Valuing Native Plantation Monocultures In

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2013 Valuing native plantation monocultures in Australia: the relationship between faunal richness and environment in Eucalypt plantations Beth Mott University of Wollongong

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Valuing native plantation monocultures in Australia: the relationship between faunal richness and environment in Eucalypt plantations

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

Doctor of Philosophy

from

The University of Wollongong

Beth Mott, B. Sc. (Hons)

School of Biological Sciences

2013 CERTIFICATION

I, Beth Mott, declare that this thesis, submitted in fulfilment of the requirements for the award of Doctor of Philosophy, in the School of Biological Sciences, University of Wollongong, is wholly my own work unless otherwise referenced or acknowledged.

The chapters in this thesis constitute original research carried out by myself during the period of candidature. I was the main contributor of the study design, and was solely responsible for data collection. I wrote the first draft of each manuscript and was responsible for responding to the editorial suggestions of those asked to comment on manuscripts.

The document has not been submitted for qualifications at any other academic institution.

Beth Mott July 2013.

Table of Contents

Abstract………………………………………………………………………………… xii Acknowledgements…………………………………………………………………… xvi

Chapter 1: The conservation value of native tree plantations for wildlife

Introduction……………………………….…………………….……………………... 1 Methods………………………………………………………………………..…….... 6 Data extraction……...……………………...….………………………...……. 6 Data Analysis…...……………………………………………………...……..………. 9 Results…………………………………………………………………………………. 13 Discussion……………………………………………………………………..……… 18 Continuing work in context……………………………………………………..……. 23

Chapter 2: The relationship between thermal and structural characteristics in managed forests: implications for microclimate diversity

Introduction……………………………….…………………….……………………... 26 Methods………………………………………………………………………..…….... 28 Study area…………………………………..……………………………………... 28 Forest types………………………………………………..…………………………..…. 29 Vegetation structure………………………………………………………………. 34 Thermal environment……………………………………………………………... 35 Data Analysis……………………………………………………………………… 38 Results………………………………………………………..………...……..………. 40 Vegetation structure………………………………………………………………. 40 Environmental temperature……………………………………………………… 41 Age related thermal differences within plantations……………………………. 42 Relating vegetation structure and thermal environment……………………… 46 Discussion……………………………………………………………………………... 47 Vegetation structure………………………………………………………………. 48

Environmental temperature……………………………………………………… 49 Plantation aging and effects on environmental temperature…………………. 52 Management recommendations and research………………………………… 53

Chapter 3: Faunal response to young afforested and old reforested Eucalypt plantations: do consistent patterns exist across animal taxa?

Introduction……………………………….…………………….……………………... 55 Methods………………………………………………………………………..…….... 57 Habitat descriptions………………………..……………………………………... 58 Sampling methodologies………………………………..…………………………..…. 59 Species characteristics..…………………………………………………………. 62 Data Analysis……………………………………………………………………… 63 Results………………………………………………………..………...……..………. 64 Birds………………………………………………………………………………… 66 Herpetofauna……………………………………………………………………… 67 Mammals…………………………………………………………………………... 68 Cross-taxa comparisons of guild structure……………………………………... 69 Discussion……………………………………………………………………………... 71 Patterns of richness………………………………………………………………. 71 Assemblage structure and ecological function………………………………… 73 Birds……………………………………………………………..…………………. 74 Ground fauna……………………………………………………………………… 76 The role of plantations in offsetting species loss………………………………. 77

Chapter 4: The influence of invertebrate resource heterogeneity on small ground-dwelling vertebrates

Introduction……………………………….…………………….……………………... 80 Prey availability……………………………………………………………………….. 81 Predators, prey and plantations…………………………………………………….. 82 Methods………………………………………………………………………..…….... 84

Invertebrate sampling……………………..…………….………………………... 84 Invertebrate hardness………………………..………..…………………………..…. 86 Relating predators and prey……………………………………………………... 87 Data Analysis……………………………………………………………………… 91 Results………………………………………………………..………...……..………. 92 Insectivores………………………………………………………………………… 92 Invertebrate prey…………………………………………………………………… 94 Discussion……………………………………………………………………………... 97 Predator abundance……………...………………………………………………. 98 Small predators……………………………...………………………………… 98 Medium predators…………………………………………..…………………. 100 Large predators..……………………………………………………………… 101 Grazing…………………………………………….………………………………. 102 Conclusions…………………………………………………………………………… 103

Chapter 5: Matrix quality and ground fauna diversity in young plantations Young native plantations in the landscape mosaic…..…….……………………... 105 The matrix and matrix quality.……………………………………………………….. 106 Methods………………………………………………………………………..…….... 110 Habitat descriptions……………………..…………….………………………...... 110 Vegetation structure………………………..………..…………………………..……... 112 Fauna trapping..…………………….…………………………………………….. 113 Data Analysis……………………………………………………………………… 115 Results………………………………………………………..………...……..………. 116 Vegetation structure….……………………………………………….…………… 116 Fauna…….…….…………………………………………………………………… 117 Discussion……………………………………………………………………………... 120 Matrix quality……….……………...………………………………………………. 120 The effects of grazing on plantations.…...………..………………………… 120 Age structure……...…………………………………………………………… 122 Conclusions…………………………………………………………………………… 123

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Chapter 6: The relative roles of structure and thermal environment in determining habitat choice by lizards in young plantations

Introduction……………………………….…………………….……………………... 125 Thermal environment and habitat choice……...……………………………….. 126 Thermal physiology of reptiles...………………………..……………………….. 127 Constraints of plantation environments on lizards…………………………….. 128 Methods………………………………………………………………………..…….... 131 Habitat choice enclosures………………..………..…………………………..…. 132 Data analysis….……………………………………………………….. 137 Results……………..…………………………………………………... 138 Feeding study…………………………………………………………………. 143 Data analysis…………………………………………………………... 145 Results…………………………………………………………………. 146 Discussion……………………………………………………………………………... 157 Preferred temperature range and habitat specialization……………………… 157 Substrate preference and habitat choice……………………………………….. 159 Rates of digestion….…………………………………………..…………………. 161 Conclusions…………………………………………………………………………… 164

Chapter 7: General Discussion

The value of plantations as habitat – an overview.…..…….……………………... 166 This research in context.…………………………………………………………….. 167 Woodland birds.……………………..…………….………………………...... 169 Forest species…………………………..………..…………………………..……... 171 Small mammals..….…………….…………………………………………….. 172 Food availability……………………………………………..………...……..………. 173 Matrix quality…………….……………………………………………….…………… 174 Plantations and the thermal environment………………………………………….. 177 Plantations and climate change…………………………………………………….. 178 The way forward………………………………………………………………………. 179

Conclusions…………………………………………………………………………… 183

References………………………………………………………………………….. 185

List of Tables

Table 1.1 Studies examined in this meta-analysis comparing forests and native

plantations………………………………………………………..…………….. 7

Table 1.2 Q tests for meta-analyses testing for differences in effect sizes (E+)

with changes in species richness and abundance between forests and 16

plantations…………………………………………………………………………..

Table 2.1 Vegetation parameters and the methods used to record them from four

forest types on the Mid North Coast of NSW………………..…………………. 36

Table 2.2 Results of RMANOVA of change in environmental temperature with

forest type …………………...…………………………………….……………….. 43

Table 2.3 Results of ANOVA and regression analyses relating vegetation

structure to solar radiation and environmental temperature levels…………… 45

Table 3.1 Means (± 1se) of vegetation variables sampled from replicated 15m2

quadrats in old growth native forest, logged native forest, old and young

plantations……………………………………………………………………… …. 59

Table 3.2 Results of ANOSIM analysis comparing assemblage structure in four

eucalypt forest types for four vertebrate taxa on the mid north coast of New

South Wales, Australia……………………………….…………………………… 66

Table 3.3 ANOVA results comparing the richness, abundance and habitat

specialization of birds, mammals and herpetofauna sampled in the study

area………………………………………………………..….………..……..…….. 67

Table 3.4 Results of ANOSIM analysis comparing the distribution of feeding

guilds in four eucalypt forest types for birds, mammals and herpetofauna…. 71

Table 3.5 Summary of the ecological character of birds, mammals, reptiles and

frogs in eucalypt plantations on the mid-north coast of New South Wales….. 75

Table 4.1 Mean gape size and prey length category used by each predator

species sampled…………………………………………………………………… 88

Table 4.2 ANOSIM results testing for changes in small medium and large

insectivore abundance in forests and plantations…….. ………………………. 93

Table 4.3 Results of χ2 analyses and associated pairwise comparisons for

changes in the frequency of preferred, available and unavailable prey

between forest types ……………………………………………………….…….. 95

Table 5.1 Vegetation variables measured to quantify structural differences

between grazed native forest and young native plantation……………….…… 113

Table 5.2 Results of paired t-tests for changes in vegetation structure between

grazed native eucalyptus woodland and adjacent native plantations…..……. 117

Table 5.3 Results of paired t-tests for changes in the richness and abundance of

reptile and mammal assemblages in plantations and native forests…………. 118

Table 6.1 Body temperatures and habitat associations of target species in

experimental choice boxes on the Mid North Coast of New South Wales…... 139

Table 6.2 Results of mixed model analysis for change in body temperature with

position in experimental choice boxes…………………………………………… 140

Table 6.3 Estimates of fixed effects for mixed models analysis of appetite and gut

passage rate from laboratory experiments for each lizard species…………... 156

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List of Figures

Fig 1.1 Forest plots of standardized differences in species richness and

abundance between forests and native plantations...... 17

Fig. 2.1 A map of the study 50 x 80km study area on the Mid North Coast of

NSW………………………………………………………………………………. 31

Fig 2.2 Diagrammatic representation of the structural complexity of vegetation in

plantations………………………………………………………………………… 32

Fig 2.3 Principal co-ordinates biplot showing the distribution of vegetation

variables in relation to forest types……………………………………………. 42

Fig 2.3 Circadian profiles for mean environmental temperatures available in the

shade and sun throughout the course of the day from 6am to 6pm, in each

of four forest types………………………………………………………………. 44

Fig 2.4 Circadian profiles for mean environmental temperatures available in

early maturity and young plantations in the sun and shade throughout the

course of a day…….…………………………………………………………….. 45

Fig 2.5 Scatter plots depicting the relationship of radiant energy/ environmental

temperature variables with vegetation variables……………………………… 47

Fig 3.1 Species richness of birds, mammals and herpetofauna sampled from

forest and plantation sites on the Mid North Coast of NSW in 2001……….. 65

Fig 3.2 Mean species richness and abundance of vertebrate taxa grouped by

habitat preference……………………………………………….……………….. 68

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Fig 3.3 Distribution of mammal, reptile, bird and frog feeding guilds between

forest types……………………………………………………………………….. 70

Fig 4.1 Cumulative abundances of small medium and large insectivores

sampled from old growth and logged native, and old and young native

plantations on the Mid North Coast of New South Wales………………..….. 93

Fig 4.2 The frequency of preferred, available and unavailable prey present in

each forest type for each predator group……………………………………. 96

Fig 5.1 Two six-year-old hardwood plantations showing typical connection to

other landscape elements………………………………………………………. 108

Fig 5.2 Mean abundance of forest, woodland and generalist fauna and the age

structure of samples populations in grazed woodland and adjacent young 119

native plantations…………………………………………………………………

Fig 6.1 Choice enclosures in situ on the Mid North Coast of NSW. A. Large

enclosures; B. Diagrammatic representation of the relative amount of

ambient radiant energy and substrate present in each compartment. C.

An example of the complex (left) and simple (right) substrates lizards

could select……………………………………………………………………… 134

Fig 6.2 Diurnal profiles of levels of insolation and environmental temperatures

available to lizards in field enclosures throughout the course of the day;

with preferred temperature range overlaid on available

temperatures …………………………………………………………………… 141

Fig 6.3 Thermal profiles of available environmental temperatures in young

plantation habitats, overlaid by mean preferred temperature ranges of

‘hot’ and ‘cold’ lizards.…………………………………………………………. 142

Fig 6.4 Appetite and gut passage rates of for the four species tested

ix experimentally at (200C) spring, (250C) spring-summer and summer

(300C) temperatures available in young plantations on the Mid North coast of New South Wales……………………………………………………. 156

List of Appendices

Appendix 1: Table 1 Summary statistics and species list for all bird species recorded in the trapping period between 2001 and 2003 from the North Coast Bioregion…………………………………………………………………………..…226

Table 2 Summary statistics and species list for all mammal, frog and reptile species caught in the trapping period between 2001 and 2003 from the North Coast Bioregion………………..…………………………………………….227

Appendix 2

Summary statistics for all herpetofaunal and mammal species recorded

from young and old plantations, logged and old growth native forests

in twenty field sites on the Mid-North Coast of NSW between 2002 -

2003…………………………………………………………………………………. 228

Appendix 3

Summary statistics for all herpetofaunal and mammal species recorded from

young plantations and adjacent native forests ten field sites on the Mid-North

Coast of NSW between 2002 and 2004…………………………………………..229

Abstract

Native forests throughout the world are disappearing at rate of approximately 13 million hectares a year. One of the ways forest managers are attempting to halt deforestation is by growing trees in plantations to provide wood products. Industrial tree plantations will occupy 10-20% of the total forested land area in the world by 2024. Research into the suitability of plantations for wildlife has grown rapidly in the last fifteen years, but it has focused primarily on exotic plantations. Comparatively little work has focused on native tree plantations, which may better benefit biodiversity in forested landscapes if they act like forests. A literature review of existing work identified that age, size, level of management and the type of forest adjacent to plantations can all affect the diversity of faunal assemblages using native plantations, and that monocultures are generally species poor. This thesis extends the small existing body of work on managed native plantations by comparing faunal use of young and old native plantations with logged and old growth native forests in the North Coast Bioregion of sub-tropical south-eastern

Australia. This thesis used trapping to identify whether birds, mammals, and herpetofauna used plantations differently from forests, and ecophysiology, landscape and community ecology theories to explain why they do so.

Plantations collectively, differed from forests in vegetation structure and available microclimate at the ground level. The open canopy and lack of sub

xii canopy complexity in young and old plantations resulted in higher insolation at the ground in plantations, which increased environmental temperature up to

200C above that of old growth native forests in summer during the day. Young plantations particularly presented a challenging environment to potential colonizers sensitive to microclimate. Large daily fluctuations in temperature and irradiance, coupled with the residual effects of management-associated soil compaction may have explained the loss of forest organisms and the persistence of simple understorey in old plantations, despite 30 years of growth and more than six years since disturbance by fire or harvesting.

Early maturity (spar) plantations differed from young and old plantations in having similarly simple understorey structure, but closed canopies, producing a cool thermal environment with low light penetration. Although spar plantations are a short-lived stage in the plantation cycle before the canopy is opened by subsequent harvesting, they could represent a valuable habitat resource for forest fauna that prefer cool thermal environments but are tolerant of simple structure. From a management perspective if this stage of plantation is promoted within the landscape mosaic at forest-plantation boundaries, it may provide forest species with additional habitat, and buffer forest from the microclimatic extremes experienced in young and old plantations.

Despite large thermal and structural differences herpetofauna, birds and mammals were equally speciose in old growth and logged native forests and young and old plantations. Assemblage structure differed significantly, and plantations were dominated by generalists and woodland species. Small ground

xiii mammals were less abundant in young plantations. Thus while afforested plantations provided habitat where habitat was previously absent, in both young and old plantations the ecological resilience of communities may have been compromised by lower species richness. From a landscape perspective plantations offered habitat to declining woodland birds, and increased the functional range of woodland species in agricultural environments.

Using small lizards as a model and conducting feeding and behavioral choice experiments, this thesis demonstrated that the thermal environment in young plantations was physiologically limiting to forest species. While temperatures optimal for digestive efficiency could be achieved, they required the species using this forest type to exhibit flexibility in both activity time and substrate choice. The forest species that did not possess this flexibility could not use young plantations. Those species that had the physiological capacity to exploit plantations were behaviourally constrained, and would only be likely to use young plantations if targeted understorey enrichment were provided.

The low abundance of insectivorous ground fauna identified from young plantations in this research was not driven by a lack of invertebrate prey, which was equally available in young and old plantations and native forests. Low mammal abundance was likely to have been related to the simplicity of the plantation environment rather than a decline in forest matrix quality associated with cattle grazing. However for herpetofauna, grazing may have increased the permeability of the forest matrix, with a resultant influx of thermophilic species into plantations.

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Collectively the results of this research stress that: 1) young and old native plantations can support faunal assemblages as diverse as those in native forests, but for small mammals in particular, abundance in plantations is low; 2) cooler spar plantations have the potential to act as thermal stepping stone which may promote incursions of forest species into plantation mosaics; 3) young and old plantations can support an invertebrate prey base as diverse as that in adjacent native forests, with positive benefits on the species richness of insectivores; and 4) the role of grazing in the forest matrix requires further investigation in order to clarify its role in influencing species exchange between plantations and forests.

Acknowledgements

So where does one begin…….at the beginning I guess. Thanks go to all those people that helped this piece of work come together. Peter Levitske (SFNSW

Wauchope) provided great initial info. Andrew Marshall (NPWS Port Macquarie) provided an Anabat . Mike Thompson at USYD generously lent me his data loggers.

Brad Law from SFNSW Pennant Hills trustingly shared his data.

Never will I forget the people who came and dug holes just for the fun of it including: Adrian Ferguson, Ittai Renan (who loved phasmids with me), Stephen and

Tarn (who dug many many holes), and Johann who laughed in the face of bats one very, very hot Christmas week.

My supervisors deserve much thanks for support and dedication. Kris French was unendingly patient, and made sure I was not lost in the forests literally and figuratively; Bill Buttemer gave comments to improve my science. Frank Lemckert just was.

John and Janette Cook get a very special mention for lending me their luxurious shed and a site for field enclosures. Also for sharing their lives with me for what seems like only a minute, but was actually nearly 4 years. May they be blessed by the many gods for being so open-hearted, for sharing stories, letting me shovel woodchip, camp with their redheads down the back and jump in the dam one lovely New Year. My thanks endure.

Elsewise and northwards there were all the rest of my people. Simon gifted invaluable science talk, encouragement and laboured through drafts like a man possessed. Pod gave statistics and smiles; Sam and Leonie reminded me of sunshine.

Kathleen talked turtles and fish from afar and Fiona kept me dreaming of snow. Of my southern folk I must thank my lab, in particular Tanya who remains ever lovely and

xvi whose professional attitude was inspiring. Also thanks go to Craig who was fun.

Gerhard talked good science throughout. Mats gave me hatchlings to love. I hope you feel me on this one people. I am beyond grateful for all your support.

My family deserves special mention. Mum bent fence poles for days at the beginning of this project, sent money and help when it was needed, scrubbed traps multiple times, built field enclosures whilst dodging snakes, babysat and cheered. If I can imitate one quarter of the strength, beauty and generousness of this woman then I’ll be doing well. Stephen painted my truck and drove north to rescue mountains of gear at the end and did everything else in between. Tarn shared the forest with me, talked muesli, and believed. James gave moments of joy, and missed me. You’ve all heard my thanks, but hear it again now. You’re incomparable. Along with these people I must mention the Frenchman. His patience has been enduring and his love boundless.

Bunya made fencing fun and dreamed on my feet while I wrote. My exquisite daughters have given me their patience and a heart overflowing. I am so blessed to share them.

Finally I want to thank this beauteous Earth. She provided me with the great joy

I took from this project, and lent me her people for a moment. I am deeply honoured.

Now I just have to work out how to pay that one back.

Much peace.

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Chapter 1: Introduction

Chapter 1 The conservation value of native tree plantations for wildlife

Introduction

Recognition of 2011 as The International Year of The Forests highlights the current interest in the world’s forests, with growing awareness of their vital role in maintaining a stable global climate. Native forests throughout the world are being converted to other land uses at a rate of approximately 16 million hectares a year (FAO 2010), in conjunction with a rapid decline in biodiversity (Perfecto & Vandenmeer 2010). One widely adopted international initiative to halt global forest decline involves establishing tree plantations to provide a source of wood products to the industrial systems which use more than ninety percent of the timber harvested from forests (Kanowski 1997). This approach is working and although global forest loss is not offset by the current rates of afforestation, it is mitigated, reducing the total per year loss to 5.2 million hectares a year

(FAO 2010).

Plantations currently represent 6.6% of the world’s forests (FAO 2011), and estimates of global expansion project that this will increase to between

10 and 20% in the next 15 years (Sample 2003). While industrial plantations have conservation significance because they reduce timber harvesting in native forests (FAO 2011), international bodies such as the Forest

Stewardship Council recognize that plantations have inherent biodiversity conservation value that should be promoted, especially where plantations

1 Chapter 1: Introduction

dominate the local landscape (Mercer & Underwood 2002). Promoting plantations as a management practice, associated with sustainable forest management, concerns some conservationists (Spellerberg 1997, Clapp

2001), primarily because there is little agreement about how plantations do benefit the environment (Bremer & Farley 2010). While timber plantations can play a significant role in conserving biodiversity by reducing timber harvesting in native forests (Sedjo & Botkin 1997, Brockerhoff et al. 2008), their value for flora and fauna as new habitat on afforested lands relies strongly on the impact establishment has on the landscape in which they are embedded (Bremer & Farley 2010) as well as the context of the plantation in the landscape mosaic (Felton et al. 2010). How well plantations can compensate for natural forest loss by acting as natural habitats is directly affected by the similarity between native forests and plantations in tree species composition (Brockerhoff et al. 2008, Bremer & Farley 2010), the structural and physical environment this engenders, and the management regime applied (Brockerhoff et al. 2012).

How well plantations support biodiversity is not always clear, often because the term ‘plantation’ can refer to a large range of stand types and tree species mixtures (Hartley 2002), which are often exposed to very different management regimes depending on the end use of the plantation, establishment goals, ownership, management practice and wood product end use. For the purpose of this paper a plantation is defined following the global plantation forests thematic study (FAO 2006) which identifies productive plantations as “forests of introduced and in some cases native species, established through planting or seeding mainly for the production of

2 Chapter 1: Introduction

wood products”. Typically large-scale tree plantations are planted and managed as monocultures to homogenise the quality of fibre inputs for pulp production (ABARE 1999a) and to ease mechanised harvesting. Unlike natural forests, managed monocultures lack structural complexity; instead being characterised by evenly aged and distributed stands of trees (Evans

1997, Turnbull 1999) and open and simple canopies. Specifically, they lack the horizontal and vertical structural complexity associated with multilayered canopies, logs and snags and uneven tree sizes (McCullough 1999). The understorey in monoculture plantations is typically either missing or highly simplified, however, without ongoing management, plantation understoreys can develop high vegetation diversity depending upon levels of crown closure and propagule availability (Bowen et al. 2007, Archibald et al. 2010). Thus, how different plantations are from forests can vary greatly with the type of plantation examined.

The value of plantations for fauna has been reviewed several times by

Christian et al. (1998), Moore and Allen (1999), Lindenmayer and Hobbs,

(2004), Kanowski et al. (2005), Carnus et al. (2006), Gardner et al. (2007a),

Stephens and Wagner (2007), Brockerhoff et al. (2008), Munro et al. (2009),

Hartmann et al., Felton et al. and Nájera and Simonetti (2010). These reviews find no general consensus that plantations either promote or decrease faunal diversity. Such a lack of consensus is unsurprising given that vegetation species, climate, site conditions, management, scale, age and species composition can all affect faunal responses to plantations

(Hartley 2002, Lindenmayer & Hobbs 2004, Gardner et al. 2007b), and these reviews combine results across plantation ages and differing landscape

3 Chapter 1: Introduction

contexts. Further, the lack of consensus between reviews may also be compounded by a failure to differentiate different types of plantations, which occurs in 80% of past reviews that assess the suitability of plantations as faunal habitat (e.g. exotic and native -Felton et al. 2010; timber plantations and other tree crops such as rubber -Náejera & Simonetti 2009). Pooling of plantation types in research can obscure particular types of plantations that may better represent faunal habitat. The overarching conclusion of past reviews is that plantations support lower total species diversity than forests, but that particular groups of fauna may be abundant. Individual studies have identified that plantation faunal assemblages are often dominated by generalists, edge specialists and exotics (Bengtsson et al. 2000, Davis et al.

2000, Boorsboom et al. 2002, Gardner et al. 2007b). Whether different sorts of plantations can support a diverse array of fauna and by extension, and how well plantation faunal assemblages provide ecosystem services (e.g. pest control, pollination) is unclear (Carnus et al. 2006). Without this sort of knowledge, advocating that plantations generally have conservation value for fauna or that they can extend faunal habitat (Paquet & Messier 2010, FAO

2011), must be qualified to certain faunal groups.

Plantations of native tree species may have the potential to better represent faunal habitat than exotic tree plantations (e.g. Farwig et al. 2008,

Bremer & Farley 2010) because they provide familiar microenvironments, food and habitat resources for vertebrate and invertebrate fauna (Hartley,

2002). Understanding the value of native plantations is rapidly becoming important as native species planting rates increased globally by 49% between 2005 and 2010, and in Australasia the establishment rate of native

4 Chapter 1: Introduction

plantations currently equals those of exotic species (FAO, 2010). In the last ten years a large body of literature examining the value of native plantations for biodiversity has emerged. This literature primarily focuses on plantations generated for protective functions, whilst research focussing on timber production plantations remains a narrow field, even though industrial timber plantations comprise >60% new plantations established globally (FAO 2010).

This chapter examines the conservation value of managed timber plantations of native tree species for vertebrate fauna. To assess whether there is a literary consensus that native plantations can act as reservoirs of for diverse faunal assemblages, we used meta-analysis techniques to determine whether stand-level faunal species richness and abundance was lower in native timber plantations than native forests, and how consistent this effect was across animal taxa. The effect of simplified understorey complexity on species richness and abundance was also examined, as understorey complexity has previously been recognised as a major determinant of diversity for all animal groups in plantations (birds- Nájera & Simonetti 2010, mammals - Ecke et al. 2002, reptiles – Brown & Nelson 1993, invertebrates –

Humphrey et al. 1999). We hypothesized that the richness and abundance of ground mammals, reptiles, birds and invertebrates would decline from forests to native plantations with complex understories, and that plantations with simple understories would support the least rich and abundant faunal assemblages.

5 Chapter 1: Introduction

Methods

Data extraction

We searched the electronic databases ISI Web of Science, Google and Google Scholar for literature examining faunal species diversity with plantations. Additional information was obtained from public and private reports and by back referencing published studies on faunal use of plantations (Moore & Allen 1999, Hartley 2002, Lindenmayer & Hobbs 2004,

Carnus et al. 2006, Stephens & Wagner 2007, Brockerhoff et al. 2008,

Nájera & Simonetti 2009).

While literature addressing physio-chemical properties of native plantations and their role in reafforestation and agroforesty (particularly palm oil, coffee and cacao) was common, there were few studies comparing managed native plantations with forests of the same or closely-related species. 224 studies investigated native monoculture plantations but only 35 examined fauna, and of these only 21 presented quantitative data on species richness and/or abundance. Most studies were excluded from analysis after failing to adequately define either the age of the plantation under investigation, the condition of the understorey or the level of management experienced in the plantation. 55% of cases focussed on birds, 35% focussed on invertebrates, and 15% on reptiles or mammals (Table 1). The spread of research was adequate and originated from temperate and tropical

South America, the Mediterranean, the Boreal North, temperate and tropical

Oceania, Central Africa and North America. Australian research contributed

52% of all records. Qualifiers were necessary to avoid confounding sources

6 Chapter 1: Introduction

Table 1. Studies examined in this meta-analysis comparing forests and native plantations.

Publication Dominant tree species Plantation age Understorey complexity Taxon investigated Country cases Farwig et al.2008 Maesopsis eminii old complex birds Africa 2 Klomp & Grabham 2002 Eucalyptus spp. young simple birds Australia 1 Loyn et al. 2007 Eucalyptus globulus young simple birds Australia 1 Loyn et al. 2009 Eucalyptus globulus young simple birds Australia 1 Proença et al. 2010 Quercus spp. old simple birds Portugal 2 Repenning & Labisky 1985 Pinus palustris old complex birds North America 3 Volpato et al. 2010 Arucaria angustifolia old complex birds South America 1 Zurita et al. 2006 Araucaria angustifolia old complex birds South America 1 Hsu et al. 2010 Eucalyptus pilularis young simple birds Australia 1 Boorsboom et al. 2002 Eucalyptus spp. old, young complex birds, mammals, reptiles Australia 8 Kavanagh et al. 2005 Eucalyptus spp. old, young simple, complex birds, mammals, reptiles Australia 18 Fonseca et al. 2009 Arucaria angustifolia old complex birds, mammals, invertebrates South America 3 Bentley et al. 2000 Araucaria cunninghamii old complex mammals Australia 1 Kanowski et al. 2005 Araucaria cunninghamii young simple reptiles Australia 2 Cunningham et al. 2005 Eucalyptus globulus young simple invertebrates Australia 1 Schnell et al. 2003 Eucalyptus punctata young simple invertebrates Australia 1 Taboada et al. 2008 Pinus sylvestris old, young complex invertebrates Spain 3 Tattersfield et al. 2001 Maesopsis eminii old complex invertebrates Africa 1 Yu et al. 2008 Larix mastersian old simple invertebrates China 1 Grimbacher et al. 2007 Araucaria cunninghamii old, young simple, complex invertebrates Australia 4 Gries et al. 2012 Eremanthus erythropappus young simple invertebrates South America 2

7 Chapter 1: Introduction

of variability when comparing forests and plantations. Research was included in this meta-analysis if it met the following criteria:

Plantations:

- Plantations of trees native to a landscape but planted where they were not

endemic, were deemed native. All non-timber plantation commodities were

excluded.

- This meta-analysis did not include studies in which clear-cutting and

subsequent natural regrowth resulted in even aged forests. Plantations were

defined as stands of trees specifically planted for timber production.

- Only plantations with a documented management history were included.

- Understorey was classified as structurally complex or simple as reported by

the research author/s. Where understorey complexity was not specifically

reported but vertical foliage density was quantified, understorey was scored

as simple or complex based upon results of analysis by the individual author

(e.g. Farwig et al. 2008).

- Plantation age was categorized as young (2-10 years old) or old (+20 years

old), as it is recognized as strongly affecting species diversity in plantations

(Hartley 2002), primarily through changes in canopy cover and understorey

complexity.

Forests:

To avoid including research biased by island effects (Gascon et al. 1999) forests had to meet the following criteria of being:

- More than ten hectares in size;

8 Chapter 1: Introduction

- Unlogged for a period of at least 20 years prior to sampling;

- Situated within 15 kilometers of plantations.

Fauna:

- Exotic species were excluded from species lists and only species recorded

during formal surveys were included.

- Mammal data were restricted to small ground mammals and invertebrate

data excluded soil dwelling and volant species, as records for large

mammals, soil fauna and volant species of all taxa were inconsistently

reported between studies.

Each research article was scored once per plantation-forest comparison for species richness and/or abundance of invertebrates, mammals, birds or reptiles.

Studies that reported several sets of results were scored more than once if the data presented were obtained from independent experiments, or where multiple ages of plantation fitting the age criterion above were investigated. This created a correlated data structure for some studies.

Data analysis

Data analysis involved compiling estimates of species richness/abundance calculated from species lists when not presented directly.

The corresponding estimates of standard deviation were back-calculated from sample size and estimates of standard error, using standard mathematical techniques. These were measured from graphs if not directly presented. The data set available for this meta-analysis faced two challenges; it was small and the data structure was correlated where multiple responses to a single control

9 Chapter 1: Introduction

were recorded. Whilst averaging the responses of the correlated data or randomly choosing a single response from a correlated set is a solution to correlation (Lipsey & Wilson 2001), it necessarily reduces the power of an analysis (Marín-Martínez & Sánchez-Meca 1999, Hochachka et al. 2007). In this meta-analysis such averaging would have also confounded plantation age categories. Instead, effect sizes which measure the magnitude of effect present in each study were generated by calculating Hedge’s g, a metric commonly used in ecological studies (e.g. Osenberg et al. 1997; Mason et al. 2009).

Hedge’s g calculates standardized mean differences around treatment means, by dividing the difference between the mean of the treatment and control groups by the pooled standard deviation of both groups (Hedges & Olkin 1985), and uses a generalized least squares framework which can be modified to address violations due to non-independence of data. However it is known to be biased towards Type I errors at small sample sizes (Hedges & Olkin 1985).

Whilst Response Ratio (RR) is a metric known to have better statistical properties for small data sets, Lajeunesse and Forbes (2003) identified that whilst RR generated fewer Type I errors (detecting an effect when none is actually present) than Hedge’s g when there were less than 15 studies, its precision did not improve with sample size above ten studies. Whilst a multiple lines of response approach to analyzing data, using both RR and Hedge’s g has been adopted by other researchers with small data sets (see Van Zandt &

Mopper 1998, Kopper et al. 2009) this method can lend itself to selective interpretation of results. For this meta-analysis, Hedge’s g was generated to

10 Chapter 1: Introduction

measure effect size, with the application of correction factors for small sample size and for correlation. Correction for correlation multiplying g by the shrunken correlation coefficient (rs) (Campbell 1980) for each correlated study. Correction by rs weighted the strength of the relationship between means by the number of variables and the number of data points (Kufs 2011), and so reduced the within- study variance, increasing the precision of the estimate of effect size (Hedges &

Olkin 1985). Rs were calculated as:

( ) rs = 2 1−�1−푟 � 푛−1 푛−2 However, for 50% of the correlated cases used in this meta-analysis, and for many published studies, data allowing calculation of correlations are often unavailable (Wampold et al. 1997). Where shrunken correlation could not be calculated a correlation of 0.5 was chosen to aggregate the effect sizes. This was assumed to be a reasonable estimate of correlation, as where rs could be calculated from the studies included in this meta-analysis, values fell between

0.49 and 0.57.

Subsequently, Hedge’s g was derived from:

g = for uncorrelated data 푋�푇 − 푋�퐶 and from Sp 퐽

g = for correlated data 푋�푇 − 푋�퐶 푠 Sp∗� 1−푟 � √ 퐽 with

11 Chapter 1: Introduction

( ) ( ) S J 1 p = 2 2 and = ( ) 푛1−1 푠푇+ 푛2−1 푠퐶 3

� 푛1+ 푛2 − 2 − 4 푛1+ 푛2−2 −1 where was the mean plantation species richness or abundance, was the

푇 퐶 mean forest푋� species richness or abundance, Sp was the pooled standard푋� deviation, J was a correction factor for small sample bias, , , and were

1 2 1 2 the forest and plantation sample sizes and standard deviations푛 푛 respectively.푠 푠 The degree of deviation from zero represented the strength of an effect, and negative values of effect size reflected an increase in species richness/abundance in forests. Effect size distributions were normalized before analysis by removing the study by Gries et al. (2012) and two cases from the study by Grimbacher et al. (2007), which were identified in

normal quantile plots as highly non-normal.

After examination of the data, analyses of plantation age were abandoned for all analyses except bird richness, as the data available for old plantations with simple understory was too small to allow meaningful statistical comparisons. All other analyses were pooled across plantation ages. Random- effects models were used to identify any differences in species richness or abundance between plantations and forests for all fauna. Random effects models identify heterogeneity in effect sizes by estimating the mean of a distribution of true effect sizes, and so account for within-study heterogeneity in the data originating from moderator variables (understorey complexity) and

12 Chapter 1: Introduction

between-study heterogeneity stemming from differences in methodology and design (Borenstein et al. 2009). For mammals the lack of heterogeneity in the abundance data required the use of a fixed effects model, as no data were available for mammals in plantations with simple understories.

Confidence intervals around corrected g estimates used to compare the effect of understorey complexity on richness, were calculated using resampling techniques with 20 000 iterations. Overlap between confidence intervals can be as informative as p-values themselves in meta-analysis (see Durlak 2009), and non-parametric estimates are less biased by sample size (Gliner et al. 2003). All analyses were carried out using the MetaWin program, version 2.1 (Rosenberg et al. 1997). Effect sizes were considered significant at the p < 0.05 level, and mean effects were rated as small (0.2). medium, (0.5) or large (≥0.8) following

Cohen’s rule-of-thumb (1988). In forest plots generated to depict the spread of effect sizes, a shift towards to the left of zero was interpreted as a stronger effect in forests, whilst a shift towards the right of zero indicated larger effect sizes in plantations (see Rosenberg et al. 1997).

Results

Hypothesis: plantations with simple understories support less rich or abundant faunal assemblages than plantations with complex understories or forests.

In opposition to the hypothesis posited, native plantations and forests supported similarly rich faunal communities (Table 2). Negative effect sizes indicated a consistent non-significant trend for species richness to be higher in

13 Chapter 1: Introduction

forests, as indicated by the skew of mean effect sizes to the left of Figure 1a.

Whilst understorey condition did not affect overall significance it did moderate species richness. When understories were complex bird, mammal and reptile richness increased. Bird richness was equivalent between forests and plantations with complex understories (confidence intervals bracket zero), and between young and old plantations, but became more variable in plantations with simple understories (Table 2). These trends were strong (mean effect sizes >

0.701) and consistent across both young and old plantations. As for birds, reptile richness was most similar between plantations with complex understories and forests as is reflected by the confidence intervals around the mean effect size approaching zero (Table 2), and the clumped distribution of mean effects in

Figure 1a. This result was considered to be moderately strong (mean effect size

= -0.594). For invertebrates an effect of either plantation understorey development or overall forest type was not detectable as high variability between studies obscured trends (Fig. 1). A strong mean effect (-0.830), was offset by a high mean study variance ratio (2.859) suggesting the response of invertebrate richness to changes in forest structure was affected by unmeasured variables. For mammals, whilst Figure 1a shows a spilt between simple and complex plantations and a trend towards complex plantations supporting higher richness, a weak effect size (E+ = -0.31, mean variance ratio = 1.594) suggested that accurate conclusions for this group required a greater sample size.

14 Chapter 1: Introduction

In contrast to species diversity, the abundance of reptiles, invertebrates and birds differed significantly between forests and plantations. Plantation understorey complexity influenced abundance in complex ways for each animal taxon. For birds, invertebrates and mammals when plantation understories were complex species richness was equivalent in forests and plantations (confidence intervals for mean effect sizes bracketed zero, Table 2). Where plantations had simple understories bird abundance was significantly (p < 0.000) and consistently (mean study variance ratio = 0.833) reduced. Reptiles were similarly abundant in forests and plantations with simple understories

(confidence intervals bracket zero), but abundance was highest in plantations with complex understories (Table 2, mean effect size = -1.075, Fig 1b). As for richness, invertebrate abundance showed the highest variability in response to forest change of the four taxa examined in this meta-analysis (mean study variance ratio = 1.067). Invertebrates were the only group in which abundance was strongly negatively associated with complex understorey (mean effect size

= 2.168, Table 2).

15 Chapter 1: Introduction

Table 2. Q tests for meta-analyses testing for differences in effect sizes (E+) with changes in species richness and abundance between forests and plantations. Analyses are repeated for birds, mammals, invertebrates and reptiles. Degrees of freedom = 1 for all random analyses and 8 for the fixed analysis of mammal abundance. n = the number of comparisons between plantations with simple (s) and complex (c) understories. OP = old plantations, YP = young plantations.

Understorey Bootstrap CI Faunal group condition n Q p E+ Mean E+ lower upper Species richness Birds OP c 10 3.278 0.070 -0.310 -0.728 -0.879 - 0.101 s 5 -1.397 -2.780 - -0.419 YP c 3 0.256 0.613 -0.674 -0.911 -2.157 - 0.856 s 5 -1.079 -1.786 - -0.556 Mammals c 6 0.097 0.755 -0.223 -0.311 -0.845 - -0.786 s 4 -0.388 -0.867 - 0.099 Invertebrates c 5 0.112 0.738 -0.573 -0.830 -2.157 - 0.857 s 5 -1.053 -4.025 - 1.123 Reptiles c 6 0.114 0.736 -0.540 -0.594 -0.901 - -0.071 s 8 -0.637 -0.852 - -0.328

Abundance Birds c 11 11.640 <0.000 0.181 -0.226 -0.249 - 0.558 s 5 -1.347 -2.314 - -0.660 Mammals c 8 7.870 0.446 -0.089 -0.938 - 0.794 Invertebrates c 3 4.257 0.039 -0.119 0.562 -1.014 - 0.783 s 2 2.168 1.259 - 4.492 Reptiles c 8 5.356 0.021 -0.127 -1.075 -1.708 - -0.901 s 2 -1.281 -0.390 - 0.134

Whilst abundance in was equivalent in forest and plantations with complex understories, it increased significantly in plantations with simple understories

(mean effect size = 2.168, Table 2, Fig. 1b). Similarly, mammals were equally abundant in old plantations with complex understories and forests (Table 2, Fig

1b), however, this result was considered weak (mean effect size = -0.089), and the distribution of effect sizes was bimodal (Fig. 1b).

16 Chapter 1: Introduction

Fig.1. Forest plots of standardized differences in (a) species richness and (b) abundance between forests and native plantations based on 20 independent studies. The vertical line represents no difference. Error bars show means and associated 95% confidence intervals. Points falling to the left of the zero line indicate a stronger effect in forests, whilst the points falling to the right indicate a stronger effect in plantations.

17 Chapter 1: Introduction

Discussion

Managed native timber plantations can support an array of fauna equally as species rich and abundant as native forests, and that this result was consistent across faunal groups. Understorey complexity was not significantly associated with increased species richness in plantations, but strongly moderated faunal abundance. When plantation understories were complex bird and mammal abundances were high and equivalent between forests and plantations. However reptiles and invertebrates were more abundant in plantations than forests. Plantation age did not affect bird richness. Collectively these results provides support for the idea that native plantations can better benefit fauna than exotic plantations (Hartley 2002), and highlight that directed meta-analyses can clarify states of knowledge when research findings from multiple studies are inconsistent. The equivalence of results from this meta- analysis and earlier meta-analyses that specifically compare native forests with native plantations (Stephens & Wagner 2007), suggest that native plantation monocultures with simple understories situated in close proximity to forests, can play a role in conserving faunal diversity in forests at a local scales.

Native plantations support diverse faunal assemblages

This meta-analysis identified a consistent trend in the literature for managed native plantations to support a forest-like richness of birds, mammals, reptiles and invertebrates at the stand-level. This contradicts those of individual research (Hobbs et al. 2002, Boorsboom et al. 2002), which identify less diverse native plantation faunal assemblages compared to those in forests. Brown et al.

18 Chapter 1: Introduction

(2001b) suggest that species diversity can be maintained within narrow limits provided resources are constantly available and systems are open to migration, even though species composition may vary greatly. Further, Pardini et al. (2009) identify that low contrast landscape matrices better support high faunal richness and abundance. In this meta-analysis native plantations of endemic tree species necessarily provided low-contrast to adjacent forested environments, and qualifiers for the inclusion of data specified that forests were of good quality and that plantations were within close proximity to forests in the landscape. Hence the consistently high species richness in plantations identified in this meta- analysis may result from an enhanced potential of forests to supply recruits to plantations.

Age has been recognised as an important determinant of faunal diversity in plantations, primarily because it is often equated with successional development in both unmanaged (Bremer & Farley 2010) and managed plantations (Cummings & Reid 2008). In opposition to reviews pooling exotic and native tree species (Nájera & Simonetti 2009) and individual studies

(Boorsboom et al. 2002), this meta-analysis identified that bird richness was insensitive to plantation age, suggesting that young plantations in proximity to native forests either undergo rapid species replacement, or experience highly dynamic use by birds. It is probable that managed young plantations can attract a diverse array of bird species because native tree species provide familiar foraging substrates and food resources (Hartley 2002), and abundant invertebrate populations facilitate incursions into young plantations, despite low

19 Chapter 1: Introduction

structural diversity. It is important to qualify that the data available for this analysis of plantation age included Australian research only. To more accurately quantify the role of young native plantations in supporting fauna generally, a more comprehensive global assessment needs to be undertaken as this data arises. It is probable that both understorey development and resource availability will vary considerably in young plantations depending on the productivity of environment in which they are established, and so influence both bird diversity and overall faunal use of young plantations.

Understorey complexity moderates faunal abundance

Understorey complexity has been consistently identified as a strong predictor of animal diversity, and in both native and non-native plantations.

Previous reviews have associated simply structured understories in native plantations with reduced faunal diversity and abundance (Moore & Allen 1999,

Lindenmayer &Hobbs 2004, Carnus et al. 2006, Nájera & Simonetti 2010,

Brockerhoff et al. 2012). The results from this meta-analysis agree with former reviews. In contrast to previous reviews however, reptiles were as abundant in these plantations as they were in forests, and invertebrates (predominantly beetles) were far more abundant. It is likely that for reptiles and ground invertebrates young native plantations offer increased thermal opportunities and abundant food resources which support high abundance. Whilst there was no data available allowing a meta-analysis of small mammal response to young plantations, individual authors identify lower diversity in young native plantations than in forests (e.g. Boorsboom et al. 2002, Ramírez & Simonetti 2011), and

20 Chapter 1: Introduction

shifts in assemblage composition towards generalist species (e.g. Mitchell et al.

1997), as mammals are strongly dependant on habitat complexity (Holland &

Bennett 2007).

The variation between the responses of animal taxa is driven by the need for different taxa to exploit habitat differently and is common in many cross-taxa comparisons (Lawton et al. 1998, Schulze et al. 2004, Felton et al. 2010). Whilst the results of this meta-analysis highlight the importance of complex understories for supporting high faunal abundance in plantations, they also identify that ground fauna respond to changes in plantation understory condition very differently to more mobile animals like birds. As Felton et al. (2010) suggest, the need for a case-specific assessment may be necessary to both reliably predict the responses of ground fauna to simplifying plantation understories, and to understand the persistence of diverse and abundant faunal assemblages in plantations over time.

Plantations as habitat

Whilst an assessment of the identity of faunal communities using native plantations was not a focus of this meta-analysis, it is important to qualify that equivalent species diversity does not equate to similar species composition.

Whilst this meta-analysis identified diverse faunal communities using plantations, it is likely that these communities are not dominated by forest fauna.

The individual studies used to generate this meta-analysis repeatedly identified plantation faunal assemblages as lacking in forest specialists. Many forest species are particularly sensitive to forest naturalness (e.g. Farwig et al. 2008,

21 Chapter 1: Introduction

Proença et al. 2010), and the persistence of forest species in plantations, whilst often related to understory condition (Aubin et al. 2008), is likely to be driven by a complex array of factors. Further, the way in which fauna are sampled and subsequently grouped strongly influences the outcome of results in meta- analyses (Felton et al. 2010). For example, in this meta-analysis invertebrate samples were heavily biased towards beetles, which respond strongly to successional stages of vegetation (Pawson et al. 2009). Thus high invertebrate abundance in plantations with simple understories reflects that plantations are good habitat for beetles, but not necessarily other invertebrates.

Acknowledgement of assemblage identity is particularly important when assessing the value of native plantations as faunal habitat. Without a focussed assessment of the identity of fauna using native plantations, we can conclude that whilst native plantations in forested environments can support diverse and abundant faunal assemblages, these assemblages may be comprised of generalist, edge specialist and exotic animals (Bengtsson et al. 2000, Davis et al.

2000, Boorsboom et al. 2002) rather than forest specialists.

Stephens and Wagner (2007) argue that the true conservation value of a forest is determined by comparing it to the land use it replaces. In this case the literature examined often failed to identify any pre-plantation land use, and thus this meta-analysis necessarily became a comparison of plantations with forests.

The results of this meta-analysis identify native plantations as a feasible means to augment and support native forest diversity, if not the diversity of forest specialist species. Whilst we acknowledge the small sample size of the overall

22 Chapter 1: Introduction

data set, we identify that the very strong effect of understorey complexity on faunal abundance in plantations identified in this meta-analysis, suggests developing or retaining a complex understorey in plantations should be recognised as a key management concern if maintaining diverse faunal communities is a goal. Further investigation of the role of ecological management (e.g. Fischer et al. 2006, Hsu et al. 2010) may provide guides to developing high faunal diversity in managed native plantations, without compromising timber production.

Continuing work in context

This thesis is an original body of work that increases knowledge of monoculture hardwood plantation habitat use by fauna in Australia. This meta- analysis has identified that despite their growing presence in the global plantation estate very little research is focussed on faunal diversity in managed native tree plantations. Particularly, our knowledge of young plantations as habitat is often context specific and generally poor, particularly for ground fauna.

In this thesis I extend knowledge of young plantations as habitat, and specifically focus on ground fauna. I address the ecological context of faunal assembly in native plantations in comparison to native forests, and assesses whether plantations are physiologically limiting for ectotherms. Specifically I compare faunal diversity between industrial monoculture Eucalyptus plantations

(Blackbutt - Eucalyptus pilularis) and adjacent native forests, and investigate some of the potential reasons for the associations identified between vegetation

23 Chapter 1: Introduction

structure and faunal habitat use by exploring the relationship between shifts in habitat structure and faunal habitat use. I further explore some of the mechanisms that generate the observed relationships using thermal physiology for reptiles, and landscape context and prey availability for ground fauna. My approach to understanding faunal use of modified habitats such as managed forests and plantations incorporates broad sampling across all animal groups including birds, small mammals, herpetofauna and invertebrates, an approach that is still relatively new for plantations generally. This is also the first Australian study that has comprehensively compared the thermal environments available to wildlife in plantations and native forests. I aim specifically to:

1. identify differences in vegetation structure, and thermal environment (light,

air temperature) between plantations and native forests and within

plantations of different ages (Chapter 2)

2. examine how these environmental differences alter patterns of habitat

use by multiple taxa of vertebrate fauna (Chapter 3)

3. determine whether there are differences in food resource availability for

insectivorous small mammals and herpetofauna between plantations and

forests (Chapter 4)

4. determine whether the landscape context of plantations influences habitat

use by small mammals and herpetofauna (Chapter 5)

5. experimentally examine how the dependence on specific structural or

thermal attributes in an environment influence habitat use by one faunal

24 Chapter 1: Introduction

group (scincid lizards), and to determine the physiological basis of habitat

preference in this group (Chapter 6)

This thesis is presented as a series of chapters that address the aims above.

Due to use of the same sites and capture methodologies for more than one aim and the need to fully explain methodologies in each paper, the introductions and methods in some chapters may be extensive.

25 Chapter 2: Plantation microclimate diversity

Chapter 2 Thermal and structural characteristics in native plantations: implications for microclimate diversity

Introduction

Available microclimates are a valuable, yet often poorly quantified resource for biotic communities. They control plant distributions

(Stoutjesdijk& Barkman 1992), and are a recognised habitat selection cue for ectothermic (invertebrates – Retana & Cerdà 2000; Pereboom & Beismeijer,

2003; Ouedraogo, 2004; amphibians - Halverson et al., 2003; fish - Wherly et al., 2003; reptiles – Huey & Slatkin, 1976; Hertz et al., 1994; Webb & Shine,

1998; Sartorius et al., 1999), and endothermic animals (small mammals-

Lagos et al. 1995, Pienke & Brown 2003, Sedgeley 2001, Chruszcz &

Barclay, 2002; birds - Wolf & Walsberg, 1996). Microclimates are produced by the interaction between vegetation structure, heat and light. Canopy openings increase the diversity of microclimates experienced by organisms at the forest floor (Van Calster et al. 2008) by allowing greater penetration of solar radiation (Hanley 2005, Weng et al. 2007), so increasing light, air and surface temperatures, (Clinton 2003, Martius et al. 2004, Rambo & North

2008) and modifying soil temperatures and humidity (Kanowski et al., 1992;

Palik & Engstrom, 1999). Secondarily both leaf shape dictated by tree species (Yirdaw & Luukkanen 2004, Strobl et al. 2011) and sub-canopy vegetation complexity influence light attenuation and heat penetration to the forest floor (MessierPorté et al. 2004), and so the distribution of microclimates available for organisms to exploit.

26 Chapter 2: Plantation microclimate diversity

Industrial tree plantations are a forest type exposed to high levels of management-induced canopy and understorey disturbance. In native eucalypt plantations management practices including slashing, grazing and thinning produce simpler and more open tree canopies and sub-canopies than forests and typically a less structurally complex understorey (Kanowski et al. 2003, Brockerhoff et al. 2008). As a consequence plantation understories are often exposed to higher levels of light, solar radiation

(Florence 2004) and air mixing (MacDonald & Thompson 2003, Laurance

2004, Wright et al. 2010) than forests. In exotic plantations these management practises result in increased litter, air and soil temperatures and decreased moisture availability (Waters et al. 1994, Lemenih et al. 2004,

Wright et al. 2010, Yuan et al. 2013). Whilst native tree plantations are likely to experience similar physical effects associated with canopy opening, microclimates at the forest floor may more closely approximate those in forests, as understorey buffers microclimates (Szarzynski & Anhuf 2001), and native plantations better regenerate simplified native understories than exotic plantations over time (Keenan et al. 1997).

Whilst sub-canopy microclimate has been examined many times in exotic plantations, few studies have directly quantified the relationship between indigenous vegetation, plantation structure and ground-level microclimates in managed native plantations (Porté et al. 2004), or examined how these change with plantation age. Such knowledge is important for foresters and conservationists, as microclimate directly influences tree health through its effects on soil and (Davidson et al. 2007), native tree plantations are promoted as low-contrast buffers that reduce edge effects in forests (Denyer

27 Chapter 2: Plantation microclimate diversity

et al. 2006, do Nascimento et al. 2011; Prieto-Benez & Mendez 2011), and manipulating plantation microclimates by changing vegetation structure is a management technique currently being investigated as a means to directly increase the conservation value of managed native plantations for plants and animals (Carey 2003, Cummings & Reid 2008, Yuan et al. 2005, 2013). This study aimed to compare the similarities between available microclimates in young and old native plantation monocultures and adjacent managed logged and old growth forests in the peak growing period of Spring-Summer, to assess available thermal environment as a habitat selection cue.

Methods Study area

Vegetation structure and associated thermal environments were investigated in the Camden Haven and Macleay-Hastings river catchments, on the mid-north coast of New South Wales, Australia, from October 2001 to

December 2003. The land in the area is a series of coastal mountain ranges with maximum elevations of approximately 400 m in the south to over 1000 m in the north. Mean annual rainfall is 1548mm at Wauchope in the approximate centre of the study area (Fig 1). Temperatures ranged between

5.3° - 25.8°C between July and January. Native forests in the area vary from sub-tropical rainforests to dry open hardwood forests, but dry eucalypt forests predominate (Harden, 1991).

The study area contained hardwood plantations aged between 6 and

32 years old. The oldest plantations within the study area were monoculture plantings of the endemic, dominant canopy tree species, Blackbutt

28 Chapter 2: Plantation microclimate diversity

(Eucalyptus pilularis), whilst young plantations were simple polycultures of three species; (Eucalyptus pilularis, E. cloeziana, E. grandis). Twenty 200 x

300m field sites were spread over a 55 x 80km area of woodland, (max.

55.2km west of the coast; 31017’S 152035'E - 31042’S 152031'E), and were matched in time since fire and logging, altitude, aspect and dominant canopy tree species. Sites were selected to provide a replicated non-orthogonal statistical design that standardized plantation age, planting density and distance to native forest as much as possible within the constraints of the existing plantations. Whilst native sites were adequately spread throughout the study area, young and old plantation sites were necessarily clumped in the Northern expanse of the study area due to the need to standardise plantation age within a limited number of closely distributed plantations in the landscape. Pseudoreplication was ameliorated by sampling at least two separate plantations and by positioning all sites as far apart as possible within an individual plantation. All plantation sites (except for spar plantations) were separated by ≥1km of plantation matrix and were established in separate stands of plantings. All sampling within sites occurred at ≥50m from any edge, including snig tracks, dirt roads or plantation stand boundaries.

Forest types

Five forest types associated with timber production were identified in the study area. These were old growth native forest, logged native forest, old plantation, young plantation and early maturity (spar) plantation.

Young plantations (Fig. 2a) were situated in two 40 hectare plantations established on land historically cleared as pasture, then afforested with plantation in 1996-97. Establishment procedures included deep ripping, weed

29 Chapter 2: Plantation microclimate diversity

control and fertilization, and tree density was ~1000/ha. Since establishment, no fires or thinning had occurred.

Young afforested plantations

Young plantations were comprised of three tree species E. pilularis (80%), E. grandis (5%) and E. cloeziana (10%) planted in monoculture blocks.

Sampling in young plantations was restricted to stands of E. pilularis only.

The canopy was open with an understorey of dense stands of Imperata cylindrica and patches of bracken (Pteridium esculentum). 40% of the plantation margin showed broad connection to disturbed (grazing, fire) native forest. All the sites were exposed to low intensity cattle grazing.

Spar plantations Early maturity (also termed ‘spar’ by Florence, 1996) plantations (Fig.

2b) were small plantations of three species polycultures dominated by

Eucalyptus pilularis in similar percentages as outlined for early young plantations. All sites in young and spar plantations were restricted to E. pilularis stands only. The five replicate sites within spar plantations were necessarily small (100 x 100m) due to the poor representation of plantations of this age in the Mid North Coast plantation estate at the time of sampling.

Sampling in these plantations was restricted to microclimate only. The two 20 ha plantations in this treatment were planted in 1991 and underwent establishment procedures similar to those in both young and old plantation treatments. Sites in this forest type were exposed to low intensity cattle grazing, and had not been thinned since establishment. Spar plantations were unmanaged after the first five years of growth and were undisturbed by

30 Chapter 2: Plantation microclimate diversity

Fig. 1. A map of the 55 x 80km study area on the mid-north coast of New South Wales. Sites are denoted by circles. YP = young plantations, OP = old plantations, LN = logged native forests, ON = old growth native forests, SP = spar plantations. The rectangular inset on the map (sourced from Gavran 2013) at the top left identifies the North Coast bioregion in which the study was conducted.

31 Chapter 2: Plantation microclimate diversity

(a)

(b)

(c)

(d)

(e)

Fig. 2. Diagrammatic representation of the structural complexity of vegetation in (a) young plantation, (b) spar plantation, (c) old plantations, (d) logged native forest, and (e) old growth native forest.

fire, creating a forest type with a dense closed canopy. All plantation treatments were matched in initial planting density.

Old reforested plantation

Sites were established in two 80 hectare monospecific plantations of E. pilularis on land initial forested, then cleared and replanted in 1974 at a

32 Chapter 2: Plantation microclimate diversity

density of 150-200/hectare. In the first five years these plantations underwent weed control and fertilization associated with accelerating tree growth.

Remnant shrubby vegetation was retained within the plantation along dry gullies as two to five metre wide strips. Seed trees were not retained in these riparian buffers, but shrubby vegetation was often well developed. Old plantation sites did not support sub-canopy or mid-storey layers outside riparian strips. Typically there was a high (+30m) canopy layer, and an understorey ranging from very dense swathes of Blady grass (Imperata cylindrica) to bare soil patches (Fig. 2c). Small, sparse patches of Acacia sp. regrowth typically did not exceed 3m in height. The plantation was last thinned in 1994 resulting in a very open canopy. Low intensity wildfire was last recorded in the plantations in 1996. Seventy percent of the plantation margin was broadly connected to managed native forest used for low intensity cattle grazing, and thirty percent abutted logged native forest.

Logged native forest

Logged native forests represented the dominant forest in the landscape. Sites had been subjected to recent (1996) and historic selective logging. Tree species mixtures were dominated by E. pilularis. This forest had been thinned by both small coup removal and selective logging, resulting in a patchy, open forest canopy (Fig. 2d). Areas within this forest type where thinning was less intense were characterised by large numbers of Casuarina sp. The mid-storey was sparse where present, but the understorey, particularly in old logging coups and along snig tracks had dense sedge

Lomandra longifolia, and Imperata cylindrica grass. As for old plantations,

33 Chapter 2: Plantation microclimate diversity

thinning and fire were excluded after 1996. Sites were wholly within extensive forested areas with a similar management history.

Old growth native forest

Native forests throughout the region had a long history of selective logging which spanned the last 100 years (Pitt, 2001). Commercial logging throughout this region commenced in the late 1800s and increased in intensity throughout and following the 1940s. The old growth native forest sampled in this study had not been logged for a minimum of 40 years, and represented the oldest closed-canopy forest within the study area. The forest contained a mixture of sub-tropical and wet coastal woodland and rainforest plants. Sites within this forest type were surrounded by logged native forest.

The trees in order of dominance included Eucalyptus pilularis, E. microcorys, and Syncarpia glomulifera. There were well developed sub-canopy and midstorey layers (Fig. 2e) and the forest floor was typically bare with patches of fallen, mossy logs, and mixed ferns and sedges (Lomandra longifolia, Gahnia sp.).

Vegetation structure

Vegetation structure was quantified at five sites in young and old plantations and logged and old growth forests. At each site vegetation characteristics were measured from sixteen 15m2 quadrats along four haphazardly placed transects spaced 50m apart. Quadrats were spaced 70m apart along transects. Four quadrats from four transects in each of five sites resulted in a total of 80 quadrats per forest type. All sampling was conducted between

November 2001 and April 2002. The vegetation characteristics measured are shown in

Table 1.

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Thermal environment

The amount of global solar radiation available at ground level was assessed using a pyranometer (model LP 02L, Campbell Scientific) that measured a combination of ultraviolet, near-infrared and incident wavelengths of light.

The amount of insolation at the ground level was recorded from five sites in each of the five forest types from three 50m transects spaced 30m apart in an area at the centre of each site. Maximum insolation was sampled by always choosing the brightest sun patch within a three metre radius of each transect point, between 11:00am and 2:00pm. Samples were collected from five points at ten metre intervals along each transect, three times during four sequential, uniformly sunny days. A total of 45 measurements per site were recorded from September 2001– February 2002. Brief snapshots of light environments such as this have previously been shown to accurately predict light environments within a season (Dignan & Bren 2003). The pyranometer was positioned 20cm above the ground, and was allowed to equilibrate for one minute before each measurement was taken. The pyranometer was calibrated against a laboratory standard LI-COR quantum sensor and data were adjusted accordingly before analysis.

Environmental temperatures measured as a combination of insolation and air temperature were recorded from the hottest (full sun daily) and coolest (full shade daily) microenvironments in all forest types, to determine whether there were any differences in the daily range of available temperature, and any seasonal shifts in this range.

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Table 1. Vegetation parameters and the methods used to record them from four forest types on the Mid North Coast of NSW.

Variable measured Definition Measurement Method Whole environment characters number of trees Trees are defined as any vegetation taller than 5m high count tree DBH tree girth at approx 1.4m above the ground length tape measure midstorey height height to the midstorey canopy below the sub-canopy layer height range finder number of saplings count vertical complexity an estimate of how much vertical space betw een the ground and the canopy categorical follow ing McDonald et al. 1990 is occupied by foliage foliage density between 0-1m above grnd. Foliage refers to any plant part and does not differentiate percentage visual estimate foliage density between 1-2m above grnd. betw een species or grow th forms percentage visual estimate foliage density between >2m above grnd. percentage visual estimate

Canopy level characters crown height height to the canopy top but excluding height range finder emergent trees canopy cover amount of horizontal space covered by canopy foliage percentage standardised photographs see McDonald et al . 1990 Ground level characters understorey height height of any vegetation up to 1.5m high height ruler shrub cover shruby grow th forms only percentage visual estimate grass cover grasses only percentage visual estimate litter cover percentage visual estimate litter depth depth of leaves from the soil to the top of the leaf pack depth ruler (cms) twig density how many tw igs < 1cm thick cover a 10 x10cm square percentage visual estimate rock cover percentage visual estimate bare ground percentage visual estimate # fallen logs Logs w ere defined as any dead w ood on the ground surface w ith count diameter >5cm and length >15cm log cover the total percentage of a quadrat covered by logs percentage visual estimate log decomposition a count of any holes/crevices in a log of >5cm long and 2cm w ide count modified from Fletcher (1977) number of stumps included to quantify the availability of vertical perches/basking points count soil moisture* Laboratory study involving drying soil samples. weight change gravimetric determination Soil dried for 48hrs and change in w eight is equated w ith soil moisture content.

*soil samples w ere collected from 1 random point in each quadrat using a 5-cm diameter soil corer inserted to a depth of 10 cm after removal of the litter and humus layer.

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Thermocron iButtons (model DS1921L, Dallas Semiconductors) are microchip- driven data loggers sensitive to 0.250C fluctuations in temperature. IButton loggers have stainless steel casing, and in this study were plastic wrapped and painted with neutral-coloured paint (following Walsberg & Weathers, 1986) to prevent heat reflection influencing recorded temperatures.

Four loggers were placed haphazardly within each microhabitat in each site, such that no sampled microhabitat was less than 30m from any edge, no logger was less than 50m from another and loggers remained wholly within the microhabitat they were sampling. Loggers recorded temperature one centimetre above the leaf litter surface every hour, for a period of three months throughout the period of the study in which vegetation structure was measured (September

2001– February 2002) resulting in twenty samples per forest type. Loggers were moved within microenvironments once per month to better sample the variability associated with each microhabitat. To sample any temperature differences associated with plantation aging four loggers were placed in sun and shade microhabitat in each of four sites in spar and young plantations, resulting in 16 samples per microhabitat. Methods for logger placement followed those described previously. Temperature was sampled from October 2002 to

November 2003.

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Data Analysis

Vegetation structure

Changes in vegetation structure associated with management history in young and old plantations, logged and unlogged native forests were explored using nonparametric multivariate analysis of variance, conducted on Bray-Curtis dissimilarity matrices calculated on standardized data using the PRIMER analysis package (Clarke 1993). Prior to analyses, multicollinearity between vegetation variables was assessed, and the model run using only the least correlated (≤r = 0.3) variables. Data was averaged over quadrats to create site- level means before analysis. Structural differences between young and old plantations and forests were then tested iteratively by analysis of similarity

(ANOSIM). Vegetation variables driving the differences between forest types were identified using similarity analysis (PRIMER - Clarke 1993). These data were represented graphically in a principal co-ordinates biplot, using the Bray-

Curtis dissimilarity semi-metric as the distance measure as it deals well with zero values, is less sensitive to skew and non-normality than the Euclidean distance metric, and it implicitly standardizes data measured on different unit scales (McArdle 1999).

Thermal Environment

Seasonal shifts and circadian differences in mean temperature were analysed using fixed effects (forest type, month) two-way repeated measures analysis of variance (RMANOVA) with contrasts specified to test for differences

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between young and old plantations, and logged and old growth native forests.

Data was averaged within sites then pooled into hourly time bins, generating nine temperature recordings in the day (06:00- 18:00) and twelve at night

(18:00- 6:00) in each forest type. Sun and shade microhabitats were analysed separately. Within subjects degrees of freedom were adjusted using a lower bound test where the assumption of sphericity was violated (SPSS, 2003), and so deviated from the expected k-1, N-k format. Oneway RMANOVA with the same design protocols as above was used to identify any changes in temperature between young and spar plantations. All data were log (x+1) transformed before analyses. Circadian profiles were generated to visualise any differences in mean available temperature between young and old plantations and forests, and between young and spar plantations.

Linear regression was used to relate canopy cover, number of saplings, grass area, shrub cover and foliage cover up to one metre above the ground to thermal environment in plantations and forests. Subsequently, one-way analysis of variance (ANOVA) was used to compare variation in mean maximum temperature and insolation between forest types, and explain any variation present. Bonferroni corrections for multiple comparisons were applied. The vegetation variables analysed in regressions were those identified by similarity analyses as characterising forest types, and that were likely to directly influence both temperature and solar radiation penetration to the ground. Temperature data were restricted to daytime measurements from sun microhabitats between

December and January, when temperature differences between forest types

39 Chapter 2: Plantation microclimate diversity

were maximal to avoid confounding seasonal with treatment effects. For all analyses temperature and solar radiation data were pooled over sites then transformed by square root or log (x+1) as necessary to satisfy test assumptions before analysis. Relationships between thermal parameters and vegetation structure were illustrated using partial regression and scatter plots.

Results Vegetation structure

Vegetation structure differed significantly between forest types and management histories within forest type (global R= 0.955, p=0.01). Structural similarity was highest between old growth and logged native forests (average dissimilarity [a.d] = 13.9%), whilst young and old plantations were less similar

(a.d. = 30.4%). The majority of the variance in the data was explained by differences between plantation and forest vegetation structure (Fig. 3, Dim 1 =

69.63%). Old growth native forests had the highest vertical complexity of all forest types above two metres. Logged native forests were characterised by high foliage density between 1-2m and dead wood on the ground (Fig. 3). Young plantations lacked sub-canopy complexity and had a grassy ground layer interspersed with rocky and bare patches (Fig 3). Mean canopy cover was the same in plantations (20%) and increased (50%) in logged native forests. In particular the lack or development of canopy cover, foliage cover 0-1m above the ground, number of saplings and number of trees was important in defining individual forest types.

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Environmental temperature

In addition to circadian fluctuations in temperature there was a global increase in temperature of ~100C degrees in all microhabitats between

September and February. During the day mean temperatures in sun (F3,16=

336.710 p<0.001) and shade (F3,16= 20.458 p<0.001) microhabitats differed between forest types. Plantations were thermally distinct from forests between

10am – 5pm in the sun microhabitat, and logged forests were intermediate between plantations and old growth forests. Young plantations were consistently ~100C hotter than logged forests, ~200C hotter than old growth forests, and ~ 50C hotter than old plantations in the hottest three hours of the day from November - January (Fig. 4 a - c). At night old growth forests remained between 2 - 50C cooler than any other forest type in shade microhabitats (F3,16= 4.795 p=0.014), but temperatures were equivalent in sun microhabitats in all forest types (F3,16= 0.431 p= 0.734). Young plantations were exposed to the greatest temperature range throughout the period of measurement, as daytime mean maximums moved from ~30 - 480C between

September and January. In comparison old plantations and logged forests experienced temperature increases of ~100C and old growth forests

~80C (Fig, 4 a-c).

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Fig. 3. Principal co-ordinates biplot showing the distribution of vegetation variables in relation to forest type using Bray Curtis semi metric distance separation on transformed vegetation data. Only the top 45% of the explanatory vectors are shown. Symbols represent: filled circles = old growth native forest, open circles = logged native forests, filled squares = old plantations, open squares = young plantations.

Age-related thermal differences in plantations

Spar plantations differed from young plantations by having taller trees and a very dense closed canopy, but an understorey structurally similar to young plantations, with patchy grass cover on the ground and no shrub or sub-canopy layers. Whilst young and spar plantations experienced similar mean environmental temperatures in the morning, young plantations were

0 significantly ~8 C) hotter between 10am and 5pm (F1,30= 6.373 p= 0.017) (Fig.

5a). This temperature difference did not persist in shade microhabitats during the day (F1,30= 2.642 p= 0.115), or at night (sun: F1,30= 0.501 p= 0.484, shade:

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0 F1,30= 0.1.917 p= 0.176) despite young plantations cooling to ~2 C cooler than spar plantations (Fig 5b-d).

Table 2. Results of within-subjects tests for the repeated measures analysis of variance test on environmental temperature data from old growth and logged native forests and old and young plantations. ‘Time’ expresses changes in temperature over the course of a day. Time of day F df p 6AM - 6PM Sun time*forest type 9.502 3,16 0.001 month 483.418 1,16 <0.001 month*forest type 3.173 3,16 0.053 month *time 10.597 1,16 0.005 Shade time*forest type 5.191 3,16 0.011 month 202.245 1,16 <0.001 month*forest type 2.693 3,16 0.081 month *time 2.396 1,16 0.141 6PM - 6AM Sun time *forest type 2.247 3,16 0.734 month 222.363 1,16 <0.001 month*forest type 2.221 3,16 0.125 month *time 1.883 1,16 0.189 Shade time *forest type 12.917 3,16 <0.001 month 301.828 1,16 <0.001 month*forest type 1.743 3,16 0.199 month *time 2.364 1,16 0.144

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Fig. 4 Circadian profiles for mean environmental temperatures available in the sun and shade throughout the course of the day from 6am to 6pm, between September and January in forests and plantations. Data is pooled across sites within microhabitats and averaged within time categories. Graphs (a) – (c) depict available temperature in sun microhabitats, whilst graphs (d) – (f) depict those in shade microhabitats. Error bars represent +/- 1 SE. Symbols represent the four forest types: filled circles = old growth native forest, open circles = logged native forests, filled squares = old plantations, open squares = young plantations.

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Fig. 5. Circadian profiles for mean environmental temperatures available in spar and young plantations in the sun and shade throughout the course of the day. Graph (a) = sun microhabitat, 6am to 6pm, (b) = shade microhabitat, 6am to 6pm (c) = sun microhabitat, 6pm-6am, (d) = shade microhabitat, 6pm-6am. Error bars represent ± 1SE. Open squares = young plantation, filled triangles = spar plantations

Table 3. Results for analyses relating vegetation variables with maximum solar radiation levels and maximum available environmental temperatures occurring in young and old plantations, logged and old growth native forests between 6am – 6pm. Significant values are in bold. Notations are: OP = old plantation, YP = young plantation, LN = logged native forest, ON = old growth native forests.

AN O VA insolation (Wm-2) temperature (0C) F df p F df p overall 74.945 3,16 <0.001 42.384 3,16 <0.001 pairwise OP v YP 0.003 0.95 OP v LN <0.001 <0.001 OP v ON <0.001 <0.001

2 2 Regressions r r model 0.837 14.375 5,14 <0.001 0.861 17.308 5,14 <0.001 canopy cover <0.001 0.033 grass area 0.495 0.792 sapling density 0.524 0.991 shrub cover 0.249 0.049 foliage cover 0-1m 0.284 0.451

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Relating vegetation structure and thermal environment Overall ground-level microclimates received significantly less solar radiation and were cooler as canopy cover increased from young plantations to

2 old growth native forests (solar radiation:F5,14 = 14.375, p<0.00, r = 0.837;

2 temperature: F5,14= 17.308 p<0.001, r = 0.861, Fig 6a-b). Insolation levels differed between each forest type, with old plantations receiving the highest level of insolation (Fig. 6 c-d). However, whilst available temperatures also differed between plantations and forests (Fig. 6 a-b, Table 3) they were similar between young and old plantations, suggesting non-linear correlation between temperature and insolation. Canopy cover was the only vegetation parameter tested that significantly affected both insolation levels and available temperature at the ground, and shrub cover significantly affected available temperatures at the ground during the day.

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Fig. 6. (a) and (b) are scatter plots depicting the distribution of canopy cover with forest type. (c) and (d) are partial regression plots showing the strength of the linear relationship between the transformed radiant energy and environmental temperature variables and canopy cover, with the other variables associated with the multiple regression accounted for. Unbroken lines represent the linear line of best fit. Broken lines represent the 95% confidence intervals around the line of best fit.

Discussion This study demonstrated that historical management can have a strong and ongoing influence on vegetation structure in native timber plantations and forests, and that in plantations canopy opening associated with management can dictate the thermal environment available at the forest floor. The structural uniqueness of plantations and forests in this study was generated by differences

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in canopy cover, sapling density, shrub and grass cover. Unsurprisingly, the greatest degree of structural difference lay between native forests and plantations, and strong fidelity within plantations and forests as groups was driven by specific structural variables: grass in plantations and vertical complexity forests. Opening of the tree canopy significantly increased daytime temperatures, and plantations experienced lighter, hotter and less stable microhabitats at the ground than native forests during the day. While insolation was controlled by canopy cover alone, shrub density moderated available temperatures at the ground.

Vegetation structure in old plantations and logged native forests

Previous land use can strongly affect vegetation communities (e.g. Ito et al.

2004, Gachet et al. 2007). Whilst young timber plantations necessarily present a very different structural environment to forests, in this study plantations of 30 years old supported much less structural complexity in the understorey and were so distinct from logged native forests that had experienced equivalent time since fire and harvesting, equivalent grazing exclusion and equivalent access to the original forest seedbed and to seed dispersal from adjacent forests. The lack of understorey regeneration in old plantations despite these similarities suggests that a combination of abiotic and biotic factors was likely to have interacted to reduce vegetation complexity. Fire was unlikely to be the cause of the lack of complexity in old plantation understories as low intensity fires mobilize soil nutrients and so stimulate plant growth (Gill et al. 1981). In this study management associated with pole harvesting or thinning had been excluded for

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a similar time prior to study commencement. However, lower canopy cover in old plantations (20% versus 50% in logged native forests) suggests that the old plantations had experienced harvesting more frequently than logged native forests, and it is likely that relictual effects of soil compaction associated with harvesting may have affected seedling germination and growth (Kozlowskia

1999). Further, the high density of Blady grass (Imperata cylindrica) which is often associated with degraded soils and high temperature and light environments, and is known to be allelopathic (Coile & Shilling 1993), may have inhibited native regeneration through competition.

Environmental temperature

Temperature is a dynamic property of forests (Saunders et al. 1998), and this study highlights that temperature is dynamic at both macro and micro scales.

Environmental temperatures differed significantly between all the forest types assessed in this study in both sun and shade microhabitats during the middle of the day, but were cooler in shaded microhabitats in forests at night. Higher daytime temperatures in plantations than forests have been identified by Denyer et al. (2006) at plantation edges, and by Malcom (1998) and Weng et al. (2007) in thinned exotic plantations. However Porté et al. (2004) failed to find this difference between old plantations and those thinned 10 years prior to his study, and explain this as an effect of temperature buffering by the forest. Such buffering is also likely to drive temperature differences at night.

As plantations heated and cooled rapidly throughout the day, there was an expectation that they would cool rapidly during the night, and potentially

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reach a lower minimum temperature than forests as heat was lost through the open canopy (e.g. Clinton et al. 2003). However in this study, as for that of those of Denyer et al. (2006) and Baas and Mennen (1996), night temperatures were equal in forests and plantations in open microhabitats, but were cooler in forests in shaded microhabitats. Manipulative studies have identified that sub-canopy structural complexity buffers air temperatures and may prevent the temperature fluctuations associated with large canopy openings (Carlson & Groot, 1997).

Thus whilst shaded microhabitats in the old growth forests received the least insolation and were coolest, vegetative buffering meant that at night they were retained the low temperatures achieved during the day, rather than equalizing temperatures as was occurring in the more open sun microhabitats, and on a larger scale in the very open plantations. Baas and Mennen (1996) highlight that temperature differences are often seasonal, and that in summer night may be too short to allow temperatures to drop from high daytime levels.

Young native plantations possessed distinct thermal environments in the study area, and so had high potential to influence to potential colonizers.

Microclimates associated with young plantations were likely to exclude those forest organisms that lacked a tolerance for large fluctuations in temperature and irradiance. Forest understorey plants which represent the readiest source of propagules for young native plantations abutting native forests were not exposed high daily temperature fluctuations. For many plants high light high temperature environments may limit germination, growth and survivorship (e.g.

MacDonald & Thompson 2003, Dang & Cheng 2004). Thus plants colonising

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young plantations may be limited to those species that can withstand relatively large diel and seasonal temperature shifts.

Thermal environment and vegetation structure

Gradients defined by solar radiation and environmental temperature only loosely followed the structural complexity of vegetation in this study. Although there was a general decrease in insolation and temperature as structural complexity increased, levels of insolation in young and old plantations were similar, as were temperatures in old plantations and logged native forests, despite significant structural differences between these three forest types.

Saunders et al. (1998) and Pritchard and Comeau (2004) identify that temperature structure is often distinct from vegetation structure in managed forests. In this study there was a complex, non-linear relationship between vegetation structure and thermal environment, which was probably related to a combination of vegetation removing specific elements of solar radiation by shading (Holl, 1999; Kotzen, 2003) and wind. Various studies recognize that wind and understorey vegetation influence forest temperatures (Morecroft et al.,

1988; Palik and Engstrom, 1999; Rambo & North, 2008). Understorey complexity can reduce wind flow in forests and allow air mixing (e.g. Chen et al.,

1995; Chen & Franklin, 1997; Novak et al., 2000) which results in higher and more stable air temperatures. In this study such mixing may have stabilized temperatures in old growth forests, while the lack of mixing explains the labile nature of plantation temperatures.

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The similar circadian profiles of environmental temperatures in logged native and old plantations are likely to represent a convergence of thermal conditions generated by different means, rather than a similarity of the mechanisms producing them. Old plantations received very high irradiance at the ground resulting in high environmental temperatures. In logged native forests high canopy cover decreases irradiance at the ground level, but complex understorey increases air mixing, which may result in buffered air temperatures approximating those in old plantations. Results support this idea: it is only during summer that environmental temperatures in plantations and logged native forests converge, when all the forest types are exposed to seasonally higher irradiance. Similar insolation in young and old plantations is more linearly related to canopy cover (e.g. Yirdaw &Luukkanen, 2004). Plantations supported simple understorey and had equally open canopies, which resulted in equivalent amounts of light attenuation in both forest types.

Plantation aging and effects on environmental temperature

Spar (12 years old) plantations had increased canopy cover and were colder than young (6 years old) plantations. This result documents the typical thermal sequence observed in plantations, of cooling at canopy closure and associated low understorey diversity (Florence, 1996). Spar plantations represent a short-lived stage in the plantation cycle before the canopy is opened by subsequent harvesting, but they may have large ramifications for fauna. For those forest fauna that prefer cool thermal environments but are tolerant of simple structure (e.g. Mott et al., 2010) spar plantations could represent a

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valuable habitat resource. From a management perspective if this stage of plantation is promoted within the landscape mosaic at forest-plantation boundaries, it may provide forest species with accessible habitat, as well as buffering forest from the microclimatic extremes experienced in young and old plantations.

Management recommendations and research

Importantly this study indicates that monoculture and simple polyculture hardwood plantations support a warmer and more varied microclimate than do native forests, even native forests that have been heavily disturbed. For animals, the labile nature of the environmental temperatures in logged native forests and plantations are likely to have ramifications for those species that use them, particularly species of low mobility. The large magnitude of differences in both structure and thermal environment in plantations are likely to impact strongly on fauna and plants as habitat selection cues, and may exclude species from plantations seasonally. Further, because these differences persist in plantations as they age and are exacerbated by current management practices, plantations may not represent faunally diverse habitats throughout the length of a rotation.

In order to increase the conservation value of plantations, managers will need to support the microclimatic needs of specific groups of organisms, and modify management practices to provide both structurally and thermally attractive habitat features for these organisms. Focusing further research on seasonal shifts in microclimate would allow managers to better understand thermal regime in plantations, and so target management strategies to increase diversity. For

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example, Kluber et al. (2009) identify that different log sizes offer a variety of thermal refuges that allow seasonal persistence of salamanders in thinned forests. If these measures were adapted and adopted in Australian plantation forests they may result in significant biodiversity gains. A focused analysis of the biota using spar plantations needs to be conducted and should be prioritized, as much of the Australian hardwood plantation estate is now entering this growth stage. If adopting conservation and management practices that promote and maintain functionality is important to forest managers, thermal environments in native plantations need to be specifically recognised as having a strong potential to influence biodiversity.

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Chapter 3 Faunal response to young and old eucalypt plantations: do consistent patterns exist across animal taxa?

Introduction

Investigating patterns of response to habitat change that are consistent across different faunal groups is a unifying approach to fauna conservation in disturbed landscapes. Vegetation structure in forests heavily influences physical and chemical environments (e.g. Leite et al. 2010, Zhao et al. 2011). If forest structure changes with land use and management, faunal communities associated with specific forest environments may also change. Often this change is a decline in cross-taxa species richness with increasing habitat simplification (e.g. Dunn 2004, Schulze et al. 2004, Scott et al. 2006).

Native monoculture tree plantations are a highly simplified and rapidly proliferating (FAO 2010) environment which may more closely approximate native forest than exotic plantations (Brockerhoff et al. 2012), and so may better support fauna (Sala et al. 2000, Lindenmayer & Hobbs 2007). There is a growing body of research that suggests that native plantations better meet the ecological needs of native species than do exotic plantations (Potts et al. 2001,

Becker et al. 2007). As such native plantations represent a topical model system in which to investigate faunal response to forest simplification across animal taxa.

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Studies that compare animal species richness between simplified native plantation environments and forests often provide conflicting results. Whilst bird, plant and invertebrate species richness have been reported as declining in native plantations (Hobbs et al. 2003), equivalent richness generated by compensatory responses of fauna (e.g. Frederickson & Frederickson 2002,

MacDonald et al. 2002, Woinarski & Ash 2002, Barbaro et al. 2005, Fonseca et al. 2009) are also reported. Further, while forest specialists are often lost from native plantations (e.g. Moore & Allen 1999, Kanowski et al. 2005), in the presence of high amounts of mature forest in the landscape and matrices of low contrast, large numbers of forest species can persist (e.g. Pardini et al. 2009,

Volpato et al. 2010). Whilst there is no clear consensus that reduced structural complexity in plantations decreases species richness across multiple animal taxa, it is generally accepted that planted forests are less valuable for biodiversity than natural forests (Brockerhoff et al. 2012). Further, whilst young plantations are thought to be less likely to support fauna due to the immediate effects of plantation generation, there is some research suggesting that young plantations may support faunal diversity (e.g. Sullivan et al. 2009). This research addresses the role of age in influencing faunal use of managed plantation habitats. It aims to identify whether there are any consistent patterns of faunal response to young afforested and old reforested native tree monocultures across animal taxa by:

1) comparing species richness and assemblage structure of fauna using native eucalypt forests of differing management age, with those using plantations of

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endemic tree species of different age but equivalent time since management, and

2) examining the difference in ecological function of plantation assemblages by comparing the distribution of feeding guilds across mammal, bird and herpetofaunal taxa, between plantations and native forests.

Materials and Methods

Faunal communities were surveyed in parts of the Camden Haven and

Macleay-Hastings river catchments in the North Coast Bioregion of New South

Wales, Australia, from September 2001 to March 2002 (Fig.1). Mean annual rainfall in the area was 1548mm at Wauchope in the approximate centre of the study area over the period of sampling. Temperatures achieved a mean minimum of 5.3° C in July and mean maximum of 25.8° C in January. Native forests in the area varied from sub-tropical rainforests to dry open hardwood forests, but dry eucalypt forests were dominant (Harden 1991). The study area contained hardwood plantations aged between 6 and 32 years old. The 32 year old plantations in the study area were monoculture plantings of the endemic, dominant canopy tree species, Blackbutt (Eucalyptus pilularis), whilst 6 year old plantations were simple polycultures of three species; (Eucalyptus pilularis, E. cloeziana, E. grandis). Habitat descriptions for each of these forests types are presented in chapter two. The landscape mosaic surrounding plantations consisted of cleared, grazed pasture, grazed E. pilularis -dominated woodlands,

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tall closed woodlands, and closed-canopy, sub-tropical rainforest. All sites were established on Triassic tuff soils in the study area. E. pilularis was the dominant canopy-level or emergent tree species at all sites. As described in chapter two, twenty 200 x 300m field sites were spread over a 55 x 80km area of woodland,

(max. 55.2km west of the coast; 31017’S 152035'E - 31042’S 152031'E), and were matched in time since fire and logging, altitude, aspect and dominant canopy tree species. Sites were selected to provide a replicated non-orthogonal statistical design that standardized plantation age, planting density and distance to native forest as much as possible within the constraints of the existing plantations.

Habitat Descriptions

Four forest types with differing disturbance histories were identified: old growth native forest, logged native forest, old plantation established on cleared and subsequently reforested eucalypt woodland and young plantation established on reclaimed pasture. Chapter two provides details of each forest treatment. Analysis of vegetation structure (chapter two) identified that each forest type was structurally distinct, with native forests having dense tree canopies and sub-canopy complexity, while young and old plantations lacked sub-canopy complexity and had grassy ground layers. Table 1 outlines some of the defining structural characteristics of each forest type.

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Table 1. Means (± 1se) of vegetation variables sampled from 16 x15m2 quadrats in twenty sites in old growth native forest, logged native forest, old plantation and young plantation forest types, in NSW Australia between 2001 and 2002.

Time since tree¹ crown foliage² grass³ leaf³ canopy4 Forest Age management (y) density height (m) cover cover litter cover Old Growth native +100 +50 195 ± 8.5 39 ± 4.7 40 ± 4.6 2 ± 1.7 96 ± 5.4 1175 ± 27 Logged native +70 6 166 ±10.4 27 ± 1.1 63 ± 5.4 30 ± 5.2 58 ± 3.0 816 ± 63 Old plantation 32 6 51 ± 7.6 28 ± 0.6 83 ± 5.0 75 ± 5.5 24 ± 5.4 277 ± 27 Young plantation 6 6 216 ± 4.4 10 ± 0 95 ± 0.7 86 ± 1.2 6 ± 1.2 283 ± 21

¹ number of trees per 15m2 ² % cover of foliage up to 1m above the ground, excluding foliage associated with saplings and trees ³ % mean cover of grass within each quadrat 4 % of canopy covered by tree crown identified using standardised photographs (McDonald et al. 1990) All vegetation measuresments are presented ± 1 se.

Sampling methodologies

Five 200 x 400m sites (as described in chapter 2) were established within each forest type at least 50m from any roads, and in plantations 50m from dry gullies with retained riparian vegetation or remnant trees. All sites were approximately 1 kilometer apart to minimize the possibility that ground fauna captured would travel between sites, and so acted as independent replicates.

Surveys were conducted over spring and summer (Aug. 2001 to Mar. 2002), and sampling effort was spread evenly across forest types to accommodate for any seasonal shift in faunal activity. Data was collected for one season and so could not contribute to questions about annual fluctuations in assemblage structure.

Pitfall Trapping Pitfall traps were used to sample ground-living terrestrial vertebrate fauna.

A single trapping grid composed of three trapping arrays was installed at each site. A trapping array consisted of four buckets arranged along the compass axes around a fifth central bucket. Traps (20L buckets) were buried level with

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the ground surface. Distal buckets were placed five metres from the central bucket. Two 35cm high, 10m long fences of black polythene were erected along the north-south and east-west axes of each array so that they each stretched across three buckets, and intersected over the central bucket. The bottoms of fences were buried into the ground so animals could not pass underneath.

Arrays were situated at least 150m from each other within a site, to minimize the possibility of sequentially recapturing individual animals. Animals caught in pit traps were measured and individually, inconspicuously paint-marked to identify recaptures.

Traps were initially opened for five nights, and re-opened six weeks later for a second five night trapping period (December 2001 and March 2002). Three arrays totaling 15 traps, for each of five sites, opened for 10 nights resulted in a total of 750 trap days per forest type. Traps were cleared once per day in the mornings and, after processing, all animals were released at the site of capture.

Diurnal Bird Census

I conducted all surveys over a five month spring/summer period

(September 2001 to January 2002) for four non-consecutive days at each site.

Birds censuses involved continuous slow walking (~1.4km/hr) along a 1.5 km randomly oriented line transect through the midpoint of each site. At each site two surveys beginning at 05:30h, and two at 06:45h were conducted, to coincide with the maximum pre- and post-dawn bird activity. All birds identified visually and aurally within a ten metre band either side of the transect line were recorded.

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Each forest type was surveyed for 23 hours in total (4.6 hours for each of five sites per forest type).

Small Mammal Trapping

A grid of Elliott traps (each 35x12x15cm) was established in each site.

Thirty traps were laid out in a grid of five transects spaced 80m apart, with each transect bearing six traps spaced at 40m intervals. Traps were placed in positions that facilitated their attractiveness to small mammals (rock crevices, against logs) and baited daily with a mixture of peanut butter, oats, honey, salami and vanilla essence. Traps were opened for five consecutive nights each trapping period. Traps were cleared in the early morning, and animals caught were identified, measured and individually marked before release at the site of capture. Thirty traps opened for five nights in five sites resulted in a total of 750 trap nights in each forest type. Trapping was staggered in time between replicate sites to avoid bias associated with the onset of the wet season.

Trapping commenced in November 2001 and finished in March 2002.

Spotlighting

Spotlighting surveys were conducted on separate nights to record the presence of nocturnal birds, mammals and herpetofauna. Four, foot-based spotlighting surveys were conducted along tracks within each site, and involved continual slow walking at ~0.75km/hr along a 1.5km transect. Tracks meandered within sites but were consistently directed east south-east. All animals identified within 15m either side of the transect line were recorded. Surveys were

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conducted between December 2001 and March 2002 at an effort of 20 survey nights per forest type.

Species characteristics

To identify any direction of change in ecological function between forest types and across taxonomic groups, primary diet and foraging strata were compiled from species accounts in the literature (Birds Australia 1991-2007,

Strahan 1995, Cogger 1996, Greer 2005, Barker et al. 1995). Firstly, to identify the character of the fauna using each forest type, each species was assigned to a group defined by major habitat preference. ‘Forest’ specialists were restricted to closed forest, ‘woodland’ specialists used woodland and open forest environments and ‘generalists’ used all forest types. Habitat categories were purposefully broad to encompass ecological characters that occurred consistently across animal taxa. Feeding resource guilds were then defined by combining two variables; primary diet (seven categories: carnivore, herbivore, nectarivore, omnivore, granivore, insectivore, frugivore) and foraging stratum

(six categories: all strata, all strata excluding the ground, canopy/subcanopy only, midstorey (>10m above the ground)-canopy, ground-low (0-2m above the ground, ground). Foraging stratum was defined as the stratum in which the majority of foraging behaviour occurred. Unlike other guild-based analyses, guilds were defined by combining diet and foraging characteristics because the interaction between these more accurately represented how a species was using habitat, and so generated a guild structure that was adequately descriptive across taxonomic groups. This resulted in twelve unique bird feeding guilds, five

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herpetofaunal guilds and four mammal guilds (Appendix 1). Reptiles and frogs were pooled for guild analyses because diet and foraging stratum overlapped strongly for both taxa. The two possum species and one megabat representing the nectarivore/herbivore-arboreal guild were excluded from data sets to restrict analyses to ground mammals only.

Data analysis

After satisfying assumptions one-way analysis of variance (ANOVA) was the used to test for differences in the mean richness and abundance of birds, mammals , reptiles and frogs in old and logged forests, young and old plantations using SPSS v 21 (SPSS, 2012). All data was pooled across transects, averaged over sites and log (x+1) transformed. Two-way ANOVA was used to identify any change in the richness of habitat specialists (closed forest, open forest, generalists; Appendix 1) with forest type. Pairwise comparisons with

Bonferroni corrections were generated to test for differences between forest types. Analysis of similarity (ANOSIM, PRIMER: Clarke, 1993) was used to identify any change in assemblage structure between young and old plantations and logged and old growth native forests for each faunal group using presence- absence transformed, standardized abundance data with a species by site matrix. A guild-by-site matrix was then generated and ANOSIM repeated to identify which, if any, guilds were lost in plantations. Changes in the relative abundance of guilds between forest types were analyzed separately for birds and mammals and herpetofauna using presence-absence transformed, standardized species richness data. All ANOSIMs used Bray-Curtis dissimilarity

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matrices and were iterated 2000 times. Pairwise comparisons between forest types were tested as part of ANOSIM. All analyses were considered significant at the p=0.05 level.

Results

Fauna sampling produced 3781 individual records of 2962 birds from 70 species, 157 frog captures from 5 species, 388 mammal records from 12 species and 453 reptile captures from 26 reptile species. Appendices one and two provide a species list for these captures. Collectively the total number of species recorded represented 28% of the total 408 terrestrial species recorded from the bioregion by the NSW Atlas of Wildlife (NSW NPWS, 2001).

Distinct bird, mammal and herpetofauna assemblages of equivalent species richness and abundance were identified from each forest type sampled

(Tables 2 + 3) despite significant differences in vegetation structure between forest types (Chapter 2). All taxa displayed non-significant trends towards lowest mean species richness and abundance in plantations generally and in young plantations in particular (Fig. 1a, b), apart from birds whose mean abundance was lowest in old plantations. For birds non-significant differences in abundance between forests and plantations (Fig. 1b) were driven by high variability in abundance in native forests.

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Figure 1. Mean raw species richness (A) and abundance (B) of birds, mammals and herpetofauna sampled from four forest types on the Mid North Coast of New South Wales, Australia. Bars represent old growth forest (black bars), logged native forest (grey bars), old plantation (stippled bars) and young plantation (open bars). All error bars represent the 95% confidence interval around the mean.

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Table 2. Results of ANOSIM analysis comparing the assemblage structure of birds, mammals and herpetofauna sampled from twenty field sites in four eucalypt forest types on the Mid North Coast of New South Wales. Global R values and p values indicate whether assemblage structure differs between forest types and associated pairwise comparisons identify where differences lie. Values in bold are p>0.05. Abbreviations for forest types are ON = old growth native forest, LN = logged native forest, OP = old plantations, YP = young plantations.

Birds Herpetofauna Mammals Global R p≤ Global R p≤ Global R p≤ 0.726 0.001 0.665 0.001 0.556 0.001 pairwise comparisons ON vs LN 0.406 0.016 0.828 0.008 0.520 0.008 ON vs OP 0.926 0.008 0.948 0.008 0.564 0.008 ON vs Y P 0.980 0.008 0.808 0.008 0.720 0.008 LN vs OP 0.558 0.008 0.464 0.016 0.524 0.008 LN vs Y P 0.828 0.008 0.728 0.008 0.740 0.008 OP vs YP 0.728 0.008 0.548 0.008 0.380 0.040

Habitat specialization

Birds

Forest birds comprised half of the 70 bird species identified in this study, and dominated forested habitats. Whilst young and old plantations supported 53% and 51% of all bird species, significantly fewer forest birds were recorded in plantations, and forest species were replaced by woodland specialists and generalist birds (Table 3, Fig. 2a). Interestingly habitat specialists (as defined in

Appendix 1) occurred in young plantations (17%) with similar frequency to old growth forest (19%) and at higher frequency than in old plantations (5%), suggesting that young plantations represented preferred habitat for some species, particularly grass and canopy feeders (Appendix 1).

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Table 3. Results of ANOVAs comparing log transformed count data for species richness, abundance and species by habitat specialization for birds, mammals and herpetofauna from twenty sites in four forest types on the mid north coast of New South Wales, Australia. Values in bold are significant at the p <0.05 level.

F d.f. p≤ Richness Birds 3.063 3,16 0.073 Mammals 0.611 3,16 0.617 Herpetofauna 1.53 3,16 0.251 Abundance Birds 2.722 3,16 0.448 Mammals 3.067 3,16 0.144 Herpetofauna 2.156 3,16 0.288 Richness x specialisation Birds 13.482 6,60 0.001 Mammals 14.183 6,60 0.001 Herpetofauna 13.428 6,60 0.001

Herpetofauna

Significant differences in herpetofaunal assemblage structure between all forest types (Table 2) were driven by the strong association of individual species with particular forest types. 43% of all species were restricted to old growth forests and only 8% of all species occurred in four forest types. Young plantations failed to support 89% of forest specialist herpetofaunal species, and in both young and old plantations forest specialists were replaced by generalists

(Fig.2b). This result contrasted that of birds, where woodland specialists rather than generalists replaced forest species in plantations. Notably, while young plantations supported 44% of all reptile species, only one species occurring at high abundance was restricted to this forest type (Appendix 1).

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Figure 2. Mean species richness of birds, mammals and herpetofauna grouped by habitat preference sampled from twenty sites in four forest types on the Mid North Coast of New South Wales, Australia. Bars represent forest specialists (black bars), generalists (open bars) and woodland specialists (stippled bars). All error bars represent the 95% confidence interval around the mean. Total mean richness on the y axis varies for each animal taxon.

Mammals

The distribution of mammalian habitat specialists differed significantly amongst forest types (Table 2, Fig. 2c). A diverse suite of forest mammals were restricted to old growth forests (Appendix 1). Woodland specialists were absent

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from old growth forests and most species rich in plantations. Both young and old plantations supported similar mammal richness. Similar to birds, assemblage fidelity was generally low for mammals, with 23% of all mammal species being trapped in all four forest types. The only mammal restricted to young plantations was the exotic house mouse (Mus musculus).

Cross-taxa comparisons of guild structure

Bird guilds differed significantly in species richness between all four forest types sampled (Table 3). Forests consistently supported a higher a richness of arboreal and ground-feeding insectivores, and carnivorous species foraging at all levels. Old plantations were dominated by canopy-feeding insectivores, and young plantations by generalist and ground-feeding insectivores and canopy- feeding frugivore/omnivores (Fig 3a). In opposition to birds, herpetofaunal feeding guilds differed only between logged forests and young plantations

(Table 4), primarily due the higher diversity of ground-feeding carnivores

(snakes) and the lack of insectivores, primarily arboreal snakes and geckos, in young plantations (Fig. 3b). Despite small numbers of mammal species trapped, there was a consistently greater richness of ground-feeding insectivores in old plantations than in logged native forests (Table 4, Fig 3 c). In contrast young plantations supported a marginally higher (but non-significant) richness of ground-feeding omnivores than old plantations (Fig. 3c).

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Figure 3. Total species richness of A) bird, B) herpetofauna and C) mammals guilds sampled from twenty field sites on the Mid North Coast of New South Wales. Each guild is defined by foraging stratum and primary diet. Foraging strata are A = all strata, G = forages at 0m, G-L = using all understorey vegetation from 0-2m, C = using primarily tree canopy/subcanopy only, Ar = uses any strata except the ground. Diets are C = carnivore, C/I = carnivore/insectivore, O = omnivore, F/I= frugivore/insectivore N/I = nectarivore/insectivore, N/F/I = nectarivore/frugivore/insectivore, I= insectivore, F/O = largely frugivorous but some omnivory, G = granivore. Bars represent old growth forest (black bars), logged native forest (grey bars), old plantation (stippled bars) and young plantation (open bars). 70 Chapter 3: Plantations and fauna

Table 4. Results of ANOSIM analysis comparing the diversity of feeding guilds of birds, mammals and herpetofauna sampled from twenty field sites in four eucalypt forest types on the Mid North Coast of New South Wales. Global R values and p values identify whether guild structure differs between forest types and associated pairwise comparisons identify where differences lie. Values in bold are significant at the p>0.05 level. Abbreviations for forest types are ON = old growth native forest, LN = logged native forest, OP = old plantations, YP = young plantations.

Birds Herpetofauna Mammals Global R p≤ Global R p≤ Global R p≤ 0.565 0.001 0.218 0.029 0.296 0.001 pairwise comparisons ON vs LN 0.596 0.008 0.140 0.159 0.468 0.032 ON vs OP 0.320 0.048 0.398 0.056 0.380 0.032 ON vs Y P 0.772 0.008 0.324 0.087 0.118 0.206 LN vs OP 0.604 0.016 0.006 0.444 0.560 0.016 LN vs Y P 0.780 0.008 0.344 0.024 0.280 0.087 OP vs YP 0.452 0.008 0.120 0.214 0.204 0.135

Discussion

This research identified that simplifying native forest structure through logging and plantation development did not decrease the overall species richness of birds, mammals, or herpetofauna at the stand scale, but significantly decreased the species richness of forest fauna from all taxa. Whilst this research concludes that young native plantations can be as species diverse as old growth native forests, it is important to recognize that this statement is untrue for forest specialist fauna, and for this faunal element, both young and old plantations may represent poor habitat.

Patterns of richness

Old growth native forest was equivalent in mean richness to logged native forest, 32 year old reforested and six year old young afforested plantation, and

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this response to habitat simplification was consistent across taxonomic groups.

This result supports those of Borsboom et al. (2002) whose research in small scale eucalypt plantations in moderate proximity to forests identified similar richness between plantations older than four years and forests. Faunal richness is likely to increase in plantations where the dominant tree species is endemic, if plantations present an environment that closely approximates some aspects of native forest (e.g. biophysical environment, feeding substrates, food resources, leaf litter structure - Strauss 2001). In this research the relatively small size of plantations, the use of local endemic tree species, broad connection between plantations and forest, and the retention of key habitat features are all likely to have promoted high species richness in young and old plantations. The juxtaposition of these elements may represent a scenario in which native plantations can achieve high faunal diversity. However as the work of Hobbs et al. (2003) and Kavanagh et al. (2005) in native plantations in fragmented forest systems in Australia attest, in the presence of a regional species pool negatively affected by native forest fragmentation, even large-scale native plantations remain depauperate in comparison to forest remnants.

Plantation age is a recognized determinant of diversity (Munro et al.

2007), and previous research has consistently identified young plantations as poor habitat for rainforest reptiles (Kanowski et al. 2006), birds and arboreal mammals (Borsboom et al. 2002, Kavanagh et al. 2005, Munro et al. 2007). But in this study, neither age nor land use history decreased mean richness in any animal taxon. This was unexpected for birds and herpetofauna, as both taxa are

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highly sensitive to changes in habitat structure (Brown & Nelson 1993, Boone &

Krohn 2000, Garden et al. 2007), and birds are documented as responding negatively to managed plantations less than five years old (Baguette et al. 1994,

Klomp & Grabham 2002, Kavanagh et al. 2007). Similar species richness between forests and young plantations have previously been documented only where there is a developed under/mid-storey (Loehle et al. 2005a, Loyn et al.

2007), and both Zurita & Bellocq (2010) and Sax (2002) suggest that bird species richness may often be temporally and spatially stable regardless of the dominant vegetation type in the presence of understorey complexity. However, in this research plantation understories were not highly complex. Instead habitat enrichment measures which have been demonstrated to increase species diversity in native plantations (Law & Chidel, 2002, Dent & Wright 2009, Hsu et al. 2010), are likely to have interacted with forest proximity to increase the attractiveness of plantations, and hence overall species richness. For frogs, reptiles and ground mammals which have lower mobility and strong reliance on small-scale site characteristics, in this study where both landscape context and site-specific characteristics promote habitat use, young plantations supported diverse faunal assemblages.

Assemblage structure and ecological function

In this research forest species were lost from all taxa in both young and old plantations. This loss commonly drives changes in the distribution of guilds (birds - MacArthur et al. 1962, Pearman 2002, Díaz et al. 2005; mammals- Fox 1985; reptiles - Attum et al. 2006, Urbina-Cardona et al. 2006).

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Plantations consistently lost arboreally foraging birds, mammals and herpetofauna (Table 5). There was a replacement of forest species with woodland species and specialists with generalists across all taxa in response to habitat simplification and increase in canopy openness. This response has previously been well documented as a response to plantations by birds (Hansen

1995, Christian et al. 1998, Munro et al. 2007), reptiles (Gardner et al. 2007b) and mammals (Jenkins et al. 2003, Holland & Bennett 2007), as driven by species-specific preferences for habitat structure. Reduced guild diversity directly reduces the resilience of systems (Luck et al. 2003), as well as having flow on effects on nutrient cycling, pollination, insect population dynamics and seed dispersal, processes which play a strong role in vegetation regeneration in agricultural landscapes (e.g. Archibald et al. 2011).

Birds

In this study habitat generalists represented a high proportion of the mammal and particularly reptile species using young plantations. However, even after 25 years of growth and management old plantation reptile assemblages were still heavily dominated by generalists, and birds were the only taxon to experience significant species replacement with woodland species rather than generalists in young and old plantations.

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Table 5. A summary of the ecological character of birds, mammals, and herpetofauna sampled from ten field sites in eucalypt plantations on the mid-north coast of New South Wales. Ecological character Birds Herpetofauna Mammals Species richness declines in No No No plantations Abundance declines in No No No plantations Site fidelity Low High Low

Forest species lost from Yes Yes Yes plantations Arboreal species lost from Yes Yes Yes plantations Increased dominance of Yes Yes Yes generalist species in plantations. Increase in woodland species in Yes Yes Yes plantations

Loyn et al. (2007) also identified native plantations as supporting woodland birds, which is positive news for woodland birds that are currently in decline in many parts of Australia (Paton & O’Connor 2010). However, if native plantations make landscapes more permeable to bird dispersal (Gascon et al. 1999, Lindenmayer

& Franklin 2002, Gardner et al. 2007b) this may increase competition between forest birds and generalist or woodland birds moving into forested environments.

Whilst highly competitive species such as Noisy Miners (Manorina melanocephala) and Indian Mynas (Acridotheres tristis) were not recorded in this research, these issues need consideration before the impact of the native afforestation on birds can be properly understood.

This research and that of Borsboom et al. (2002) identified that old plantations supported canopy-feeding insectivorous birds that were absent from young plantations, but lacked high abundances of frugivores or nectarivores

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common in native canopies. This suggests simplification of resource-availability in the canopy has long-term negative effects on guild structure, despite the high mobility of birds and the close proximity of old plantations to remnant native forest.

Ground Fauna

The association of different reptile and mammal assemblages with specific forest types is likely to be driven food or refuge availability for mammals and by thermal environment for herpetofauna (Zhao et al. 2006). Herpetofaunal assemblages in this study were distributed in relation to the thermal microclimates available in colder old growth native forest and hotter young plantations (Chapter 2). The young plantation assemblage included thermophilic snakes and lizards, while lizard species requiring high habitat complexity and cool thermal environments were common in old growth forests. The lack of cryptozoic species in old plantations reflected the lack of structural complexity on the ground resulting from long-term plantation management, and suggests that the retention of complex features like coarse woody debris may increase the diversity of both forest and woodland species in old plantations.

The separation of mammal species into different assemblages with change in habitat structure is common (e.g. Mengak & Guyunn 2003, Kavanagh

& Stanton 2005). Whilst mammals have been documented as occurring at lower abundance and species richness in exotic plantations (Lindenmayer et al.1999,

Sax 2002), in ecologically-managed native plantation monocultures that retain preferred habitat features, small mammals can be as diverse as they are in

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forests (e.g. Fonseca et al. 2009). Holland and Bennett (2007) and Ramírez and

Simonetti (2011) suggest that species specific-preference for vegetation structure can be more important than forest type in determining the presence of specific mammal species. In this study small mammal richness and abundance in plantations equaled that of forests, as species replacement by incursions of woodland and generalist small mammals with preferences for less complex understories balanced forest species loss.

Whilst forest mammals were abundant in old plantations, their abundance was significantly reduced in in young plantations. Mitchell et al. (1995) suggested that in their comparison of pine forest-pine plantation ecosystems, forest species maintained a presence in young plantations due to local extinction and recolonisation. In this study at least two forest mammals were considered resident in old plantations as they were observed carrying young, although given the low abundance of most forest species it is probable that local population extinction as suggested by Mitchell et al. (1995), or seasonal use of young plantations, was occurring. Clarifying the terms of residency of ground fauna in young plantations requires a longer-term investigation in young plantations, and should include an assessment of seasonal change.

The role of plantations in offsetting species loss

This study demonstrated that young native monoculture plantations established on previously cleared farmland can increase local biodiversity, and in the presence of biological legacies and close proximity to forests, young afforested plantations can support a diverse array of species. In this scenario,

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young plantations show potential as a forest management practice that can increase the local biodiversity of vertebrate fauna. Notably, in this research plantations supported assemblages of birds and mammals listed by the NSW

Department of Environment and Heritage as vulnerable in the Macleay-Hastings sub-region of the Northern Rivers Catchment Management Authority region.

Given the current development of broad-scale afforestation with native plantations, this study suggests that as the native plantation estate matures it may provide significant positive benefits to woodland fauna. However, advocating that plantation monocultures have conservation value must be tempered by recognition that landscape context strongly influences faunal use of plantations. Further, whilst young and old eucalypt plantations in this study supported diverse faunal assemblages of birds, mammals and reptiles, many species occurred at very low abundance, and plantation assemblages were consistently dominated by generalists and woodland specialists. The environment available in plantations is likely to provide both physiological and behavioral barriers that disadvantage many forest species, and so plantations may have little value when the conservation of forest species, particularly ground fauna, is a priority.

Quantifying the effect of historical land use on plantation environments is an inherent part of assessing the conservation significance of native plantations for fauna. Land use history influences vegetation structure (Raymond 2008,

Bremer & Farley 2010) and faunal use of native forest regrowth (Bowen et al.

2007, Cunningham et al. 2007), and so may have strong potential to affect

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species diversity in monoculture plantations. It is probable that in plantations managed for timber production, management will often outweigh the effects of land use history in the short term, as management has an immediate effect on vegetation structure and thus habitat quality (e.g. Klomp & Grabham 2002). Prior land use may affect current faunal use of plantations if historical isolation has compromised the quality of biological legacies as habitat, these habitat features play a focal role in promoting faunal diversity in plantations (e.g. Pharo &

Lindenmayer 2009, Hsu et al. 2010). Clarifying the effects of prior land use may only be possible as the plantation estate matures, but needs to include an assessment of the role of biological legacies (e.g. Lindenmayer et al. 2009,

Barrientos 2010) in increasing plantation diversity.

79 Chapter 4: Invertebrate resources and ground fauna

Chapter 4 The influence of invertebrate prey availability on ground-dwelling insectivores

Introduction

Plantation forests are an important part of the forest estate in many countries, and are increasingly recognized as important to faunal diversity in forested landscapes. However, our ability to predict the responses of ecological communities and individual species to environmental changes, like plantation development, remains a key issue for ecologists and conservation managers (Williams et al. 2010, Planque et al. 2011). Research in native plantations has been dominated by particular fauna, notably those useful as biological indicators such as ants, beetles, birds and small mammals. Despite the fact that many of the vertebrates using native plantations are insectivorous, research focused on the invertebrate prey base available in these forests is scarce. One reason for this research gap may be that, as for vertebrates, invertebrate responses to the plantation environment can vary among and within orders and species, making general predictions difficult.

Generally, plantation forests are expected to support less diverse invertebrate communities than natural forests, due to the low plant species diversity associated with monocultures (Knops et al.1999, Cunningham et al.

2005) and the scarcity of preferred habitat features like coarse woody debris on the ground (Carey & Harrington 2001, Owens et al. 2008). But how this relates to ecological processes that influence vertebrate use of plantations is not clear. Research linking insectivores with prey availability (e.g. Lunney et

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al. 2001) and quantifying how both prey base and predator abundance changes in response to habitat alteration is essential if we want to identify the quality of plantations as habitat for insectivores.

Prey Availability

Prey availability is one of the primary indicators of habitat quality for many animals. It has been demonstrated to affect community structure directly by allowing survivorship (e.g. James 1994) and indirectly by influencing predator body size (e.g. Laparie et al. 2010), niche breadth (e.g.

Costa et al. 2008, Garcia-Barros & Benito 2010), life history (Forsman 1996), competitive ability and ultimately fitness (e.g. Dickman 1988). The ability of predators to adapt to changes in prey availability depends on the degree to which they can vary their diets to accept other prey (Hodgkison & Hero 2003), which in turn is constrained by behavioural flexibility (e.g. Lescroёl et al. 2010) and morphological constraints like bite force (Herrel et al. 2007).

Both prey size and intractability, defined by Evans and Stanson (2005) as the biomechanical properties of a food that confer structural strength, stiffness and toughness, may strongly influence the attractiveness of prey, and so indirectly influence habitat use. Prey size affects feeding decisions by individuals by constraining predator attack rates (Hjelm & Persson 2001), while at the community level it can determine population structure by defining trophic links (Troost et al. 2008). There is a general trend for larger animals to eat larger prey (e.g. Fisher & Dickman 1993, Lima et al. 2000) because bite force and gape increase with head size (Herrel et al. 2001, Anderson et al. 2008). Similarly, as prey size increases so does intractability. However

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intractability is an important attribute of prey choice in its own right because it affects chewing (Evans & Stanson 2005) and swallowing. Intractability has been examined to understand the evolution of phenotypic differences (Herrel et al. 2009) and niche partitioning (Dumont et al. 2009) within populations of insectivores, but is less often cited as a factor that may influence habitat occupancy. However, because intractability could constrain both passive

(gape limited predators) and active prey selection mechanisms if it decreases food profitability by increasing handling time, it has the potential to influence the types and numbers of species that may use a habitat. Thus understanding the intractability of an invertebrate prey base as well as its abundance, may help to define the resource quality of a habitat, and hence its attractiveness to insectivores.

Predators, prey and plantations

The composition and distribution of epigaeic invertebrates can be influence the diversity of insectivores such as small mammals, reptiles, birds, and microbats, and as such provide a means to link insectivores from several taxa. As for vertebrates, epigaeic invertebrate communities are often strongly linked to available habitats defined by plant diversity (e.g. Hunter et al. 2000,

Taniguchi et al. 2003). The microhabitats associated with commercial hardwood plantations differ from native forests due to tillage and maintenance procedures that change soil, sub-soil profiles and simplify understorey structure (Marcos et al. 2007, chapter 1), and changes in canopy structure that influence thermal environments (Martius et al. 2004, Chapter 2).

Because of this, the potential of plantations to support a different community

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of invertebrate prey is high. In the last decade work exploring invertebrate ecology in plantations has identified that invertebrate communities do not respond predictably to habitat simplification (Bird et al. 2004, Barbaro et al.

2005, Yu et al. 2008, Oxbrough et al. 2010). Whilst community structure changes, communities using simplified habitats can be as diverse as those in undisturbed habitats (e.g. Hodge et al. 2010), or can be dominated by a few abundant insect orders with many other species occurring at low abundance

(e.g. Anderson & Death 2000, Bonham et al. 2002, Ratsirarson et al. 2002,

Cunningham et al. 2005). Typically, forest specialists (primarily beetles) decline in young plantations, while generalists or open-area users like spiders increase (Oxbrough et al. 2010). This shift in prey type coupled with a trend towards increased body size in plantations (for beetles at least –

Cunningham & Murray 2007), suggests that plantations may not offer food resources of a similar quality to those in forests. From a feeding ecology perspective the potential of hardwood plantations to support vertebrate insectivores will result from the breadth of predator prey preferences, and the composition and palatability of the invertebrate community in residence.

However, to the present date there has been very little formal investigation of what sort of prey base native plantations support in comparison to native forest, and how this relates to vertebrate diversity in native plantations.

Chapter three of this thesis identified that the structurally simple plantation environments assessed in the study area supported reptile and mammal assemblages that were equally as species rich as forest assemblages, but that showed trends towards reduced abundance. This result contrasts those of Hobbs et al. (2003), Kanowski et al. (2006) and

83 Chapter 4: Invertebrate resources and ground fauna

Ramírez and Simonetti (2011) which identified managed native plantations as supporting significantly fewer mammal and reptile species at less abundance than forests. This chapter relates the availability of invertebrate food resources in young and old plantations and logged and old growth native forests with the abundance of insectivorous mammals and herpetofauna, as an explanation for high vertebrate richness in plantations

Methods

Invertebrate sampling

Pitfall traps were used to collect ground-dwelling invertebrates from logged and old growth native forest and old and young plantations between

November and February in 2001 and 2002. See chapter two for a description of the study area the four forest types sampled. Although pitfall traps are a poor measure of absolute abundance and density of invertebrates (du Bus de

Warnaffe & Dufrêne 2004), they are equally biased towards mobile epigeal invertebrates, and when trapping is conducted over a small time scale, they provide an effective means of measuring the relative importance of invertebrates for comparative studies (Neumann 1991).

A 40 by 50 m grid of six transects each containing five traps (30 traps/site), was installed at the geographic centre of each of five 375 x 400m sites in each treatment. Traps were opened once in each site for four consecutive days resulting in 600 trap days per forest type in total. Each four day period constituted one sample. Sampling effort was interspersed between sites types and distributed across the eight months of the sampling to minimize temporal bias in results. Pitfall traps (10cm diameter, 15cm depth)

84 Chapter 4: Invertebrate resources and ground fauna

were buried level with the ground surface and protected from incident rain by a 25 cm diameter round plastic plate suspended 15 cm above the trap. Traps were filled with 80ml of 80% ethylene glycol as preservative and 2ml of detergent to lower the surface tension of the water. Invertebrates were removed from the preservative using a 400µm sieve, and were subsequently stored in 70% ethanol until sorting. Trapping occurred at the time of peak invertebrate activity, as some invertebrates were likely to be quiescent under bark or in logs or soil in the winter. Further some of the vertebrates recorded from the study area are torpid in cooler months of the year, and as I wanted to relate prey availability to vertebrate diversity it was necessary to trade off accurate richness estimates against an examination of seasonal changes in the abundance of food resources. Although seasonal changes in resource abundance are important because they can lead to diet switching and changes in competition (Becker et al. 2007), assessing the quality of plantations as a winter refuge was outside the scope of this study.

To augment sampling and minimise any bias associated with trap size, additional invertebrates were collected daily by hand from 15 x 20L, 35cm diameter pitfall traps for four days per trap over two trapping periods in 2001 and 2002 (trapping described in chapter 3). All invertebrates longer than two millimetres were collected. These samples were analysed separately to the small pit trap samples to identify any effect of trap size.

Invertebrates were identified to order. The level of identification was appropriate for this study because I was interested in the value of a species in terms of its physical properties rather than its identity. The length of each individual was measured from the tip of the frons without including any

85 Chapter 4: Invertebrate resources and ground fauna

protrusive mandibles, spines or antennae, to the posterior tip of the abdomen using a used a stereomicroscope (Leica MS5) with a graticule (limit of measurement 0.01mm). Larger specimens like several species of

Gryllacridoidea were measured using vernier calipers. Identification followed

Zborowski and Storey (2003) and CSIRO (1991). As per Lima et al. (2000) body length was considered to indicate prey attractiveness as it can constrain capture rates (e.g. Fisher & Dickman 1993) and swallowing times (e.g. Lima et al. 2000).

Invertebrate hardness

Intractability was categorized by assigning invertebrates a hardness score based on cuticle thickness. Cuticle thickness was measured for those specimens larger than four millimeters from a flat 2 x 2 mm section of sclerotized cuticle dissected from the hardest/ thickest part of the body

(elytron or thorax). Five individuals from each morphospecies from the twelve dominant orders were measured using a digital micrometer (±1µm) (as per

Evans & Stanson 2005). The mean of these values was rounded to the nearest single figure. Intractability was measured for the twelve abundant orders including the Amphipoda, Arachnida, Blattodea, Coleoptera,

Diplopoda, Diptera, Hemiptera, ants, Orthoptera, Dermaptera and

Lepidoptera adults and larvae. Results were grouped into classes; 1: 10-

30µm, 2: 30-60µm 3: 60-90µm 4: 90-110µm 5: ≥110µm. All Acarina were granted an intractability score of one because they were too small to measure cuticle thickness in, and were likely to represent soft prey given their relative size to the insectivores sampled.

Vertebrate sampling

86 Chapter 4: Invertebrate resources and ground fauna

Small mammals were sampled using pitfall traps and metal live traps

(Elliott traps – 38 x 12 x 11 cm). Live trapping occurred in conjunction with pitfall trapping between December and March in 2001, and was repeated between October and January 2003. Chapter three describes the methods used for pitfall and live trapping in detail. Traps were opened between 16:00 and 08:00EST for five consecutive nights, resulting in a trapping effort of 750 trap nights/forest type. Traps were baited with a mixture of rolled oats, peanut butter, honey, vanilla essence and salami. All animals captured were released at the trap site after paint marking the bottoms of feet or to identify recaptures. The pitfall trapping procedures and effort (750 trap nights x forest type) followed those described in chapter 3.

Relating Predators and Prey

Fourteen of the 35 insectivore species trapped in this study were excluded from analyses as they possessed kinetic jaws, were only insectivorous at juvenile life stages or had strong preferences for forest habitat features precluding their use of plantations (Appendix 2). Of the omnivores, the Northern Brown Bandicoot Isoodon macrourus was excluded as invertebrates were unlikely to control its distribution due to high levels of mycophagy, but house mice (Mus musculus) and native rats (Rattus fuscipes,

R. lutreolus) were included as they were known to be insectivorous during

Spring-Summer when this study was conducted (Watts & Braithwaite 1978,

Cheal 1987).

87 Chapter 4: Invertebrate resources and ground fauna

Table 1. The distribution of mean gape sizes and prey size category used by insectivores sampled from four forest types on the Mid North Coast of New South Wales. The maximum insectivore gape was calculated from snout-vent lengths of field-caught individuals, and determined the prey size range available to each species. Diet preference data come from Fisher & Dickman 1993, Dickman 1988, Green 1989, Lima et al. 2000, Lunney et al. 1989, 2001, Vinyard and Payseur 2008, Williams et al. 2009. All measurements are in millimeters. Preferred forest lists the forest type in which each species was most abundant. In this study Abbreviations for forest type are OP- old plantation, YP – young plantation, LN – logged native forest, ON – old growth native forest.

Predator Snout-Vent Mean Preferred prey Available prey Dominant Preferred Taxa Group Species Length gape size range size range Feeding Mode Forest Abundance Herpetofauna Small Adrasteaiscincus amicula 30.00 3.22 1-5 5-10 Insectivore YP 59 Small Lampropholis delicata 40.00 4.68 1-10 10-15 Insectivore OP 265 Small Lampropholis guichenoti 40.00 4.68 1-10 10-15 Insectivore YP 175 Small Crinia signifera 23.00 8.72 1-15 15-20 Insectivore OP 15 Small Pseudophryne coriacea 23.00 8.72 1-15 15-20 Insectivore OP 248 Small Uperolia fusca 25.00 9.48 1-15 15-20 Insectivore OP 23 Medium Amphibolurus muricatus 80.60 10.61 5-20 1-5 Insectivore YP 24 Medium Calyptotis ruficaudata 47.40 5.76 5-15 1-5 Insectivore LN 51 Medium Ctenotus robustus 100.00 13.44 2-20 20-25 Insectivore YP 18 Medium Lerista muelleri 58.00 7.31 1-10 10-15 Insectivore ON 26 Large Cyclodomorphus gerrardi 200.00 28.04 25-50 15-25 Insectivore ON 9 Large Limnodynastes peroni 65.00 24.68 25-40 15-25 Insectivore YP 35 Large Tiliqua scincoides 300.00 42.64 25->50 15-25 Omnivore OP 14

Weight (g) Mammal Small Planigale maculata 6.58 15.00 1-15 15-40 Insectivore OP 20 Small Sminthopsis murina 18.00 22.40 5-25 25->50 Insectivore OP/YP 27 Medium Antechinus stuartii 23.43 17.40 15-25 5-10,25->50 Insectivore ON 205 Medium Antechinus swainsonii 51.30 28.40 15-35 5-10,35->50 Insectivore LN 9 Small Mus musculus 15.53 12.50 5-15 1-5 Omnivore, granivore YP 67 Large Rattus fuscipes 96.07 23.00 20-30 5-20 Omnivore, mycophage ON 427 Large Rattus lutreolus 120.45 31.00 20-40 5-20 Omnivore OP/YP 69

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The species included in the data set were grouped by body size into the groups: small, medium and large herpetofauna, small and medium dasyurids, small and large rodents.

Decisions about what length of invertebrate represented a potential prey item were made by referring to published accounts of prey size for the species encountered (Green 1989, Lunney et al. 1989, 2001, Littlejohn et al.

1993, Barker et al. 1995, Wapstra & Swain 1996, Lima et al. 2000, Greer

2005). When this information was lacking predator gape size was used to determine which invertebrates were available as potential prey. This approach was used rather than a more direct analysis of faecal pellets, as preliminary analyses determined that invertebrate fragments in pellets of small lizard species (0.3-0.5g) were too small to infer prey length (e.g.

Angelici et al. 2007, Bos & Carthew 2007). For herpetofauna gape was calculated from the modal snout-vent lengths of all individuals of each frog and lizard species included in this study. Gape size was adjusted to account for body shape using the equations of Lima et al. (2000) who have generated correction factors for biometric gape size-body size relationships in small

Australian frogs and lizards. Mammal prey size preference of the same species or of species with similar body size and habits were determined in previous research by Fisher & Dickman 1993, Dickman 1988, Green 1989,

Hanson et al. 2007, Vinyard and Payseur 2008 and Williams et al. 2009.

Invertebrates were assigned to mutually exclusive categories as being either preferred (preferentially eaten), available (outside of the preferred prey range) or unavailable (not able to be eaten) on the basis of insectivore gape size. For herpetofauna preferred prey were defined as those whose length

89 Chapter 4: Invertebrate resources and ground fauna

were 50% of the maximum gape, which in frogs and lizards is the portion of the gape in which significant bite force is applied (Druzinsky & Greaves 1979).

Whilst rats chew their food (Cox et al. 2012) bite force is optimized at 40-50% of the maximum gape (Williams et al. 2009), so preferred invertebrates were defined as those whose lengths were 40% of the recorded maximum gape, where published mean and maximum prey lengths did not exist. For all insectivores available prey was defined as any invertebrate length that was

10% less than the minimum preferred prey size, and that was between the top end of the preferred prey range and the maximum gape. However, because dasyurids orient, chew and dismember their food all prey lengths represented available prey for all species except Planigale maculata which was estimated to be constrained to prey less than 40mm long due to its small size. After defining the preferred and available prey ranges for each species the size classes of invertebrates falling within these ranges were assigned to each predator group (Table 1). Predator defense mechanisms like distasteful secretions or poison were not accounted for because literature accounts of species-specific preferences for certain prey types were often contradictory.

An unavailable prey was either below the recorded minimum preferred prey size or above the maximal gape size.

90 Chapter 4: Invertebrate resources and ground fauna

Data analysis

Insectivores

As data could not be transformed to meet parametric test assumptions and so allow posthoc testing, the relative abundance of small, medium and large insectivores (Appendix 2) was compared between logged and old growth forests and young and old plantations using the non-parametric analysis of similarity (ANOSIM) procedure in Primer (Clarke 1993). Bray-

Curtis similarity matrices were generated using log x+1 transformed, column- centred abundance data. Multiple comparisons of each forest level were performed on data bootstrapped 2000 times. All data were analysed at a significance level of p = 0.05.

Invertebrates

After preliminary analyses showed that small and large traps consistently sampled the same suite of invertebrates each year, both year and trap size were removed as factors in the statistical models. Pitfall trapping yielded 10653 individuals from 28 invertebrate orders. Orders that occurred at less than 20 individuals in any forest type were excluded from the data set because they were not considered abundant enough to drive patterns of habitat use by insectivores . The 14 excluded orders accounted for 2 percent of the total abundance.

To assess any differences in the abundance of invertebrates of varying intractability between forest types and orders a two-way Analysis of

Variance (ANOVA) with forest type as fixed factors and order as a covariate was generated. This analysis was repeated to assess differences in the modal size of invertebrates in each forest. Data was normalised to meet test

91 Chapter 4: Invertebrate resources and ground fauna

assumptions using log x+1 transformation and Bonferroni correction was applied to all posthoc tests for differences between forest types. The available prey base in each forest type was assessed by analysing the frequency of preferred, available and unavailable prey sizes using Chi- squared tests for independence. Invertebrate data were grouped by insectivore body size to allow meaningful comparisons across species. This resulted in resulting in seven groups: small, medium and large herpetofauna, small and medium dasyurids and small and large rodents. A separate Chi- squared was performed for each group after verifying the assumptions of

Chi-squared were upheld. Bonferroni corrections were applied to identify significant differences between forest types.

Results

Insectivores

Two years of trapping yielded a total 1978 captures of 5 frog, 21 reptile and 9 mammal species (Appendix 2). Plantations collectively supported a marginally richer insectivore community than native forests (old growth 20 spp., logged native 17 spp., young plantation 20 spp., old plantation 21 spp.). After the exclusion of forest specialists a reduced data set of 1786 captures from 20 species captures were analysed. The abundance of large insectivores declined significantly moving from plantations to forests, but both small and medium insectivores were similarly abundant in logged and old growth forests and significantly more abundant in plantations than forests (Table 2, Fig. 1).

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Table 2. ANOSIM results testing for changes in small medium and large insectivore abundance in each forest type. Significant differences at p < 0.05 are in bold. ON = old growth native, LN = Logged native, OP = old plantation, YP = young plantation.

Small Medium Large Global R p≤ Global R p≤ Global R p≤ 0.67 0 0.573 0.001 0.429 0.001 pairwise comparisons ON v LN 0.224 0.079 0.220 0.016 0.076 0.325 ON v OP 0.508 0.016 0.432 0.016 0.406 0.048 ON v Y P 1.000 0.008 1.000 0.008 0.908 0.008 LN v OP 0.504 0.008 0.156 0.056 0.342 0.056 LN v Y P 1.000 0.008 0.956 0.008 0.840 0.008 OP v YP 0.852 0.008 1.000 0.008 0.078 0.238

Figure 1. Cumulative abundances of insectivores from small, medium and large body size groups sampled from old growth and logged native, and old and young native plantations on the Mid North Coast of New South Wales. Letters denote significant differences between forest types.

93 Chapter 4: Invertebrate resources and ground fauna

In young plantations large bush rats were replaced by swamp rats (Rattus lutreolus) which occurred at very low abundance and were co-dominant in this habitat with medium-sized house mice (Mus musculus). Mice were only sampled from young plantations and comprised 18% of the individuals captured in this forest type. Patterns within the dasyurids varied between sizes. Only medium-sized Brown Antechinus (Antechinus stuartii) were significantly more abundant in old growth forests than in either plantation forest types. Small dasyurids were similarly abundant in logged native and plantation habitats but avoided old growth forests, whilst large dasyurids were generally rare and occurred only in logged native forests (Fig 1).

Young and old plantations were dominated by small herpetofauna, particularly lampropholine litter skinks, but this assemblage was replaced in old growth native forest by less abundant assemblages of cryptic skinks

(Appendix 2). Although large herpetofauna were rare they occurred at greater frequency in young plantations than any other forest type. Generally dasyurids, occurred most frequently in old plantations where they comprised

15-17% of the total number of individuals. In young plantations dasyurids represented only 8% of the total number of captures.

Invertebrate prey

Whilst invertebrate hardness differed between invertebrate orders

(F13,54 = 119.03 p < 0.001) it did not differ between forest types (F3,54 = 0.039 p < 0.990) and so was subsequently dropped from all further analyses.

Similarly the mean size (forest - F3,54 = 0.059 p = 1.088, order – F1,54 = 0.078,

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p = 1.443) and abundance (forest - F3,54 = 0.572 p < 0.248, order – F1,54 =

0.552, p = 0.548) of invertebrates within orders did not differ between forests and plantations. There were non-significant trends for young plantations to support high abundances of ants, millipedes, beetles, amphipods and grasshoppers whilst native forest assemblages were dominated by mites, beetles, flies, bugs and ants. Only hemipterans and ants showed trends towards higher abundance in logged native and old plantation forests.

However, whilst the mean size and abundance of invertebrates did not differ between plantations and forests, the frequency of invertebrates available to and preferred by insectivores differed significantly between forest types

(Table 3). Preferred prey occurred most frequently in young and old plantations for small and medium herpetofuana, medium dasyurids and small rodents (Fig. 2, Table 3). Available prey for small dasuyrids and large rodents were also trapped more frequently in plantations, however plantations supported very little preferred or available prey for large herpetofauna (Fig. 2).

Table 3. Results of Chi-squared analyses and associated pairwise comparisons for changes in the frequencies of preferred, available and unavailable prey sampled from old growth and logged native forests, and eucalypt plantations on the Mid North Coast of NSW. Abbreviations for forest types are ON = old growth native, LN = Logged native, OP = old plantation, YP = young plantation.

Predator group χ 2 d.f. p Small herpetofauna 1025.327 6 <0.001 Medium herpetofauna 983.442 6 <0.001 Large herpetofauna 1084.933 6 <0.001 Small dasyurid 540.179 6 <0.001 Medium dasyurid 172.201 6 <0.001 Small rodent 983.442 6 <0.001 Large rodent 143.805 6 <0.001

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Fig. 2. The frequency of preferred, available and unavailable prey present in each forest type for the seven predator groups analysed. Each subscript letter denotes how the frequency of column proportions within each forest category subset differs significantly at the .05 level. Bars represent old growth forest (black), logged forest (grey), old plantation (stippled) and young plantation (white). 96 Chapter 4: Invertebrate resources and ground fauna

Discussion Young native plantations in this study supported a diverse epigaeic insectivore community and significant food resources for ground-dwelling insectivores. They supported similar abundances of invertebrates as logged and old growth native forests, and higher frequencies of preferred prey sizes for all insectivores than old plantations and native forests. Whilst this distribution of preferred prey benefited small and medium-sized insectivores, which were significantly more abundant in plantations than forests, large insectivores did not effectively exploit high prey densities in plantations. For large rodents in particular were uncommon where preferred and available prey was abundant. If the habitats investigated in this study were saturated and insectivores conformed to an ideal free distribution, individuals should have aggregated in each forest type proportionately to the amount of resources available in each (Fretwell & Lucas 1970). In this study insectivore abundance was not so simply distributed. Many studies recognise the importance of multiple cues for habitat selection, which often vary with species (e.g. Fischer et al. 2004, Radford & Bennett 2007, Price et al. 2010).

In this study while the disjunction between prey availability and large insectivore abundance may have been influenced by intrinsic factors including breadth of habitat choice, diet, and preference for foraging optimally, a variety of extrinsic cues including structural and thermal microhabitat complexity and predation risk, may impinge more strongly on medium and large, rather than small, species in plantations. This is because high levels of management-associated disturbance in plantation environments are likely to more directly impose constraints on the resources large predators require. In

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particular, retreat sites which need to be large enough to provide refuges adequate for predator avoidance and thermoregulation in the hotter (Chapter

2) plantation environment, are lost in plantations by active understorey management and through grazing (Chapter 5). As several authors have identified increased abundance in large ground mammals in plantations in association with increased vegetation complexity at the ground level (coarse woody debris- Loeb 1999, shrubs- Loyn et al. 2008), the lack of retreat sites definitive of plantations in this study (Chapter 2) may limit the species richness of large predators.

Predator Abundance

Small Predators

Prey availability was unlikely to constrain habitat use by small insectivores in any of the four forest types examined in this study. Prey preferred by small herpetofauna and dasyurids occurred frequently in all forest types. Instead, other factors such as habitat preferences or predation rates were likely to overshadow prey in driving patterns of habitat use. While the abundance of small predators did not differ between forests, there were strong trends towards plantations supporting higher abundances of small lizards. Breadth of habitat choice, in particular, preferred temperature range, may explain much of this pattern as Lampropholine lizards were the dominant small predators. The most ‘generalist’ of these, Lampropholis guichenoti and L. delicata are thermophilic (Spellerberg 1972, Greer 1980,

Chapter 6). Similarly despite the assertion of Wells (2010) that the smaller- bodied Adrasteia amicula is associated with cooler habitats, in both this study

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and that of Boorsboom et al. (2002), A. amicula was restricted to native plantations and logged native forests, suggesting that it too may be thermophilic, or strongly associated with open canopy, or both.

In this study small dasyurids (common dunnarts and planigales) were not trapped in old growth forests despite an equal abundance of prey in all forest types. While common dunnarts are likely to have been excluded from native forests by a preference for dense grassy understoreys (Stokes et al.

2004), the absence of common planigales in old growth native forests despite the high availability of palatable prey may result from competitive exclusion by the medium-sized brown antechinus. Competitive exclusion resulting in the displacement of smaller species is known to occur between small dasyurids (Dickman 1988), unless diverse foraging microhabitats allow co- existence. In this study logged native forests supported the most diverse foraging microhabitats which included shrub layers and high stem density lacking in old growth native forests (Chapter 2),and were the only forest type to concurrently support small (Planigale maculata, Sminthopsis murina) and medium (Antechinus stuartii, Antechinus swainsonii) dasyurids. This result is particularly important for plantation management as the Common Planigale is currently listed by the New South Wales Department of Environment and

Conservation as vulnerable in the Macleay-Hastings catchment sub-region. If management efforts aim to benefit this species in favoured habitats like young and old plantations, retaining or protecting preferred habitat features like grassy understoreys will be necessary at harvest or thinning.

99 Chapter 4: Invertebrate resources and ground fauna

Medium Predators

While medium-sized prey was abundant in plantations, medium-sized dasyurids were rare, and were effectively replaced by the medium-sized rodent, the introduced house mouse. The high abundance of house mice in plantations is likely to result from an omnivorous diet and strong association with highly cultivated landscapes (Singleton 1995) rather than the intrinsic abundance of invertebrate prey in plantations. This numeric dominance by mice over other rodents of similar body size was also identified by

Boorsboom et al. (2002) who found strong association between grassy understorey, young plantations and house mouse abundance. Both

Boorsboom et al. (2002) and Caughley (2001) identified that mouse abundance was much higher in winter, than summer, which suggests some potential for the Brown Antechinus to be seasonally excluded from young plantation, although Singleton (1995) suggests mice are poor competitors with herbivores of similar size (e.g. Archibald et al. 2011). A winter influx of mice into plantations may have knock-on effects for small lizards in plantations, as mice were observed preying upon lizards on several occasions in this study.

The disjunction between prey availability and predator abundance also occurred in the medium-sized dasyurid, the Brown Antechinus and may have resulted from a scarcity of retreats or of useable foraging substrates. While

Brown Antechinus did occur in young plantations their total abundance was significantly higher in old plantations where their preferred prey was less abundant, suggesting that the young plantation environment was less than optimal for this species. If predators were to forage optimally they should

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consume the biggest prey possible for the best energetic return (see review by Pyke1984). Whilst a preferred prey range exists, the available prey range for the generalist Brown Antechinus was large due to its ability to process food by chewing and stretching (Fisher & Dickman 1993) and generalists will readily exploit prey sizes outside the preferred range to avoid competition with other insectivores (e.g. Dickman 1988). As Brown Antechinus are strongly opportunistic foragers, and prey switching in this species is common

(e.g. Green 1989, Lunney et al. 2001), it is likely that this species was successful in plantations because it had the ability to exploit abundant prey outside its preferred prey range.

Large Predators

Plantations supported valuable food resources for large herpetofauna but these were underutilized. The very low abundance of large herpetofauna in plantations is not likely to be driven by prey availability alone because pink-tongued skinks are cryptic and semi-arboreal and prefer dense ground layers (Cogger, 1996) not available in young plantations. Livestock grazing is known to negatively affect the abundance of reptiles of similar body size as the ‘large’ lizards in this study (Kutt & Woinarski 2007, Brown et al. 2008) by reducing the structural complexity of the understorey vegetation. The low abundance of large lizards may also have been a more direct response to plantation structure, because while food is abundant in plantations, retreat sites are lacking in the plantation matrix itself, as hollow stumps, logs and snags are only retained in habitat islands, and whilst Blue-tongued skinks will tolerate open habitats, they require retreats to avoid predators. The importance of refuges in maintaining ground fauna has demonstrated by

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Webb and Shine (1998) who found that without suitable retreat sites, thermal regimes forced snakes to switch habitats. Similarly the many studies that advocate retained biological legacies like coarse woody debris a means to increase faunal diversity (see Stevens 1997, Lindenmayer & Hobbs 2004 for reviews), identify the importance of refuges in allowing species to persist in managed habitats. A final explanation for low large lizard abundance in plantations may be that large lizards are generally poorly represented in systematic trapping surveys (Eyre et al. 1999, Borsboom et al. 2002) and so may have been under-sampled by the trapping methods employed.

Whilst large rodents were significantly more abundant in native forests, their preferred prey was more abundant in young plantations. However invertebrate prey may not be the only dietary habitat selection cue for large rodents, as other studies have identified that Bush rats (Rattus fuscipes) and

Swamp rats (Rattus lutreolus), the two rat species resident in plantations, use invertebrate prey in summer to differing degrees (Cheal, 1987, Carron et al.

1990). While Bush rats are “selective” generalists that seasonally consume fungi (Vernes & Dunn 2009), Swamp rats eat a large amount of plant material (Watts & Braithwaite 1978, Cheal 1987). Thus for all the large predators vegetation structure is likely play a strong role in contributing to habitat choice.

Grazing

While grazing is likely to influence the quality of habitats at a local scale in plantations, it is also has potential to be a landscape-level driver of ground-level plant diversity and thus structural complexity. Livestock grazing

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is common in young native plantations and is encouraged in this study as a multi-use land strategy by plantation managers. While it is well documented as negatively affecting specific ground fauna species by broadly reducing habitat heterogeneity (e.g. Kutt & Woinarski 2007, Brown et al. 2008, Price et al. 2010, Read & Cunningham 2010), its action at the landscape scale

(Fisher et al. 2004) means grazing is likely to influence regional species pools. In this study while plantations have broad connection to closed native woodlands, these woodlands have a long history of intermittent grazing at low stocking density, suggesting that the negative effects of grazing may be intensified by the position of plantations in the landscape mosaic.

We might expect this response to be strong for ground fauna of all sizes, particularly small lizards and frogs, as dispersal ability in this faunal group is often relatively limited (Brown et al. 2008). However in this study grazed young plantations are more speciose than old growth native forests for every taxon, suggesting that a complex interaction between grazing, habitat simplification and dispersal is occurring. Prey availability has an important role to play in this interaction as it may compensate for the lack of structural diversity as a habitat selection cue, allowing small insectivores that aren’t as explicitly reliant on retreats, to use young plantations.

Conclusions

This study has identified that young and old plantations support significant food resources for small, medium and large epigaeic insectivores.

However medium and large-sized insectivores appeared to be responding to habitat selection cues independent of prey availability, and patterns of habitat

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use by these insectivores did not follow the abundance of preferred prey. If species are foraging opportunistically, prey availability is likely to be an important habitat selection cue for fauna using plantations, however in plantations that are a thermally stressful habitat (Chapter 6), prey availability appears to be less important in predicting where a species will be abundant than extrinsic cues like retreat availability or substrate complexity. In addition to prey availability, landscape composition and individual species habitat requirements will need to be considered on a species-by-species basis in order to predict the persistence of ground-fauna in plantations. Further research needs to assess the residency of fauna in young plantations, and particularly to identify whether management designed to enrich plantations like retaining coarse woody debris acts as a habitat selection cue for species of various body sizes, so increasing biodiversity across food chains. In my study area this could have immediate implications for the management of vulnerable species like Common Planigales, and should be addressed in order to generate effective management for plantations currently being established.

104 Chapter 5: Matrix quality and ground fauna

Chapter 5 Matrix quality and ground fauna diversity in young plantations

Young native plantations in the landscape mosaic

Native plantations are increasing rapidly and within Australia, in the last ten years, they have increased by 51% to an area of 0.99 million hectares, representing 49% of the total plantation estate (Gavran 2013). Recent statistics confirm that The Mid-North Coast forest region of New South Wales currently supports 93% (86 296 ha) of the state’s hardwood plantation estate, and is one of four areas in Australia with the fastest rate of increase in total plantation area

(mainly hardwoods) since 2005 (Gavran 2013). Thus young plantations (<10y.o.) represent a significant and growing landscape feature in the region, and in

Australia as a whole. Despite the differences in species combination, topography and proximity to native landscapes, all managed new plantations have the potential to represent a cohesive landscape element because their youth and the typically simple structure produced by standard establishment procedures and subsequent management, creates a similar forest type where plantation occur. Understanding how this expanding forest type is used by fauna is particularly important in North-eastern New South Wales as this area is one of the most biologically diverse regions of Australia (DEWHA 2009), and is home to endemic and threatened species, as well as species at the margins of their distributional ranges (Pressey et al. 1996).

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One dominant paradigm of conservation in managed landscapes stems from metapopulation and island biogeography theory (Ricketts 2001) and consists of viewing the landscape as a fragmented series of habitat patches, dominated by an extensive matrix habitat that has a strong influence on both fauna and ecological processes (McIntyre & Hobbs 1999, Radford & Bennett

2007). This is extended by the landscape continuum model (see Lindenmayer &

Franklin 2002) which suggests fragment dwelling fauna can disperse in the matrix along suitable food, shelter or climatic gradients. While applying these models may be an appropriate approach to understanding the dynamics of native forest patches within large-scale industrial exotic plantations, it may be a less accurate way to predict diversity in young native plantations as it necessarily brands them with the status of ‘fragment’. However, native plantations in New South Wales are established on reforested pasture land, often in broad connection to both existing forest types and cleared farmland.

Thus it may be misleading to view young native plantations as fragments. Given their presence and connectance to the forest matrix, young plantations may be better visualized as a skirt of compartmentalized blocks of simple forest (the variegated landscape condition of McIntyre & Hobbs 1999), which border (and may buffer) native forest from cleared land, and which may complement and extend, rather than fragment, native forests.

The matrix and matrix quality

In the Macleay-Hastings management area of the North Coast Bioregion the landscape mosaic is dominated by closed and open eucalypt forest and

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woodland (Fig. 1), retained as part-forested agricultural land and a chain of public reserves which link 25 national parks and 13 state forests between the coast and the foothills of the Great Dividing Range (Pitt 2001). While young plantations are present in the landscape, they are currently not extensive enough to fragment native forests, but instead represent simplified habitat patches containing small islands of complex structure (retained habitat features) embedded in and adjacent to an extensive native forest matrix (Chapter 2).

Understanding how this forest matrix influences plantations is necessary to understand the value of plantations for fauna, as the adjacency and quality of the matrix can profoundly effect specific groups of fauna (Lindenmayer &

Franklin 2002, Loyn et al. 2007, Popescu & Hunter 2011).

The importance of the surrounding matrix in influencing plantation biodiversity cannot be underestimated (Gascon et al. 1999, Lindenmayer &

Hobbs 2004, Umetsu & Pardini 2007; reviews by Brockerhoff et al. 2008,

Perfecto & Vandermeer 2010). As a matrix exotic plantations can ameliorate species vulnerability to fragmentation in remnant native forest (e.g. Henle et al.

2004, Umetsu & Pardini 2007, Brockerhoff et al. 2008, Tomasevic & Estades

2008), facilitate dispersal between fragments (e.g. Marchesan & Carthew 2008,

Predevello & Viera 2010), and supplement species habitat or resources at a landscape level (e.g. Pawson et al. 2008).

107 Chapter 5: Matrix quality and ground fauna

Fig 1. Two six-year-old hardwood plantations in the Mid North Coast NSW study area showing typical connection to other landscape elements. A large retained habitat feature is present in the right hand side photograph and both pictures show grazed eucalypt woodland in broad connection to the edges of the plantation compartments.

At the stand scale exotic plantations may buffer fragments from adverse environmental microclimates (e.g. Wright et al. 2010) and bias community composition towards generalist species (Ewers & Didham 2006). However, while a large body of work has looked at the role of native neighboring landscapes in increasing diversity in native remnants (Munro et al. 2009, Archibald et al. 2011), there is currently no published work investigating whether native matrices have positive effects on fauna in native plantations.

Generally we might suppose that native plantations adjacent to the native forest matrix will exhibit high diversity due to an influx of dispersers from source populations, and that this diversity will be tempered internally by conditions imposed by the plantation environment and externally by matrix quality. If plantations are acting as a habitat sink, the age structure of populations within plantations may be biased towards juveniles as plantations receive recruits from native forests in close proximity (Jansen & Yoshimura 1998). Further to this, we

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might expect that the simple environment of plantations would support more generalist species than the surrounding matrix (Chapter 3), although this may depend on matrix quality. Thus understanding matrix quality and its permeability to fauna is important in predicting plantation diversity.

While matrix permeability may explain some of the high species diversity in young plantations identified in this study for all fauna, and for generalist species in particular (Chapter 3), it does not explain the lower abundance of many species, particularly small mammals, in young plantations (Chapter4). If low abundance is unrelated to food availability (Chapter 4), then it may be related to the quality of the surrounding matrix (e.g. Perfecto & Vandenmeer

2002). In my study area the forest matrix adjacent to plantations is cattle-grazed.

Grazing has been demonstrated to negatively affect understorey complexity

(Yates et al. 2000) and associated ground fauna in several biomes (woodlands:

Bromham et al. 1999, James 2003, Driscoll 2004, Kutt & Woinarski 2007; steppes: Beever & Brussard 2004; shrubland: Read & Cunningham 2010), although responses to grazing can be highly species specific (Woinarski & Ash

2002, Fischer et al. 2004). Using standardized surveys this chapter evaluates forest matrix use by herpetofuana and small mammals in comparison to adjacent young plantations. I focus on ground fauna because they occur at low abundance in plantations (Chapter 4), they are known to be negatively affected by grazing, and their relatively low vagility means that landscapes are generally less permeable for this faunal element, and so matrix quality may affect them strongly. I test the theory that matrix quality is compromised by long-term

109 Chapter 5: Matrix quality and ground fauna

grazing. I hypothesize that if grazing has negatively affected the species pool recruiting to young plantations then:

1) plantations should support a species richness similar to the forest matrix

regardless of the more complex simpler vegetation structure in

plantations

2) plantation population structures should be skewed towards a larger

number of juveniles if simplified habitat structure is generating a

population sink. If young plantations were a population sink young

dispersers moving into plantations from the forest matrix should bias the

population structure of plantation assemblages towards juveniles.

I test these hypotheses by comparing vegetation structure in grazed forest and young plantations, the species richness and abundance of ground fauna in both forest types and the population age structure of fauna in young plantations and adjacent native forests.

Methods

Habitat Description

A general description of the study area and the young plantation environment is presented in the study area section of Chapter 2. Sites within the young plantation treatment were the same as those surveyed for fauna as described in Chapter 3. In summary, young plantations at eight years old were open-canopied, with a canopy height of approximately six metres, and a patchy, grassy understorey dominated by Blady grass (Imperata cylindrica). A very

110 Chapter 5: Matrix quality and ground fauna

patchy shrub layer of the exotic Lantana camara was occasionally present.

Young plantations lacked logs and shrubby understorey. Establishment procedures in young plantations included the retention of tree-bearing riparian vegetation buffers, and aggregations of seed and habitat trees within the plantation. Young plantations had been exposed to cattle grazing for two years prior to the commencement of the study, and prior to plantation establishment, the land had been grazed as pasture for at least 40 years. Forty percent of the plantation edge abutted the native matrix forest, and was separated from it by a four-metre-wide fire trail. Plantations were a three-species polycluture but all sampling within them took place only in stands of Blackbutt (Eucalyptus pilularis).

The forest matrix adjacent to plantations in the study area was closed woodland dominated by Blackbutt (Eucalyptus pilularis), and containing a mixture of Tallowood (E. microcorys) and White mahogany (E. acmenoides), typical of the region. A diverse subcanopy layer was present in places, and the ground layer was patchily grassy and dominated by Blady (Imperata cylindrica) and Basket (Lomandra longifolia) grasses. Fire had been excluded from all field sites sampled in native forest for the eight years since plantation establishment, as managers prioritized fire control in native forest due to the proximity of plantations and the forest matrix (pers. comm., P. Levitske, SF Wauchope).

However fire scars on tree trunks and the abundance of stumps suggested that forests had experienced at least one medium-intensity fire before 1996 when plantations were planted. The forest matrix was grazed at low stocking density

111 Chapter 5: Matrix quality and ground fauna

for at least 40 years prior to the establishment of plantations (pers. comm., P.

Levitske).

Sites were selected to maximize distance apart within the plantations

(~1km), to standardize aspect, and to minimize distance between plantations and the forest matrix. Each plantation site was paired with an adjacent matrix site. All sites were situated 100 m from edges with cleared pasture as there is a significant edge effect at the pasture-plantation boundary that persists for up to

50m (Chen et al. 1993, Wright et al. 2010). Five sites in each forest type were sampled for fauna and vegetation structure. Sites were considered as true replicates as other studies have shown that the small mammals sampled in this research typically move less than 1km (Pires et al. 2002).

Vegetation structure

Vegetation structure was quantified in young plantations and adjacent native forests by measuring characteristics from sixteen 20m2 quadrats along four haphazardly placed transects spaced 150m apart, in each of five 200x600 sites. Vegetation variables were a subset of those measured in chapter 2 with the potential to influence habitat use by ground fauna. Quadrats were spaced

70m apart along transects. Four quadrats from four transects in each of five sites resulted in a total of 80 quadrats per forest type. All sampling was conducted in October 2003. The vegetation characteristics measured were known to influence habitat use in ground fauna (Table 1).

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Table 1. Vegetation variables measured to quantify structural differences between grazed native forest and young native plantation forest types on the Mid north coast of NSW. Sampling effort was 80 quadrats from five replicate sites in forests and adjacent young plantations. Paired t-tests identify significant differences between matrix and young plantation for each vegetation variable.

Variable measured Definition Measurement Method Whole environment characters number of trees Trees are defined as any vegetation taller than 5m high count number of saplings count count vertical complexity an estimate of how much vertical space between the ground and the categorical McDonald et al. 1990 canopy is occupied by foliage crown height height to the canopy top but excluding emergent trees height range finder canopy cover amount of horizontal space covered by canopy foliage percentage standardised photographs# midstorey height height to the midstorey where present but excluding shrubs and subcanopy height range finder Ground level characters trunk cover the amount of the quadrat coverd by tree truks of gretaer than 15cm diameter percentage understorey height height of any vegetation up to 1.5m high height ruler shrub cover shruby growth forms only percentage visual estimate grass cover percentage visual estimate sedge cover sedeg were primarily Lomandra longifolia which formed patchy clumps percentage visual estimate litter cover percentage visual estimate litter depth depth of leaves from the soil to the top of the leaf pack depth ruler (cms) broadleaf cover an estimate of the surface cover of fallen broad leaves from any species. percentage visual estimate Broad leaves were ≥4cm in diameter. rock cover percentage visual estimate bare ground percentage visual estimate log cover* the total percentage of a quadrat covered by logs percentage visual estimate log decomposition a count of any holes/crevices in a log of >5cm long and 2cm wide count modified from Fletcher (1977) number of stumps included to quantify the availability of vertical perches/basking points count presence of retreats including sedge baskets, holes >3cm diameter in logs/ground/stumps count or log cracks >10mm wide and 5mm deep *Logs were defined as any dead wood on the ground surface with diameter >5cm and length >15cm # Standardised photograhps of canopy cover classes are provided in McDonald et al. 1990.

The percent cover of broad leaves and sedges were included as a specific character of habitat for ground fauna as broader leaves can be specifically attractive to thermophilic lizards (e.g. Valentine et al. 2007), and clumps of sedges represented complex retreat sites for small species where the ground was sparsely grassed. Litter depth was included because deep leaf litter packs are necessary for daytime retreats and breeding in direct developing frogs such as the Red-backed Toadlet (Pseudophyrne coriacea).

Fauna Trapping

Trapping included the use of standard procedures for sampling ground fauna detailed in chapter three. Pitfall trapping was conducted over two years between October 2002 - February 2003 and October 2003 - February 2004.

113 Chapter 5: Matrix quality and ground fauna

Live trapping was conducted from October 2003 - February 2004 in conjunction with pitfall trapping. Pitfall trapping data from 2002-03 was a subset of data analyzed in chapter 4, whilst live trapping data was independent of any other trapping data presented in this thesis. Young and matrix forest types were sampled by thirty live traps (Elliott traps, type “A”) laid in a 375 x 400 m (traps 50 m apart) grid in each site. Traps were opened between 16:00 and 08:00EST for five consecutive nights, resulting in a trapping effort of 750 trap nights/forest type. Traps were baited with a mixture of rolled oats, peanut butter, honey, vanilla essence and salami. All animals captured were released at the trap site after paint marking the bottoms of feet to identify recaptures. Recaptures were excluded from data analyses.

Pitfall trapping arrays were established as per the methods described in chapter three. Traps were initially opened for five day/nights, and re-opened at four and eight weeks later for a second and third five day/night trapping period in each site. Three arrays totaling 15 traps, for each of five sites, opened for 15 nights resulted in a total of 1125 trap nights per forest type. Traps were cleared once per day in the mornings and, after measuring and individually paint marking mammals all animals were released at the site of capture. Skinks were uniquely identified by tail-regenerated tail and snout-vent length measurements, and did not require marking. Detailed biometrics were recorded for mammals to identify any recaptures between sampling periods within sites, and avoid overestimating abundance. Although the time over which sites were trapped was somewhat limited, it was considered adequate to estimate the distribution and

114 Chapter 5: Matrix quality and ground fauna

relative abundance and age structure of populations during the spring-summer activity period for the most common herpetofaunal and small mammal species in the study area, and encompassed the times in which juvenile herpetofauna and mammals were recruiting into habitats

Data Analysis

Vegetation

A paired t-test was used to identify any differences in individual vegetation characters between matrix and plantation forest types. Prior to analysis data was averaged over quadrats to create site-level means, and for nominal variables the modal value was applied.

Fauna

Fauna were classified as either generalist or specialist on the basis of strong preferences for diet or specific habitat features (defined in Chapter 4).

While this is a coarse predictor of expected habitat use, it can provide some indication of the negative response of a species to habitat alteration, as many studies identify that abundance of generalists increases in modified landscapes

(e.g. Devictor et al. 2008). Individuals were allocated into either adult or juvenile age classes based on morphometrics. Adult mammals were identified by pes length and head width as per published records, or by reproductive condition.

Adult reptiles were defined as those having a snout-vent length of at least four fifths of the mean snouth-vent length for each species (using an average of all

115 Chapter 5: Matrix quality and ground fauna

trapping records for that species throughout the study). This means that the juvenile category included some sub-adults, which for the purposes of this chapter are discussed as juvenile. Herpetofauna were analyzed as a group because given the small body sizes and cryptic habitats of many of the snake and lizard species trapped, their dispersal ability was assumed to approximate that of frogs. Mammals were analyzed separately. Juvenile and adult captures were pooled for all analyses except those focused specifically on age effects..

Paired samples t-tests were used to compare the abundance and species richness of small mammals and herpetofauna among forest types using SPSS v21.0 (SPSS 2012). Before analysis data were pooled to site level and log (x+1) transformed, to satisfy assumptions and homogenize variance among forest types. Two-way Analysis of Variance (ANOVA) was used to test whether the abundance of species with specific habitat preferences differed between forest types, and repeated to test whether adults and juveniles were significantly more abundant in plantations and matrix habitats. Bonferroni correction was applied to all estimates of significance to reduce error rates associated with multiple analyses.

Results Vegetation structure

Vegetation structure differed significantly between young plantations and adjacent native forests. Native forests had greater leaf litter cover with deep leaf pack, a taller understorey, more saplings and shrubs, and higher log cover with logs that contained retreats. These were almost completely lacking in

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plantations. Whilst plantations contained higher tree density, the percentage of a quadrat covered by tree trunks was higher in forests due to greater tree girth.

Plantations had a greater cover of rocks and bare ground than forests, but both habitats had similar grass cover. Whilst broad leaf cover did not differ between forests and plantations in forests leaves were native, whilst in plantations they originated primarily from exotic Lantana camara (pers obs).

Table 2. Paired t-tests for differences between grazed native forest and young plantation vegetation variables measured from 80 quadrats in five replicate sites on the Mid North Coast of NSW. Significant values are denoted in bold.

Mean Variable measured Matrix Plantation df t p Whole environment characters number of trees 6.267 13.475 4 -11.088 >0.001 number of saplings 8.636 0.775 4 4.505 0.011 vertical complexity 0.200 1.013 4 1.137 0.319 crown height 31.592 10.000 4 21.913 >0.001 canopy cover 43.333 17.713 4 11.186 0.006 midstorey height 13.962 0.024 5.349 >0.001 Ground level characters trunk cover 3.933 1.763 4 2.908 0.044 understorey height 39.844 4.938 4 4.177 0.014 shrub cover 2.615 15.187 4 -3.993 0.022 grass cover 66.572 94.200 4 -2.159 0.097 sedge cover 0.894 0.000 4 22.011 >0.001 litter cover 51.244 5.750 4 -4.124 0.013 litter depth 5.113 2.264 4 3.622 0.022 broadleaf cover 21.125 0.525 4 2.238 0.089 rock cover 0.350 1.138 4 -7.188 0.002 bare ground 0.975 15.188 4 -1.297 0.264 log cover 8.950 0.013 4 7.024 0.002 log decomposition 3.011 0.213 4 12.033 >0.001 number of stumps 0.789 0.000 4 4.105 0.015 presence of retreats 0.942 0.000 4 25.924 >0.001

Fauna In total 475 individuals from 26 species, from seven mammal, four frog and 15 lizard species were captured (Appendix 3). Young plantations and adjacent native forests each supported 20 species. Woodland-associated fauna

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comprised 35.2% of capture, whilst forest and generalist species comprised 13.3 and 51.4% respectively. Despite large differences in vegetation structure between the forest matrix and young plantations, mammal and herpetofaunal abundance and species richness was similar in both forest types (Table 3).

Table 3. Summaries for paired t-tests examining differences in ground faunal species richness and abundance from native eucalypt woodland and adjacent native plantations on the Mid north coast of NSW. Sampling effort was 750/1125 trap nights for live/pitfall trapping from five replicate sites.

Mean Variable measured Matrix Plantation SD df t p mammal abundance 46.6 27 41.789 1.049 4 0.353 mammal richness 9.8 11.6 17.021 -0.236 4 0.825 herpetofaunal abundance 8.8 6.2 4.037 1.44 4 0.223 herpetofaunal richness 1.6 2.6 1.732 -1.291 4 0.266

However forest mammals and herpetofauna were significantly less abundant in young plantations than matrix forests (herp: F2,24 = 9.200 p = 0.002, mammals:

F2,15 = 12.485 p = 0.001, Fig. 1a). House mice (Mus musculus), Swamp Rats

(Rattus lutreolus), Common Dunnarts (Sminthopsis murina), Jacky Dragons

(Amphibolurus muricatus) and Robust Striped Skinks (Ctenotus robustus) replaced Bush rats (Rattus fuscipes), Red-tailed Calyptotis (Calyptotis ruficaudata) and Red–backed toadlets (Pseudophryne coriacea) (Appendix 3).

Adult mammals and herpetofuana were significantly more abundant than juveniles, but the abundance of juveniles did not change in matrix forests and plantations (Table 3, Fig. 1b).

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Fig.2 Mean abundance of (A) forest, woodland and generalist species and (B) of adults and juvenile mammals and herpetofauna sampled from native forest matrix and adjacent young plantation forest types from five replicates sites on the Mid North Coast of NSW. Error bars are 95%CI.

119 Chapter 5: Matrix quality and ground fauna

Discussion

In this study in coastal New South Wales grazed native forests adjoining grazed young eucalyptus plantations supported equally rich and abundant populations of ground fauna. Whilst generalist species dominated plantations, the species richness of ground fauna identified in this study approximated that of old growth native forests in the region (Chapter 3). The first hypothesis posited in this chapter suggested that equivalent species richness in the matrix and young plantations may indicate poor quality of the matrix habitat. However, the high species richness of ground fauna in in both forest types and the presence of abundant forest-associated species, identifies that both that young plantations and grazed native forest matrices act as habitat in the study area.

Matrix quality

The effects of grazing on plantations

Pastures are known to rarely support reptiles (Driscoll 2004, Kavanagh et al. 2005, Kanowski et al. 2006) or small mammals (Downes et al. 1997), and in the study area are documented as lacking small mammals (Law et al. 2002) .

Thus, if pastures provide an effective barrier for small mammals, faunal assemblages within plantations adjacent to pastures and forests are likely to originate from immigration through the forest matrix. However, in this study plantation herpetofaunal assemblages were not subsets of the forest assemblage, but included the heliothermic Striped Skinks (Ctenotus robustus) and Jacky Dragons (Amphibolurus muricatus), which were not trapped in the

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forest matrix. Whilst this result may arise from the snapshot nature of sampling in this study, it could also suggest that grazing in the forest matrix can increase the permeability of the matrix for woodland species by creating open spaces in the understorey (e.g. Yates et al. 2000). Without a direct comparison of faunal diversity in pastures, plantations and forests, my explanation for high abundances of woodland species in plantations but not matrix habitats, must remain hypothetical.

The effect of grazing in the forest matrix on mammals is less clear.

Chapters three and four of this thesis consistently identified young plantations as supporting < 50% of the mammal abundance of undisturbed forests, yet in this study plantation and matrix mammal abundance is similar. This study records that the matrix supports the lowest mammal abundance recorded in three years of trapping. Whether such low abundance results from reduced matrix quality or is related to decreasing rainfall during the course of this study is uncertain. Even if matrix quality was unaffected by grazing, this may not have resulted in species gains in adjacent plantations, as young plantations simply did not embody enough forest-like features to be attractive to other forest mammal species known from the regional pool. Combined, these results indicate that mammal and herpetofaunal responses to habitat simplification through grazing can vary considerably. If a cessation of grazing in the matrix resulted in an increase in the structural heterogeneity of forests (e.g. James 2003) this may decrease matrix permeability for lizards by physically obstructing dispersal (Prevedello et al.

2010), or by altering the availability of thermal resources which may have knock-

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on negative consequences for reptile fauna. However, if the lack of transfer of mammal species between forests and young plantations is typical for mammals, in woodland environments whilst a cessation of grazing in the matrix might benefit local diversity, it would be unlikely to affect plantation diversity. Clarifying the role of grazing and its interaction with species transfer between forests and plantations will require further study, and may benefit from a case-by-case basis as has been stressed previously by other authors (Fischer et al. 2004, Felton et al. 2010).

Age structure

The age structure of mammal and reptile populations in plantations were not skewed towards juveniles relative to those in the native forest matrix, suggesting that species abundance in young plantations was unlikely to have been be maintained by recruitment. However, the alternative hypothesis that reptiles reproduce well in plantations but persist in a strongly adult-biased population due to a loss of young through predation or emigration is equally plausible, although to support this hypothesis, trapping would have been expected to reflect higher numbers of juveniles. Both ideas hinge on the effectiveness of sampling for juveniles in this study. While sampling took place at the time when new recruits should be evident in populations, relatively poor capture rates for juveniles of both faunal groups suggests either that standard sampling methods for ground fauna are biased towards adults, or that naive recruits really were simply experiencing high predation pressure in young

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plantations where the availability of retreats was much lower. A targeted study focusing on the recruitment of reptiles through the plantation-matrix boundary and particularly on the persistence of reptiles in young plantations after recruitment is necessary to clarify what drives such age-biases.

Conclusions

The grazed forest matrix adjacent to young native plantations in this study had the potential to represent different habitat quality for herpetofauna and mammals. In this study the grazed native forest matrix had some potential to influence the diversity of herpetofauna in young native plantations adjacent to, and connected with, the matrix by acting as a source of recruits and a permeable conduit for reptiles (e.g. Kupfer et al. 2006). The quality of this matrix for ground mammals was more ambiguous. In contrast to reptiles, while the abundances of mammals were equivalent in the matrix and young plantations, they were significantly lower than those supported by ungrazed native forests sampled in other parts of the study area (Chapters 3 + 4). Until the role of grazing in the forest matrix and its effect on the recruitment of ground fauna into plantations is understood, managers should consider situating new plantations in the landscape mosaic in proximity to grazed and ungrazed forests. Where management targets vulnerable species in the region like the Common

Planigale (Planigale mauclata), which uses plantations and ungrazed native forests (Chapter 3) but is absent from grazed forests (this chapter), grazed matrices may be less valuable as a source of recruits. Future research efforts

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that focus on understanding immigration and residency in plantation fauna and on their relationship with surrounding agricultural matrices, will clarify the role of native matrices in contributing native plantation biodiversity.

124 Chapter 6: Lizard habitat choice

Chapter Six

The roles of structural and thermal factors in determining habitat choice by lizards in young plantations

Introduction

Understanding how habitat choice affects local species distribution and abundance is one of the factors necessary for the development of effective management plans for biodiversity conservation. However, knowledge of the ecological requirements and habitat relationships of many species, especially reptiles (Michael et al. 2010), is often poor. Reptiles are in decline globally (Gibbons et al. 2000), and as poor dispersers they are vulnerable to landscape scale changes (Gardner et al. 2007b, Brown et al.

2008) which change vegetation structure and thus thermal habitats. The availability of appropriate microhabitats with suitable thermal ranges has been long accepted as an important habitat selection cue for reptiles (e.g.

Grant & Dunham 1988), because available temperatures constrain activity

(Magnuson et al. 1979, Tracy & Christian 1986, Dunham et al. 1989, Díaz

1997). Whilst a growing body of research attempts to identify how reptile habitat selection changes in human-modified landscapes (e.g. Row & Blouin-

Demers 2006, Todd & Andrews 2008, Wanger et al. 2009) very few studies have examined the thermal properties of plantation habitats and their impacts on reptiles (e.g. Becker et al. 2007, Mott et al. 2010).

125 Chapter 6: Lizard habitat choice

Thermal environment and habitat choice

An animal’s thermal environment is the result of complex interactions between many physical characteristics of a specific location, which include the amount and direction of solar radiation, air and substrate temperatures, and convective characteristics. Because of the dominant influence of solar radiation on diurnal microclimates, complex vegetation can markedly influence the thermal potential of a given habitat by limiting the amount of direct sunlight and providing a greater range of convective and radiative conditions than occur in more open habitats. The importance of specific microclimates as habitat selection cues is well recognized for both diurnal and nocturnal ectotherms (e.g. fish – Wehrly et al. 2003, Ward et al. 2010; invertebrates –Pereboom and Beismeijer 2003, Ahnesjö & Forsman 2006; herpetofauna – Cowles & Bogert 1944, Tracy & Christian 1986, Adolph

1990, Webb & Shine 1998, Sabo 2003, Goldsborough et al. 2006, Row &

Blouin-Demers 2006, Bancroft et al. 2008, Hoare et al. 2009). In addition to the thermal influence of daytime light levels on microclimate, light may also affect habitat choice through its effects on visual systems and associated communication and predator avoidance (Carrascal et al. 2001, Leal &

Fleishman 2004).

Previous studies investigating the roles of microclimate and structure on animal habitat selection have inferred thermal environments from light penetration through tree canopies (Pringle et al. 2003, Langkilde et al. 2003) or have used qualitative measures of structural complexity (Schultz 1998,

Skelly et al. 1999, Halverson et al. 2003, Pineda & Halffter 2004). Such measurements are biased to visible wavelengths, and consequently ignore

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ultraviolet and infrared components of the light spectrum. However, non- visible wavelengths of light are known to influence habitat selection in birds, mammals and reptiles (Walsberg et al. 1997, Carrascal et al. 2001, Huey

1991, Christian et al. 1996), and are important for foraging and social behaviours in many birds, reptiles and amphibians (e.g. Fleishman et al.

1997, Honkavaara et al. 2002). Infrared wavelengths are particularly important for behavioural thermoregulation in heliothermic reptiles (Christian et al. 1996, Belliure & Carrascal 2002) as this part of solar radiation constitutes up to half of the energy received at the ground during the day

(Brown & Gillespie 1995, Kotzen 2003). Because of the highly variable nature of vegetation in modifying particular wavelengths of sunlight and so changing the thermal and photic characteristics of a particular microhabitat

(e.g. Yang et al. 1999, Dölle & Schmidt 2009), it is very difficult to predict the consequences of vegetation restructuring on biodiversity. Such predictions require an understanding of how resident animal species respond to changes in irradiance and habitat complexity as well as how changes to vegetation structure affect these interactions.

Thermal physiology of reptiles

Temperature is known to strongly affect physiological processes associated with energy gain and expenditure (e.g. Harlow et al. 1976, Huey

1982, Alexander et al. 2001, Pafilis et al. 2007). While moderate to high body temperatures can maximize performance (Huey & Kingsolver 1989) including behaviours such as predatory efficiency (Díaz 1995) and rates of digestive processes (e.g. McConnachie & Alexander 2004), the actual body temperature required for optimization of particular functions varies markedly

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between species, and often reflects their natal environment (e.g. Bilcke et al.

2006, Aubret & Shine 2010). Consequently, the extent to which animals can exploit new environments depends on their behavioural capacity to attain optimal body temperatures in the new habitat, or their potential to adjust thermal optima for physiological functions through acclimation (Van Damme et al. 1990). Many ectotherms behaviourally thermoregulate to attain a limited range of preferred body temperature ranges. Maintaining preferred temperatures increases fitness by optimizing growth (Sinervo & Adolph

1994), performance (e.g. Huey & Kingsolver 1993, Downes & Shine 1998,

Pitt 1999) and reproduction (Shine 2005). By contrast, habitat generalists that occupy a broader range of thermal environments may trade off performance optima at particular temperatures, for lower performance over a broader range of temperatures.

Constraints of plantations environments on lizards

Whilst recognition of the conservation value of native plantations for fauna is increasing (Brockerhoff et al. 2008) most studies have examined the structural complexity of vegetation (see Brockerhoff et al. 2008 and

Hartmann et al. 2010 for reviews), plantation size and landscape context

(Kavanagh et al. 2001) as drivers of change in diversity between hardwood plantations and native forests. The role of thermal and light environments and their influence on habitat choice in native plantations remains largely unexplored. Monoculture tree plantations provide an excellent opportunity to address these questions about the role of thermal environments as their uniformity in tree species, size and distribution can generate specific thermal

128 Chapter 6: Lizard habitat choice

microhabitats, and these microhabitats change in predictable directions as plantations age. Given that there are recognized differences in the attractiveness of plantations of different ages to fauna (e.g. Borsboom et al.

2002, Hobbs et al. 2003) and plantation management goals often include increasing faunal diversity, differentiating thermal from structural habitat cues may provide a key to understanding plantation faunal diversity.

In artificial environments such as managed plantations, where planting structure facilitates early canopy closure, the relationship between light, heat and vegetation structure is not always linear. As a consequence, high canopy density, low light and cool environmental temperatures can be linked to low, rather than high, understorey structural complexity (Chapter 2). This has ramifications for biodiversity within plantations if thermal and structural habitat selection cues differ in importance among animal species. The extent of this effect will depend on the behavioural and physiological flexibility of each species, which ultimately determines whether a species can accept thermal conditions outside of the preferred temperature range (see Heatwole

1977, Huey 1982, Heatwole & Taylor 1987, Blouin-Demers & Nadeau 2005 and Vickers et al. 2011 for more discussion).

This thesis has identified that native tree plantations on the Mid North

Coast of New South Wales had more open canopies, more simply structured sub-canopy vegetation, received higher insolation at the ground, and attained higher and more variable diurnal environmental temperatures than native forests (Chapter 2). Lizard assemblages responded to these shifts in available microhabitat. The hot, simple plantation environments supported

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distinct assemblages dominated by small-bodied open forest-woodland and generalist species and heliotherms (Chapter 3), a shift previously identified between plantation and forest environments for tropical reptiles in exotic plantations (Gardner et al. 2007a) and temperate reptiles in native plantations (Ryan et al. 2002, Kanowski et al. 2006). Young native plantations conspicuously lacked forest specialist herpetofauna, and large- bodied species were rare despite an abundance of palatable invertebrate prey (Chapter 5).

This chapter aimed to identify whether forest specialist lizards were absent from young native plantations due to physiological constraints associated with the thermal environment in young plantations, or due to behavioural constraints driven by preferences for specific structural attributes of the environment unavailable in plantations. To apply these ideas the physiological consequences of habitat choice on forest specialist, generalist and plantation-associated lizards were examined experimentally. Appetite and gut passage rates and habitat selection preferences were compared between species at temperatures common to young plantations at the peak of lizard activity in summer.

I hypothesized that:

1. generalist species occurring in both plantations and forests would have a broader preferred temperature range than habitat specialists

2. because coadaptation amongst traits that strongly affect fitness should be favoured (Mayr, 1963) specialist species would choose habitat based on a preference for substrate type, and would thermoregulate narrowly

130 Chapter 6: Lizard habitat choice

3. because preferred temperature maximizes enzyme function (Harlow et al. 1976, Hochacha & Somero 1984), gut passage rate would be slower in specialist species in temperatures common to habitats outside their own, where preferred temperatures would be harder for lizards to achieve.

Methods

Study System

Due to the influence of phylogeny on preferred body temperature

(Garland et al. 1991), I restricted species for comparisons to diurnally active skinks from the sub-family Lygosominae. All species were morphologically similar, but differed somewhat in body mass and limb:body ratio, and all were carnivorous, eating a typical scincid diet of spiders, coleopterans, insect larvae, gastropods and ants, lizards (Wilson & Knowles 1988). The species used in experiments in this chapter were those identified previously as being habitat restricted (Chapter 3). They included three diurnal, surface-active species the plantation-restricted Adrasteia amicula, the old growth native specialist Eulamprus murrayi and the wide-ranging generalist Lampropholis guichenoti ; and two cryptozoic species, the forest-associated Calyptotis ruficaudata, and the woodland-associated Lerista muelleri. The species in order of size were Eulamprus murrayi (SVL: 87mm, mass: 15.70 ± 3.12g),

Lerista muelleri (SVL: 56mm, mass: 1.66 ± 0.40g), Calyptotis ruficaudata

(SVL: 42mm, mass: 1.16 ± 0.30g), Lampropholis guichenoti (SVL: 35mm, mass: 0.88 ± 0.24g) and Adrasteia amicula (SVL: 15 mm, mass: 0.52 ±

0.26g). All of these species are heliothermic (use the sun as a heat source),

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but E. murrayi is a sedentary a posturing heliotherm and regulates body temperature by altering the preferred angle of the body to the sun

(Hutchinson 1993). Whilst C. ruficaudata and L. muelleri are cryptozoic species, they were included in this study because retreat choice in cryptozoic species is influenced by thermal characteristics (e.g. Shah et al. 2004) and preliminary field observations identified surface basking for both species. L. guichenoti and A. amicula are shuttling heliotherms, and control their body temperature almost entirely by actively moving between sun and shade.

Habitat choice enclosures

To separate behavioural preferences based on structure from those of thermal microclimate, four enclosures with particle-board walls were built in the central part of the study area near Wauchope on the Mid North Coast of

New South Wales. These were placed in an open paddock in cleared farmland that was unshaded by trees. Enclosures were built to expose lizards to a series of specific thermal and structural microenvironments. They offered a choice of simple and complex substrates, built to mimic those available in plantations or native forests, along a gradient of solar irradiance. Three large enclosures (7.5 x 3 m, Fig 1a), partitioned internally into five 1.5 x 1.5 x 1m compartments, and connected to the two adjacent by 15 x 15cm openings

(Figure 1a). Enclosures were oriented along the solar path to minimize internal shading by the partition walls, and while a small amount of shading occurred, this area was consistent among treatments within each box. A solar energy gradient was generated by roofing each compartment with shade cloth which allowed 96%, 70%, 50% and 10% of incident sunlight respectively (Fig 1b), based on readings from a calibrated pyranometer

132 Chapter 6: Lizard habitat choice

(model LP 02L, Campbell Scientific). Monofilament bird netting allowing 100% radiant energy penetration was placed over the unshaded treatment, to prevent predation and standardize disturbance levels when checking the position of animals. A fourth and much smaller enclosure (2.8 x 0.9m) was divided into ten compartments (56 x 45 x 50cm) to examine habitat choice in

Adrasteia amicula because of its much smaller adult size (0.5g). Substrate types were the same in all enclosures,

One of each pair of compartments at each level of irradiance was randomly assigned a substrate, either ‘complex’: representing the complex ground layer common to forests and containing a complete covering of forest leaf litter, five logs with a minimum diameter of 15cm and a log pile of three additional logs, or ‘simple’: a ground layer common to plantations mimicked using six vertically positioned bunches of dry native grass collected in the study area (Fig 1c), built to approximate the tussocks of Blady grass

(Imperata cylindrica) common to plantations in the study area. Up to 60% of the substrate floor in the simple treatment was bare, in accordance with the conditions common to plantations in the study area, but in both choice boxes

10% of the grass or logs were placed against the shadowed wall of the box, to allow lizard a choice of sheltered temperature ranges. Prior to applying substrates, enclosures were floored with weed matting to prevent animals escaping and were covered with a layer of mixed river sand and local soil

~8cm deep. Air temperatures did not vary significantly between compartments of the enclosure as a result of the access ports between compartments. These ports were large enough to permit lizards to readily view and assess adjacent compartments.

133 Chapter 6: Lizard habitat choice

C S S C C

S C C S S

100 90 50 30 4 % Radiant energy penetration

Fig. 1. Choice enclosures in situ on the Mid North Coast of NSW. A. Large enclosures; B. Diagramattic representation of the relative amount of ambient radiant energy and substrate present in each compartment, (c = complex, s = simple). C. An example of the complex (left) and simple (right) substrates lizards could select.

Operative environmental temperatures (Tenv; Bakken 1992) within compartments were measured using iButton data loggers (model DS1921L,

Dallas Semiconductors). These were plastic wrapped and painted with neutral-coloured paint (following Walsberg & Weathers 1986) to closely match lizard coloration. Data loggers were placed in the hottest (full sun) and coolest (full shade) microenvironments in both substrate treatments in each

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compartment of all enclosures. Prior to introduction of lizards, preliminary tests confirmed that small and large boxes and complex and simple substrates sharing the same shade treatment did not differ in levels of insolation received or in Tenv through the diel cycle. To quantify the relationship between insolation and body temperatures (Tb) in experimental enclosures, insolation readings were collected hourly from each box each day of sampling using a pyranometer (LP 02L, Campbell Scientific) that measured a combination of ultraviolet, near-infrared and incident wavelengths of light.

Sampling of young plantation thermal environments was conducted in conjunction with preferred temperature experiments to provide an estimate of available temperature in young plantations, to compare with experimentally determined preferred temperature ranges. Environmental temperature (Thab) in young plantations was sampled in shaded and unshaded sites using iButton data loggers (model DS1921L, Dallas Semiconductors). Preparation, replication and placement of these data loggers followed methods described in chapter two. Thab measurements indicated the range of temperatures available to lizards in young plantations at the appropriate spatial scale. Field

Thab was recorded from September 2002 – March 2003 (spring - summer), corresponding to the period of sampling in the field enclosures, and maximal activity in lizards. To verify that environmental temperatures (Tenv) available in the enclosures fell within the range of those available in the wild, iButtons were also placed in sun and shade microhabitats within enclosures at each level of shading.

135 Chapter 6: Lizard habitat choice

Scincid lizards were hand captured by active searching and pitfall trapping in the forest types described in chapter two during October and

November 2002 and January 2003. A minimum of fifteen individuals of each species were sampled (Table 1). Lizards were introduced singly into choice boxes within two hours of capture and allowed to familiarize themselves with the enclosure for 24 hours. Several mealworms (Tenebrionid sp.) and eight commercially available crickets (Gryllus sp.) of appropriate size were provided ad libitum in each box as was water via a dish and twice-daily spraying of the enclosure walls. All species of lizards were observed using these resources during the period of familiarization. Naturally occurring prey

(ants, moths, flies, beetles) also entered enclosures and were eaten by lizards.

Sampling involved scoring each individual for position, substrate type and behaviour every 30 minutes between 0800h and 1700h, times associated with most daily activity. Specific behaviours were recorded, and were subsequently divided and then pooled into ‘active’ and ‘inactive’ categories for analysis. To score ‘active’ a lizard was basking, foraging, fleeing, exploring or retreated under shelter but moving and alert. To score

‘inactive’ a lizard was buried in the substrate, or retreated under shelter but not alert. Cryptozoic species were considered active even when retreated under logs, but were scored as inactive when eyelids were closed or when they were buried in the substrate. Cryptozoic species were located by tracks and pinpointed by uncovering only enough of a retreat to allow sighting of an individual, after a pilot study verified that this level of disturbance did not cause them to change position. In addition, lizards were captured every 1.5

136 Chapter 6: Lizard habitat choice

hours, and external groin temperature (Tb) was recorded using a calibrated quick reading Schultheis thermometer within five seconds of capture, as an infrared thermometer could not differentiate the small size of the lizards against the substrate to allow measurement. Groin temperature was measured by pressing the thermometer bulb against the junction of the hindlimb and abdomen. This method was previously established to closely approximate cloacal temperature and to be much quicker and less stressful for scincids than repeatedly sampling cloacally. Body temperatures were not recorded for any lizard that fled for more than 15cm from the point of first observation, to avoid any temperature bias associated with sprinting or forced change of microhabitat. After sampling all lizards were returned to point of capture within the enclosure, and substrate temperature at the site of capture was recorded. All experimental trials were conducted on sunny days.

All lizards were returned to their original capture site within 24 hours of cessation of the enclosure experiments, and each lizard was tested only once.

Data Analysis

To identify how insolation affected microclimate throughout the day linear mixed-effects repeated measures models were used. This type of model better handles correlated data and unequal variances than general linear models associated with analysis of variance, by accounting for residual errors which are correlated within each subject, but independent across subjects (see Verbeke & Molenbergs 2000 and Demidenko 2004 for more discussion of these models). To identify differences in Tenv between shading levels, time and position in the gradient (fixed factors) and data logger

137 Chapter 6: Lizard habitat choice

(repeated factor) were analyzed using log transformed data. This analysis was repeated using the same fixed factors for insolation data but replacing data logger with day as the repeated factor. To identify any change in mean

Tb within time of day a third mixed model replacing day with individual identity was analyzed using log transformed data, separately for each lizard species, as no transformation could normalize the data for a species-combined data set. Because this test identified a consistent diel rhythm in mean Tb typical of diurnal species (Heatwole 1987), a two-way analysis of variance (ANOVA) testing for differences in Tb between species included time as a co-variate.

All test assumptions were satisfied before analysis. For graphical purposes the preferred temperature range was defined as the central 68% of Tb recorded in enclosures. This range is considered to represent the typical Tb that a species prefers during activity (Huey 1982). Substrate preferences were described qualitatively as choice was highly skewed with all species preferring complex substrates more than 90% of the time, and three of the five species only using complex substrates. For all models tested an alpha level of p< 0.05 was considered significant and Type III sums of squares were used. Analyses were performed using SPSS version 16 (SPSS 2007).

Results

The insolation received at the ground varied with time of day and level of shading (shade*time - F40,151.1 = 6.475, p < 0.001) as did the associated

Tenv (shade*time - F60,13543.0 = 116.666, p < 0.001). Whilst shade treatments 0 and 10 consistently received higher insolation throughout the day (Fig. 2a), they were only hotter in the morning, with Tenv equalized across all levels of shading after 1100h EST (Fig. 2b). Despite these fluctuations lizards

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thermoregulated carefully throughout the day. After heating to their preferred temperatures between 0800h and 0900h EST all species maintained mean

0 Tb of 29 – 30 ± 1.5 - 2.5 C (Table 1, Fig. 2c) apart from the forest specialist

0 Eulamprus murrayi which maintained a significantly cooler Tb of 24 ± 3.5 C throughout the day (F4,376 = 52.055, p<0.001; Fig. 2c).

Whilst Tb fluctuated with time of day for all species it was independent of location alone (Table 2), and each species used location in the gradient differently according to their behavioural mode. Lampropholis guichenoti and

Adrasteia amicula were the most active species while the cryptozoic woodland and forest-associated Calyptotis ruficaudata were relatively inactive, and the forest-specialist Eulamprus murrayi fell in between these groups.

Table 1. Body temperatures and habitat associations of target species in experimental choice boxes on the Mid North Coast of New South Wales. Mean temp is pooled over positions in enclosures and excludes the first temperature recording between 8 and 9am where lizards were heating to preferred temperature. Range encompasses the central 68% of the body temperatures measured.

Mean Behavioural Species n Temp SD Occurrence Mode L. guichenoti 23 29.71 2.54 all habitats shuttling heliotherm A. amicula 19 29.96 1.73 young plantation shuttling heliotherm L. mulleri 15 30.49 2.01 forests, old plantation cryptozoic C. ruficaudata 17 30.78 1.6 forests, old plantation cryptozoic E. murrayi 18 24.41 3.48 old growth forest posturing heliotherm

Two distinct activity patterns emerged with the shuttling heliotherms

Lampropholis guichenoti and Adrasteia amicula being active throughout the day, and using the less shaded end of the gradient preferentially for the first three hours of measurement. This behaviour occurred even though temperatures in the gradient at locations 0 and 10% shade exceeded mean preferred temperature range for all species between 0800 and 0110h (Fig.

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2b). In the afternoons this pattern switched and A. amicula became less active and was associated with locations having the least irradiance. The more sedentary woodland Lerista muelleri and forest–associated Eulamprus murrayi and Calyptotis ruficaudata were consistently recorded from the more shaded end of the gradient, but differed in activity patterns. Both L. muelleri and C. ruficaudata were unimodal in activity pattern and were active until

11am, and whilst they were both inactive for the latter part of the day, C. ruficaudata used all positions between 50-96% shade, while L. muelleri occupied retreats primarily in 96% shade. E. murrayi fell between these two groups and while it showed sporadic activity throughout the day, it was restricted almost exclusively to 96% shade to maintain its low preferred temperature.

All lizards strongly preferred complex to simple substrates in the field enclosures (percentage of use of complex substrate for each species: guichenoti = 95%, amicula = 92%, muelleri = 100 %, ruficaudata = 97% murrayi = 99% - from 1077 observations of substrate choice in total), however, whilst L. guichenoti and A. amicula the two species using plantations were expected use simple substrates, the forest-associated C. ruficaudata also tolerated simple substrates whilst inactive.

Table 2. Results of mixed model analyses for change in body temperature with position in experimental choice boxes. TOD = time of day, Loc = position in the gradient corresponding to level of insolation. Significant values are in bold.

Species Factor df F p L. guichenoti Loc*TOD 19,57.0 2.543 0.003

A. amicula Loc*TOD 10,30.4 1.819 0.049

L. mulleri Loc*TOD 5,14.3 7.288 0.001

C. ruficaudata Loc*TOD 10,30.4 3.166 0.007

E. murrayi Loc*TOD 7,34.8 4.575 0.001

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- 2) - 2)

Insolation (Wm Insolation (Wm

16 16 15 14 11 10 9 15 14 11 10 9 13 12 13 12 - - 10 10 ------17 17 16 15 12 11 16 15 12 11 14 13 14 13

B. Time of day Position 0 10 50 70 96

C) C) C) C) 0 0 0 ( ( ( env env env env env T T T

Time of day C. L. mulleri C. ruficaudata

A amicula L. guichenoti

0 E. murrayi ( b

8 13 14 9 11 16 - - 9 10 - - - - 14 15 12 17

Time of day Fig 2. A. Profiles for levels of insolation and environmental temperatures available to lizards across the gradient of shading generated using shade cloth covers over the field enclosures throughout the course of the day. B. Profiles for Tenv in choice boxes. Preferred temperature range (mean ± 68%CI) for the hot (all species except E. murrayi – red bar) versus cold (E. murrayi – blue bar) lizard groups are overlaid on available temperatures Tenv. C. Overall body temperatures, Tb, used by lizards in experimental choice boxes, including the first hour of measurement in which lizards were heating to preferred temperatures. All error bars represent 95% confidence intervals.

141 Chapter 6: Lizard habitat choice

Sep - Oct

C) 0 (

hab T Nov - Dec

Jan - Feb 6-7 7-8 8-9 9-10 10-11 11-12 12-13 13-14 14-15 15-16

Time of day

Figure 3. Thermal profiles of environmental temperatures (Thab) available between sun and shade in young plantation habitats throughout the course of a day in spring (Sep – Oct), spring-summer (Nov – Dec) and summer (Dec – Jan). Upper profile is sun temperature, lower profile is shade temperature. All error bars represent ± 2SE.Profiles are overlaid by mean preferred temperature ranges of ‘hot’ (all species except E. murrayi – red bar) and ‘cold’ (E. murrayi – blue bar) lizards determined in experimental choice boxes.

Overlaying preferred Tb and available Thab temperature ranges in figure four identifies that achieving preferred temperatures in young plantation environments would be challenging for all species that did not exhibit behavioural flexibility. For all species except Eulamprus murrayi, Tb fell within

Thab ranges, but preferred temperatures were available only during the middle of the day in Spring (Fig. 3). For Eulamprus murrayi whilst springtime Thab overlapped preferred temperatures, by January young plantation Thab was above the preferred temperature range at all times of the day (Fig. 3).

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Feeding study

A laboratory experiment assessing gut passage time was used to examine whether preferred temperatures identified in field enclosures were physiologically optimal for processing food. Previous studies have shown that food-passage rates are often temperature dependent (Huey 1982, Beaupre et al. 1993) and to consequently influence habitat choice (e.g. Pafilis et al.

2007, Homyack 2010). In January sixteen adults (7♀ and 7♂) of each species used in the choice experiments were collected from the Mid-North

Coast, transported to Wollongong University, and allowed to adjust to captivity for three weeks. Lizards were housed individually in a controlled temperature room with a circadian fluctuation of 24-270C and photoperiod

L:D 12:12h. The smaller species were placed individually in 30 x 25 x 15cm boxes and larger E. murrayi in 60 x 45 x 50 boxes. All boxes had a substrate of natural leaf litter and wood shavings. Thermal gradients for basking were generated by an overhead lamp above and a coiled heating wire (Resun

HTC-750, 55W ) beneath one end of each box on a 10:14 L:D photoperiod.

Water and food (Tenebrio larvae and Gryllus sp.) were available ad libitum.

Trials during the pilot study identified that L. muelleri would not adjust to experimental enclosures, and this species was summarily excluded from experimental testing. All females were palpated before testing to ensure they were not gravid.

Testing for temperature effects on gut passage rate involved placing lizards in 40 x 13 x15cm plastic tubs in controlled temperature cabinets set to

0 temperatures of 20, 25 or 30 C, the mid-point of Thab available to lizards in young plantations in spring, spring-summer and summer. Temperatures

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representing mid-winter and mid-summer (15 and 350C) were initially included in a pilot study but were abandoned after establishing that 350C was above the critical thermal maximum for Calyptotis ruficaudata, and was lethal for Eulamprus murrayi, and at 150C <10% of individuals of all species besides E. murrayi fed. Temperatures in cabinets varied <10C around the set point temperature during trials. Tubs contained two retreats and were floored with river sand. Once in the cabinets lizards were provided with a cold light source and were left to acclimate for two hours prior to testing. A pilot study using crickets marked with a fluorescent pigment (#JS-S03019, sunset orange, Radiant Color Inc, Richmond) diluted in acetone in a 1:15 ratio and painted onto crickets (which were then heated under a lamp to facilitate evaporation of acetone), showed that gut passage was completed within 92h in all test temperatures. It was assumed that the first faeces contained the remains of prey consumed in that trial, as lizards were fasted for 96h prior to feeding trials. During testing lizards were filmed in real time using a video camera (JVC Everio), downloading to a video tape player (Sony LVX811).

These tapes were later reviewed and times of feeding and defecation recorded using a stopwatch. Appetite was inferred from the total number of prey items consumed. Gut passage rates were defined as the time between the complete disappearance of the first prey into the mouth of the lizard, and the first appearance of faeces. While a variety of other behaviours were initially recorded, only gut passage time and appetite were finally analyzed as they were not influenced by the novel environment (substrate, motion) in experimental cages, whereas other behaviours often analyzed as indicative of feeding efficiency (e.g. prey handling time, number of attempts etc.) rely on

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subjective observations of ‘normal’ behaviour and had the potential to be compromised in the experimental enclosures.

Each test involved introducing five crickets (Gryllus sp.; each no more than 3.0 ± 0.5% of the test lizard weight to standardize appetite against body size (Bilcke et al. 2006) into tubs ten centimeters from the lizard via a small tube. Additional crickets were introduced singly until an individual did not eat or attempt to catch any further crickets for 10 minutes. After this point any remaining crickets were removed, and lizards were maintained in their tubs at the test temperature until faeces were produced. Individuals were allocated in a random sequence to temperature treatments. In between trials, lizards were returned to controlled temperature rooms for eight days of rest during which they were fed ad libitum for four days and then fasted for four days prior to the next testing. Each individual was tested once at each temperature.

Data Analysis

A mixed-effect repeated measures model was used to identify any effect of temperature on gut passage time, using ‘species’ and ‘temperature’ as fixed effects and ‘lizard’ as the repeated factor. A repeated measures design was necessary as each lizard was measured three times; once per temperature. All of the continuous response variables were square-root transformed so that the residuals of the final models were normally distributed. This model was also used to identify whether appetite was suppressed at temperatures to which lizards would not normally be exposed.

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Results

Temperature interacted with species to affect their appetite (F = 12,133

= 23.583, p < 0.001) and gut passage time (F = 9,112 = 12.729, p < 0.001) in the laboratory experiments. Gut passage rates and appetite were lowest at the spring temperature of 200C for all species. At 250C, Adrasteia amicula had the greatest appetite and fastest rate of gut passage (Fig. 4), despite this

0 being lower than its preferred Tb of 30 C in the field enclosures.

Lampropholis guichenoti differed by showing greatest appetite at 300C, in accordance with its mean preferred temperature identified in choice boxes, while its gut passage rates did not differ between 25 and 300C. Whilst field enclosure data suggested Eulamprus murrayi should perform best at the physiologically optimal temperature of 250C, there was no difference in either gut passage rate or appetite between 25 and 300C (Fig. 4). Similarly, the forest-associated Calyptotis ruficaudata was expected to show highest rates of gut passage at 300C, but appetite did not differ between 25 and 300C, although gut passage rates were fastest at 300C (Fig. 4).

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Table 3. Estimates of fixed effects for mixed models analysis of appetite and gut passage rate for each lizard species. Results shown are for overall model for each species. Significant values are in bold.

Appetite Gut passage rate Species df t p df t p L. guichenoti 133 6.679 0.000 112 2.695 0.008 A. amicula 133 0.216 0.834 112 0.090 0.031 C. ruficaudata 133 16.762 0.023 112 4.888 <0.001 E. murrayi 133 0.772 0.014 112 9.049 <0.001

200C 0 25 C 300C

No. prey eaten No. prey eaten

Hours

Temperature (0C)

Fig 4. Appetite and gut passage rates of for the four species tested experimentally at (200C) spring, (250C) spring-summer and summer (300C) temperatures available in young plantations on the Mid North coast of New South Wales. All error bars are ±2SE.

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Discussion

This study demonstrates that habitat choice in the small scincid lizards examined in this study is constrained by behaviour and that this can influence temperature-dependent physiological processes such as food assimilation rates. Such outcomes can have flow-on consequences for the diversity of lizard assemblages found in young native plantations. The summertime thermal environment in young plantations appear to be physiologically stressful for all lizard species, and while temperatures optimal for digestive efficiency can be achieved, they require the species using this forest type to exhibit flexibility in both activity time and substrate choice. Those species that did not possess this flexibility were excluded from young plantations. I discuss reasons for this in relation to the three hypotheses initially proposed.

Hypothesis 1- habitat generalist species have a broader preferred temperature range than habitat specialists

The breadth of preferred temperature ranges are ultimately constrained by the temperature-dependence of physiological processes

(Huey 1991, Blouin-Demers & Nadeau 2005). However for heliothermic reptiles, achieving the preferred temperature range relies on the availability of preferred thermal microenvironments in a habitat, the flexibility of behavioural mechanisms which adjust rates of heat exchange, and the need to balance predation risk with the need to bask (Heatwole 1977, Huey & Slatkin 1976,

Heatwole & Taylor 1987, Vickers et al. 2011). In opposition to my initial hypothesis the generalist Lampropholis guichenoti did not have a broader

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preferred temperature range than the less widely distributed skink species examined in this study. While young plantations could thermally satisfy L. guichenoti’s 300C physiologically optimum temperature (Fig 5.), to achieve the narrow preferred temperature range it showed in the habitat-choice enclosures, L. guichenoti would need to restrict activity to the middle of the day in the sun in spring, but by summer would need to remain exclusively in shaded microhabitats or confine it’s activity period to mornings and afternoons (Fig. 5). L. guichenoti is known exhibit flexibility in behaviour

(Prosser et al. 2006) and substrate choice (Penn et al. 2003, Fischer et al.

2005). Such flexibility explains its wide range generally, and its use of a variety of habitats with different structures in this study (Chapter 2). In addition to young plantations providing optimal temperature, the persistence of L. guichenoti in the hot environment of young plantations throughout an annual cycle may also be related to its tolerance of dry hydric conditions in the nest environment. Such conditions are known to affect the body mass of hatchlings, but to have little effect on other phenotypic traits related to fitness such as sprint speed or growth rate (Du & Shine 2008).

The high preferred body temperatures identified in this study are typical of diurnal species like Lampropholis guichenoti, but cryptozoic species are often characterized as having low preferred and optimal temperatures

(Huey & Bennett 1987). However, the cryptozoic Lerista muelleri and

Calyptotis ruficaudata had preferred temperature ranges matching that of the generalist L. guichenoti. For Lerista muelleri this is unsurprising, as high preferred temperatures are typical of the Eyrean (central Australian) radiation of Sphenomorphine skinks (Spellerberg 1972b, Greer 1980, Bennett & John-

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Alder 1986, Skinner et al. 2008), and the majority of species of both Lerista and its sister genus Ctenotus are thought to have radiated in association with the historical aridification of Australia (Rabosky et al. 2007). While the plantation environment is not physiologically limiting for L. muelleri in spring and early summer , its rigid activity times and preferences for low levels or irradiation coupled with a strong preference for complex substrate, is likely to hamper the use of young plantations by these species. Calyptotis ruficaudata, while also a member of the diverse Sphenomorphine skink group (Reeder

2003), radiated from a Bassian (coastal southeastern Australia) lineage which typically prefer lower temperatures (Huey & Bennett 1987). Despite this phylogenetic tendency, the forest-associated Calyptotis ruficaudata in this study displayed optimal and preferred temperatures similar with those of the woodland L. muelleri and L. guichenoti. While the plantation thermal environment is not physiologically limiting for either L. muelleri or C. ruficaudata in spring and early summer. L. muelleri’s rigid activity times and preferences for low levels of irradiation coupled with a strong preference for complex substrates is likely to hamper its use of young plantations.

Hypotheses 2 - habitat-specialist species should show stronger preference for particular substrate types than habitat-generalist species

Specialist species exhibited two patterns of habitat use in this study. In support of hypothesis two, the forest specialist Eulamprus murrayi did not separate preferences for thermal and structural environment, and chose habitats that were consistent with a strong preference for low irradiance and complex structure. Such limited use of cues that link with particular habitat

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choices are a typical trait of habitat-specialist species (Greenville & Dickman

2009). Unlike the situation for other lizard species examined, the mid- summer operative temperatures in plantations exceeded the critical thermal maximum E. murrayi and would preclude its use of this habitat outside of spring and autumn. This coupled with the sedentary nature of this species and it’s rigid preference for low-light environments and complex substrates, suggests that neither young nor old native plantations could act as habitat for

E. murrayi, even if plantation management included diversifying ground layer complexity, as plantations are typically light-saturated due to canopy disturbance.

However, in refute of hypothesis two, the plantation specialist

Adrasteiascincus amicula, did not choose habitat based on a preference for substrate type. While A. amicula consistently preferred complex substrates in experimental enclosures its acceptance of simple substrate in young plantations in the field reflects high behavioural flexibility. While A. amicula is not truly a ‘specialist’ to young plantations, its preference for these forests in the study area (Chapter 3, 5) is unusual given its typical association with moist closed woodland and heath (Cogger 2006). Its failure to use old plantations or logged native forest in the study area is unlikely to be mediated by competition, as no other skink species of equivalent size were detected in the study area. A more likely explanation relates to its opportunistic ability to exploit the ‘new’ habitat young plantations represent, which is promoted by its small size, and thus ability to find refuges in the plantation understorey that has typically very little leaf litter depth. It may also be related to predation pressure, which has been demonstrated to force the use of suboptimal

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microclimates in lizards (Díaz et al. 2005), and it is notable that the diversity of invertebrates and the abundance and diversity of lizard predators

(e.g.dasyurid mammals) increases in old plantations and logged native forests.

Hypothesis 3 – habitat specialists should have the most efficient gut passage time in temperatures common to the habitats in which they specialize

This study did not identify any association between habitat specialization and gut passage time. For Adrasteiascincus amicula gut passage time was maximal at 250C, while for Eulamprus murrayi it did not differ between 25 and 300C. While it seems counterintuitive that habitat specialists do not perform optimally at the temperatures they behaviourally select, partial co-adaptation between behaviour and digestive efficiency may account for the disjunction between preferred and optimal temperatures.

Huey and Bennett (1987) believe partial coadaptation to result from a faster rate of evolution of preferred temperature than optimal temperature. This produces disjunction between the temperatures that maximize fitness, and those which allow an individual to be active in a habitat. There is no consensus of how common partial coadaptations are among lizards,

(Angilletta et al. 2002a), and whether such conditions typify Australian skinks

(Garland et al. 1991), but Angilletta et al. (2002b) suggests that the outcome of this trade-off is that thermoregulatory behaviour favours optimal physiology over performance due to the direct fitness costs associated with not assimilating energy efficiently. The results of the present study do not agree with those of Angilletta et al. (2002b) but identify A. amicula as maintaining

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preferred temperatures that are suboptimal for digestive rate. While many ectotherms do behaviourally select suboptimal temperatures, Hutchison &

Maness (1979) and Martin and Huey (2008) argue that they may do so to avoid the often fatal consequences of attaining body temperatures only a few degrees above their physiological maxima. The persistence of A. amicula in young plantations through the summer (Chapter 2) when thermal conditions are persistently above those that are physiologically optimal suggests that the benefits they gain by exploiting empty niches in the ‘new’ habitat created by plantations, outweighs the costs of suboptimal physiological performance.

Similar situations are described by Huey and Bennett (1987) as “adequacy rather than optimality guiding the evolution of thermal performance and behaviour”.

Such disjunctions between performance and physiology can arise when optimal temperatures are evolutionarily static (e.g. Van Damme et al.

1990, Angilletta et al. 2002a, Rodríguez-Serrano et al. 2009). In this study low optimal temperatures probably represent the ancestral state of A. amicula, a temperate, south-eastern Australian endemic (Wells 2010). The high preferred temperatures that have allowed A. amicula to radiate from closed woodland into young plantations may represent an evolutionary pressure that will, in the future, shift optimal temperatures upwards through co-adaptation. Further, because all physiological processes do not share the same optimal temperature (Van Damme et al. 1991, Du et al. 2000,

Ojanguren et al. 2001, Angilletta et al. 2002b), it is possible that the preferred temperature range identified for A. amicula optimizes some other measure of performance that was not investigated in this study (the ‘multiple optima’

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hypothesis, Bustard 1967, Pough 1980, Huey 1982). Temporal modifications in behaviour to meet multiple optima are common for ectotherms (Angilletta et al. 2002b), and as an active shuttling heliotherm with very small size (SVL

30mm), and thus very little thermal inertia (Bell 1980, Garrick 2008), A. amuicula’s ability to rapidly change body temperature via behaviour to ensure optimal physiological status is probably easy for this species to achieve, and could foster its persistence in young plantations. That A. amicula experiences multiple optima and adjusts preferred temperature accordingly is supported in this study by the shift in activity observed from the active morning period, to the inactive afternoon period at the cooler end of the gradient, and the associated decline in body temperature, towards the digestively optimal 250C.

Such non-random retreat selection is recognized as important to determining habitat selection in reptiles (e.g. Grant & Dunham 1988, Pringle et al. 2003,

Goldsbrough et al. 2006, Andersson et al. 2010, Martín & López 2010, Pike et al. 2011), and offers a good explanation for the disjunction between preferred and optimal temperatures exhibited by A. amicula.

In contrast to Adrasteiascincus amicula, the mean preferred temperature range for Eulamprus murrayi corresponds with its optimal temperature for food passage and appetite. As a rainforest endemic

(O’Connor & Mortiz 2003), this low optimal temperature was expected for E. murrayi, but the invariance of response to temperature in both physiological parameters measured was not. While researches have often identified strong inverse correlation between temperature and gut passage time (e.g.

McConnachie & Alexander 2004, Bilcke et al. 2006), there are cases where gut passage time (Alexander et al. 2001) and other measures of

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performance such as sprint speed (Angilletta et al. 2002b) do not increase with increasing temperature. It seems that for E. murrayi and for the other skink species examined, the optimal temperature range for digestion is broad.

This breadth may reflect the historic radiation of these skinks either out of mesic habitats, in the case of E. murrayi (O’Connor & Mortiz 2003) and L. amicula, or out of arid habitats into mesic ones in the case of Lerista muelleri.

Conclusions

This study demonstrates that young native plantations can offer a highly seasonal resource for small skinks, and that in the system studied low spring and high summer temperatures could preclude use of this habitat by lizard species with rigid preferences for particular body temperatures and physiological performance, if thermal refuges do not exist. Experimental tests of thermal and substrate preference reveals the simple substrate found in young plantations is not preferred by any lizard species examined. This suggests that substrate enrichment is needed to increase skink diversity, and, by association, the diversity of other species as well. While we might expect to see species of low mobility persisting in young plantations by exploiting microclimatic variability associated with habitat edges (Chen et al. 1993,

Laurance 2004), such seasonal shifts in habitat use of low mobility fauna remain poorly documented in terrestrial environments (Seebacher & Alford

2002), and are largely unexplored in plantations. Understanding these sorts of seasonal movements may be important however, as they have ramifications for the way habitat enrichment in plantations could be achieved.

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The management of slash is currently being investigated as a means of enhancing understorey complexity in plantations (Matthews et al. 2010).

Enhancing habitat complexity by simply leaving slash is unlikely to generate thermal refuges cool enough to allow persistence throughout the annual cycle. My results suggest that, habitat enrichment protocols that would need to include coarse woody debris piles, cover boards or some other measure of ground cover able to decrease irradiance and reduce microhabitat temperatures by more than five degrees in summer to permit their year-round residence, and attract cryptozoic species. More thermally targeted refugia such as stacked log piles are likely to better benefit these cryptozoic species, in particular C.ruficaudata, which shows some tolerance of higher irradiance, and so may use young plantations with the provision of necessary habitat selection cues. The case for attracting L. muelleri to young plantations is less certain, primarily because other species of Lerisa show strong preferences for uncompacted substrates that reduce the energetic costs of movement

(Greenville & Dickman 1999), and soil compaction associated with machines and cattle’s grazing is typical of managed plantations. The success of those species already common in young plantations is attributed to high behavioural flexibility which allows these species to exploit the mismatch between ecologically and physiologically optimal temperatures.

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Chapter 7: General discussion

Chapter 7 General Discussion

The value of plantations as habitat – an overview

As for all countries globally, Australia’s unique biota are under threat from changes in land use, altered fire regimes and hydrology, and the spread of invasive species, pathogens and diseases. These threats have generated a loss of biodiversity in the last 200 years that encompasses the disappearance of up to 30 per cent of small mammals, as well as numerous plants, birds, reptiles, fish and amphibians. Despite the adoption of global and national policy to slow current rates of species decline, Australia is still losing biota at an increasing rate (DEWHA 2009). This rate of loss is likely to be accelerated by climate change which is expected to place already stressed and vulnerable species at greater risk (Beeton et al. 2006).

There is recognition that national reserve systems are inadequate to protect biodiversity (Rodrígues et al. 2004). Reserve size is often insufficient to buffer ecosystems against large-scale disturbances such as fire and climate change, and reserve connectivity is insufficient to accommodate large-scale animal movements often associated with these disturbances

(Paton & O’Connor 2010). Native plantations can play many roles in mitigating biodiversity loss, if they enhance and extend native forests. Further, afforestation with plantations has the potential to address one of the crucial issues in rural biodiversity conservation; the restoration and maintenance of fauna in partly cleared rural landscapes (Loyn et al. 2008). This research

166 Chapter 7: General discussion

identified that providing habitat for fauna is one role native plantations can fulfill.

This research in context

This research conducted in on Australia’s East coast in small-scale, afforested and reforested native plantation monocultures in close proximity to forests, identified that young and old managed plantations can support vertebrate assemblages as species rich as those in old growth forests.

Borsboom et al. (2002) conducting research in a similar plantation system in south-east Queensland, identified lower vertebrate richness in plantations less than four years old in comparison to adjacent logged native forests, but similar richness between forests and native plantations 4-40 years old. Birds only, showed lower richness in young plantations than forests at the taxon- level. This thesis extended Boorsboom et al.’s (2002) research by including comparisons of faunal richness in old growth forests of high faunal diversity with logged native forests and young and old plantations, and by targeting herpetofaunal and ground mammal responses to plantations. This research found that the mean species richness and abundance of birds, mammals and herpetofauna was statistically similar in logged and old growth forests and in structurally simpler young and old plantations. Whilst similar cross-taxa responses to plantations have been documented previously in old, ecologically managed plantation monocultures of Araucaria angustifolia in

Brasil (Fonseca et al. 2009), this thesis is the first to identify similar species diversity responses to commercially managed young plantations across animal taxa, and between old growth forests and native plantations in

167 Chapter 7: General discussion

forested environments. This result contradicts taxon-specific meta-analyses of bird (Nájera & Simonetti 2009) and mammal (Ramírez & Simonetti 2011) use of native plantation monocultures, which find that species richness in plantations increases with structural complexity. Whilst structural complexity did influence abundance in this research, species richness was maintained by a high species turnover in young and old plantations, where forest species loss was offset by incursions of woodland and generalist species. This result contrasts that of Hobbs et al. (2003) whom identify young native plantations in large-scale plantation systems in fragmented forest landscapes in Western

Australia as supporting less species rich assemblages than forest remnants.

Faria et al. (2007) also found that in old, ecologically-managed Brazilian cacao plantations reduced forest cover in the landscape mosaic generated impoverished plantations irrespective of the biological group considered. The retention of biological legacies (Tews et al. 2004) and high diversity in the regional species pool (Tscharntke et al. 2012) all influence local species diversity, and were likely to have promoted species replacement in plantations in this research. The differences in results from this research and that of Hobbs et al. (2003) suggests that whilst commercially managed, young native plantations can provide habitat for a variety of species, landscape context and immigration rate is likely to be equally as important as local vegetation heterogeneity in influencing the development of diverse vertebrate assemblages.

In broad connection to native forest, without accessory management to enhance biodiversity beyond the retention of legacy trees and riparian strips, and with the impact of cattle grazing, in this research native

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plantations six years old supported diverse faunal assemblages including woodland birds and open-habitat herpetofauna. While this result appears positive, many authors (Dunning et al. 1992, Didham et al. 1996, Peterson et al. 1998, Barbour et al. 2003) stress that ecosystem function, rather than species richness (which may respond to disturbance slowly - Vellend et al.

2006), is a better indicator of ecosystem health. Both this research (Chapter

3) and that of other authors (Borsboom et al. 2002, Lindemnmayer & Hobbs

2004, Kanowski et al. 2006, Zurita et al. 2006, Brockerhoff et al. 2008) agree that assemblages shift towards generalist species in native plantations, and that forest-specialist species from all faunal groups are poorly represented.

The lack of nectivorous guilds, sub-canopy specialists, and the dominance of generalist ground feeders in both young and old plantations points to persistently compromised pollination and plant dispersal systems, and suggests that even in older plantations where faunal assemblages are diverse, ecosystem function is unlikely to operate as it does in forests.

Woodland birds

In a nationwide review of Australian bird diversity, Paton & O’Connor

(2010) identified monocultures generally as providing poor habitat for birds.

This thesis and other research comparing bird diversity between forests and monoculture plantations in both fragmented forest landscapes in Australia

(Loyn et al. 2007) and continuous forested landscapes in Australia

(Boorsboom et al. 2002), South America (Zurita et al. 2006) and Africa

(Farwig et al. 2008), contradict Paton and O’Connor’s (2010) statement. This body of research collectively identifies that native plantations can support high species richness of canopy and ground feeding woodland birds. Loyn et

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al.’s (2007) results find that stand-level habitat features are stronger predictors of bird richness than landscape context. If commercially managed plantations typically retain open sub-canopy structure throughout the rotation cycle, as was identified in this research (Chapter 3), and landscape context is less important than local vegetation structure in influencing bird species richness, then it is likely that native plantations can promote incursions of woodland birds into forested environments.

In this research plantations supported diverse assemblages of woodland birds after a relatively short six-year time period. This result suggests some potential for woodland birds to expand their ranges into forested environments using native plantations as a conduit. This range expansion may be facilitated by the vegetation structure and thermal environment in managed young plantations, which can remain hot and open after 30-years of growth and management (Chapter 2). Reino et al.’s (2009) research examining the invasion of afforested plantations by open-country birds suggests that in a Mediterranean forest-farmland system, a mosaic of young and old plantation ages influences the persistence of woodland birds in the landscape as woodland species respond more positively to hard (old) than soft (young) edges. Certainly exotic plantations are known to attract open-habitat birds both in Australia (e.g. Curry et al. 1991) and in temperate ecosystems overseas (e.g. Paquet et al. 2006), and birds in native plantation systems in Australia use plantation edges differently to interiors (e.g. Hobbs et al. 2003). Thus Reino et al.’s (2009) assertion that plantation age mosaics foster incursions of woodland birds into forested landscapes by providing favoured edge habitats is plausible for Australian plantation systems.

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Woodland bird incursions into plantations are positive if they increase regional diversity, but negative if plantations provide corridors for invaders which change community dynamics by increasing local levels of competition or predation (e.g. Poulin et al. 2011), or disrupting local adaptations by allowing incursions of novel genes (see Bennett 1998, 2003 Table 4-3 for a review). While these issues have been addressed extensively in the fragmentation literature, the role of native plantations as corridors or matrix has only recently begun to be addressed by researchers (e.g. Bentley et al.

2000, Kavanagh et al. 2005, Barlow et al. 2007a, Brockerhoff et al. 2008,

Lindenmayer et al. 2009, 2010). The impact of plantations as conduits for woodland species invasion needs investigation before predictions about the impact of native plantations on birds in adjacent forests are possible. Such predictions will require long-term studies as the global native plantation estate matures.

Forest species

General statements in the plantation literature identify native plantations as poor habitat for forest species (Lindenmayer & Hobbs 2003,

Brockerhoff et al. 2008). The results of this thesis (Chapter 3) suggest this statement needs qualification. In this research native plantations in broad connection with forests supported 46% of forest bird species as well as 40% of mammals, and 18% of herpetofauna (Chapter 3). Whilst the mean richness of forest species was lower in plantations than forests for all faunal groups, old plantations supported similar abundances of forest fauna from all taxa. Lindenmayer et al. (2003) suggest that knowledge of species-specific preferences for habitat attributes is more important in predicting how a

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species will use habitat than anthropogenic concepts like ‘forest’ or ‘non- forest’. From this perspective the results of this research suggest that plantations only negatively affected those forest species intolerant of low- floristic diversity and structural complexity. Certainly, for the plantation system investigated in this study, a generalization about the effect of plantations on forest species must be qualified to state that only a percentage of the forest species in the species pool were excluded from plantations.

Small mammals

This research identified that even without complex understorey condition, young and old native plantations in close proximity to native forests can support assemblages of small mammals equally as diverse as those in old growth forests (Chapter 3). This result contrasts those from a study in fragmented Araucaria vine forests and native Araucaria plantations in southeastern Queensland (Bentley et al. 2000) which found an increase in mammal richness and abundance with increasing sub-canopy complexity. A meta-analysis by Ramírez and Simonetti (2011) identified that small mammals increase in response to structural complexity independent of plantation type in commercially managed-landscapes. Similarly Holland &

Bennett ‘s (2007) species-based analysis of landscape-scale habitat use by small mammals in fragmented eucalypt forests also found that species richness increased with vegetation complexity. Boorsboom et al. (2002) found in their analysis of native plantations in broad connection to forests that small mammal communities in simply-structured young plantations were depauperate compared with those in older plantations and forest remnants.

My results contradict these findings; whilst young plantations showed trends

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towards lower forest mammal richness and lower small mammal richness overall, richness did not differ significantly between plantations and old growth forests (Chapter 5). This result supports work by Mitchell et al. (1995) and Mengak and Guyunn (2003) from native pine plantations in broad connection to continuous pine forests in the United States, which identifies that the availability of preferred microhabitat features influences mammal diversity more than stand age or canopy structure. Significant taxonomic replacement occurred in both Mengak and Guyunn’s (2003) study, and in my research (Chapter 3) where there was a shift from rodent-dominated forest assemblages to dasyurid-dominated plantation assemblages and species preferring simpler vegetation structures. Whilst local vegetation structure was likely to have ultimately determined species diversity in plantations, the species pool available in an unfragmented forest landscape was large enough to allow species replacement, resulting in the development of diverse ground mammal assemblages in plantations. Taxonomic shifts were notably lacking in the studies by Ramírez and Simonetti (2011), Boorsboom et al.

(2002) and Bentley (2000) which compared plantations with fragmented forests, stressing that as for birds, landscape context and pathways for immigration mediated by retained vegetation or proximity to forests are likely to be important factors promoting small mammal diversity in native plantations.

Food availability

Decreased body size as a result of reduced precipitation is a global phenomenon only recently being investigated in relation to organism

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development and growth (Sheridan & Bickford 2011). In commercial plantation systems that have lowered soil moisture compared to forests

(Chapter 2) and are generally structurally simple and exposed to high levels of disturbance, there is high potential for invertebrate fauna of larger body- sizes to be lost, so reducing available prey, and potentially insectivore abundance. This research identified significant decreases in invertebrate body size moving from old growth native forests down the gradient of structural complexity to young plantations (Chapter 4). Jelaska et al. (2011) identify a similar response to plantation age by invertebrates in native beech plantations in Croatia. This decrease in invertebrate size occurred in concert with a lower abundance of large and medium insectivores in young plantations. However, both this research and that from exotic plantation monocultures (Cunningham et al. 2005, Samways et al. 1996, Prieto-Benítez

& Méndez 2010) identified that invertebrate assemblages were equally diverse and abundant in forests and young plantations. Whilst invertebrates were smaller and less abundant in plantations than forests, they were equally as speciose and palatable, and insectivores using young plantations could access a broader range of preferred prey types than were available in old plantations or native forests. Hence lower insectivore abundances in young plantations were more likely to be driven by the scarcity of refuges than prey availability.

Matrix quality

Current plantation research recognizes that the quality of matrix habitat and its management influences local species richness and incidence

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and so can mediate plantation diversity (Ricketts 2001). How matrix habitats influence diversity depends on whether the matrix is defined as the dominant form of vegetation or as the area that is most disturbed (see Kupfer et al.

2006 for a discussion). In landscapes dominated by continuous forests where in plantations are relatively small, forests are the matrix. Comparisons of forest matrices and exotic plantations in both large and small-scale tropical and temperate forest systems have identified that plantations often support a lower species richness of beetles (Yu et al. 2006, Henríquez et al. 2009) birds and ground mammals (Gjerde & Sætersdahl 1997, Lindenmayer et al.

2000, Barlow et al. 2007a).

Chapter five of this thesis analyzed the potential of the grazed forest matrix to act as a source of recruits into abutting native plantations. My results identified that despite contrasting vegetation structure between young plantations and the adjacent forest matrix, small mammal and reptile assemblages were equally species rich in both habitats, and that plantation and forest assemblages were dominated by generalist species. Similar richness between forest and matrix has been observed previously when comparing low contrast habitats for ants in the Amazonia savanna

(Vasconcelos et al. 2006), and for ground mammals in pine-afforested

Afromotane grasslands in South Africa (Johnson et al. 2002). Bihn et al.

(2010) assert that for ants in Atlantic forest remnants in Brazil, a lack of specialist species in assemblages despite high species richness overall, indicates that habitat quality is poor. This research disagrees with Bihn et al.’s 2010 assertion, and finds that whilst cattle-grazed forest matrices lacked

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forest specialist species they supported forest generalists, and ground faunal assemblages equally as rich as those in old growth forests in the region.

Cattle-grazing in the forest matrix more strongly influenced small- mammals than herpetofauna, and influenced the pool of species available to recruit into young plantations (Chapter 5). For mammals whilst grazing may have reduced mammal diversity in the forest matrix itself, strong preferences for complex habitat features were more likely to preclude forest species recruiting to young plantations. For herpetofauna, grazing may have increased the permeability of the matrix, with a resultant influx of thermophilic species into plantations. The disparity in diversity benefits for herpetofauna and mammals suggests native plantations need to be positioned in the landscape in proximity to both grazed and ungrazed forests to maximize recruitment across multiple taxa. Whilst implementing such a design will rely on the site-specific factors, it is a worthy aim for inclusion in the process of afforesting pastures with native plantations, as current research identifies fragmentation as a driver that may render Eastern Australian eucalypt forests susceptible to large changes in ecosystem function, and ultimately biodiversity decline (Laurance et al. 2011). By understanding whether plantations differ in permeability with individual species choices and proximity to source populations, managers can attempt to minimize afforestation that fragments forest faunal assemblages, and in doing so generate plantation systems that provide both local and regional benefits to forest fauna.

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Plantations and the thermal environment

Floristics, vegetation structure and plant orientation all play a role in determining microclimatic diversity at the ground level. While plantations of endemic trees species may be more likely to approximate thermal microclimates at the ground level than exotic species, canopy cover is the strongest determinant of thermal environment (e,g, Porté et al. 2004, Rambo

& North 2009). Reducing canopy complexity in forest necessarily leads to an increase in the range of microclimates experienced by organisms at the forest floor, which in turn may increase in biodiversity (Hanley 2005, Weng et al. 2007). This research is the first to identify that, as for exotic plantations, the vegetation structure and associated microclimate in young and old native monocultures and logged and old growth native forests follows levels of canopy reduction. The greater the loss of canopy, the higher the maximum temperature at the ground level (Chapter 2), which in summer results in both young and old plantations having a more variable microclimate than forests and generates a temperature difference approximating 200C at its greatest point. This temperature differential may be physiologically limiting to plants and animals using plantations, particularly forest species most likely to recruit to plantations from the abutting forest matrix. Chapter six adopted an ecophysiological approach to identifying physiological limits to lizard diversity in young plantations, using small skinks as a model.

This research is the first to conclusively demonstrate that young plantations can be physiologically limiting to some forest species. Numerous studies identify that habitat choice in reptiles (e.g. Vitt et al. 2005,

Goldsbrough et al. 2006, Pike et al. 2011), mammals (e.g. Hill 2006, Issac et

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al. 2008) and frogs (e.g. Vallan 2002) is influenced by available thermal microclimate. Habitat choice in the small scincid lizards examined in this study was constrained by both behavior and physiology and this had flow- through consequences on lizard species richness in young native plantations

(Chapter 6). The summertime thermal environment in young plantations was stressful for all lizard species, and while temperatures optimal for digestive efficiency could be achieved, they required the species using this forest type to exhibit flexibility in both activity time and substrate choice. Those species that did not possess this flexibility could not use young plantations. Whether these results apply widely to other ectotherms is unknown, but this chapter suggests that identifying whether a species is able to separate preferences for habitat structure from those for thermal environment, will allow researchers to predict the potential for those species to use limiting habitats like young plantations.

Plantations and climate change

Climate change is predicted to be one of the greatest drivers of ecological change in the coming century (Lawler et al. 2009). This research

(Chapters 2 and 6) identifies that young plantations are likely to be physiologically stressful for those biota that prefer cool microclimates and for all species during the summer. Thus whilst plantations under current climatic conditions limit some species, the potential for them to become even less suitable with climate warming is high. For ground fauna of limited mobility that are already living at the extreme of physiologically tolerable limits

(Chapter 6), increasing temperature may have directly negative consequences for their persistence in native plantations, and thus for

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plantation diversity as a whole. This is particularly true of species which currently persist in plantations by exploiting microclimatic variability associated with habitat edges (Chen et al. 1993, Laurance 2004). However, for herpetofauna microclimatically-driven species losses in plantations may be offset by an increase of thermophilic species, which in the area studied can permeate through the forest matrix (Chapter five). While there is very little work available that examines the role of microclimate in influencing habitat selection in native plantations, such research is increasingly important as the range shifts associated with climate change occur.

The way forward

Developing plantation management practices that concurrently conserve biodiversity and maintain profitability is a key component of multi- use forest management, and is becoming increasingly important as the plantation estate expands. Identifying the variables that enhance the occurrence and survival of plants and animals in plantations is a mandatory step toward developing effective multi-use plantations (Tews et al. 2004,

Stephens & Wagner 2007, Nájera & Simonetti 2010). This thesis combined landscape ecology, physiology and population ecology to identify that in proximity to continuous forests commercially managed, small-scale, young and old Eucalyptus plantations can support equivalent faunal richness to old growth forests. However, the lack of nectarivores and arboreally-feeding insectivores suggests the ecosystem services these functional groups provide were absent from plantations.

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The correlation between reduced abundance and understorey complexity for birds, mammals and forest reptiles in plantations provides an opportunity to employ a set of management practices which could enhance future plantation diversity (Florence, 2004). Managed young plantations in particular, have a greater potential to become functionally diverse habitats for fauna with changes in plantation design and management that increase the heterogeneity of thermal environments. The retention of coarse woody debris

(reviewed by Stevens 1997) is known to increase faunal diversity in plantation landscapes. While the influence of overstorey vegetation structure on understorey microclimates need further research (Parrotta et al. 1997), increasing ground-level complexity will directly diversify understorey microclimates, which in turn may increase herpetofaunal diversity in plantations with knock-on effects for predators. Including preferred microhabitat features like coarse woody debris and targeting enrichment towards linear strips of low, shrubby vegetation increases forest mammal recruitment into young (e.g. Holland & Bennett 2007) and old plantations

(e.g. Moser et al. 2002), and can promote mammal movement within plantations matrices (e.g. Predevello & Viera 2010). Christian et al. (1998) identified such enrichment increased bird diversity in poplar plantations in the northeastern United States, and birds are positively associated with coarse woody debris in Southeast Australian floodplain forests (Mac Nally et al.

2001). Experimental manipulation of habitat enrichment techniques to understand the both costs to production and the value to fauna is an avenue of value for further investigation for plantation managers.

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Methods for improving faunal diversity in plantation environments have been discussed comprehensively by Kerr (1999), Hartley (2002),

Lindenmayer and Hobbs (2004) and Brockerhoff et al. (2012). This thesis identifies further avenues of research in small-scale native plantations that will advance understanding of the capacity of native plantations to increase faunal diversity. These include :

1) quantifying the of degree of faunal exchange between plantations and forests and

the capacity of disturbance in adjacent habitats to influence faunal diversity in

plantations. Lindenmayer et al. (2009) identify that management of pine stands in

large-scale systems where pine comprises the landscape matrix negatively effects

faunal diversity in adjacent remnant forest patches. Understanding whether

Lindenmayer et al.’s (2009) results apply in small-scale native plantation systems

that experience dynamic movement of fauna, will allow managers to gauge the

impacts of end-rotation harvesting on fauna in retained plantation stands and so

assess the importance of managing plantations for multiple age classes.

2) identifying the potential of native plantations to act as ecological traps through higher

predation or lower breeding success rates. Ecological traps occur when animals

choose habitat in a maladaptive way because they do not perceive differences in

habitat quality, which eventually leads to population extinction (Delibes et al. 2001).

Battin (2004) states that ecological traps are more common in anthropogenically

generated habitats that arise quickly. Mid-rotation native plantations may act as

ecological traps because they resemble forests and so encourage immigration, but

have the potential to promote higher mortality risks and poorer breeding conditions

than forests due to their lack of sub-canopy structure and are likely to supply high

temporal variation in plant resources. As Gilroy & Sutherland (2007) state, activities

that mitigate the effects of ecological traps could help to minimize the impact of

habitat change on natural environments. For native plantations this would entail

reducing management throughout the plantation cycle allowing the development of

some level of understory and adopting longer rotations which would allow

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populations to either evolve new habitat preferences or adapt to changed

environmental conditions. As the nature of ecological traps is often transient, future

research to clarify whether native plantations act as ecological traps needs to be

staggered across plantation ages and should include an assessment of proximity to

forest, which will influence the potential for immigration for all animals.

3) investigating the role of spar plantations in buffering forest from

microclimatic extremes and increasing the functional connectivity of

plantation networks for forest species. Knowledge of animal use of spar

plantations is particularly limited in native plantation systems. However, if

spar plantations can provide habitat for those species using young

plantations, they have strong potential to provide alternative habitat when

staged management occurs in plantations. Spar plantations are cooler

than plantations and so may represent thermal barriers to ectotherms and

competitive extra-regional species moving between plantations and

forests. Clarifying how effective spar plantations are as barriers will

provide better understanding of the role of spar-aged stands as part of the

plantation age mosaic. Such knowledge is currently valuable for

Australian systems as 22.5% of the national hardwood plantation estate

will mature to spar age in the next five years (Gavran 2013). This

research is likely to increase in importance as range shifts associated with

climate change escalate.

4) clarifying the role of retained habitat features and ground-level complexity

in increasing alpha diversity in both young and old plantations. Whilst a

number of studies have investigated positive diversity benefits of including

retained habitat features in plantation designs (e.g Hsu et al. 2010,

riparian strips, Law & Chidel 2002 – habitat trees, Cummings & Reid

182 Chapter 7: General discussion

2008- slash), there is little Australian research into faunal responses to

less cost-intensive management strategies such as retaining/burning

slash, that can be implemented throughout the rotation cycle. Retained

coarse woody debris is known to positively benefit fauna and offset

disturbance arising from mid-rotation management in studies from North

America (e.g. Stevens 1997, Owens et al. 2008). Replicating such

research in Australian plantation systems and identifying the management

costs that occur in conjunction with retaining coarse woody debris could

identify a cost effective and easily achievable means of increasing native

plantation biodiversity.

Conclusions

This thesis provides a response to the call for greater knowledge of whether small-scale locally endemic tree plantations are diverse habitats for fauna (Brockerhoff et al. 2008). Fonseca et al. (2009) identify that when monoculture plantations are managed ecologically, high species diversity can result. This thesis provides evidence that with retained habitat features and in close proximity to native forests monoculture native plantations can enhance and extend native forests and can increase both local and regional diversity.

However, young native plantations offer few benefits for larger species of ground fauna or forest-specialists, and support extra-regional species that may negatively affect forested environments. How applicable these small- scale results are to plantations in forested environments generally is uncertain, although the mechanisms identified as driving species responses

183 Chapter 7: General discussion

to plantations are not location-specific and are likely to apply in many environments.

While management to enhance biodiversity at the stand level and landscape levels is now employed in many commercial native plantations in

Australia, for the privately owned small-scale plantations that comprise more than 8% the plantation estate such extra management aimed at specifically increasing biodiversity is often too costly to implement (Alig 1998, Armsworth et al. 2004). In commercial systems valuing native plantations as faunal habitat and managing them as multi-use landscapes will incur some level of economic cost in establishment, extra management, reduced timber production and planning. Thus the decision to incorporate ecological concepts into the planning, establishment and management of native plantations must be undertaken with a mindfulness that the relative costs and benefits of such extra expense will vary with the ecological context of the plantations themselves. In the case where ecological management does not supply great benefits to biodiversity, it may provide a better conservation outcome to direct efforts and funding to other conservation measures.

184 References

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Appendices

Appendix 1

Table 1. Summary statistics for all bird species recorded from young and old plantations, logged and old growth native forests from twenty field sites on the Mid-North Coast of NSW between 2001 and 2002. Guild membership is specified by strata commonly used and diet. Notations are A= all strata, C=canopy, Gr= ground, G-L= ground-low, M-C= midstorey-canopy, G= granivore C= carnivore, H=herbivore, N= nectarivore, O= omnivore, I= insectivore, F= frugivore. Habitat associations are F = forest, W = woodland, G = generalist. Species indicated by * are excluded from analyses as they were only documented overflying field sites.

Family Species Old Growth Native Logged Native Old Plantation Young plantation Habitat Guild captures mean se captures mean se captures mean se captures mean se Acanthizidae Brown gerygone 13 2.6 0.32 ------F Ar, N/I Acanthizidae Brown thornbill 43 8.6 0.83 12 2.4 0.23 1 0.2 0.05 - - - F Ar, N/I Acanthizidae Striated thornbill 7 1.4 0.37 1 0.2 0.05 ------F C, N/I Acanthizidae White-browed scrubwren 25 5 0.30 6 1.2 0.20 2 0.4 0.11 6 1.2 0.21 F G-L, I Acanthizidae Yellow thornbill ------58 11.6 1.38 30 6 1.15 W C, F/O Acanthizidae Yellow-throated scrubwren ------1 0.2 0.05 F G-L, I Accipitridae Brown goshawk 7 1.4 0.14 ------F A, C/I Alcedinidae Azure kingfisher 2 0.4 0.07 ------G G-L, C Apodidae White-throated needletail* 1 0.2 0.05 - - - 9 1.8 0.32 5 1 0.27 W C, N/I Cacatuidae Glossy black cockatoo - - - 4 0.8 0.21 ------W C, G Cacatuidae Yellow-tailed black cockatoo 3 0.6 0.11 7 1.4 0.26 - - - 2 0.4 0.11 F C, G Campephagidae Black-faced cuckoo shrike 1 0.2 0.05 3 0.6 0.16 3 0.6 0.16 8 1.6 0.22 W C, F/O Campephagidae White-bellied cuckoo-shrike ------2 0.4 0.11 W C, N/I Caprimulgidae White-throated nightjar 1 0.2 0.05 5 1 0.12 11 2.2 0.20 1 0.2 0.05 W G-L, I Climacteridae Brown treecreeper 1 0.2 0.05 5 1 0.21 ------W Ar, N/I Climacteridae White-throated treecreeper 12 2.4 0.40 36 7.2 0.71 17 3.4 0.31 - - - F Ar, N/I Columbidae Brown pigeon 150 30 1.34 132 26.4 1.76 23 4.6 0.35 79 15.8 0.58 F M-C, N/I/F Columbidae Peaceful dove ------3 0.6 0.07 W G-L, G Columbidae Wompoo pigeon 2 0.4 0.11 ------F C, F/O Columbidae Wonga pigeon 9 1.8 0.13 2 0.4 0.11 ------F G-L, G Corvidae Australian Raven* ------2 0.4 0.11 6 1.2 0.21 W G-L, O Cracticidae Australian magpie - - - 3 0.6 0.11 1 0.2 0.05 14 2.8 0.41 W G-L, O Cracticidae Pied butcherbird - - - 3 0.6 0.11 13 2.6 0.29 4 0.8 0.15 W G-L, O Cracticidae Pied currawong 10 2 0.40 16 3.2 0.23 18 3.6 0.39 - - - W A, O Cuculidae Channel-billed cuckoo - - - 3 0.6 0.16 7 1.4 0.23 - - - W C, F/O Cuculidae Horsfield's bronze cuckoo - - - 3 0.6 0.11 - - - 6 1.2 0.21 W Ar, N/I Cuculidae Pheasant coucal - - - 1 0.2 0.05 - - - 6 1.2 0.15 W Gr, I Estrildidae Red-browed finch ------31 6.2 0.73 26 5.2 1.19 G G-L, G Halcyonidae Laughing kookaburra 4 0.8 0.10 7 1.4 0.31 22 4.4 0.61 8 1.6 0.20 W G-L, C Maluridae Red-backed wren ------16 3.2 0.62 G G-L, I Maluridae Superb wren 9 1.8 0.48 59 11.8 1.35 25 5 0.25 75 15 1.18 G G-L, I Maluridae Variagated wren 5 1 0.27 ------W G-L, I Megapodidae Australian brush turkey 3 0.6 0.07 5 1 0.15 ------F G-L, O Meliphagidae Bell miner ------6 1.2 0.26 W M-C, N/I/F Meliphagidae Eastern spinebill 4 0.8 0.21 1 0.2 0.05 - - - 2 0.4 0.07 F Ar, N/I Meliphagidae Lewins honeyeater 75 15 0.89 61 12.2 1.12 30 6 0.42 57 11.4 0.73 F M-C, N/I/F Meliphagidae Little friar bird 7 1.4 0.37 48 9.6 1.17 37 7.4 0.72 - - - F C, N/I Meliphagidae New holland honeyeater - - - 11 2.2 0.58 0 0 0.00 - - - W Ar, N/I Meliphagidae White-eared honeyeater - - - 2 0.4 0.11 ------F M-C, N/I/F Meliphagidae White-naped honeyeater 5 1 0.27 1 0.2 0.05 ------F Ar, N/I Meliphagidae Yellow-faced honeyeater 18 3.6 0.55 30 6 0.24 22 4.4 0.61 9 1.8 0.24 W C, F/O Menuridae Superb lyrebird 13 2.6 0.35 10 2 0.28 ------F Gr, I Monarchidae Black-faced monarch 29 5.8 0.41 21 4.2 0.48 - - - 2 0.4 0.11 F C, N/I Monarchidae Restless flycatcher - - - 1 0.2 0.05 ------W Ar, N/I Monarchidae Satin flycatcher 0 0 0.00 1 0.2 0.05 1 0.2 0.05 4 0.8 0.15 F C, N/I Nectariniidae Mistletoebird 6 1.2 0.20 0 0 0.00 10 2 0.40 7 1.4 0.25 W C, F/O Oriolidae Olive-backed oriole 6 1.2 0.26 1 0.2 0.05 2 0.4 0.07 10 2 0.17 F C, N/I Orthonychidae Logrunner 4 0.8 0.15 1 0.2 0.05 2 0.4 0.07 - - - W Gr, I Pachycephalidae Crested Bell bird - - - 3 0.6 0.16 7 1.4 0.37 - - - W C, N/I Pachycephalidae Golden whistler 93 18.6 0.80 56 11.2 0.57 8 1.6 0.36 3 0.6 0.11 F Ar, N/I Pachycephalidae Grey shrike thrush 25 5 0.47 23 4.6 0.38 11 2.2 0.18 24 4.8 0.65 F A, C/I pachycephalidae Rufous whistler 36 7.2 1.22 1 0.2 0.05 30 6 0.42 8 1.6 0.14 W C, N/I Paradisaeidae Paradise riflebird 2 0.4 0.07 ------F C, N/I Pardalotidae Spotted pardalote 2 0.4 0.11 ------W C, N/I Petroicidae Jacky winter ------7 1.4 0.31 - - - W G-L, I Petroicidae Eastern yellow robin 26 5.2 0.18 46 9.2 0.57 33 6.6 0.37 53 10.6 0.38 F G-L, I Petroicidae Rose robin - - - 1 0.2 0.05 ------F Ar, N/I Petroicidae Scarlet robin 26 5.2 0.85 - - - 4 0.8 0.21 15 3 0.57 W G-L, I Psittacidae Crimson rosella 13 2.6 0.42 10 2 0.47 ------F C, G Psittacidae King parrot 6 1.2 0.21 4 0.8 0.13 2 0.4 0.11 - - - F C, G Psittacidae Rainbow lorikeet 35 7 1.23 111 22.2 2.95 21 4.2 0.69 9 1.8 0.41 G C, N/I Psittacidae Scaly-breasted lorikeet 0 0 0.00 2 0.4 0.07 ------F C, N/I Psophodidae Eastern whipbird 60 12 1.06 13 2.6 0.20 17 3.4 0.34 3 0.6 0.11 F Gr, I Ptilonorhychidae Green catbird 14 2.8 0.32 1 0.2 0.05 ------F M-C, N/I/F Ptilonorhychidae Satin bowerbird 8 1.6 0.20 7 1.4 0.16 2 0.4 0.11 - - - F A, O Rhipiduridae Grey fantail 90 18 2.45 22 4.4 0.26 32 6.4 0.49 36 7.2 0.41 F Ar, N/I Rhipiduridae Rufous fantail 71 14.2 1.11 36 7.2 0.65 4 0.8 0.15 1 0.2 0.05 F G-L, I Strigidae Southern boobook owl 5 1 0.08 1 0.2 0.05 2 0.4 0.07 - - - G A, C/I Strigiformes Sooty owl 2 0.4 0.11 ------F A, C/I Zosteropidae Silvereye 0 0 0.00 ------64 12.8 0.90 G M-C, N/I/F 226

Appendices

Table 2. Summary statistics for mammal, frog and reptile species recorded from young and old plantations, logged and old growth native forests from twenty field sites on the Mid-North Coast of NSW between 2001 and 2002, and analyzed in chapter 3. Guild membership is specified by strata commonly used and diet. Notations are A= all strata, C=canopy, Gr= ground, G-L= ground-low, M-C= midstorey-canopy, G= granivore C= carnivore, H=herbivore, N= nectarivore, O= omnivore, I= insectivore, F= frugivore. Habitat associations are F = forest, W = woodland, G = generalist. Species indicated by * are excluded from analyses. Values represent total numbers of unique captures.

Taxon Species Old Growth Native Logged Native Old Plantation Young plantation Habitat Guild captures mean se captures mean se captures mean se captures mean se AssociationSpecification Mammals Antechinus stuartii 37 7.4 0.95 15 3 0.46 20 4 0.84 9 1.8 0.56 F Gr, I Antechinus swainsonii 1 0.2 0.13 4 0.8 0.52 0 0 0.00 0 0 0.00 F Gr, I Isoodon macrourus 1 0.2 0.13 4 0.8 0.38 6 1.2 0.32 4 0.8 0.24 W Gr, O Melomys cervinipes 8 1.6 0.39 ------F Gr, H Mus musculus ------7 1.4 0.33 G Gr, O Planigale maculata - - - 6 1.2 0.38 9 1.8 0.59 3 0.6 0.16 G Gr, I Pseudocheirus peregrinus* 7 1.4 0.48 ------F A, H/N Pteropus scapulatus* 10 2 0.82 12 2.4 0.72 ------G A, H/N Rattus fuscipes 76 15.2 1.53 59 11.8 1.34 32 6.4 1.54 1 0.2 0.13 F Gr, O Rattus lutreolus ------22 4.4 0.97 15 3 0.68 W Gr, H Sminthopsis murina ------7 1.4 0.44 5 0.6 0.16 W Gr, I Trichosurus vulpecula* - - - 2 0.4 0.16 4 0.8 0.32 - - - W A, H/N

Amphibians Crinia signifera - - - 1 0.2 0.20 2 0.4 0.24 - - - G Gr, I Limnodynastes peronii - - - 5 1 0.55 5 1 0.45 3 0.6 0.60 G Gr, I Mixophyes fasciolatus 2 0.4 0.24 ------F Gr, I Pseudophryne coriacea 28 5.6 4.13 53 10.6 2.69 41 8.2 5.58 - - - W Gr, I Uperolia fusca 3 0.6 0.24 4 0.8 0.37 8 1.6 0.93 2 0.4 0.24 F Gr, I

Reptiles Adrasteia amicula ------17 3.4 0.55 W Gr, I Amphibolurus muricatus ------2 0.4 0.11 3 0.6 0.11 W G-L, I Cacophis k reffti 1 0.2 0.09 ------F Gr, C Calyptotis ruficaudata 8 1.6 0.30 12 2.4 0.22 15 3 0.20 1 0.2 0.09 F Gr, I Cryptophis nigrescens ------1 0.2 0.09 G Gr, I Ctenotus robustus ------1 0.2 0.09 4 0.8 0.09 W Gr, I Cyclodomorphus gerrardi 3 0.6 0.11 ------F G-L, I Demanisa psammophis ------1 0.2 0.09 G Gr, C Drysdalia coronoides - - - 3 0.6 0.18 1 0.2 0.09 - - - G Gr, C Egernia mcpheei 4 0.8 0.32 ------F Gr, O Eulamprus murrayi 46 9.2 1.11 ------F Gr, I Eulamprus quyoii 1 0.2 0.09 ------F Gr, I Eulamprus tenuis 1 0.2 0.09 ------F G-L, I Hoplocephalus stephensi 2 0.4 0.11 ------F A, C-I Hypsilurus spinipes 6 1.2 0.21 3 0.6 0.18 ------F A, C-I Lampropholis delicata 16 3.2 0.35 21 4.2 0.54 26 5.2 0.38 4 0.8 0.16 G Gr, I Lampropholis guichenoti 7 1.4 0.26 12 2.4 0.41 40 8 0.83 15 3 0.44 G Gr, I Lerista muelleri 7 1.4 0.30 2 0.4 0.11 1 0.2 0.09 - - - W Gr, I Lialis burtonis ------2 0.4 0.11 G Gr, C Morelia spilota 4 0.8 0.26 ------F A, C-I Physignathus lesueurii ------2 0.4 0.11 - - - F Gr, O Pseudechis porphyriacus - - - 1 0.2 0.09 2 0.4 0.11 5 1 0.24 G Gr, C Ramphotyphlops nigrescens 3 0.6 0.18 7 1.4 0.11 ------W Gr, I Rhinoplocephalus nigrostriatus 0 2 W Gr, I Saltuarius swainii 8 1.6 0.18 ------F A, C-I Tiliqua scincoides - - - 1 0.2 0.09 1 0.2 0.09 2 0.4 0.11 G Gr, O Vermicella annulata 3 0.6 0.11 ------G Gr, C

227

Appendices

Appendix 2

Summary statistics for all herpetofaunal and mammal species recorded from young and old plantations, logged and old growth native forests from twenty field sites on the Mid-North Coast of NSW between 2002 and 2003 (as described in chapter 4). Species indicated by * were excluded from analyses. Values represent total numbers of unique captures.

Old growth Logged native Old plantation Young plantation Species N1 N2 N3 N4 N5 LN1 LN2 LN3 LN4 LN5 OP1 OP2 OP3 OP4 OP5 YP1 YP2 YP3 YP4 YP5 Mammals Antechinus stuartii 10 12 11 12 9 - 6 5 8 20 21 16 25 19 21 3 - - 3 4 Antechinus swainsonii - - 2 - - - 4 - - 3 ------Isoodon macrourus* - 1 - - - - 1 2 - - - 1 3 1 1 - 2 - 2 - Melomys cervinipes* 2 6 - 6 2 - 1 ------Mus musculus ------18 6 17 11 15 Planigale maculata - - - - - 1 3 2 - - 3 1 1 4 1 - 3 - - 1 Rattus fuscipes 28 32 28 35 56 27 26 35 30 38 9 - 21 25 23 - 7 7 - - Rattus lutreolus ------6 8 12 8 - 12 5 2 7 9 Sminthopsis murina ------4 7 3 2 - 4 4 3 - Reptiles Adrasteia amicula ------5 5 3 - 11 5 8 2 20 Amphibolurus muricatus ------2 3 - - 3 - 11 - 5 Cacophis kreffti* - 2 1 3 1 ------Calyptotis ruficaudata 3 3 5 2 - 2 5 4 3 3 4 7 5 4 1 - - - - - Concinnia tenuis* - - 1 1 ------Ctenotus robustus ------3 - - - - - 7 2 3 3 Drysdalia coroniodes* ------2 1 - 3 1 1 - - - - - 1 Egernia mcpheei* - - - 2 1 ------Eulamprus quyoii* 1 - - 1 1 ------Eulamprus murrayi* 23 12 10 10 11 ------Hemisphaerodon gerrardi* 6 - - - 2 - - - - - 1 ------Hypsilurus spinipes* 5 2 3 5 4 - 4 1 2 ------Lampropholis delicata 5 15 9 8 7 11 13 17 5 7 19 18 29 11 12 8 21 14 5 31 Lampropholis guichenoti ------14 37 6 - 11 44 14 6 43 Lerista mulleri 3 5 6 1 1 3 - 3 - - - - 1 - - - - - 1 2 Lialis burtonis* ------1 1 2 - - Pseudechis porphyriacus* ------1 - 2 - - - - Ramphotyphlops nigrescens* 1 1 - - 1 - 1 2 3 - - - 1 - - - 1 1 2 2 Rhinoplocephalus nigrescens* ------1 - 2 Saltuarius swainii* 3 1 2 2 8 ------Tiliqua scincoides ------1 - 2 - 4 - - 2 2 3 - - Amphibians Crinia signifera ------3 - - - 6 4 - - - 2 - - - - Limnodynastes peroni ------3 - 5 - 3 5 5 - - - 9 3 - 2 Mixophys fasciolatus* - 1 1 - 1 ------Pseudophryne coriacea 9 10 10 - - 15 24 16 17 17 15 26 61 12 8 - - - - 8 Uperolia fusca 1 ------2 3 2 - 6 3 - 2 - 2 1 - 1

228

Appendices

Appendix 3

Summary statistics for all herpetofaunal and mammal species recorded from young plantations and adjacent native forests ten field sites on the Mid-North Coast of NSW between 2002 and 2004 (as described in chapter 5). Values represent total numbers of unique captures.

Taxon Species Matrix Young Plantation Habit Mammals M1 M2 M3 M4 M5 YP1 YP2 YP3 YP4 YP5 Antechinus stuartii - - 1 6 6 - - - - 1 F Isoodon macroura ------2 - - - W Mus musculus - - - - - 12 10 10 6 3 G Planigale maculata ------1 - - - G Rattus fuscipes - - 4 1 25 - - 1 - - F Rattus lutreolus ------5 - 2 - W Sminthopsis murina - - 1 - - - - 3 2 - W Reptiles Adrasteia amicula 11 1 12 16 4 - 3 8 2 17 W Amphibolurus muricatus ------6 - 3 W Cacophis kreffti - - 2 - 3 - - - - - F Calyptotis ruficaudata 2 1 - 1 ------F Ctenotus robustus ------1 1 1 1 W Cyclodomorphus gerrardi - - 1 ------F Demansia psammophis - - - 1 1 - - - - 1 G Lampropholis delicata 12 2 14 3 2 - 3 6 1 15 G Lampropholis guichenoti 16 2 30 15 14 - 24 4 1 22 G Lerista mulleri 4 - 9 1 12 - 1 - 1 1 W Lialis burtonis - - 1 1 ------G Pseudechis porphyriacus - - - 1 ------G Ramphotyphlops nigrescens - - - 1 0 - 2 - - - W Rhinoplocephalus nigrescens - - - - 2 - - - - 2 W Tiliqua scincoides - - 4 1 1 - - 2 - - G Amphibians Crinia signifera 1 - - - 0 - - - - - G Limnodynastes peroni - - 2 - 2 - 1 1 - 1 W Pseudophryne coriacea 13 1 3 5 1 - v - - 1 W Uperolia fusca - - 1 - - - 1 1 - - F Species richness n=20 n=19

229