 birds and 30 mammals. Good populations of 152 bush rats R. fuscipes were surveyed at 21 sites during the

2001 report D. Andrew (personal communications 27 April 2010).

The vegetation communities that were surveyed in this report are grouped into vegetation community profiles based on Sydney Metropolitan Catchment Management Authority

(SMCMA) region (DECC 2009) and listed in Table 2.1 & Table 2.2.

2.2.3 Method of Survey site selection

The Atlas of NSW Wildlife was used to provide full reports of selected areas for detailed information from the Office of Environment & Heritage database (www.bionet.nsw.gov.au).

Information of geographical areas from the database showing numbers of both high and low bush rat density and habitat types was used to identify field sites. High density populations are areas where 10 or greater bush rats were captured in recent trap line sampling. Low density areas where less than 5 bush rats were captured in recent trap samplings. The main area of observation work was associated with site characteristics, vegetation structure, plant species and plant communities (refer appendix Table 2.3 Vegetation Survey Field Sheet). The field work involved setting up quadrats at all sites within Ku-ring-gai, The Royal and

Heathcote National Parks. Surveys were carried out at 22 quadrats, 10 m2. Selections of quadrats were based on map location and GPS readings from National Parks Records accessed from their data base. Sites were selected randomly within areas where bush rat populations were known to exist in the three National Parks (Fig 2.2 & Fig 2.3). I verified the precision and accuracy of the GPS coordinates by checking the coordinates on topographic and orthophoto maps 1:25000 of each trap site. Locations were then checked with information on the National Parks data base where a short description of site locations could be verified to ground truths. These locations were chosen from NP&WS data to represent a sample of the vegetation within the home range of many bush rats trapped over a 10 year period. The 10m² survey sites were chosen due to the abundance of vegetation at these recorded sites and to provide a detailed description of the habitat where bush rats are

17 recorded. A larger scale may not have provided the plant species and structure detail. Future research may look at defining suitable spatial scale for small mammal habitat.

2.2.4 Field methods

Site selection was based on the findings that there were few if any sites within the central parklands habitats where bush rats have not been found, but in the sites where they are found some sites supported high densities and others low densities. Vegetation surveys were conducted during the Australian autumn and winter months (April through August).

Vegetation surveys were conducted during this period due to the suitability of time to spend collecting field data. Flowering times vary between species and seasonal conditions. Many plants in autumn still have seeds from previous spring season and many Australian native plants flower in winter. Plants with seeds and flowers assist identification. This period also allowed time to return in spring if further identification was needed. March and June are said to be Sydney’s most reliable rain period with rainfall highest in June (Australian Bureau of

Meteorology 2013).

Overall I selected 22 field sites, 14 of these sites supported high density bush rat populations, and 8 supported low density bush rat populations (Précis Table 2.4 & 2.5).

Table 2.4 Précis record of Ku-ring -gai Chase National Park data for complete T able 2.4 refer appendix.

average of 4 Light Total Height class Leaf % High / readings at each level No. No. No. % no. litter Fallen Soil Site Low bush plant plant plant Leaf bush depth timber pH rat no. species genus families litter rats cm >5cm 2003- ground 1.2m 2m lower mid upper 2010 level

K1 High 45 30 341 530 26 21 12 1 3 7 65 3 40 4.5

K6 High 15 142 160 238 32 26 15 1 3 15 50 20 35 6

K10 High 44 43 50 125 16 15 10 2 3 5 50 40 30 4.5

K15 High 25 29 334 514 26 21 14 2 3 15 30 15 45 5

G3 High 48 44 184 158 41 28 23 1 5 7 50 20 25 4.5

G20 High 49 16 40 57 27 23 14 0.5 3 7 45 40 10 4.5

G27 High 18 38 61 59 24 21 15 1 2 15 40 15 25 5

G28 High 31 116 210 242 28 23 12 1 5 15 50 20 45 4.5

G34 High 43 37 85 103 29 26 17 1 5 25 45 20 50 5

Table 2.5 Précis record of The Royal National Park & Heathcote National Park data for complete Table 2.5 refer appendix.

average of 4 Light Height class High / Total Leaf % readings at each level No. No. No. % Low no. litter Fallen Soil Site plant plant plant Leaf bush rat bush depth timber pH ground species genus families litter no. rats 1.2m 2m lower mid upper cm >5cm 2003- level 2010

1(12) Low 1 11 33 89 17 12 8 1 3 15 15 2 15 5

2(16) Low 1 37 180 273 18 13 7 1 3 15 70 2 5 4.5

3(17) Low 1 16 95 609 16 13 9 1 2 3 15 1 0 5

4(15) High 32 3 4 5 35 21 13 1 3 15 45 4 30 4.5

5(11) High 39 3 4 5 23 21 21 1 5 25 70 3 25 4.5

6(10) Low 1 23 149 152 24 23 20 1 5 25 40 10 20 5

7(8) Low 1 6 8 9 21 18 14 1 5 25 35 8 20 6

8(4) Low 1 48 260 234 44 30 12 0.5 2 15 10 2 15 4.5

9(7) Low 1 7 6 12 21 17 15 1 2 7 45 2 20 6

10(5) Low 1 17 53 273 41 31 17 0.5 3 15 30 10 40 5

11(5C) High 10 15 20 22 29 27 22 0.5 3 10 45 15 20 4.5

12(7A) High 13 24 27 22 19 18 17 1 3 20 30 10 30 5

13(8) High 10 17 36 25 23 10 1 5 0 70 40 20 5.5

Site Characteristics

The site characteristics measured at each quadrat consisted of: aspect, contour level, land element, distance from the nearest creek, percent bare earth, percent rock cover, percent leaf litter, leaf litter depth, percent fallen timber, soil pH and soil texture (A complete version of

Table 2.4 & Table 2.5 can be found in the appendix). That is for each variable bare earth, rock cover, leaf litter, fallen timber a standard area of 1m² in the quadrate was determined and a percentage was estimated from this reference point. To standardise the measurement only one person was used to measure all sites to give consistence of results. Leaf litter depth was measured with a field measuring tape at various locations in each quadrate and averaged.

Soil pH was measured using a Manutec pH Soil Test Kit (Manutec, South Australia).

Sampling of pH can be difficult to interpret as pH is very sensitive to plant differences as the pH under one plant may be quite different from that under another. To avoid this sampling was taken at 4 different locations in each quadrate. Soil texture was determined at each site

19 by forming a soil bolus from 4 different locations in each survey quadrate and measures from the reference categories using the Field Handbook (McDonald et al. 1990).

Vegetation layers

Plant height was used to determine vegetation layer. A grouping of 3 vegetation layers was used to break down the quadrate in a 3 dimensional form by height to determine plant community and plant species present (appendix Table 2.3 Vegetation Survey field sheet).

This grouping consisted of;

a) Lower <0.25m, 0.25-0.5m, 0.5-1m

b) Middle 1-2m, 2-3m, 3-5m

c) Upper layers 5-10m, 10-20m, 20-35m

Three height layers or growth forms of vegetation (lower, middle and upper layers) were further divided into layers as no absolute height class for all growth forms occur in Australia

(McDonald et al. 1990). This analysis helps group cover into height classes that can be used to define vegetation structure. This is based on visual estimates of horizontal vegetation layers. For each of these vegetation layers plant species and plant communities were identified. At the 3 different vegetation layers plants were identified and keyed out to genus species and family within each layer a cover score of 1 to 6 was given where the value for each score was 1(<5%), 2(5-10%), 3(10-30%), 4(30-50%), 5(50-70%), 6(70-100%).

Each layer has an important influence on the amount of light that is transmitted through the layer. To measure the effect of filtered light on plant growth a simple test was carried out at 3 levels: ground level, 1.2 meters and 2 meters on the 4 corners of each quadrate. In the field experiment I measured light only at a single point in time as a guide to understand the

20 influence of light reaching the two lower stratum levels of the quadrates. Light intensity was measured using a LiCorr 2000 hand held light meter.

2.2.5 Vegetation Community Profiles

Each of the survey site plant lists were compared and grouped accord to the floristic profiles from The Native Vegetation of the Sydney Metropolitan SMCMA region profile sheets

(DECC 2009) Table 2.1 and 2.2 list the vegetation classification at all 22 survey sites. All vegetation layers at each survey site were initially given a range of percent cover then grouped to a cover score between 1 and 6.

Table 2.1 The Royal National Park landscape and vegetation characteristics based on SMCMA region (2009 DECC) Vegetation Community Profiles

Location Survey site Vegetation class (SMCMA region) Royal National Site 1 Carlot Coastal Sandstone Heath-Mallee Park Sydney S_HL08 Site 2 Flat rock ck Coastal Sandstone Sheltered Peppermint – Apple Forest S_DSF09 Site 3 Mt. Bass trail Coastal Sandstone Heath-Mallee S_HL08 Site 4 Sir Bertram Stevens Coastal Sandstone Exposed Drive Scribbly Gum Woodland S_DSF05 Site 5 Bola Ck. Coastal Warm Temperate Rainforest S_RF03 Site 6 Upper Causeway Coastal Warm Temperate Rainforest S_RF03 Site 7 Palm Gully Coastal Warm Temperate Rainforest S_RF03 Site 8 Kingfisher Ck. Coastal Sandstone Sheltered Peppermint-Apple Forest S_DSF09 Site 9 Otford Coastal Headland Littoral Thicket S_RF08 Site 10 Stanwell Tops Coastal Sandstone Exposed Scribbly Gum Woodland S_DSF05 Site 11 Lilyvale Track & Coastal Sandstone Exposed Coast Track Scribbly Gum Woodland S_DSF05 Site 12. Lady Carrington Coastal Warm Temperate Drive Rainforest S_RF03 Site 13. Garie Rd. & Coastal Sandstone Sheltered Garawarra Hill Rd. Peppermint – Apple Forest S_DSF09

Table 2.2 Ku-ring-gai Chase National Park landscape and vegetation characteristics based on SMCMA region (2009 DECC) Vegetation Community Profiles

Location Survey site Vegetation class (SMCMA region) Ku-ring-gai Site 1 (K1) Centre track Coastal Sandstone Sheltered National Park Peppermint – Apple Forest Sydney S_DSF09 Site 2 (K6) West Head Rd Coastal Enriched Sandstone 800 m from gate LHS at Sheltered Forest S_DSF04 bandicoot sign

Site 3 (K10) Towlers track Hornsby Sandstone Heath- 5m from West Head rd Woodland S_DSF12 Site 4 (K15) Chiltern Track Hornsby Sandstone Heath- 50m from McCarrs Ck rd Woodland S_DSF12 Site 5 (G3) Ryland Track Coastal Sandstone Sheltered RHS 1k from gate. Peppermint-Apple Forest S_DSF09 Site 6 (G20) Heath Track Hornsby Sandstone Heath- Woodland S_DSF12 Site 7 (G27) Ryland / Coastal Sandstone Sheltered Cooyong Trail 10m from gate Peppermint – Apple Forest S_DSF09 Site 8 (G28) Ryland / Hornsby Enriched Sandstone Cooyong 300m from gate Exposed Woodland S_DSF10 Site 9 (G34) Cascade track Coastal Sandstone Sheltered Peppermint-Apple Forest S_DSF09

2.2.6 Data analysis

I used simple logistic regression analysis to test the significance of each environmental variable in predicting poor or good bush rat habitat. I then analysed the significance of all variables in a stepwise logistic regression model. This model was selected as I wanted to assess the successive contributions of environmental variables to predicting bush rat habitat where it appeared that, following the simple logistic regression, only a small number of environmental variables were significant. To control for increased variation sometimes associated with stepwise logistic regression, I also ran a forced entry regression model. The results between the two multiple regression models did not differ. In addition, to control for

23 biases in the results to correlations between predictor variables, we assessed the pair wise relationships between variables using Pearson correlation coefficients. Where two variables were significantly correlated, the weakest predictor variable was removed from a new stepwise logistic regression model. All analyses were conducted using SPSS version 21

(IBM Statistics, NY).

2.3 Results

I found the only significant factor in predicting bush rat population density was percent fallen timber (Logistic Regression Model P<0.05, Table 2.6. and Fig. 2.4a).

Overall, the environmental variables were not correlated with one another. Some variables

(percent rock versus percent bare ground; height of the upper canopy versus site elevation; species richness versus height of the lower canopy) were correlated. Removing correlated variables from the stepwise logistic regression model did not change the significance of the other predictor variables.

Table 2.6 Stepwise Logistic Regression Model showing the significant variable Percent timber that affects the predictive power of the model. Model statistics include the constant

(B), Standard Error (S.E.), Wald’s test statistic, degrees of freedom (d.f.), P, and the regression exponent (Exp).

Variables in B S.E. Wald d.f. P Exp(B) equation

Percent timber -0.236 0.105 5.013 1 0.025 0.790

Constant 1.660 0.955 3.023 1 0.082 5.261

Step 1a Variable(s) on Step 1: Percent fallen timber

High

Bush rat density rat Bush

Low

0 10 20 30 40 % Fallen Timber

High

Bush rat density rat Bush

Low

0 20 40 60 80 100 % Leaf Litter

Fig. 2.4 a) & b) Logistic regressions demonstrating a significant (P = 0.025) relationship between bush rat density and percent fallen timber cover across sample plots (a), and a marginally significant (P = 0.05) relationship between bush rat density and percent leaf litter across sample plots (b). Symbol size is presented relative to the number of overlapping data points for a given value of percent fallen timber or leaf litter (smallest points represent a single sample plot and largest point’s represent four overlapping sample plots). The “noise” is just due to the variable cover in the habitats. The logistic regression quite clearly shows a

25 trend (a step) in bush rat presence at relatively high percent fallen timber and percent leaf litter.

Table 2.7 Stepwise Logistic Regression Model shows variables that do not significantly affect the predictive power of the model but ranks each variable in order of importance to bush rat habitat.

Variables not in the multiple logistic regression Equation

Variables Score d.f. P

percent leaf litter 3.836 1 0.050

soil pH 3.771 1 0.052

distance from water 2.024 1 0.155

percent rock 1.963 1 0.161

lower layer 1.148 1 0.284

no. family species 0.376 1 0.540

percent bare earth 0.343 1 0.558

upper layer 0.255 1 0.614

elevation 0.098 1 0.755

light reading at 2m 0.016 1 0.900

mid layer 0.014 1 0.906

light reading at ground 0.005 1 0.942

no. of plant species 0.000 1 0.990

Overall Statistics 13.437 13 0.415

Step 1

The remaining variables (Table 2.7) were not significant in predicting bush rat populations.

However, leaf litter was (P = 0.05, Fig. 2.2b) with soil pH (P = 0.052).

Vegetation structure variables and cover score

Ku-ring-gai Chase National Park lower layer ground cover score varied from 3(10-30%) to 6

(70-100%) (Table 2.4 showing précis record of Vegetation Structure Variable & Cover Score of complete Table 2.4 in Appendix). While the lower layer ground cover score at The Royal

& Heathcote National Parks varied from 2 (5-10%) to 6 (70%-100%). Ku-ring-gai Chase

National Park is renowned for high bush rat numbers and survey sites consisted mostly of

Sandstone Heath Woodland to Forest Vegetation (Table 2.2). The Royal survey sites varied from Rainforest to Mallee Heath Vegetation (Table 2.1). The upper layer canopy in Ku-ring- gai Chase National Park overall cover score varied between 2 (5-10%) and 5(50-70%) while

The Royal and Heathcote varied from 0 to 6 (70-100%). These values indicate that only in 1 of the sites surveyed no upper canopy was measured or had less than 5% canopy cover.

Mid layer in Ku-ring-gai National Park showed little variation between sites with cover score varying from 3 (10-30%) to 6 (70-100%) while The Royal and Heathcote gave very similar results of 2(5-10%) and 6 (70%-100%).

Table 2.4 Précis records of Vegetation Structure Variable and Cover Score.

<5 5- 10- 30- 50 - 70 - Percentage Vegetation Cover 0 % 10% 30% 50% 70% 100% Cover Score Lower Layer 0 1 2 3 4 5 6 Ku-ring-gai Chase National Park 0 0 0 1 1 2 5 The Royal National Park 0 0 2 4 3 2 2

<5 5 - 10 - 30 - 50 - 70 - Percentage Vegetation Cover 0 % 10% 30% 50% 70% 100% Cover Score Middle Layer 0 1 2 3 4 5 6 Ku-ring-gai Chase National Park 0 0 0 3 2 3 1 The Royal National Park 0 0 1 1 4 3 4

<5 5 - 10 - 30 - 50 - 70 - Percentage Vegetation Cover 0 % 10% 30% 50% 70% 100% Cover Score Upper Layer 0 1 2 3 4 5 6 Ku-ring-gai Chase National Park 0 0 1 2 3 3 0 The Royal National Park 1 1 1 3 1 2 4

2.4 Discussion

2. 4.1 Bush rat Habitat

The prevalence of fallen timber was the only significant indicator of bush rat density across forest areas in the Sydney region (Table 2.6). My results suggest that very specific aspects of understory complexity such as percent timber with a diameter greater than 5cm provide cover to support sustainable bush rat population. Other variables though not significant may be potentially important for example leaf litter p=0.05 and pH p=0.052 highlighting the need for future analysis to determine their merit to bush rat habitat. The study sites are in vegetation communities typical of natural native bush areas ideal habitat to study the bush rat’s ecological niche. Bush rats were never completely absent in these sites. The low density sites tended to only have recorded a single bush rat while the high density sites recorded 10 or greater bush rats. While bush rats were found throughout the sampled areas, and certainly numerous factors such as food sources and plant community type will likely influence their distribution. Also different scales of the environment or hierarchy preferences (Johnson 1980) could impact on the distribution of the bush rat in different ways. Their home range tends to be less than half a hectare with small territories due to abundant local food (Breed & Ford

2007). Male bush rats travel longer distances than females which may be up to 1km. Bush rats are omnivores there food source consists of around 6 types of foods including seeds, green plant material, terrestrial invertebrates, fruit, nectar and fungi (Dickman et al. 2000).

However, within the habitats tested I found a strong relationship between bush rat densities with fallen timber and to a lesser degree leaf litter in the understory (Table 2.7 & Fig. 2.4b).

Habitat is the physical area where an animal lives and survives, with persistence of habitat the key to persistence of species (Fortey 2012). The study sites are important because they represent 10 years of established bush rat habitats in 3 National Parks surrounded by

Sydney’s urban growth. Bush rats are not threatened or a rare species and commonly trapped

28 in annual fauna surveys so age, sex or numbers of juveniles are not recorded. Urban development in Sydney has obviously shifted or moved bush rat populations to remaining habitats surrounding Sydney.

2.4.2 Ground cover density

Ground cover is particularly important for small mammals as it provides a microhabitat to avoid predator and other species (Cox et al. 2000). The formation of the ground layer is also essential for shelter and food supply. To support a community, ground cover needs to have a survival value (Villee et al. 1985). This survival value means it provides adequate food supply, nesting material, cover for young and protection from general predators & other species, leading to persistence in the habitat. The understanding of cover complexity, structure and density is important to assist management of on-ground restoration projects and translocation of species. As habitat quality is rarely straightforward, (Armstrong 2005) measurement of variables need to fit with species concerned at individual sites to have meaning for habitat quality. Implementing the right strategies can be critical for success to quality habitat establishment.

The results of my study demonstrate that percent timber is ecologically significant enough to provide a suitable habitat for bush rat numbers. The variable percent timber used in conjunction with other variables would assist land managers to evaluate presence of bush rats. The presence of timber on the ground has to be influenced by upper and mid canopy cover and as measurements of timber was >5cm, most of these branches would be fallen branches from shrubs and trees. Fire frequency could also influence bush rat distribution, but fire was not investigated in this thesis. Fire sensitivity is a factor to consider with management of species but how sensitive populations are needs further research.

Leaf litter results showed a pattern of association and may also have a positive input into bush rat habitat (Fig 2.4b). A balance of leaf litter accumulation would be important as too much litter would block seedling recruitment and species movement. The depth also has implications for management as unfortunately leaf litter and timber are usually the first thing to be cleared from an area near urban population due to fuel loads and the possibility of an out of control fire in difficult to access sites (Sturtevant et al. 2009; Penman et al. 2013). Leaf litter is important for cover but also for food supply, providing biodiversity for insects and fungal growth and litter dependent fauna (Goldstein 1984). Eucalyptus leaf litter can decompose quite rapidly depending on climate conditions and may provide ideal conditions for fungal growth. The bush rat is known to feed on fungi, insects and seeds depending on season and availability (Carron et al. 1990; Tory et al. 1997).

Bush rats have been associated with woodland habitats (Maitz & Dickman 2001) which may result from a need for protection from above and a dense ground cover for protection on the ground. Past research work has show identification of habitat association with bush rats was at the lower strata layer implying that this layer was s factor of bush rat habitat. Factors varied such as, cover and friable soil (Taylor & Calaby 1988), vegetation complexity in lower canopy (Dickman & Steeves 2004), low leaf litter, low ground cover low understorey cover

(Lindenmayer et al. 2005; Spencer et al. 2005), a strong relation with leaf litter rather than vegetation density and time since last fire (Fox & Monamy 2007), and logs (Claridge et al.

2008).

2.4.3 Other structural variables

The 4 structural variables that showed no pattern of significant association were % rock, lower layer, upper layer and mid layer and formed a decreasing list of importance (Table 2.7).

Each structural variable though not significant has merit for bush rat habitat as each variable also helps form the structure and complexity of cover density.

The presence of rocks forms a structural feature for protection and camouflage but from my results not critical for bush rats. Generally rock cover found at all sites was low with only one site found to have high percentage of rock cover this may be a reason why rock cover was not found to have an association. Rocks and leaf litter provide a forest floor microhabitat for invertebrate’s (Hickerson et al. 2012). Invertebrates were not investigated in this research and may be an area of future research. Rocks do provide protection for burrowing or tunnels to burrows (Fleming 2013). There was also no association between the lower layer and bush rat density but obviously an important variable in habitat quality. Dense ground cover has advantages for wildlife but disadvantage for managers of urban vegetation due to the fear of fire and the Australian bush. In summary the majority of the sites had thick upper and mid canopies providing material for a complex ground cover. Vegetation at Ku-ring-gai had generally heath type vegetation in mid and upper layers while The Royal and Heathcote vegetation varied, with rainforest, woodland and heath sites.

None of the vegetation layers had an effect on bush rat density. The lower level ranked higher than the upper and mid layers in association. Dense vegetation assists ground-dwelling animals, with food, nesting opportunities and protection from agonistic conspecific and heterospecific encounters (Monamy & Fox 2000). The upper layer was found to be more important than mid layer this may be due to protection and a source of understory complexity. Analysis of this characteristic is complex but illustrates the theory that bush rats are associated with woodlands (Maitz & Dickman 2001) and if there was a sparse upper layer limited timber and leaf litter would be found on the ground layer.

2.4.4 Floristic variables

Plant family richness and plant species richness showed no association in predicting the presence of bush rat populations. Similarly, Garden et al. (2007) and Monamy and Fox

(2000) did not find significant associations between small mammal populations and plant taxonomic diversity. Average numbers of plant taxa per survey site for Ku-ring-gai N.P. were

15 plant families, 23 plant genera and 28 plant species. While numbers at The Royal &

Heathcote N.P. were similar 14 plant families, 21 plant genera and 26 plant species per site.

When comparing total plant numbers for all sites surveyed in The Royal & Heathcote

National Park the lower canopy had a diverse range of 50 plant families. The mid layer of total plant numbers had a substantial drop down to 19 families with the upper canopy consisted of just 11 families.

A detailed identification of all plant species was recorded at every survey site (appendix

Table 2.8 & 2.9). Floristic variables also had no pattern of associated and may be due to the fact that seeds are not a major component of bush rat diet. Bush rats are omnivore with around 6 types of food consumed including seeds, green plant material, terrestrial invertebrates, fruit, nectar and fungi (Dickman et al. 2000). My surveys involved detailed florist identification but I was looking at general plant species not specific species as a component of bush rat diet.

2.4.5 Environmental variables

The six environmental variables assessed were soil pH, distance to water, % bare earth, elevation, light readings at 2m, and at ground level. Soil pH may influence vegetation (Begon et al. 1995) though no pattern of association was found with bush rat presence. Soil pH can be an indicator of plant communities due to available nutrients in the soil. The pH scale is a measure of soil acidity and alkalinity. Soil pH influences availability of nutrients to plants,

32 and is a major factor controlling the distribution of many plant species, pH is also related to plant community diversity. Therefore pH can be related to the presence of species used as a food source or cover for bush rats. While soil pH is an environmental factor which varies due to space and time and may be changed by the presence of plants, temperature and humidity

(Begon et al. 1995).

Studies in Ku-ring-gai National Park (Le Brocque & Buckney 2003) examined soil physical and chemical characteristics in relation to species richness and found no single measure but a combination of environmental variables that could explain variations in total species richness of shrub, ground cover and to a lesser degree tree species. Reduced nutrient availability in the communities on Hawkesbury Sandstone soils correlates with high shrub and ground cover richness. Tree species richness was thought to be a function of historical factors such as fire and climatic patterns. Garden et al. (2007) found soil compaction was associated with mammal species occurrence but if too hard, compacted soils influenced vegetation floristic.

Soil compaction was not tested in this study only pH results analysed. In south western

Victoria, bush rats are mycophagous, (Tory et al. 1997) while little is known or researched about mycorrhiza symbiosis it is important for Eucalyptus forest where mammals such as the bush rat are thought to play a role in dispersal of mycorrhiza spores for forest diversity.

Soil pH tested across all National Parks sites had a range of 4.5 to 6 indicating soils strongly acid (pH up to 5.5) to acid (pH 5.5 to 6.5), typical of Hawkesbury Sandstone. Soils found with a pH lower than 5.5 are generally low in nutrients especially nitrogen and phosphorus

(Corbett 1969). By world standards Australian soils are particularly low in phosphorus and may contain as little as .02% much of which is unavailable to plants (Corbett 1969). Le

Brocque (2003) studied Ku-ring-gai soils and found no single environmental variable could adequately explain the variation in total species richness but the nature and distribution of the

33 vegetation was strongly related to geology soil drainage and aspect. All study sites in Ku- ring-gai Chase National Park over the 3 years (2010-2012) have recorded high bush rat population where each year the annual trapping is carried out to determine northern limit of the Southern Brown Bandicoot M. Hall (personal communications 15 September 2011

Ranger Ku-ring-gai Chase National Park).

Light availability was fairly constant across the lower levels of the canopy. This is probably due to much of the available light being filtered by the middle and upper canopies. This variable was difficult to estimate as a control was impossible to maintain between sites due to inconsistent weather conditions. The plant species in these communities keep their leaves year-round, and the light levels in a given community relative to other communities tend not change through the year. Thus, the one-off light readings are an appropriate tool to estimate light interception in through a plant canopy. Research by Le Brocque & Buckney (2003) found competition among species for light may lead to the exclusion of species, and reduction in species richness where light availability is low. Species richness at all sites was recorded in the different layers of vegetation but statistical analysis did not show this variable to be significant. The lower layer in the majority of sites I surveyed showed greater richness than the mid and upper layer. This is an area for further research as Le Brocque & Buckney

(2003) also found tree species richness at the local scale to be a function of historical factors such as fire and or climate patterns that also influence diversity at a larger scale. The use of light meter readings at ground level was used to show vegetation density which was expected to be an indicator of small mammal presence (Kearney et al. 2007; Fox & Monamy 2007).

However, density of shed branches and litter were more important in predicting bush rat populations than was the density of canopy cover.

2.4.6 Implications for the reintroduction of bush rats

The loss of foraging mammals such as bilbies, bandicoots and potoroos in Australia since

European settlement was linked to ecosystems decline (Fleming 2013). Several native mammal species have suffered sharp declines since European settlement in Australia which holds the unenviable status of being one of the world's worst countries for mammal extinctions (Firth et al. 2010). Some animals do survive close to human habitation, such as bandicoots, as they adapt and manage to survive predators such as cats and dogs. However, other species don’t adapt to changed habitats and introduced predators (McDonald &

Woinarski 2011).

Do urban bushland remnants provide adequate food supply, nesting material, cover for young and protection from predators? Bush rats are reported to survive close to human habitation but we are currently uncertain of the long-term viability of reintroductions into more urban areas such as Sydney Harbour National Park. One of the aims of the Sydney Conservation program to return the native bush rat to the Sydney region was to test at local level previous indications (Maitz & Dickman 2001; Stokes et al. 2009a) that bush rats could be competitive and compete with black rats. Understanding case studies is important to determine habitat parameters especially in changing ecosystems where habitat is under constant threat by invading species and competition for managing urban space.

In 2010 a project was trailed to reintroduce bush rats to areas of Northern Sydney where habitat complexity still exists. The project offered a novel solution by reintroduction of a native species the bush rat to fill vacated niches and repel re-invaders such as the black rat.

The project also advances a new paradigm of “reinvasion biology” and the fundamental process of population recovery, making significant and novel contributions to population ecology and conservation theory (Banks 2010).

My research linked with bush rat reintroduction by looking at habitat characteristics of the bush rat. The dynamics of the project of “reinvasion biology” have shown successful breeding and provided ongoing monitoring over 3 years. Results found in this study have identified key factors influencing bush rat habitat and discusses the complexity of habitat quality. The persistence of bush rats in reintroduction areas will depend on strategic management of the disturbed habitat. If optimal habitat conditions are not maintained for bush rats in a disturbed and fragmented environment they will be exposed to predators due to lack of suitable cover. Reintroduction scale would depend on habitat suitability, food supply, breeding potential and predators. My research found essential to reintroduction would be preliminary research into suitable habitat before scale of reintroduction could be determined.

2.5 Conclusion

Reintroduction trials are being carried out to test “reinvasion biology” in disjunct habitat fragments where vacated niches are hoped to be filled by bush rats to test population ecology.

I found that there is a relationship between the presence of high density bush rat populations and dense ground cover of fallen timber and leaf litter on the ground. This density of ground cover is needed as a survival value to protect bush rats from predators, and may help to provide an adequate food supply, nesting material, and cover for young. Landscape managers will need to maintain a mosaic approach to manage habitat cover where bush rats are located to achieve the habitat survival value needed to provide persistence of habitat and persistence of the species. Understanding key habitat attributes are essential for the ecology of landscapes and restoring vegetation for conservation translocation or reintroduction programs.

(a) Site 5 (11) Bola Cr. Lady Carrington Drive.

 High density of bush rats.

 High % fallen timber.

 High % leaf litter.

 Coastal Warm Temperate Rainforest

(S_RF03).

(b) Site 4(15) St. Bertrum Stevens Drive.

 High density of bush rats.

 High % fallen timber.

 High % leaf litter.

 Coastal Sandstone Exposed Scribbly

Gum Woodland S_DSF05.

 High density of bush rats.

 High % fallen timber.

 High % leaf litter.

 Coastal Warm Temperate

Rainforest S_RF03.

Plate 1. High bush rat density in The Royal National Park & Heathcote National Park (a) Site 5(11) St. Bertrum Stevens Drive. (b) Site 4 (15) Bola Creek Lady Carrington Drive. (c) Site 12(7A) Lady Carrington Drive (photo Wendy Kinsella).

(a) Site 2(16) Flat rock creek.

 Low density bush rats.

 Low % fallen timber.

 High % leaf litter.

 Coastal Sandstone Sheltered

Peppermint-Apple Forest S_DSF09.

(b) Site 3(17) Mt. Bass fire trial.

 Low density bush rats.

 Low % fallen timber.

 Low % leaf litter.

 Coastal Sandstone Heath-Mallee

S_HL08.

 Low density bush rats.

 Low % fallen timber.

 Low 5 leaf litter.

 Coastal Sandstone Sheltered

Peppermint-Apple Forest S_DSF09.

Plate 2. Low bush rat density survey sites The Royal & Heathcote National Parks (a) Site 2(16) Flat rock creek. (b) Site 3(17) Mt. Bass fire trial. (c) Site 8(4) Kingfisher Creek.

(a) Site 1(K1) Central track.

 High density bush rats.

 High % fallen timber.

 High % leaf litter.

 Coastal Sandstone Sheltered

Peppermint-Apple Forest S_DSF09.

(b) Site 8(G28) Ryland/Cooyong 300m from

gate.

 High density of bush rats.

 High % fallen timber.

 High % leaf litter.

 Hornsby Enriched Sandstone

Exposed Woodland S_DSF10.

 High density of bush rats.

 High % fallen timber.

 High % leaf litter.

 Coastal Sandstone Sheltered

Peppermint-Apple Forest S_DSF09.

Plate 3. High bush rat density survey sites Ku-ring-gai Chase National Park. (a) Site 1 (K1) Centre track. (b) Site 8 (G28) Ryland/Cooyong 300m from gate. (c) Site 9 (G34) Cascade / Cambourne.

Chapter 3 - Competition interactions with black rats limit bush rat distribution within its native range

3.1 Introduction

3.1.1 Understanding establishment of Communities and Competitors

Competition between species is a major factor controlling populations defining patterns of diversity and distribution (Tilman 2004). Competitors limit the distribution of many mammal species (Downes et al. 1997; Holland & Bennett 20011). Patterns of competition between species may have important implications on the diversity and conservation of native species.

The loss of native species from a habitat following land use changes can promote the invasion of non-native species, and these species can be superior competitors in these modified habitats (Downes et al. 1997; Moles et al. 2008). The presence of a competitor can be particularly problematic to reintroduction efforts, since differences in competitive ability in favour of the invasive or unwanted species will likely drive the desired re-introduced species to extinction (Sax et al. 2007). Translocating species into areas with potential competitors, particularly if the competitors are similar in form and resource requirements, can be complex and limit success of establishment (Griffith 1989). With substantial loss of 27 mammal species in Australia over the past 200 years (Richie et al. 2013) understanding competition and habitat use will enhance our knowledge to protect mammal species.

3.1.2 Reintroduction and impacts from competitor species

Could the successful reintroduction of native bush rats be limited by a competitive invasive species in fragmented habitats impacted by development? The aim of translocation is to decrease the risk of extinction by increasing the range of a vulnerable species. The intention also is to yield a measurable conservation benefit at the level of population, species and the

40 ecosystem (IUCN 2013). Poor release site quality increases the difficult challenges of species adjusting to new environments by decreasing fitness and reducing its chances of survival.

Lack of habitat assessment and low knowledge of the impact of invasive competitive species can limit the success of species reintroductions (Griffith et al. 1989; Short 2009; Sheean et al.

2011).

In this study, I examine the evidence for competitive interactions between native bush rats and invasive black rats, and if competition limits the distribution of these species. Bush rat populations have become extinct in many areas, particularly in areas associated with human development (Breed & Ford 2007; Garden et al. 2007; Bilney et al. 2010). These areas have been extensively occupied by the invasive black rat (Breed & Ford 2007; Stokes et al.

2009a). Competitive interactions between bush rats and black rats may limit reintroduction success of bush rats into their former habitats. Recent research has demonstrated that bush rats can fill vacated niches and compete with pest species that have similar niche requirements (Stokes et al. 2009a). In addition, bush rats have shown rapid habitat shifts when competing with swamp rats (Rattus lutreolus) a competitive species approximately the same size as the bush rat (Maitz & Dickman 2001). Thus, bush rats likely compete with other mammal species and these interactions likely control bush rat populations. Recolonisation of bush rat populations was also found to occur in fragmented forest landscapes (Lindenmayer et al. 2005; Holland & Bennett 2011) if access to suitable habitats were left available to assist population recovery. However, successful recolonisation likely depends on a low frequency of competitor species in these habitats.

The black rat is a highly successful introduced species native to Asia. Black rats breed prolifically in warm habitats such as Australia (Breed & Ford 2007). Black rats are a major

41 threat to native mammal diversity because of their invasive characteristics and adaptability to urban environments with impacts on biodiversity worldwide (Stokes et al. 2009b). The bush rat and black rat are similar in size and form but black rats are recorded to have potentially larger body size and larger litters more frequently (Lunney 1991; Breed & Ford 2007), resulting in high potential population growth rates.

Habitat research in New Zealand suggests the black rat is a superior competitor compared to other introduced rodents such as Norway rats (Rattus norvegicus) and Pacific rats (Rattus exulans) in all vegetation sites (Harper et al. 2005). The microhabitat preferred by the black rat is one of forest complex structure (Cox et al. 2000; Harper et al. 2005) with dense understorey and leaf litter, habitat remarkably similar to the bush rat’s preferred habitat

(Dickman & Steeves 2004; Garden et al. 2007). Black rats are very agile climbers and are found nesting in trees and associated with declines in bird populations from predation on their eggs (Cox et al. 2000; Whisson et al. 2007). The capacity for black rats to reduce the supply food sources such as invertebrates and seeds usually increases their competitive effects on other species.

Stokes et al. (2009b) found evidence that the presence of black rats can reduce the breeding success and or juvenile recruitment of bush rats. However, where bush rat populations are established, they are able to successfully compete against black rat populations (Holland &

Bennett 2007). While bush rats and black rats likely compete for limited resources, we don’t know if competition limits bush rat distribution. This is an important factor for the success of reintroduction of a species and the probability of invasive species driving native species to extinction.

Long term trap line data provides an excellent opportunity to test for how species distributions are shaped by competition. For example, distribution patterns of small mammal species in North American desert environments have been used to test for competitive exclusion between small mammal species with similar resource requirements (e.g. Bowers and Brown 1982). I used the extensive small mammal trapping datasets available on the Atlas of Living Australia. Bush rats and Black rats have an extensive range overlap in the Sydney area (appendix Fig 3.1 & 3.2) and they have a broadly similar habitat association in that they tend to select areas with some ground cover for protection from predators (Harper et al. 2005;

Stokes et al. 2009). The long term trapping datasets permit a test of observed species coexistence versus a null or random distribution. I tested the hypothesis that bush rats and black rats coexist less often than expected by chance. Low coexistence between species with similar resource requirements is consistent with the prediction that competition limits the distribution of these species. Coexistence between these species would indicate niche partitioning through differential resource use. This research has promising implications for species interaction by advancing knowledge of interspecific competition between a native species in human impacted woodlands.

3.2 Methods

3.2.1 Coexistence of Bush rats and Black rats

I used long-term small mammal trapping data to test for competitive interactions between bush rats and black rats. Data were collected within 150 km from the city of Sydney, predominantly covering the northern and southern areas of the Sydney Basin. My review included 266 sites covering 39 IBRA (Interim Biogeographic Regionalisation of Australia) locations. Data were assembled based on longitude and latitude location and trapping date of bush rats and black rats found within the 39 locations (appendix Table 3.3).

3.2.2 Data collection

I reviewed data from The Atlas of Living Australia (www.ala.org.au) data based on all records of bush rats Rattus fuscipes and black rats Rattus rattus trapped in New South Wales from 1926 through to 2010. Comparison was then made of all IBRA bioregions in NSW that recorded bush rats and black rats. To ensure I was comparing regions with overlapping bush rat and black rat distributions, bioregions that had not recorded both bush rat and black rats were eliminated. This process of elimination showed selected regions in the Sydney basin with overlapping bush rat and black rat distributions. Within the Sydney Basin region, a total of 1,233 black rats and 3,954 bush rats were recorded between 1926 and 2010.

Data were sorted grouping longitude and latitude to similar location. Google maps were then used by entering longitude and latitude to find sites with suitable distances between selected sites. Trap lines up to 1km of each other were considered within the same site. Multiple sampling did occur with some of the sites over several decades. Trap lines sampled within a two week time span were recorded as the same time. Trap lines were sampled within a consistent time frame averaged over six months apart and recorded at different times.

Replicate sites were the number of sites sampled at the same time. The presence of bush rats and black rats was recorded at each replicate site. Distribution of the bush rat and black rat was randomly surveyed to test for evidence of competitive exclusion or coexistence through niche partitioning between the two species present in the Sydney Basin area. Coexistence was estimated at the same place at the same time rather than just at the same place.

Study species

The black rat can reach an average size (95-280g), (Lunney 1991) compared to the similar omnivorous bush rat (50-225g) (Pedersen et al. 2014). Litter size of the Black rat averages 5-

10 per year while the bush rat litter size is generally only 5 per year. The bush rat is strictly nocturnal (Wood 1971) breeding in November at densities of around 10 per ha. Black rats have a strong competition capacity (Cassaing et al. 2007) are excellent climbers and highly adapted to coexisting with human populations, and are particularly common in the urban environment (Schneider 2014). Breeding of the black rat is between October and March

(Meek et al. 2011). Access to water has been found to limit the spread and distribution of black rats (Watts & Braithwaite 1978). Harper (2005) also found cold /wet conditions at altitudes with a rainfall above 3200mm per year limited black rats foraging ability.

3.3 Analysis

I used binomial probability test for calculating significant non-random associations between the presence of bush rats and black rats across overlapping parts of their distribution. For the binomial test, sample size was the number of site by time replicates with bush rats and/or black rats present. The observed frequency of coexistence (K) is the number of site by time replicates where both bush rats and black rats were present. The expected frequency sites where bush rats and black rats should be found together was calculated as the number of replicate sites occupied by bush rats relative to the total number of replicate sites multiplied by the number of replicate sites occupied by black rats relative to the total number of sites.

Total site by time replicates was estimated as both the total number of site replicates with either bush rats or black rats, and as the sites where both bush rats and black rats have been recorded, though not necessarily at the same time. This later test was done to control for

45 different habitat specialisation between bush rats and black rats that could lead to non- random species association not due to competition. In other words, the second test allows a test for species associations in sites where both species are known to occur. Binomial statistics were calculated using the online web stats tool Vassarstats (http://vassarstats.net/)

3.4 Results

Bush rats and or black rats were found in 642 replicate sites (sites where one or both species were found at the same sampling time). Bush rats were found in 513 replicate sites, and black rats were found in 263 replicate sites. I found bush rats and black rats coexisting at 134 replicate sites (i.e. sites where species were recorded at the same place at the same time).

Based on these records, I calculated an expected frequency of bush rat coexistence with black rats of 0.33 across all sites (i.e. black rats should coexist with bush rats in 210 replicate sites).

There were 330 replicate sites where bush rats and black rats were found in the same location but not necessarily at the same sampling time. Bush rats were found in 255 of these replicate sites and black rats were found in 211 of these replicate sites, I calculated an expected frequency of bush rat coexistence with black rats of 0.49 (i.e. black rats should coexist with bush rats in 162 replicate sites).

Across all sites, and for sites where bush rats and black rats have both been recorded, observed coexistence between bush rats and black rats was significantly less than the expected coexistence based on random distributions of bush rats and black rats across sites

(Table 3.1).

Table 3.1 Binomial Tests for coexistence between Bush rats and Black rats.

Black rats & Bush rats n K Expected prob. z P

Across all sites 642 134 0.33 -6.49 <0.0001

Across all sites where both 330 134 0.49 -3 0.0006 species have been recorded

Most sites had trapped more than a single bush rat and/or black rat. This suggests that a single animal does not preclude the presence of another. Thus, low coexistence is not simply an artefact of a maximum of one individual (and necessarily one species) per site.

There are remarkable cases of study sites where both bush rats and black rats are commonly found, but never at the same sampling time. For example, Wyong is located about 90 km from Sydney. Trap lines recorded data of 122 bush rats and 14 black rats in 9 random transects. One transect in the study was carried out at Chittaway Bay in 2001, 2002, 2003 and in 2009 in Wyong- South and West Region (appendix Table 3.3, Site 7) showing 7 bush rats and 5 black rats in the same place at different times.

3.5 Discussion

3.5.1 Small Mammal Communities

The following discussion of my results is possible beyond the immediate scope of this thesis but important as relates to reintroduction and translocation of bush rats.

Bush rats and black rats have overlapping distribution and were found to live in the same habitats. However, they tend not to live in the same place at the same time. Understanding communities and species use of resources is complex but important if we are to reintroduce species or translocate species from one location to another. Rodents have been extensively studied but we still understand little about assemblages of species in communities and the

47 role competition plays. Previous research suggests that if bush rats have residency status in forest areas they are likely to retain this status (Holland & Bennett 2007; Stokes 2009a). This thesis captures presence and absence of the two rodent species over a broad landscape and time frame to evaluate competition in fragmented and stable habitats.

This study provides an insight into the process of invasion of a competitor the black rat into an area originally occupied by a similar species the bush rat in a changing urban environment the Sydney Region (appendix Fig. 3.3). Population numbers of bush rats in the Sydney Basin

Region are still very health. A grouping of 266 transects at 39 locations resulted in 1528 bush rats recorded and only 484 black rats detected.

3.5.2 Community assembly and coexistence with invasive species

Understanding the process of invasion can provide an insight into competition and fragmentation of communities (Sax et al. 2007). The black rat has become established in the

Sydney basin because of human activities since its introduction 200 years ago (Cox et al.

2000). In general in eastern Australian forests, small mammal species richness can be low due to competition for limited resources (Fox 1987). In forest and woodland habitats, species richness of small mammals in the same site is typically not greater than two (Breed & Ford

2007). In desert habitats worldwide, small mammal species richness is generally 2-3 species

(Rodriguez & Ojeda 2013). Thus, a low number of small mammal species generally coexist within the same site. This trend is consistent with the idea that competition generally limits species richness and diversity across habitats. These results highlight the need for future studies using similar data to explore species richness found in these locations to understand community assemblages and coexistence especially for reintroduction or translocation of species.

3.5.3 Coexisting and available resources

Coexistence of species that use the same resource is a common phenomenon (e.g. Sharov

1997) a cyclical population growth or dynamic disturbance regimes can allow species to coexist even if they cannot coexist in stable systems and with similar resource requirements.

This argument highlights the complexity of understanding coexistence and available resources in natural dynamic ecosystems. Each location or habitat only has finite resources and space providing cover and protection from predators. Competition for these limited resources is therefore a critical factor in determining community structure of what species can coexist within a habitat. The genetic, environmental and diet flexibility of the black rat gives this rodent a wide adaptive ability to implement strong competition capacities (Cassaing et al.

2007). Competition for resources between these two species can impacts on food supply, nesting material, home range, social organisation and shelter from predators. Black rats are excellent climbers while bush rats are poor climbers and rarely found inhabiting elevated areas (Wendy Gleen personal communication 2014). Black rats also represented a danger to endemic insular populations already weakened by human activities (Cassaing et al. 2007).

Thus, coexistence between bush rats and black rats may be infrequent.

3.5.4 Community structure and competition

The effects of competition are complex and difficult to isolate but highly important if we are to understand communities and species reintroduction. Pedersen et al. (2014) found bush rats negatively related to deer (Rusa timorensis) presence on recently burnt sites in coastal heath areas in The Royal National Park. These coastal heath areas are important habitat areas for small mammals but are under threat from habitat fragmentation due to urban development.

The advantage the black rat has over the bush rat is the ability to diversify its food source and habitat and adjust to the different conditions (Harper et al. 2005; Cassaing et al. 2007; Stokes

49 et al. 2009b). The black rat is capable of persisting through habitat disturbance, and maintaining its ability to breed and increase its population numbers potentially to the exclusion of the bush rat. Species with high environmental carrying capacity (K) can have high negative effects on species of lower environmental carrying capacity (e.g. Ricklefs

2007). Thus, even if bush rats were equal in competitive ability to black rats, a shift in habitat towards increasing the carrying capacity of black rats relative to bush rats could allow black rat populations to exclude bush rats. While locations where native species richness is well established Fridley et al. (2007) found evidence that exotic invasion can be blocked by interactions of established native species and importantly if this richness is maintained.

Grassland studies (Sax et al. 2007) also show species rich plots are more difficult to invade than species poor plots. These studies may provide evidence that occupied habitats tend to resist invasion due to establishment of resident species. Establishment may depend on many factors and in disturbed and fragmented area the dominance may swing towards the adaptable or generalist species. Sampling size may be a factor to consider in future studies where more information regarding home range is readily available. Each of the trap lines were selected because of extensive robust data. Lack of coexistence found in this study may be due to under sampling. Individuals in the data set were selected as coexistence does not make predictions on population density. While the absence of a species suggests that the species is not present in a habitat and may be due to exclusion.

3.6 Conclusion

Developing a clear picture regarding competition is challenging but evidence suggests from this study competition exists between bush rats and black rats. The following discussion of my results is possible beyond the immediate scope of this thesis but relates to reintroduction of bush rats. Past research has shown bush rats are late successional species (Fox 1987;

Lindenmayer et al. 2005; Fox & Monamy 2007; Pedersen et al. 2014) especially after fire

(Pedersen et al. 2014) and have a complex community structure. If bush rats had established community structure in suitable niche habitats (Fox 1987; Holland & Bennet 2007; Kearney et al. 2007) with suitable home range and resources they may survive but in fragmented locations. The challenge is great. The black rat has the advantage that it has genetic plasticity

(Cassaing et al. 2007) allowing it to adjust to changed conditions from its ideal habitat to disturbed or fragmented habitat. If habitat areas along the east coast of Australia established by bush rats are not protected completion from the black rat will expand. Planning urban development with consideration of small mammal habitat, biodiversity and metapopulations will be an easier alternative to reintroduction or translocation of species into urban areas where building habitats and re-establishing communities is an expensive and difficult task.

The bush rat is a tough competitor and has been show to defend its territory (Fox 1987; Maitz et al. 2001; Lindenmayer et al. 2005; Holland & Bennett 2007; Stokes et al. 2009). But evidence (Bowers & Brown 1982; Fox 1987) on community assemblage patterns suggests species niche patterns are strongly influenced by diet selection. Therefore interspecific competition will effect growth and reproduction of the bush rat. Competition for resources may limit populations, where this competition between species is modest; it can also drive species to extinction. The impact of competition by black rats on bush rat reintroduction programs may show negative impacts if long term management of black rat populations are not carried out in urban remnant vegetation. Competition may be a limiting factor in determining the success of reintroduction with the probability of invasive species driving native species to extinction.

Chapter 4.

General Discussion

4.1 Why habitat matters

The loss of native species from a habitat following land use changes can promote the invasion of non-native species, and these species can be superior competitors in these changed environments (Downes et al. 1997; Moles et al. 2008). With around eighty percent of people in Australia living within 50 km of our coast, urban expansion is impacting on coastal ecosystems (Gurran et al. 2007). Understanding biological patterns of diversity especially factors that influence distribution and conservation of our wildlife can assist reintroduction or translocation of species into their former habitats or similar environments.

The challenge ahead is to achieve successful translocation program as in the past 50 years humanity has changed ecosystems more rapidly and extensively than any comparable time in human history (State of the Environment Committee 2011).

Selection of habitats in modified landscapes for reintroduction or translocation is our future challenge. To survive in urban environments small mammals must be able to disperse to areas of natural vegetation and persist long enough to reproduce (Dickman & Doncaster

(1989). Understanding the essential factors of species distribution will also assist establishing metapopulation in suitable habitats (Armstrong 2005). Quality habitat assessment is critical to understand the substantial loss of mammal species in Australia over the last 200 years (Short

2009; Richie et al. 2013). The importance of quality habitat structure is essential for the persistence of bush rat populations, particularly in habitats that may already be occupied by an invasive competitor the black rat. This research in three National Parks surrounding a large urban area highlights variables that promote stable and persistent conditions for small mammal populations.

4.2 Why determine factors that influence Rattus fuscipes (bush rat) distribution

The research behind this study was aimed to assist the reintroduction of the bush rat into vacated niches in Sydney Harbour National Park. The first goal of this research was to identify habitat characteristics associated with high density bush rat populations. The second goal or hypothesis was to test if there is a negative association between populations of bush rats and a potential competitor, the black rat.

4.3 What I found in relation to high density of bush rat populations

A significant result from the logistic regression model showed percent timber in the ground cover greater than 5cm to predict bush rat population density. Survey sites were selected to show high and low densities of bush rats. Habitat surrounded the survey sites was stable vegetation which would influence population processes where resources conditions would have been ideal. Obviously bush rats (like any organism) need essentials such as access to food and water for survival. However unless they have protection offered by complex ground cover, establish will not occur in that area. My results suggest that very specific aspects of understory complexity such as percent timber provide cover to support sustainable bush rat populations, while other variables tested were not significant to bush rat density.

The hypothesis of negative association between the bush rat and a similar size competitor the black rat is important to determine presence of species in an area. Evidence from Atlas of living Australia data base records in the Sydney Basin showed clear evidence that bush rats and black rats have overlapping distribution and even live in the same habitat but tend not to live in the same place at the same time. The value of these data base records is in future studies of species distribution and can be evaluated over time to determine species presence with changes to habitat.

Until we understand all determining factors of habitat suitability of small mammals we will not stop the reduction or extinction of small mammal species. The lack of knowledge is really associated with the dynamic factors of habitat. Habitat provides the physical area where an animal lives, and survives and persistence which is the key to persistence of species (Fortey

2012). Studying bush rats in established habitats provided an insight into essential criteria why they choose to live there and what factors supported their habitat.

One of the main problems with translocating species is assessment of critical habitat and often little research in carried out to evaluate suitable or unsuitable habitat (Griffith 1989).

Species community structure is also critical and important for establishment of species as these factors can be essential for a successful translocation into new areas. Geographical barriers may prevent establishment of species back into their once natural area (Sheean et al.

2012) or habitat quality in fragments areas will influence population numbers (Holland &

Bennett 2010). Fragment size can also influence bush rat densities. Larger fragments have the advantage of buffering uncertainty regarding habitat quality to establish stable conditions for bush rats (Holland & Bennett 2010). Home ranges of a species can vary due to habitat and food resources (Breed and Food 2007) Small habitats or minimum home range size may be a function of habitat productivity, resource distribution as well as individual energy requirements (Fisher 2000) and be influenced by competitors (Maitz and Dickman 2001).

Minimum amount of habitat requirements are not know as home ranges can fluctuate depending on seasonal conditions (Sanecki et al. 2006) and habitat relationships (Mitchell &

Powell 2004).

4.4 Variables suitable for bush rat establishment in reintroduction sites

Bush rats are found widespread in structurally dense and diverse ground cover habitats they are omnivorous and generalist in habitat use and found to occur in five habitat types

(Dickman et al. 2000). That is a species with broad diets and habitat tolerance and more likely to adapt to modified habitats but (Holland & Bennett 2010) found demography of the bush rat to be profoundly affected by forest fragmentation. My research supports the argument that structure in the lower canopy (significant percentage of timber) is associated with high densities of bush rats present. Competition from invasive non-native species especially black rats will have an advantage in disturbed and fragmented habitat favouring the invader (Cassaing et al. 2007). Though bush rats have been known to compete with black rats (Stokes et al. 2007) under certain conditions, habitat and population establishment may influence their coexistence. For coexistence to occur in a landscape one species needs to survive better in a secondary habitat than the dominant species (Rosenzweig & Abramsky

1997). Gregory & Macdonald (2009) found palatability and preferred diet a competitive advantage for an endemic rodent Nesoryzomys swarthy against the black rat (Rattus rattus) in the Galápogos Islands. Studies between two rodents Rattus lutreolus velutinus (120g) and

Pseudomys higginsi (62g) found that competition not only depended on population density, resource availability and habitat specialisation but also the phase of the population cycle (Luo et al. 1998).

Land managers will need an intricate knowledge of habitat, species community assemblages and competition to return species to their former range in urban landscapes. Understanding key habitat attributes is essential for the ecology of landscapes and restoring vegetation for translocation or reintroduction programs. The emerging field of invasion biology highlights human activity and negative consequences (Vitousek 1990; Dunstan & Fox 1996; Tait et al. 2005;

Salo et al. 2007). The more we understand about habitat distribution of native species the better the chance of their survival. The number of lost mammal species over the last 200 years in

Australia emphasis how little we know and how much is to gain from further research into key habitat features of specific species.

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Appendix Table 2.3 Bush rat (Rattus fuscipes) Vegetation Survey Field Sheet Quadrate Location description

Surveyed by Wendy Kinsella Date 2012

Site Characteristics Ground cover Land Element Bare Earth %

Ridge Aspect [Compass] [GPS] Rock %

Upper Slope Slope Leaf Litter %

Leaf litter Mid Slope Soil texture depth cm

Lower Slope Soil pH Fallen timber > 5cm =

Gully Soil Colour Moisture %

Creek Soil Total Weed distance compaction Cover

Circle Score 1 2 3 Vegetation - Lower layer Cover < 5% 5-10% 10-30% 0.25- Height Class <0.25m 0.5 0.5-1m Score 4 5 6 Plant Weed 30- 50- 70- community score Cover 50% 70% 100% Plant species

Circle Score 1 2 3 Middle Layer Cover < 5% 5-10% 10-30% Height Class 1-2 m 2-3 m 3-5 m Score 4 5 6 Weed 30- 50- 70- Plant community score Cover 50% 70% 100% Plant species

Circle Score 1 2 3 Upper layer Cover < 5% 5-10% 10-30% 10-20 20- 4 Height Class 5-10 m m 35m Score 5 6 Plant Weed 30- 50- 70- community score Cover 50% 70% 100% Plant species

Growth stage Mature Fire History Yea r of burn Cool burn Warm burn

Senescing No fire Hot burn Inferno

Table 2.4 Summary of all records collected Ku-ring-gai Chase National Park Total average of 4 Light Contour no. readings at each level level Vegetation Present bush Land Site Location Aspect Class Absent rats Element ground 2003- 1.2m 2m m 2010 level

Centre Trial site west head VES K1 S_DSF09 High 45 30 341 530 Ridge SW 180 end, 30m from gate, RHS.

800m from K6 NPWS gate LHS S_DSF04 High 15 142 160 238 Ridge WSW 140 at bandicoot sign

Towers Bay track 5m from K10 S_DSF12 High 44 43 50 125 Ridge SSW 170 West Head Road.

50m from Lower K15 S_DSF12 High 25 29 334 514 NW 45 McCarrs Ck. slope

Lower G3 5 mile track S_DSF09 High 48 44 184 158 WSW 55 slope

Upper G20 Heath track S_DSF12 High 49 16 40 57 N 150 slope

Ryland/Cooyong G27 trail (10m from S_DSF09 High 18 38 61 59 Ridge WSW 180 gate)

Ryland/Cooyong G28 trail (300 from S_DSF10 High 31 116 210 242 Ridge SW 180 gate)

Cascade /Lower Lower G34 S_DSF09 High 43 37 85 103 SE 90 Cambourne slope

Table 2.4 (continued) Ku-ring-gai Chase National Park Creek Lower Middle Upper Height class distance No. No. No. layer layer layer ite plant plant plant species genus families Cover Cover Cover M lower mid upper score score score

K1 325 1 3 7 26 21 12 4 5 4

K6 200 1 3 15 32 26 15 6 3 5

K10 300 2 3 5 16 15 10 6 3 4

K15 8 2 3 15 26 21 14 5 5 3

G3 4 1 5 7 41 28 23 5 5 3

G20 200 0.5 3 7 27 23 14 6 6 2

G27 500 1 2 15 24 21 15 6 3 5

G28 450 1 5 15 28 23 12 6 4 5

G34 200 1 5 25 29 26 17 3 4 4

Table 2.4 (continued) Ku-ring-gai Chase National Park

Leaf % Fallen Total % Bare % Rock % Leaf litter Site timber weed Soil texture Soil pH earth cover litter depth >5cm cover cm

K1 20 50 65 3 40 0 loamy sand 4.5

K6 0 0 50 20 35 0 loamy sand 6

K10 5 5 50 40 30 0 loamy sand 4.5

K15 20 20 30 15 45 0 loamy sand 5

G3 10 45 50 20 25 0 clayey sand 4.5

G20 5 10 45 40 10 0 clayey sand 4.5

G27 15 10 40 15 25 0 sandy loam 5

G28 10 5 50 20 45 0 sandy loam 4.5

G34 20 30 45 20 50 0 loamy sand 5

Table 2.5 Summary of all records collected The Royal National Park & Heathcote National Park

Total average of 4 Light Contour no. bush Vegetation Present readings at each level Land level Site Location rats Aspect Class Absent Element 2003- ground 1.2m 2m m 2010 level

Near 1(12) S_HL08 Low 1 11 33 89 Mid slope NE 125 Carlot site

Flat rock 2(16) S_DSF09 Low 1 37 180 273 Mid slope ENE 90 creek

Mt Bass Upper 3(17) S_HL08 Low 1 16 95 609 NW 150 fire trail slope St Bertrum Lower 4(15) S_DSF05 High 32 3 4 5 SW 190 Stevens slope Drive Bola Cr Lady Lower 5(11) S_RF03 High 39 3 4 5 SW 60 Carrington slope Drive

Upper Lower 6(10) S_RF03 Low 1 23 149 152 NW 40 Causeway slope

.4 km SW Lower 7(8) of Palm S_RF03 Low 1 6 8 9 NW 40 slope Gully

Lower 8(4) Kingfisher S_DSF09 Low 1 48 260 234 SW 140 slope Lower 9(7) Otford S_RF08 Low 1 7 6 12 SE 50 slope Stanwell Tops Upper 10(5) S_DSF05 Low 1 17 53 273 W 263 Hargrave slope Creek 4km south of Lilyvale 11(5C) track & S_DSF05 High 10 15 20 22 ridge SW 230 beginning of Coast track Lady 12(7A) Carrington S_RF03 High 13 24 27 22 Gully NW 20 Drive Garie Rd & Lower 13(8) S_DSF09 High 10 17 36 SW 210 Garawarra slope Hill Rd

Table 2.5 (continued) The Royal National Park & Heathcote National Park

Creek Lower Middle Upper Height class distance No. No. No. layer layer layer Site plant plant plant species genus families Cover Cover Cover m lower mid upper score score score

1(12) 300 1 3 15 17 12 8 2 6 1

2(16) 85 1 3 15 18 13 7 3 5 2

3(17) 200 1 2 3 16 13 9 3 6 3

4(15) 220 1 3 15 35 21 13 5 6 3

5(11) 20 1 5 25 23 21 21 4 4 6

6(10) 100 1 5 25 24 23 20 6 4 5

7(8) 20 1 5 25 21 18 14 3 3 6

8(4) 1 0.5 2 15 44 30 12 3 5 4

9(7) 350 1 2 7 21 17 15 2 2 6

10(5) 5 0.5 3 15 41 31 17 6 6 3

11(5C) 200 0.5 3 10 29 27 22 5 4 6

12(7A) 15 1 3 20 19 18 17 4 4 5

13(8) 400 1 5 0 25 23 10 4 5 0

Table 2.5 (continued) The Royal National Park & Heathcote National Park

Leaf % % Fallen Total % Rock % Leaf litter Soil Site Bare timber weed Soil texture cover litter depth earth >5cm cover pH cm

1(12) 1 0 15 2 15 0 clayey sand 5

2(16) 5 5 70 2 5 0 loamy sand 4.5

3(17) 5 0 15 1 0 0 clayey sand 5

4(15) 10 2 45 4 30 0 loamy sand 4.5

5(11) 35 10 70 3 25 0 loam 4.5

6(10) 2 0 40 10 20 0 sandy loam 5

7(8) 40 30 35 8 20 0 clayey sand 6

8(4) 35 5 10 2 15 0 clayey sand 4.5

9(7) 15 15 45 2 20 0 sandy loam 6

10(5) 10 0 30 10 40 0 clayey sand 5

11(5C) 5 1 45 15 20 0 sandy loam 4.5

12(7A) 10 15 30 10 30 0 sandy loam 5

13(8) 1 1 70 40 20 0 loamy sand 5.5

Table 2.8 SPECIES SUMMARY OF KU-RING-GAI CHASE NATIONAL PARK Lower Layer Family Genus Genus & Species Common name Adiantaceae Adiantum Adiantum aethiopicum Common Maidenhair Fern Apiaceae Platysace Platysace linearifolia Narrow-leaf Platysace Apiaceae Xanthosia Xanthosia tridentata Rock Xanthosia Apiaceae Actinotus Actinotus minor Apiaceae Platysace Platysace linearifolia Apiaceae Actinotus Actinotus minor Lesser Flannel Flower Apiaceae Platysace Platysace linearifolia Apiaceae Xanthosia Xanthosia pilosa Woolly Xanthosia Asteraceae Cassinia Cassinia denticulata think Casuarinaceae Casuarina Casuarina sp Casuarinaceae Allocasuarina Allocasuarina distyla Scrub She-Oak Cunoniaceae Callicoma Callicoma serratifolia Black Wattle Cyperaceae Caustis Caustis pentanda Cyperaceae Tricostlaria Tricostularis paucifolaria Needle Bogrush Cyperaceae Caustis Caustis flexuosa Curly Sedge Cyperaceae Caustis Caustis pentandra Cyperaceae Caustis Caustis Cyperaceae Gahnia Gahnia Dilleniaceae Hibbertia Hibbertia empetrifolia Trailing Guinea Flower Dilleniaceae Hibbertia Hibbertia aspera Rough Guinea Flower Dilleniaceae Hibbertia Hibbertia empetrifolia Trailing Guinea Flower Dryopteridaceae Gleichenia Gleichenia dicarpa Pouched Coral Fern Epacridaceae Epacris Epacris Epacridaceae Styphelia Styphelia tubiflora Red Five Corners Epacridaceae Epacris Epacris pulchella Epacridaceae Epacris Epacris microphylla Coral Heath Epacridaceae Epacris Epacris longiflora Native Fuschia Epacridaceae Epacris Epacris microphylla Coral Heath Epacridaceae Epacris Epacris pulchella NSW Coral Heath Epacridaceae Leucopogon Leucopogon amplexicaulis Epacridaceae Leucopogon Leucopogon microphyllus Epacridaceae Leucopogon Leucopogon setiger Ericaceae Monotoca Monotoca scoparia Fabaceae Dillwynia Dillwynia retorta Fabaceae Phyllota Phyllota grandiflora Fabaceae Pultenaea Pultenaea daphnoides Large -leaf Bush-pea

Table 2.8 (continued) Species Summary of Ku-ring-gai Chase National Park Lower Layer Family Genus Genus & Species Common name Fabaceae Pultenaea Pultenaea tuberculata formally (P. elliptica) Fabaceae Pultenaea Pultenaea linophylla need to check this one Fabaceae Phyllota Phyllota phylicoides Common Phyllota Fabaceae Bossiaea Bossiaea heterophylla Fabaceae Hovea Hovea linearis Formally (P. elliptica) Fabaceae Bossiaea Bossiaea scolopendria Fabaceae Lambertia Lambertia formosa Mountain Devil Fabaceae Aotus Bossiaea scolopendria Fabaceae Pultenaea Pultenaea Fabaceae Pultenaea Pultenaea polifolia Goodeniaceae Dampiera Dampiera stricta Haloragaceae Zieria Zieria pilosa Haloragaceae Gonocarpus Gonocarpus teucrioides German Raspwort Hardenbergia Hovea Hovea longifolia Iridaceae Patersonia Patersonia sericea Silky Purple-flag Mimosaceae Acacia Acacia suaveolens Sweet-scented Wattle Mimosaceae Acacia Acacia terminalis Sunshine Wattle Myrtaceae Darwinia Darwinia fascicularis Myrtaceae Leptospermum Leptospermum Myrtaceae Angophora Angophora hispida Dwarf Apple Myrtaceae Leptospermum Leptospermum squarrosum Pink Tea-tree Myrtaceae Darwinia Darwinia fascicularis Myrtaceae Angophora Angophora hispida Dwarf Apple Myrtaceae Leptospermum Leptospermum Myrtaceae Angophora Angophora costata Sydney red gum Myrtaceae Leptospermum Leptospermum trinervium Paperbark Tea-tree Poaceae Entolasia Entolasia stricta Poaceae Anisopogon Anisopogon avenaceus Oat Speargrass Proteaceae Hakea Hakea teretifolia Proteaceae Persoonia Persoonia pinifolia Pine -leaf Geebung Proteaceae Grevillea Grevillea sericea subsp. sericea Pink Spider Flower Proteaceae Persoonia Grevillea buxifolia Green Spider Flower Proteaceae Conospermum Conospermum longifolium Long-leaf Coneseed Proteaceae Banksia Banksia marginata Proteaceae Grevillea Grevillea speciosa Proteaceae Banksia Banksia ericifolia Heath -leaved Banksia Proteaceae Banksia Banksia oblongifolia

Table 2.8 (continued) Species Summary of Ku-ring-gai Chase National Park Lower Layer Family Genus Genus & Species Common Name Proteaceae Grevillea Grevillea buxifolia Grey Spider Flower Proteaceae Conospermum Conospermum longifolium Long-leaf Coneseed Proteaceae Hakea Hakea teretifolia Dagger Hakea Proteaceae Isopogon Isopogon anethifolius Proteaceae Lambertia Lambertia formosa Mountain Devil Restionaceae Lepyrodia Lepyrodia scariosa Restionaceae Empodisma Empodisma minus Spreading Rope-rush Rutaceae Boronia Boronia ledifolia Rutaceae Boronia Boronia serrulata Native Rose Rutaceae Boronia Boronia ledifolia Sydney Boronia Rutaceae Crowea Crowea saligna Rutaceae Phebolium Phebolium squamulosum Smilacaceae Smilax Smilax glyciphylla Native Sarsaparilla Stackhousiaceae Pomaderris Pomaderris phylicifolia? Tremandraceae Tetratheca Tetratheca ericifolia Black -eyed Susan Xanthorhoeaceae Lomandra Lomandra filiformis Wattle Mat-rush Xanthorrhoeaceae Xanthorrhoea Xanthorrhoea Grass tree Xanthorrhoeaceae Lomandra Lomandra filiformis Wattle Mat-rush Xanthorrhoeaceae Xanthorrhoea Xanthorrhoea Grass tree Lomandra filiformis or Xanthorrhoeaceae Lomandra gracilis Xanthorrhoeaceae Lomandra Lomandra obliqua

Middle Layer Family Genus Genus & Species Common Name Apiaceae Platysace Platysace linearifolia Narrow-leaf Platysace Casuarinaceae Allocasuarina Allocasuarina distyla Scrub She-Oak Casuarinaceae Allocasuarina Allocasuarina littoralis Black She-Oak Cunoniaceae Callicoma Callicoma serratifolia Black Wattle Cyperaceae Caustis Caustis Fabaceae Phyllota Phyllota grandiflora Fabaceae Dillwynia Dillwynia retorta Heathy Parrot Pea Fabaceae Pultenaea Pultenaea stipularis Fine-leaf Bush-pea Fabaceae Gompholobium Gompholobium grandiflorum Large Wedge-pea Mimosaceae Acacia Acacia ulicifolia Prickly Moses Mimosaceae Acacia Acacia suaveolens Sweet-scented Wattle Mimosaceae Acacia Acacia terminalis Sunshine Wattle

Table 2.8 (continued) Species summary of Ku-ring-gai Chase National Park Middle Layer Family Genus Genus & Species Common Name Myrtaceae Darwinia Darwinia fascicularis Myrtaceae Leptospermum Leptospermum squarrosum Pink Tea-tree Myrtaceae Leptospermum Leptospermum trinervium Flaky-barked Tea-tree Myrtaceae Angophora Angophora hispida Myrtaceae Baeckea Baeckea linifolia Proteaceae Banksia Banksia serrata Old Man Banksia Proteaceae Banksia Banksia ericifolia Heath Banksia Proteaceae Petrophile Petrophile pulchella Proteaceae Grevillea Grevillea buxifolia Proteaceae Lambertia Lambertia formosa Mountain Devil Proteaceae Hakea Hakea dactyloides Broad-leaved Hakea Proteaceae Persoonia Persoonia levis Smooth Geebung Proteaceae Hakea Hakea gibbosa Proteaceae Grevillea Grevillea buxifolia Grey Spider Flower Proteaceae Hakea Hakea dactyloides Broad-leaved Hakea Proteaceae Hakea Hakea teretifolia Dagger Hakea Proteaceae Persoonia Persoonia pinifolia Pine-leaf Geebung Rutaceae Crowea Crowea saligna

Upper Layer Family Genus Genus & Family Common Name Myrtaceae Eucalyptus Eucalyptus sclerophylla Hard-leaved Scribbly Gum Myrtaceae Eucalyptus Eucalyptus capitellata Myrtaceae Eucalyptus Eucalyptus Myrtaceae Eucalyptus Eucalyptus eximia Myrtaceae Eucalyptus Eucalyptus haemastoma Myrtaceae Angophora Angophora costata Sydney Red Gum Myrtaceae Eucalyptus Eucalyptus piperita Sydney Peppermint Proteaceae Banksia Banksia ericifolia Heath-leaved Banksia Proteaceae Banksia Banksia serrata Old Man Banksia Rough-barked Apple Mallee sp.

Table 2.9 SPECIES SUMMARY OF THE ROYAL & HEATHCOTE NATIONAL PARK Lower Layer Family Genera Genus and Species Common Name Acanthaceae Pseuderanthemum Pseuderanthemum variabile Pastel Flower Adiantaceae Adiantum Adiantum aethiopicum Common Maidenhair Adiantaceae Adiantum Adiantum formosum Giant Maidenhair Agavaceae Doryanthas Doryanthes excelsa Gymea Lily Anthericaceae Arthropteris Arthropteris tenella Apiaceae Actinotus Actinotus minor Apiaceae Centella Centella asiatica Swamp pennywort Apiaceae Hydrocotyle Hydrocotyle peduncularis Apiaceae Platysace Platysace lanceolata Native Parsnip Apiaceae Platysace Platysace linearifolia Carrot Tops Apiaceae Xanthosia Xanthosia pilosa Apiaceae Xanthosia Xanthosia tridentata Rock Xanthosia Apocynaceae Parsonsia Parsonsia straminea Araceae Gymnostachys Gymnostachys anceps Settlers Flax Arecaceae Livistonia Livistonia australis Cabbage tree Palm Aspleniaceae Asplenium Asplenium flabellifolium necklace Fern Blechnaceae Blechnum Blechnum cartilagineum Gristle Fern Blechnaceae Doodia Doodia aspera Rasp Fern Caesalpinioideae Senna Senna pendula Cassia (Weed) Casuarinaceae Casuarina Casuarina sp. Celastraceae Cassine Cassine australis Cyperaceae Caustis Caustis pentandra Cyperaceae Caustis Caustis recurvata Cyperaceae Caustis Caustis sp. Cyperaceae Gahnia Gahnia aspera Cyperaceae Gahnia Gahnia clarkei Tall Saw-sedge Cyperaceae Gahnia Gahnia melanocarpa Cyperaceae Lepidosperma Lepidosperma laterale Cyperaceae Lepidosperma Lepidosperma sp. Sword Sedge Cyperaceae Schoenus Schoenus ericetorum Dennstaedtiaceae Hypolepis Hypolepis muelleri Harsh Ground Fern Dennstaedtiaceae Pteridium Pteridium esculentum Bracken Dicksoniaceae Calochlaena Calochlaena dubia Common Ground Fern Dilleniaceae Hibbertia Hibbertia dentata Twining Guinea Flower Dilleniaceae Hibbertia Hibbertia scandens Golden Guinea Flower Dryopteridaceae Lastreopsis Lastreopsis decomposita Dryopteridaceae Lastreopsis Lastreopsis microsora Creeping Shield Fern Epacridaceae Acrotriche Acrotriche divaricata

Table 2.9 (continued) SPECIES SUMMARY OF THE ROYAL & HEATHCOTE NATIONAL PARK Lower Layer (continued) Family Genera Genus and Species Common Name Epacridaceae Astroloma Astroloma humifusum Cranberry Heath Epacridaceae Epacris Epacris microphylla Epacridaceae Epacris Epacris sp Epacridaceae Leucopogon Leucopogon ericoides Epacridaceae Leucopogon Leucopogon juniperinus Bearded Heath Epacridaceae Leucopogon Leucopogon lanceolatus Lance Beard-heath Epacridaceae Lissanthe Lissanthe strigosa Native Cranberry Epacridaceae Monotoca Monotoca scoparia Epacridaceae Woollsia Woollsia pungens Escalloniaceae Polyosma Polyosma cunninghamii Euphorbiaceae Pseudanthus Pseudanthus pimeleoides Fabaceae Acacia Acacia suaveolens Sweet Scented Wattle Fabaceae Gompholobium Gompholobium glabratum Fabaceae Gompholobium Gompholobium minus Dwarf Wedge-pea Faboideae Glycine Glycine clandestine Love creeper Goodeniaceae Goodenia Goodenia ovata Hop-Goodenia Goodeniaceae Scaevola Scaevola ramoisssima Snake-flower Haloragaceae Gonocarpus Gonocarpus micranthus Lindsaeaceae Lindsaea Lindsaea linearis Screw Fern Lobeliaceae Pratia Pratia purpurascens White Root Lomandraceae Lomandra Lomandra longifolia Mat Rush Lomandraceae Lomandra Lomandra obliqua Fish Bones Lomandraceae Lomandra Lomandra sp. Luzuriagaceae Eustrephus Eustrephus latifolius Wombat Vine Mimosaceae Acacia Acacia myrtifolia Myrtle Wattle Monimiaceae Wilkiea Wilkiea huegeliana Wilkiea Moraceae Ficus Ficus coronata Sandpaper Fig Moraceae Maclura Maclura cochinchinensis Cockspur Thorn Myrsinaceae Rapanea Rapanea howittiana Turnipwood Myrtaceae Acmena Acmena smithii Lillypilly Myrtaceae Angophora Angophora costata Sydney Red Gum Myrtaceae Callistemon Callistemon citrinus Crimson Bottlebrush Myrtaceae Eucalyptus Eucalyptus sp. Myrtaceae Kunzea Kunzea ambigua Tick Bush Myrtaceae Leptospermum Leptospermum trinervium Paperbark Tea-tree Myrtaceae Melaleuca Melaleuca hypericifolia

Table 2.9 (continued) SPECIES SUMMARY OF THE ROYAL & HEATHCOTE NATIONAL PARK Lower Layer (continued) Family Genera Genus and Species Common Name Oleaceae Notalaena Notelaea venosa Oleaceae Notalaena Notelaea longifolia Orchidaceae Acianthus Acianthus fornicatus Pixie Orchid Orchidaceae Pterostylis Pterostylis sp Phormiaceae Dianella Dianella caerulea var producta Blue Flax Lily Pittosporaceae Citriobatus Citriobatus pauciflorus Orange Thorn Poaceae Digitaria Digitaria parviflora Smallflower fingergrass Poaceae Entolasia Entolasia marginata Grass Poaceae Entolasia Entolasia stricta Grass Poaceae Imperata Imperata cylindrica Blady grass Poaceae Oplismenus Oplismenus aemulus Basket Grass Poaceae Poa Poa affinis Cone-sticks Polypodiaceae Pyrrosia Pyrrosia rupestris Rock Felt Fern Proteaceae Banksia Banksia serrata Proteaceae Grevillea Grevillea doffisa ssp. linifolia Proteaceae Grevillea Grevillea oleoides Proteaceae Grevillea Grevillea sphacelata Grey Spider Flower Proteaceae Hakea Hakea dactyloides Broad-leaved Hakea Proteaceae Isopogon Isopogon anemonifolius Drumsticks Proteaceae Lambertia Lambertia formosa Mountain Devils Proteaceae Lomatia Lomatia silaifolia Crinkle bush Proteaceae Persoonia Persoonia levis Broad-leaf Geebung Proteaceae Petrophile Petrophile pulchella Proteaceae Stenocarpus Stenocarpus salignus Scrub Beefwood Proteaceae Telopea Telopea speciosissima Waratah Restionaceae Empodisma Empodisma minus Spreading Rope-rush Ripogonaceae Ripogonum Ripogonum album White Supplejack Rubiaceae Opercularia Opercularia aspera Common Stinkweed Rutaceae Boronia Boronia ledifolia Sydney Boronia Schizaeaceae Gleichenia Gleichenia dicarpa Pouched Coral Fern Scrophulariaceae Veronica Veronica plebeia Creeping Speedwell Selaginellaceae Selaginella Selaginella uliginosa Sinopteridaeae Pellaea Pellaea falcata Sickle Fern Smilacaceae Geitonoplesium Geitonoplesium cymosum Scrambling Lily Smilacaceae Smilax Smilax glyciphylla Native Sarsaparilla Thymelaeaceae Pimelea Pimelea linifolia ssp. linifolia

Table 2.9 (continued) SPECIES SUMMARY OF THE ROYAL & HEATHCOTE NATIONAL PARK Lower Layer (continued) Family Genera Genus and Species Common Name Violaceae Viola Viola (unknown) Unidentified sp. (A. Fairley) Violaceae Viola Viola hederacea Native Violet Winteraceae Tasmannia Tasmannia insipida Bush Pepper-bush Xanthorrhoeaceae Xanthorrhoea Xanthorrhoea sp. Unknown Unknown sedge sp.

Table 2.9 (continued) SPECIES SUMMARY OF THE ROYAL & HEATHCOTE NATIONAL PARK Middle Layer Family Genera Genus and Species Common Name Agavaceae Doryanthes Doryanthes excelsa Gymea Lily Apiaceae Platysace Platysace linearifolia Carrot Top Araceae Gymnostachys Gymnostachys anceps Settlers Flax Arecaceae Livistonia Livistonia australis Cabbage-tree Palm Asteraceae Cassine Cassine australis Red-fruited Olive Plum Casuarinaceae Allocasuarina Allocasuarina distyla Scrub She-oak Casuarinaceae Allocasuarina Allocasuarina littoralis Black She-oak Cyperaceae Gahnia Gahnia clarkei Cyperaceae Gahnia Gahnia melanocarpa Cyperaceae Gahnia Gahnia sieberana Dilleniaceae Hibbertia Hibbertia scandens Climbing Guinea Flower Ericaceae Epacris Epacris longifolia Ericaceae Epacris Epacris microphylla Coral Heath Ericaceae Epacris Epacris pulchella NSW Coral Heath Ericaceae Leucopogon Leucopogon amplexicaulis Ericaceae Leucopogon Leucopogon ericoides Bearded Heath Ericaceae Monotoca Monotoca elliptica Tree Broom-heath Ericaceae Woollsia Woollsia pungens Eupomatiaceae Eupomatia Eupomatia laurina Bolwa rra, Native Guava Fabaceae Bossiaea Bossiaea heteroplylla Fabaceae Dillwynea Dillwynea floribunda Fabaceae Dillwynea Dillwynia retorta Heathy Parrot Pea Fabaceae Pultenaea Pultenaea elliptica Fabaceae Pultenaea Pultenaea stipularis Fabaceae Viminaria Viminaria juncea Native Broom Grossulariaceae Polyosma Polyosma cunninghamii Featherwood Mimosaceae Acacia Acacia floribunda Mimosaceae Acacia Acacia linifolia Flax -leafed Wattle

Table 2.9 (continued) SPECIES SUMMARY OF THE ROYAL & HEATHCOTE NATIONAL PARK Middle Layer Family Genera Genus and Species Common Name Mimosaceae Acacia Acacia longifolia Sydney Golden Wattle Mimosaceae Acacia Acacia myrtifolia Myrtle Wattle Mimosaceae Acacia Acacia rubida Mimosaceae Acacia Acacia suaveolens Mimosaceae Acacia Acacia terminalis Mimosaceae Acacia Acacia ulicifolia Prickly Moses Myrtaceae Acmena Acmena smithii Lilly Pilly Myrtaceae Backhousia Backhousia myrtifilia Grey Myrtle, Lancewood Myrtaceae Callistemon Callistemon citrinus Myrtaceae Eucalyptus Eucalyptus piperita

Table 2.9 (continued) SPECIES SUMMARY OF THE ROYAL & HEATHCOTE NATIONAL PARK Middle Layer (continued) Family Genera Genus and Species Common Name Myrtaceae Eucalyptus Eucalyptus multicaulis Wipstick Mallee Ash Myrtaceae Eucalyptus Eucalyptus obstans Port Jackson Mallee Myrtaceae Eucalyptus Eucalyptus sp. Myrtaceae Kunzea Kunzea ambigua Tick Bush Myrtaceae Leptospermum Leptospermum Myrtaceae Leptospermum Leptospermum juniperinum Prickly Tea-tree Myrtaceae Leptospermum Leptospermum rotundifolium Myrtaceae Leptospermum Leptospermum squarrosum Myrtaceae Leptospermum Leptospermum trinervium Paperbark Tea-tree Myrtaceae Melaleuca Melaleuca hypericifolia Red Honey-myrtle Myrtaceae Malaleuca Melaleuca sieberi Myrtaceae Syncarpus Syncarpus glomulifera Turpentine Oleaceae Notalaena Notelaea venosa Pittosporaceae Billardiera Billardiera scandens Apple Berry, Dumplings Polypodiaceae Platycerium Platycerium bifurcatum Elk Horn Proteaceae Banksia Banksia ericifolia Heath-leaved Banksia Proteaceae Banksia Banksia integrifolia Silver Banksia Proteaceae Banksia Banksia marginata Silver Banksia Proteaceae Banksia Banksia oblongifolia Proteaceae Banksia Banksia serrata Old Man Banksia Proteaceae Banksia Banksia spinulosa Hair-pin Banksia Proteaceae Grevillea Grevillea buxifolia Grey Spider Flower Proteaceae Grevillea Grevillea longifolia Proteaceae Grevillea Grevillea sericea Pink Spider Flower Proteaceae Grevillea Grevillea sphacelata Grey Spider Flower

Fig 3.1 Mapping occurrence records of Bush rats Rattus fuscipes in Australia. Darker colour indicates higher density of bush rats. Source The Atlas of Living Australia (www.al.org.au).

Fig 3.2 Mapping occurrence records of Black rats Rattus rattus in Australia. Darker colour indicates higher density of black rats. Source The Atlas of Living Australia (www.al.org.au).

Table 3.2 List of sites selected in Sydney Basin where each site had numerous transects. Source The Living Atlas of Australia No. sites Location Bush rat Black rat 1 Mid-Western Regional - Pt A 26 12 2 Mid-Western Regional - Pt B 17 3 3 Muswellbrook 32 8 4 Singleton 23 10 5 Port Stephens 8 4 6 Cessnock 12 26 7 Newcastle - Outer West 2 3 8 Lake Macquarie - North 47 26 9 Wyong - South and West 122 14 10 Wyong - North-East 17 26 11 Lake Macquarie - West 4 18 12 Gosford - West 34 9 13 Lithgow 61 1 14 Gosford - East 52 8 15 Hawkesbury 26 14 16 Baulkham Hills - North 8 20 17 Pittwater 20 8 18 Penrith - West 0 11 19 Hornsby - North 74 14 20 Ku-ring-gai 9 7 21 Warringah 70 5 22 Blue Mountains 25 3 23 Ryde 0 9 24 Manly 2 23 25 Blacktown - South-West 1 1 26 Mosman 0 7 27 Liverpool - West 1 10 28 Liverpool - East 0 31 29 Sutherland Shire - East 36 44 30 Sutherland Shire - West 47 10 31 Wollondilly 60 7 32 Campbelltown - South 7 4 33 Wollongong Bal 269 17 34 Wingecarribee 25 10 35 Wollongong - Inner 3 1 36 Shellharbour 3 3 37 Kiama 13 0 38 Shoalhaven - Pt B 326 54 39 Jervis Bay 38 3 1520 484

Table 3.3 Case Study Wyong (South & West) New South Wales

Site 1

total rep. Bush Black Bush Black Latitude Longitude Survey Location Date each (different rat rat rat rat site time same place) -33.10014 151.2516 Wyong - South and West 6/10/2004 1 0 1 1 0 -33.10014 151.2516 Wyong - South and West 10/09/2004 1 0 2 1 0 -33.10014 151.2516 Wyong - South and West 13/08/2004 1 0 3 1 0 -33.10014 151.2516 Wyong - South and West 12/08/2004 1 0 -33.10014 151.2516 Wyong - South and West 11/08/2004 1 0 -33.10014 151.2516 Wyong - South and West 10/08/2004 1 0 -33.10014 151.2516 Wyong - South and West 9/07/2004 1 0 4 1 0 -33.10014 151.2516 Wyong - South and West 9/07/2004 1 0 -33.10014 151.2516 Wyong - South and West 8/07/2004 1 0 -33.10014 151.2516 Wyong - South and West 7/07/2004 1 0 -33.10014 151.2516 Wyong - South and West 6/07/2004 1 0 -33.10014 151.2516 Wyong - South and West 11/06/2004 1 0 5 1 0 -33.10014 151.2516 Wyong - South and West 11/06/2004 1 0 -33.10014 151.2516 Wyong - South and West 10/06/2004 1 0 -33.10014 151.2516 Wyong - South and West 15/05/2004 1 0 6 1 0 -33.10014 151.2516 Wyong - South and West 15/05/2004 1 0 -33.10014 151.2516 Wyong - South and West 14/05/2004 1 0 -33.10014 151.2516 Wyong - South and West 14/05/2004 1 0 -33.10014 151.2516 Wyong - South and West 12/05/2004 1 0 -33.10014 151.2516 Wyong - South and West 12/05/2004 1 0 -33.10014 151.2516 Wyong - South and West 11/03/2004 1 0 7 1 0 -33.10014 151.2516 Wyong - South and West 11/03/2004 1 0 -33.10014 151.2516 Wyong - South and West 16/02/2004 1 0 8 1 0 -33.10014 151.2516 Wyong - South and West 10/01/2004 1 0 9 1 0 -33.10014 151.2516 Wyong - South and West 10/01/2004 1 0 -33.10014 151.2516 Wyong - South and West 10/01/2004 1 0 -33.10014 151.2516 Wyong - South and West 9/01/2004 1 0 -33.10014 151.2516 Wyong - South and West 9/01/2004 1 0 -33.10014 151.2516 Wyong - South and West 8/01/2004 1 0 -33.10014 151.2516 Wyong - South and West 8/01/2004 1 0 -33.10014 151.2516 Wyong - South and West 7/01/2004 1 0 -33.10014 151.2516 Wyong - South and West 5/12/2003 1 0 10 1 0 -33.10014 151.2516 Wyong - South and West 5/12/2003 1 0 33 0

Table 3.3 (continued) Site 2, Case Study Wyong (South & West) New South Wales total rep. Bush Black Bush Black Latitude Longitude Survey Location Date each (different rat rat time same rat rat site place) -33.101364 151.250744 Wyong - South and West 8/10/2004 1 0 1 1 0 -33.101690 151.251531 Wyong - South and West 8/10/2004 1 0 -33.101364 151.250744 Wyong - South and West 7/10/2004 1 0 -33.101364 151.250744 Wyong - South and West 7/10/2004 1 0 -33.101364 151.250744 Wyong - South and West 6/10/2004 1 0 -33.101364 151.250744 Wyong - South and West 10/09/2004 1 0 2 1 0 -33.101443 151.251268 Wyong - South and West 9/09/2004 1 0 -33.101443 151.251268 Wyong - South and West 9/09/2004 1 0 -33.101690 151.251531 Wyong - South and West 9/09/2004 1 0 -33.102949 151.251216 Wyong - South and West 9/09/2004 1 0

-33.101364 151.250744 Wyong - South and West 8/09/2004 1 0 -33.101443 151.251268 Wyong - South and West 8/09/2004 1 0 -33.101690 151.251531 Wyong - South and West 8/09/2004 1 0 -33.102949 151.251216 Wyong - South and West 8/09/2004 1 0 -33.103719 151.250194 Wyong - South and West 12/08/2004 1 0 3 1 0 -33.103719 151.250194 Wyong - South and West 11/08/2004 1 0 -33.101443 151.251268 Wyong - South and West 8/06/2004 1 0 4 1 0 -33.102949 151.251216 Wyong - South and West 8/06/2004 1 0 -33.101443 151.251268 Wyong - South and West 8/01/2004 1 0 5 1 0 -33.101443 151.251268 Wyong - South and West 7/01/2004 1 0 -33.101443 151.251268 Wyong - South and West 7/12/2003 1 0 6 1 0 -33.101443 151.251268 Wyong - South and West 7/12/2003 1 0 -33.101443 151.251268 Wyong - South and West 7/12/2003 1 0 -33.101364 151.250744 Wyong - South and West 5/12/2003 1 0 -33.101364 151.250744 Wyong - South and West 5/12/2003 1 0 -33.101364 151.250744 Wyong - South and West 5/12/2003 1 0 -33.101690 151.251531 Wyong - South and West 5/11/2003 1 0 7 1 0 -33.101690 151.251531 Wyong - South and West 5/11/2003 1 0 -33.102949 151.251216 Wyong - South and West 5/11/2003 1 0 -33.101690 151.251531 Wyong - South and West 4/11/2003 1 0 -33.101443 151.251268 Wyong - South and West 3/11/2003 1 0 -33.101690 151.251531 Wyong - South and West 3/11/2003 1 0 32 0

Table 3.3 (continued) Site 3, Case Study Wyong (South & West) New South Wales total rep. Bush Black (different Bush Black Latitude Longitude Survey Location Date each rat rat time same rat rat site place) -33.133969 151.21792 Wyong - South and West 12/08/2004 1 0 1 1 0 -33.133969 151.21792 Wyong - South and West 11/08/2004 1 0 -33.133969 151.21792 Wyong - South and West 10/08/2004 1 0 -33.133969 151.21792 Wyong - South and West 3/12/2003 1 0 2 1 0 -33.133969 151.21792 Wyong - South and West 3/12/2003 1 0 -33.134 151.2182 Wyong - South and West 8/10/2004 1 0 3 1 0 -33.134 151.2182 Wyong - South and West 6/10/2004 1 0 -33.134 151.2182 Wyong - South and West 10/09/2004 1 0 4 1 0 -33.134 151.2182 Wyong - South and West 9/09/2004 1 0 -33.134 151.2182 Wyong - South and West 7/07/2004 1 0 5 1 0 -33.134 151.2182 Wyong - South and West 11/06/2004 1 0 6 1 0 -33.134 151.2182 Wyong - South and West 9/06/2004 1 0 -33.134 151.2182 Wyong - South and West 16/03/2004 1 0 7 1 0 -33.134 151.2182 Wyong - South and West 15/03/2004 1 0 -33.134 151.2182 Wyong - South and West 14/01/2004 1 0 8 1 0 -33.134 151.2182 Wyong - South and West 4/12/2003 1 0 9 1 0 -33.134 151.2182 Wyong - South and West 11/01/2003 1 0 10 1 0 -33.134 151.2182 Wyong - South and West 11/01/2003 1 0 18 0

Table 3.3 (continued) Site 4. Case Study Wyong (South & West) New South Wales rep. total Bush Black (different Bush Black Latitude Longitude Survey Location Date each rat rat time same rat rat site place) -33.134747 151.21805 Wyong - South and West 7/07/2004 1 0 1 1 0 -33.134747 151.21805 Wyong - South and West 11/06/2004 1 0 2 1 0 -33.134747 151.21805 Wyong - South and West 13/05/2004 1 0 3 1 0 -33.134747 151.21805 Wyong - South and West 16/04/2004 1 0 4 1 0 -33.134747 151.21805 Wyong - South and West 16/03/2004 1 0 5 1 0 -33.134747 151.21805 Wyong - South and West 15/03/2004 1 0 -33.134747 151.21805 Wyong - South and West 13/03/2004 1 0 -33.134747 151.21805 Wyong - South and West 13/01/2004 1 0 6 1 0 -33.134747 151.21805 Wyong - South and West 12/01/2004 1 0 -33.134747 151.21805 Wyong - South and West 2/12/2003 1 0 7 1 0 -33.134747 151.21805 Wyong - South and West 2/12/2003 1 0 -33.135471 151.21822 Wyong - South and West 9/09/2004 1 0 8 1 0 -33.135471 151.21822 Wyong - South and West 13/08/2004 1 0 9 1 0 -33.135471 151.21822 Wyong - South and West 8/07/2004 1 0 10 1 0 -33.135471 151.21822 Wyong - South and West 11/06/2004 1 0 11 1 0 -33.135471 151.21822 Wyong - South and West 15/05/2004 1 0 12 1 0 -33.135471 151.21822 Wyong - South and West 12/05/2004 1 0 -33.135471 151.21822 Wyong - South and West 17/04/2004 1 0 13 1 0 -33.135471 151.21822 Wyong - South and West 17/04/2004 1 0 -33.135471 151.21822 Wyong - South and West 16/04/2004 1 0 -33.135471 151.21822 Wyong - South and West 15/04/2004 1 0 -33.135471 151.21822 Wyong - South and West 16/03/2004 1 0 14 1 0 -33.135471 151.21822 Wyong - South and West 21/02/2004 1 0 15 1 0 -33.135471 151.21822 Wyong - South and West 14/01/2004 1 0 16 1 0 -33.135471 151.21822 Wyong - South and West 4/12/2003 1 0 17 1 0 -33.135471 151.21822 Wyong - South and West 3/12/2003 1 0 -33.135471 151.21822 Wyong - South and West 11/01/2003 1 0 18 1 0 27 0

Table 3.3 (continued) Site 5. Case Study Wyong (South & West) New South Wales rep. total Bush Black (different Bush Black Latitude Longitude Survey Location Date each rat rat time same rat rat site place) -33.302529 151.44119 Wyong - South and West 15/03/2007 0 1 1 0 1 -33.302529 151.44119 Wyong - South and West 14/03/2007 0 1 -33.303545 151.43308 Wyong - South and West 14/03/2007 0 1 -33.303545 151.43308 Wyong - South and West 13/03/2007 0 1 -33.303545 151.43308 Wyong - South and West 13/03/2007 0 1 0 5

Table 3.3 (continued) Site 6. Case Study Wyong (South & West) New South Wales rep. Bush Black total (different Bush Black Latitude Longitude Survey Location Date rat rat each site time same rat rat place) -33.332729 151.412109 Wyong - South and West 24/03/2009 0 1 1 0 1 -33.331577 151.408676 Wyong - South and West 28/06/2005 1 0 2 1 1 -33.331577 151.408676 Wyong - South and West 28/06/2005 0 1 1 2

Table 3.3 (continued) Site 7. Case Study Wyong (South & West) New South Wales rep. Bush Black total Bush Black Latitude Longitude Survey Location Date (different rat rat each site time same rat rat place) -33.336456 151.423630 Wyong - South and West 11/03/2009 1 0 1 1 1 -33.336456 151.423630 Wyong - South and West 11/03/2009 0 1 -33.339035 151.420407 Wyong - South and West 15/09/2003 1 0 2 1 1 -33.339035 151.420407 Wyong - South and West 15/09/2003 0 1

-33.339035 151.420407 Wyong - South and West 18/08/2003 1 0 3 1 0 -33.339035 151.420407 Wyong - South and West 28/07/2003 1 0 4 1 1 -33.339035 151.420407 Wyong - South and West 28/07/2003 1 0 -33.339035 151.420407 Wyong - South and West 28/07/2003 0 1 -33.339035 151.420407 Wyong - South and West 28/07/2003 0 1 -33.338654 151.322641 Wyong - South and West 10/12/2002 1 0 5 1 0 -33.338966 151.415036 Wyong - South and West 1/10/2001 1 0 6 1 1 -33.338966 151.415036 Wyong - South and West 1/10/2001 0 1 7 5

Table 3.3 (continued) Site 8. Case Study Wyong (South & West) New South Wales rep. Bush Black total Bush Black Latitude Longitude Survey Location Date (different rat rat each site time same rat rat place) -33.346821 151.374084 Wyong - South and West 18/03/2009 1 0 1 1 1 -33.346821 151.374084 Wyong - South and West 18/03/2009 0 1 -33.349159 151.366496 Wyong - South and West 19/05/2004 0 1 2 0 1 -33.348051 151.420244 Wyong - South and West 25/08/2003 1 0 3 1 0 2 2

Table 3.3 (continued) Site 9. Case Study Wyong (South & West) New South Wales rep. Bush Black total (different Bush Black Latitude Longitude Survey Location Date rat rat each site time same rat rat place) -33.233000 151.283000 Wyong - South and West 3/11/1929 1 0 1 1 0 -33.233000 151.283000 Wyong - South and West 3/11/1929 1 0 2 0

Total 122 14

Fig. 3. 3. Map of the Sydney Metropolitan Area showing black rats (Rattus rattus) in urban areas and their impact into surrounding areas of bush rat (Rattus fuscipes) populations along the east coast of New South Wales. (Source Atlas of NSW accessed October 2009).