Assessing the Viability of Lowland Populations in a Fragmented Landscape

By

Emília Patrícia Medici

January 2010 To Dr. Maurício Pompilio, Dr. José Tadeu dos Santos and the entire staff of Clínica Campo Grande for saving my husband's life and ultimately allowing me to finish this thesis …

… and to the of the Atlantic Forest …

Abstract

Resumo

Acknowledgements

Table of Contents

Cha pter 1 General Introduction & Thesis Aims and Structure 1.1 General Introduction 002 1.2 Thesis Aims and Structure 013 Chapter 2 Study , Study Area, and General Methods 2.1 Introduction 017 2.2 The Lowland Tapir 017 2.3 The Atlantic Forest Biome 039 2.4 The Atlantic Forest of the Interior and the Pontal do Paranapanema Region 045 2.5 Study Site: Morro do Diabo State Park 049 2.6 General Methods 056 Chapter 3 Spatial Ecology and Intra-Specific Interactions 3.1 Introduction 059 3.2 Methods 063 3.3 Results 081 3.4 Discussion 107 Chapter 4 Determining Population Size 4.1 Introduction 122 4.2 Methods 125

4.3 Results 138 4.4 Discussion 147 Chapter 5 Spatial and Temporal Interactions between Tapirs and the Landscape 5.1 Introduction 166 5.2 Methods 169 5.3 Results 172 5.4 Discussion 187 Chapter 6 Population Viability Analysis (PVA): Using a Modelling Tool to Assess the Viability of Tapir Populations in a Fragmented Landscape 6.1 Introduction 197 6.2 Methods 199 6.3 Results 215 6.4 Discussion 222 Chapter 7 Conclusions & Conservation Recommendations 7.1 Viability of Lowland Tapir Populations in the Atlantic Forest of the Interior of the Pontal do Paranapanema Region 230 7.2 Conservation Recommendations: A First Step in Building a Regional Action Plan for the Conservation of Lowland Tapirs in the Pontal do Paranapanema Region 232 7.3 Determinant s for Long -Term Persistence of Populations across Fragmented Landscapes 239 References 243 Appendix I 270 Appendix II 271 Appendix III 273

List of Figures

Figure 2.1.

Figure 2.2.

Figure 2.3.

Figure 2.4. Figure 2.5. Figure 2.6.

Figure 2.7.

Figure 3.1. Figure 3.2. Figure 3.3.

Figure 3.4.

Figure 3.5. Figure 4.1.

Figure 4.2.

Figure 4.3. Figure 4.4.

Figure 4.5.

Figure 4.6.

Figure 5.1. Figure 5.2.

Figure 5.3.

Figure 5.4.

Figure 5.5.

Figure 5.6.

Figure 5.7.

Figure 6.1. Figure 6.2.

Figure 6.3.

Figure 6.4. Figure 6.5.

List of Tables

Table 2.1. Table 2.2.

Table 2.3. Table 2.4.

Table 2.5. Table 2.6.

Table 2.7. Table 2.8. Table 2.9. Table 2.10. Table 2.11.

Table 3.1.

Table 3.2. Table 3.3.

Table 3.4. Table 3.5.

Table 3.6. Table 3.7.

Table 3.8.

Table 3.9. Table 3.10.

Table 3.11.

Table 3.12.

Table 3.13.

Table 3.14. Table 3.15. Table 3.16.

Table 3.17.

Table 4.1.

Table 4.2. Table 4.3.

Table 4.4.

Table 4.5. Table 4.6.

Table 4.7. Table 5.1.

Table 5.2. Table 5.3. Table 5.4.

Table 5.5. Table 5.6.

Table 6.1.

Table 6.2. Table 6.3.

Table 6.4. Table 6.5.

Table 6.6.

Table 6.7.

Table 6.8.

Table 6.9.

Table 6.10.

Chapter 1

General Introduction & Thesis Aims and Structure

1.1 General Introduction 1.1.1 Habitat Fragmentation

1.1.2 Long-Term Viability of Animal Populations

1.1.3 Why is the tapir a good model?

1.1.4 Why is the Atlantic Forest of the Interior a good model?

1.2 Thesis Aims and Structure • •

Chapter 1

Chapter 2

Chapter 3

Chapter 4

Chapter 5

Chapter 6

Chapter 7

Chapter 2

Study Species, Study Area and General Methods

2.1 Introduction

2.2 The Lowland Tapir 2.2.1 and Evolution 2.2.2 Morphology

Table 2.1. Weight N AV SD Min Max ADULT FEMALES 233 ADULT MALES 208 SUB-ADULT FEMALES 200 SUB-ADULT MALES 195

Table 2.2. ADULT Females ADULT Males Corporal Measurements Significant N AV SD Min Max N AV SD Min Max Differences Neck circumference* 86 79 P=0.021

Head length 48 48

Head width 42 38

Fu ll length (dorsum)* 221 205 P=<0.001

Full length (lateral)* 202 186 P=0.036

Front leg length 70 65

Rear leg length 72 65

Thorax circumference* 144 129 P=0.004

Abdomen circumference 169 152

Front height* 108 101 P=0.024

Rear height* 110 101 P=0.008

Ear length 13 13

Distance between eyes 23 24

Table 2.3. SUB-ADULT Females SUB-ADULT Males Corporal Measurements Significant N AV SD Min Max N AV SD Min Max Differences Neck circumference 83 81

Head length 46 47

Head width 36 32

Full length (dorsum) 213 198

Full length (lateral) 165 184

Front leg length 69 64

Rear leg length 68 65

Thorax circumference 140 118

Abdomen circumference 144 142

Front height 103 97

Rear height 108 102

Ear length 16 13

Distance between eyes 23 20

Table 2.4. JUVENILES Corporal Measurements N AV SD Min Max Neck circumference 73

Head length 44

Head width 34

Full length (dorsum) 176

Full length (side) 158

Front leg length 66

Rear leg length 70

Thorax circumference 110

Abdomen circumference 132

Rear height 89

Fr ont height 89

Ear length 13

Distance between eyes 14

Figure 2.1.

Figure 2.2. 2.2.3 Geographic Distribution

Figure 2.3.

Table 2.5. General Biomes Brazilian Biomes Colombian Biomes Suriname Biomes Venezuelan Biomes

2.2.4 Ecology and Behaviour 2.2.4.1 Habitat Requirements

Figure 2.4.

2.2.4.2 Feeding Behaviour

2.2.4.3 Spatial Requirements 2.2.4.4 Population Density

Table 2.6.

Location Biome Country Method Home Range Size Reference 39.14 km² 16.4 km² 48.78 km² ≤ 4 km² 2.81 km² 2.18 km² 25-75% 0.97-3.74 km² 1.03-4.83 km² 0.50-5.78 km² 2.48 km² 32-55% 43.5% 1.06-3.86 km² 2.61 km² 0.14-0.88 km² 0.05-0.33 km² 2.86 km² 2.11 km² 0.95-3.53 km² 2.25 km² 2.53 km² 1.66 km²

Table 2.7. Location Biome Country Method Density Estimate Reference 0.41 0.47 0.30 2.20-2.50 0.11-0.52 0.62 0.23 0.5 < 0.5 5.0 0.40 0.60 0.20 0.06-0.08 0.31

3.3-3.7 0.64 0.55 0.21 0.55 0.58 2.04-2.34 0.50 0.22-0.80 0.71 0.6 0.8 0.53 1.60 1.60

2.2.4.5 Social Behaviour and Activity Patterns 2.2.4.6 Reproduction - -

2.2.4.7 Predators 2.2.5 Status and Threats

Table 2.8. General Biomes HUN HDA REX FSP CR DIS RK Brazilian Biomes HUN HDA REX FSP CR DIS RK Colombian Biomes HUN HDA REX FSP CR DIS RK Suriname Biomes HUN HDA REX FSP CR DIS RK Venezuelan Biomes HUN HDA REX FSP CR DIS RK Table 2.9. Threats Ranking

2.3 The Atlantic Forest Biome 2.3.1 Historical Outline 2.3.2 Ecological Outline

2.3.3 Current

Table 2.10.

Original Amount % of Cover Remaining Original Reference Comments Cover

2.4 The Atlantic Forest of the Interior and the Pontal do Paranapanema Region

Figure 2.5. ´

Figure 2.6.

2.5 Study Site: Morro do Diabo State Park 2.5.1 Location and Legal Protection 2.5.2 Landscape Matrix 2.5.3 Climate Table 2.11. Temperature Relative Humidity Monthly Month Min Mean Max Min Mean Max Rainfall Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total 1,347 MEAN 21.9 112 MAX MIN 2.5.4 Abiotic Factors

2.5.5 Biotic Factors

2.5.5.1 Fauna 2.5.5.2 Flora 1) Tall mature forest: 2) Low mature forest:

3) Secondary forest in early stages of regeneration: 4) Secondary forest in late stages of regeneration:

5) Cerrado (Savannah):

6) Riparian forest along the Ribeirão Bonito Stream: 7) Riparian forest along the Paranapanema River:

8) Low, dense forest in various stages of regeneration:

9) Temporary lakes: 10) Agricultural and pastoral land:

Figure 2.7.

2.6 General Methods 2.6.1 Logistics and Personnel 2.6.2 Field Methods

2.6.3 Timeframe of Study 2.6.4 Research Permits

Chapter 3

Spatial Ecology and Intra-Specific Interactions

3.1 Introduction

3.2 Methods 3.2.1 Radio-Telemetry

3.2.2 Capture Sites

2

1

4

3

Figure 3.1. 3.2.3 Capture Methods

3.2.3.1 Iron Box Traps

3.2.3.2 Pitfalls 3.2.3.3 Anaesthetic Dart Shooting

3.2.3.4 Wooden Corrals

1 2

3 4

Figure 3.2.

3.2.3.5 Trapping Effort and Trapping Success

Table 3.1. Trapping Tapir Trapping Capture Method Effort Captures Success Ir on box traps Pitfalls Anaesthetic dart shooting Wooden corrals

3.2.4 Chemical Immobilisation

Table 3.2. Parameter of Anaesthesia Monitoring N AV SD Heart rate Heart rate Rate of ventilations Rate of ventilations Oxygen saturation % Oxygen saturation % Temperature ºC Temperature ºC

3.2.5 Manipulation and Collection of Biological Samples

3.2.6 Captured Individuals

3.2.7 Telemetry Equipment

1

2

Figure 3.3.

3.2.8 Telemetry Data Collection

3.2.9 Telemetry Data Screening

3.2.10 Tapir Spatial Ecology 3.2.10.1 Home Range Size

3.2.10.2 Maximum Distance Moved (MDM) 3.2.10.3 Size of Core Areas of Use 3.2.10.4 Seasonal Home Range

3.2.10.5 Determination of Home Range Size over Time 3.2.11 Tapir Intra-Specific Interactions 3.2.11.1 Home Range Overlap

3.2.11.2 Overlap of Core Areas of Use 3.2.12 Statistical Significance

3.3 Results 3.3.1 Tapir Spatial Ecology 3.3.1.1 Home Range Size

Table 3.3. Telemetry Period of Timeframe of Individual Sex Age Class Area MDSP KDE 95% MCP 95% Locations Radio-Tracking Radio-Tracking JOANA PAULETE CHU-CHU 14.2 PATRICIA DOCINHO JAMES BOND GEORGETE 1.0 CHICO 13.2 JULIA CIDAO TONHA TINA MAMA ESPERTA 1.1 LUIZINHO XUXA JOAO BABY DEDINHO

Table 3.4. 19 tapirs Group 2 (14 tapirs) KDE 95% MCP 95% KDE 95% MCP 95% 4.9 4.1 4.7 4.4 ALL INDIVIDUALS 4.7 3.84 5.0 3.8 FEMALES 5.4 4.7 3.6 6.5 MALES 3.9 3.7 3.3 3.7 ADULTS Adult Females Adult Males 8.5 5.7 12.7 8.1 SUB-ADULTS Sub-Adult Females Sub-Adult Males 3.3.1.2 Maximum Distance Moved (MDM)

Table 3.5. Telemetry Period of Timeframe of Individual Sex Age Class Area MDSP MCP 95% MDM Locations Radio-Tracking Radio-Tracking JOANA PAULETE CHU-CHU 6,112 PATRICIA DOCINHO JAMES BOND GEORGETE CHICO JULIA CIDAO TONHA TINA MAMA 1,595 ESPERTA LUIZINHO XUXA JOAO BABY DEDINHO Mean MDM 3,233 Mean MDM Females 3,032 Mean MDM Males 3,667 Mean MDM Adults 2,949 Mean MDM Sub-Adults 4,294

3.3.1.3 Size of Core Areas of Use

Table 3.6. Timeframe of Individual Sex Age Class Area MDSP KDE 95% KDE 50% KDE 25% Radio-Tracking JOANA PAULETE CHU-CHU 2.5 (2) PATRICIA 0.9 (2) DOCINHO JAMES BOND GEORGETE CHICO JULIA CIDAO TONHA TINA MAMA ESPERTA LUIZINHO XUXA JOAO DEDINHO BABY 0.1 (3) 0.03 (1)

Table 3.7. KDE 50% KDE 25% 0.8 0.3 ALL INDIVIDUALS 0.8 0.3 FEMALES 0.9 0.3 MALES 0.6 0.2 ADULTS Adult Females Adult Males 1.5 0.5 SUB-ADULTS Sub-Adult Females Sub-Adult Males

3.3.1.4 Seasonal Home Range and Core Areas of Use

Table 3.8. Home Core Range Area Individual Tapir Season/Year KDE 95% KDE 50% KDE 25% JOANA (A;F) 0.4 0.01 PAULETE (A;F) 0.04 0.01 0.03 0.01 CHU-CHU (SA;F) 0.4 7.1 1.2 16.7 2.3 0.7 PATRICIA (SA;F) DOCINHO (A;F) JAMES BOND (A;M) GEORGETE (A;F) CHICO (A;M) JULIA (A;F) CIDAO (A;M) TONHA (A;F)

TINA (A;F) MAMA (A;F) ESPERTA (A;F) 0.3

Table 3.9.

Wet Season Dry Season

KDE 95% KDE 50% KDE 25% KDE 95% KDE 50% KDE 25% 2.5 0.4 0.1 2.6 0.4 0.1 ALL INDIVIDUALS 2.2 0.4 0.1 2.6 0.4 0.1 Females 4.0 0.6 0.2 2.3 0.4 0.1 Males 2.1 0.3 0.1 1.8 0.3 0.1 Adults 4.2 0.7 0.2 6.6 1.1 0.4 Sub-Adults

3.3.1.5 Determination of Home Range Size over Time

Table 3.10. Individual Tapir Period of Telemetry Annual Area Period Radio-Tracking Locations KDE95% JOANA (A;F) PAULETE (A;F) CHU-CHU (SA;F) 8.2 PATRICIA (SA;F) CHICO (A;M) TONHA (A;F) TINA (A;F) MAMA (A;F) DOCINHO (A;F) JAMES BOND (A;M) GEORGETE (A;F) ESPERTA (A;F) 0.4

Table 3.11. Annual Area KDE95% 2.5 ALL INDIVIDUALS 2.5 Females 2.4 Males 2.1 Adults 4.1 Sub-Adults Table 3.12. Increase in Individual Tapir Period of Estimate of Radio- Cumulative Home Period Tracking Area of Use Overlap Range Size JOANA (A;F) 3.4 1.3

0.3 5.0 PAULETE (A;F) 0.1 0.0 CHU-CHU (SA;F) 2.3 6.3

2.4 PATRICIA (SA;F) 0.7 2.7

3.0 CHICO (A;M) 2.7 2.8

0.9 TONHA (A;F) 2.1 0.1

0.1 TINA (A;F) 3.5 0.1 MAMA (A;F) 2.1 0.2 DOCINHO (A;F) 0.6 2.5 JAMES BOND (A;M) 1.5 0.1 GEORGETE (A;F) 2.2 0.0 ESPERTA (A;F) 1.0 0.2

Paulete (A;F) - WEST BORDER Esperta (A;F) - NORTHWEST BORDER 33 Months 22 Months

Home Range (ha) Home Range (ha) Number of Months Number of Months

James (A;M) - SOUTHEAST BORDER Georgete (A;F) - SOUTHEAST BORDER 29 Months 25 Months

Home Range (ha) Home Range (ha) Number of Months Number of Months

Tonha (A;F) - CENTRE Tina (A;F) - CENTRE 33 Months 27 Months

500 Home Range (ha) Home Range (ha) 0 Number of Months Number of Months

Figure 3.4.

3.3.1.6 Tapir Movements throughout the Fragmented Landscape of the Pontal do Paranapanema Region

1 km 1

2 Figure 3.5.

3.3.2 Intra-Specific Interactions 3.3.2.1 Home Range Overlap Table 3.13. Female - Female Female - Male Male - Male Overlap Overlap Overlap Overlap Overlap Overlap 33.4 2.0 34.9 1.5 43.2 1.9

Table 3.14. Adult - Adult Sub -Adult - Sub -Adult Adult - Sub -Adult Overlap Overlap Overlap Overlap Overlap Overlap 32.2 1.2 39.0 3.0 37.2 2.2

3.3.2.2 Overlap of Core Areas of Use

Table 3.15. Female - Female Female - Male Male - Male 50% Overlap Overlap Overlap Overlap Overlap Overlap 21.4 0.3 18.9 0.1 12.9 0.1 Female - Female Female - Male Male - Male 25% Overlap Overlap Overlap Overlap Overlap Overlap 23.1 0.1 19.4 0.03

Table 3.16. Adult - Adult Sub -Adult - Sub -Adult Adult - Sub -Adult 50% Overlap Overlap Overlap Overlap Overlap Overlap 17.2 0.1 32.0 0.8 22.4 0.2 Adult - Adult Sub -Adult - Sub -Adult Adult - Sub -Adult 25% Overlap Overlap Overlap Overlap Overlap Overlap

24.0 0.03 13.0 0.04

3.3.2.3 Social Organisation

Table 3.17. HR Overlap CA Overlap CA Overlap Timeframe of Female-Female Area MDSP KDE 95% KDE 50% KDE 25% Radio-Tracking HR Overlap CA Overlap CA Overlap Timeframe of Male-Male Area KDE 95% KDE 50% KDE 25% MDSP Radio-Tracking

HR Overlap CA Overlap CA Overlap Timeframe of Female-Male Area KDE 95% KDE 50% KDE 25% MDSP Radio-Tracking

3.4 Discussion 3.4.1 Use of Radio-Telemetry for Studying Tapirs 3.4.1.1 Tapir Capture and Chemical Restraint

3.4.1.2 Timeframe of Telemetry Monitoring for Tapir Studies

3.4.2 Methodological Considerations on the Estimate of Tapir Home Range Size

3.4.3 Spatial Ecology of Tapirs in Morro do Diabo State Park 3.4.3.1 Home Range Size

3.4.3.2 Home Range Internal Structure: Core Areas of Use

3.4.3.3 Seasonal Variations in Home Range and Core Areas of Use

3.4.3.4 Tapir Movements throughout the Fragmented Landscape of the Pontal do Paranapanema Region

3.4.4 Tapir Intra-Specific Interactions in Morro do Diabo State Park 3.4.4.1 Tapir Spatial Territoriality in Morro do Diabo State Park

3.4.4.2 General Inferences about Tapir Social Organisation in Morro do Diabo State Park

3.4.5 Lowland Tapir Spatial Requirements and Implications for Population Viability

Chapter 4

Determining Population Size

4.1 Introduction

4.2 Methods • • • 4.2.1 Estimate of Tapir Population Density using Radio-Telemetry Data 4.2.2 Nocturnal Line-Transect Sampling 4.2.2.1 Line-Transect Sampling Methodology

• • • • •

4.2.2.2 Line-Transect Sampling Data Collection

77

44 66 33 88 55 11 22

Figure 4.1. 4.2.2.3 Estimate of Tapir Population Density using DISTANCE Analysis

Number of Encounters of Number

Distance from the Line-Transect (m) Figure 4.2. 4.2.3 Footprint Identification Technique (FIT) 4.2.3.1 Development of the FIT Algorithm for Lowland Tapirs

1

2

3

Figure 4.3.

Table 4.1. Number Average Facility N of photos number of taken footprints/set

1 2

Figure 4.4.

89

88

87

86 ChuvaA RCV

Canonical2 85 ChuvaB

84

83

82 51 52 53 54 55 56 57 58 Canonical1

1

43

42

41 Rosinha 40

39 RCV 38 Canonical2 Chuva A Chuva 37

36

35

34 -24 -23 -22 -21 -20 -19 -18 -17 Canonical1

2 Figure 4.5.

4.2.3.2 Validation of the FIT Algorithm for Lowland Tapirs 4.2.3.3 Estimate of Tapir Population Density using FIT

4.2.4 Estimates of Lowland Tapir Population Size 4.2.5 Comparisons between Density Estimators

4.3 Results 4.3.1 Estimate of Tapir Population Density using Radio-Telemetry Data

Table 4.2. Home Range Density Population Size Overlap ESA Estimate Size KDE95% 4.7 0.21 79 Method 1 MCP95% 4.4 0.23 85 KDE95% Method 2 4.7 37% 0.34 126 KDE95% 0.33 122

Method 3 MCP95% 0.43 159

All captured tapirs 0.32 118 Method 4

4.3.2 Estimate of Tapir Population Density using Nocturnal Line- Transect Sampling Table 4.3. Density Estimate SE % CV 95% CI Population Size 1.35 499

4.3.3 Estimate of Tapir Population Density using Footprint Identification Technique (FIT)

4.3.3.1 FIT Identification of Individual Tapirs

≤ ≤ Table 4.4. Tapir Footprint Set # Date of Gender Age Class ID & Relatedness Collection 1 01 2 02 05 3 03 21 4 04 22 5 06 6 07 7 08 09 14 15 16 18 19 27 8 10 9 11 10 12 13 17 23 26 11 20 12 24 13 25 4.3.3.2 Estimate of Tapir Population Density using FIT Table 4.5. Density Population ½MMDM ESA Estimate Size Tapirs Footprinted 0.43 159 Method 1 Radio-tracked Tapirs 0.26 96 Method 2 Traps/Recaptures 0.36 133 Method 3

4.3.4 Estimate of Lowland Tapir Population Sizes in Morro do Diabo State Park and Surrounding Atlantic Forest Fragments of the Pontal do Paranapanema Region

SM SP SMSP

SM C SMM LN ASEA

PB

RB

MDSP

Figure 4.6.

Table 4.6. Population Size MDSP & Forest Fragments Area Radio -Telemetry FIT Line -Transect Morro do Diabo State Park (MDSP) 370 126 159 237

Total number of Tapirs in Forest Fragments 22 27 40

Total number of Tapirs in the Pontal do Paranapanema Region 148 186 277 (MDSP + fragments)

4.3.5 Comparison between Density Estimators Table 4.7. Density Estimator Cost Personnel Field Effort Score

0 3 6 4.4 Discussion 4.4.1 Methodological Considerations on the Estimate of Tapir Population Size 4.4.1.1 Comparing Density Estimators: Cost, Personnel, and Field Effort

4.4.1.2 Comparing Density Estimators: Invasiveness of the Method

4.4.1.3 Comparing Density Estimators: Accuracy of Estimates

4.4.2 Lowland Tapir Population Density in Morro do Diabo State Park

4.4.3 Lowland Tapir Population Sizes in the Atlantic Forests of the Pontal do Paranapanema Region and Implications for Population Viability

Chapter 5

Spatial and Temporal Interactions between Tapirs and the Landscape

5.1 Introduction

5.2 Methods

5.2.1 Radio-Telemetry

5.2.2 Habitat Composition within Tapir Home Ranges and Core Areas of Use

5.2.3 Tapir Habitat Selection 5.2.3.1 Habitat Availability 5.2.3.2 Habitat Use

5.2.3.3 Habitat Selection Index

Wi = oi /πi oi πi 5.2.4 Tapir Activity Patterns

5.2.5 Patterns of Tapir Habitat Use in Relation of Activity 5.3 Results 5.3.1 Habitat Composition in Tapir Home Ranges and Core Areas of Use

Table 5.1.

West Border Habitat HR Habitat CA Habitat CA Type KDE 95% Type KDE 50% Type KDE 25% TMF TMF RFRB AGPL RFRB TMF LMF LDFV LDFV RFRB LMF LMF SFLSR SFESR SFESR LDFV AGPL AGPL SFESR SFLSR SFLSR

Southeast Border Habitat HR Habitat CA Habitat CA Type KDE 95% Type KDE 50% Type KDE 25% RFPR RFPR RFPR TMF SFLSR SFLSR SFLSR SFESR TMF AGPL TMF SFESR SFESR AGPL AGPL TL TL LDFV LDFV LDFV TL

Centre Habitat HR Habitat CA Habitat CA Type KDE 95% Type KDE 50% Type KDE 25% TMF TMF TMF SFLSR SFLSR SFLSR SFESR SFESR SFESR LDFV LDFV LDFV

Northwest Border Habitat HR Habitat CA Habitat CA Type KDE 95% Type KDE 50% Type KDE 25% TMF TMF TMF SFLSR SFLSR SFLSR AGPL SFESR SFESR SFESR AGPL LDFV LMF LMF AGPL LDFV LDFV LMF

Table 5.2.

West Border Habitat Type HR -Dry KDE 95% Habitat Type HR -Wet KDE 95% TMF TMF AGPL AGPL SFLSR SFLSR RFRB RFRB LMF LMF LDFV LDFV SFESR SFESR

Southeast Border Habitat Type HR -Dry KDE 95% Habitat Type HR -Wet KDE 95% RFPR RFPR TMF TMF SFLSR SFLSR AGPL AGPL SFESR SFESR TL TL LDFV LDFV

Centre Habitat Type HR -Dry KDE 95% Habitat Type HR -Wet KDE 95% TMF TMF SFLSR SFLSR SFESR SFESR LDFV LDFV

Northwest Border Habitat Type HR -Dry KDE 95% Habitat Type HR -Wet KDE 95% SFLSR TMF TMF SFLSR AGPL LMF SFESR AGPL LMF SFESR LDFV LDFV

5.3.2 Tapir Habitat Selection

5.3.2.1 Habitat Availability and Habitat Use

Table 5.3. Area Area Used Area Available MDSP N (KDE95%) ½MMDM (KDE95%+Buffer) West Border Southeast Border Centre Northwest Border

Table 5.4. West Border Habitat Habitat Habitat Habitat Use Habitat Use Habitat Use Type Availability Availability Home Range Wet Season Dry Season AGPL TMF 29.6 28.0 31.2 SFLSR LMF SFESR LDFV RFRB TL 0 0 0

Southeast Border Habitat Habitat Habitat Habitat Use Habitat Use Habitat Use Type Availability Availability Home Range Wet Season Dry Season AGPL TMF SFLSR RFPR 48.7 50.2 46.9 SFESR TL LDFV 0.5 0.6 0.4

Centre Habitat Habitat Habitat Habitat Use Habitat Use Habitat Use Type Availability Availability Home Range Wet Season Dry Season TMF 72.6 78.2 69.7 SFLSR AGPL 0 0 0 SFESR LDFV 0

Northwest Border Habitat Habitat Habitat Habitat Use Habitat Use Habitat Use Type Availability Availability Home Range Wet Season Dry Season TMF 46.1 56.5 AGPL 0 SFLSR 54.3 SFESR LMF LDFV 0 0 0

5.3.2.2 Habitat Selection Index

Table 5.5. West Border Habitat Habitat Habitat Selection P Type Used Available Index Wi (0.00625) AGPL 0.000 TMF SFLSR 1.069 LMF 1.709 SFESR 2.488 LDFV 2.942 RFRB 8.043 0.000 TL Southeast Border Habitat Habitat Habitat Selection P Type Used Available Index Wi (0.00714) AGPL 0.000 TMF SFLSR 1.268 RFPR 4.416 0.000 SFESR 1.112 TL LDFV 2.206 Centre Habitat Habitat Habitat Selection Type Used Available Index Wi P (0.01) TMF 1.261 0.001 SFLSR AGPL 0.000 SFESR LDFV 0.000 Northwest Border Habitat Habitat Habitat Selection P Type Used Available Index Wi (0.00833) TMF 1.316 AGPL 0.000 SFLSR 2.181 0.000 SFESR 0.000 LMF LDFV 0.000

5.3.3 Tapir Activity Patterns

06:00-06:59 07:00-07:59 70 05:00-05:59 08:00-08:59 04:00-04:59 60 03:00-03:59 09:00-09:59 50

40 10:00-10:59 02:00-02:59 30

11:00-11:59 20 01:00-01:59

10

12:00-12:59 0 24:00-24:59 70 60 50 40 30 20 10 0 10 20 30 40 50 60 70 10

13:00-13:59 20 23:00-23:59

30 14:00-14:59 22:00-22:59 40

50 21:00-21:59 15:00-15:59 60 16:00-16:59 20:00-20:59 70 17:00-17:59 19:00-19:59 18:00-18:59

Figure 5.1.

5.3.3.1 Activity Patterns in Different Seasons

06:00-06:59 07:00-07:59 70 05:00-05:59 08:00-08:59 04:00-04:59 60 03:00-03:59 09:00-09:59 50

40 10:00-10:59 02:00-02:59 30

11:00-11:59 20 01:00-01:59

10

12:00-12:59 0 24:00-24:59 70 60 50 40 30 20 10 0 10 20 30 40 50 60 70 10

13:00-13:59 20 23:00-23:59

30 14:00-14:59 22:00-22:59 40

50 21:00-21:59 15:00-15:59 60 16:00-16:59 20:00-20:59 70 17:00-17:59 19:00-19:59 18:00-18:59

Figure 5.2.

06:00-06:59 07:00-07:59 70 05:00-05:59 08:00-08:59 04:00-04:59 60 03:00-03:59 09:00-09:59 50

40 10:00-10:59 02:00-02:59 30

11:00-11:59 20 01:00-01:59

10

12:00-12:59 0 24:00-24:59 70 60 50 40 30 20 10 0 10 20 30 40 50 60 70 10

13:00-13:59 20 23:00-23:59

30 14:00-14:59 22:00-22:59 40

50 21:00-21:59 15:00-15:59 60 16:00-16:59 20:00-20:59 70 17:00-17:59 19:00-19:59 18:00-18:59

Figure 5.3.

5.3.3.2 Activity Patterns of Different Sexes

06:00-06:59 07:00-07:59 70 05:00-05:59 08:00-08:59 04:00-04:59 60

03:00-03:59 09:00-09:59 50

40 10:00-10:59 02:00-02:59 30

11:00-11:59 20 01:00-01:59

10

12:00-12:59 0 24:00-24:59 70 60 50 40 30 20 10 0 10 20 30 40 50 60 70 10

13:00-13:59 20 23:00-23:59

30 14:00-14:59 22:00-22:59 40

50 21:00-21:59 15:00-15:59 60 16:00-16:59 20:00-20:59 70 17:00-17:59 19:00-19:59 18:00-18:59

Figure 5.4.

06:00-06:59 07:00-07:59 70 05:00-05:59 08:00-08:59 04:00-04:59 60 03:00-03:59 09:00-09:59 50

40 10:00-10:59 02:00-02:59 30

11:00-11:59 20 01:00-01:59

10

12:00-12:59 0 24:00-24:59 70 60 50 40 30 20 10 0 10 20 30 40 50 60 70 10

13:00-13:59 20 23:00-23:59

30 14:00-14:59 22:00-22:59 40

50 21:00-21:59 15:00-15:59 60 16:00-16:59 20:00-20:59 70 17:00-17:59 19:00-19:59 18:00-18:59

Figure 5.5.

5.3.3.3 Activity Patterns of Different Age Classes

06:00-06:59 07:00-07:59 70 05:00-05:59 08:00-08:59 04:00-04:59 60 03:00-03:59 09:00-09:59 50

40 10:00-10:59 02:00-02:59 30

11:00-11:59 20 01:00-01:59

10

12:00-12:59 0 24:00-24:59 70 60 50 40 30 20 10 0 10 20 30 40 50 60 70 10

13:00-13:59 20 23:00-23:59

30 14:00-14:59 22:00-22:59 40

50 21:00-21:59 15:00-15:59 60 16:00-16:59 20:00-20:59 70 17:00-17:59 19:00-19:59 18:00-18:59

Figure 5.6.

06:00-06:59 07:00-07:59 70 05:00-05:59 08:00-08:59 04:00-04:59 60 03:00-03:59 09:00-09:59 50

40 10:00-10:59 02:00-02:59 30

11:00-11:59 20 01:00-01:59

10

12:00-12:59 0 24:00-24:59 70 60 50 40 30 20 10 0 10 20 30 40 50 60 70 10

13:00-13:59 20 23:00-23:59

30 14:00-14:59 22:00-22:59 40

50 21:00-21:59 15:00-15:59 60 16:00-16:59 20:00-20:59 70 17:00-17:59 19:00-19:59 18:00-18:59

Figure 5.7. 5.3.4 Patterns of Tapir Habitat Use in Relation to Activity

Table 5.6. West Bor der Home Range Wet Season Dry Season Habitat Type %ACT Habitat Type %ACT Habitat Type %ACT TMF TMF TMF RFRB RFRB RFRB SFESR LDFV SFESR LDFV SFESR LMF SFLSR SFLSR LDFV LMF LMF SFLSR AGPL AGPL AGPL Habitat Type %INA Habitat Type %INA Habitat Type %INA TMF RFRB TMF RFRB TMF RFRB LMF LMF LMF LDFV SFESR LDFV SFESR LDFV SFESR AGPL AGPL AGPL SFLSR SFLSR SFLSR Southeast Border Home Range Wet Season Dry Season Habitat Type %ACT Habitat Type %ACT Habitat Type %ACT RFPR RFPR RFPR TMF TMF TMF SFLSR SFLSR SFLSR SFESR SFESR SFESR AGPL AGPL TL TL TL AGPL LDFV LDFV LDFV Habitat Type %INA Habitat Type %INA Habitat Type %INA RFPR RFPR RFPR TMF TMF SFLSR SFLSR SFESR TMF SFESR SFLSR SFESR AGPL AGPL AGPL LDFV LDFV TL TL TL LDFV Centre Home Range Wet Season Dry Season Habitat Type %ACT Habitat Type %ACT Habitat Type %ACT TMF TMF TMF SFLSR SFLSR SFLSR SFESR SFESR SFESR LDFV LDFV LDFV Habitat Type %INA Habitat Type %INA Habitat Type %INA TMF TMF TMF SFLSR SFLSR SFLSR SFESR SFESR SFESR LDFV LDFV LDFV Nor thwest Border Home Range Wet Season Dry Season Habitat Type %ACT Habitat Type %ACT Habitat Type %ACT TMF TMF SFLSR SFLSR SFLSR TMF SFESR SFESR SFESR LMF LMF AGPL AGPL AGPL LMF Habitat Type %INA Habitat Type %INA Habitat Type %INA SFLSR TMF SFLSR TMF SFLSR TMF SFESR SFESR SFESR AGPL AGPL AGPL LMF LMF LMF

5.4 Discussion 5.4.1 Spatial Interactions between Lowland Tapirs and the Landscape

5.4.1.1 Habitat Composition in Tapir Home Ranges and Core Areas of Use

5.4.1.2 Tapir Habitat Use and Habitat Selection

5.4.2 Temporal Interactions between Lowland Tapirs and the Landscape

5.4.2.1 Tapir Activity Patterns

5.4.2.2 Patterns of Tapir Habitat Use in Relation to Activity 5.4.3 Spatial and Temporal Interactions between Lowland Tapirs and the Landscape and Implications for Population Viability

Chapter 6

Population Viability Analysis (PVA): Using a Modelling Tool to Assess the Viability of Tapir Populations in a Fragmented Landscape

6.1 Introduction

6.2 Methods 6.2.1 VORTEX Simulation Model

6.2.2 VORTEX Input Parameters: Baseline Model

6.2.2.1 Scenario Settings • Duration of Simulation:

• Number of Iterations:

6.2.2.2 Species Description • Definition of :

• Concordance of Environmental Variation (EV) between Reproductive Rates and Survival Rates:

• Inbreeding Depression: 6.2.2.3 Reproductive System and Rates • Breeding System: .

• Age of First Reproduction:

• Maximum Age of Reproduction:

• Maximum Number of Offspring per Year:

• Maximum Litter Size:

• • Sex Ratio at Birth:

• Female Breeding Success (Percentage of Adult Females Breeding):

• Environmental Variation in Breeding:

• Density Dependent Reproduction:

• Mate Monopolisation:

6.2.2.4 Mortality Rates

Table 6.1.

Age Class Mortality Rate Standard Deviation

6.2.2.5 Population Description • Number of Populations:

• Dispersal among Populations:

• Initial Population Size (N):

• Carrying Capacity (K):

• Catastrophe:

• Harvest:

• Supplementation: Table 6.2. Parameter Baseline Value 126 159 237 ♀ ♂ ♀ ♂

6.2.3 Sensitivity Analysis

Table 6.3. Parameter Low Baseline Highest

♀ ♂

6.2.4 Minimum Viable Population Analysis

• Average probability that the population will go extinct (P(E) 100 ):

• The genetic diversity of the extant populations (GD): 6.2.5 Modelling of Alternative Scenarios

6.2.5.1 Impact of Fire

6.2.5.2 Impact of Road-Kill

1

2

Figure 6.1.

6.2.5.3 Impact of Infectious Disease

6.2.5.4 Metapopulation Scenario

SMSP

SMM LN SMC

ASEA

PB

RB MDSP

Figure 6.2. 6.2.5.5 Establishment of Corridors

Figure 6.3.

Table 6.4. Population Size MDSP & Forest Fragments Area Radio -Telemetry FIT Line -Transect Morro do Diabo State Park (MDSP) 370.0 126 159 237

Total population size in fragments where tapirs are present (A) 22 27 40

Carrying capacity in fragments where tapirs are absent (B) 15 20 30

Total Tapir Population (MDSP + fragments A) 148 186 277

6.3 Results 6.3.1 VORTEX Baseline Model 6.3.1.1 Deterministic Output

6.3.1.2 Stochastic Baseline Results

6.3.2 Sensitivity Analysis

0.08 Lowest Parameter Value Highest Parameter Value 0.06

0.04

0.02

Stochastic rStochasticvalue 0.00

-0.02

-0.04 %FB FA1R MA1R JuMort AdMort MaxAR SubMort

Parameter Figure 6.4.

6.3.3 Minimum Viable Population (MVP)

Table 6.5.

Population Size GD P(E) 100 rstoc N-extant 200 95 0 0.035 197 6.3.4 Modelling of Alternative Scenarios 6.3.4.1 Impact of Fire

0.05 Initial Population 126 Initial Population 159 0.04 Initial Population 237

0.03

0.02

0.01

0.00

-0.01 Stochastic GrowthStochastic (r) Rate

-0.02

-0.03 0 20 40 60 80 100 Frequency of Occurence of Fire in 100 Years

Figure 6.5.

6.3.4.2 Impact of Road-Kill

Table 6.6.

Population Size rstoc P(E) 100 MeanTE GD N-extant

6.3.4.3 Impact of Infectious Disease

6.3.4.4 Metapopulation Scenario

Table 6.7. 1% 0.5% Sub-Populations dispersal dispersal & 90% 100% 90% 100% Metapopulation survival survival survival survival

rstoc P(E) 100 rstoc P(E) 100 rstoc P(E) 100 rstoc P(E) 100 MDSP 0.002 1 0.003 0 0.022 0 0.022 0 METAPOPULATION 0.028 0 0.031 0 0.032 0 0.034 0 Table 6.8. 1% 0.5% Sub-Populations dispersal dispersal & 90% 100% 90% 100% Metapopulation survival survival survival survival

rstoc P(E) 100 rstoc P(E) 100 rstoc P(E) 100 rsto c P(E) 100 MDSP 0.007 0 0.007 0 0.026 0 0.026 0 METAPOPULATION 0.034 0 0.037 0 0.037 0 0.038 0 Table 6.9.

Sub-Populations rstoc & 1% 0.5% Metapopulation dispersal dispersal MDSP 0.024 0.032 METAPOPULATION 0.034 0.036

6.3.4.5 Establishment of Corridors

Table 6.10.

Population Size rstoc P(E) 100 N-extant GD WITHOUT Corridors WITH Corridors WITHOUT Corridors WITH Corridors

6.4 Discussion 6.4.1 Biological Potential of Tapirs in Morro do Diabo State Park 6.4.2 Sensitivity Analysis: Uncertainty in Input Parameters

6.4.3 Minimum Viable Population (MVP): Morro do Diabo State Park 6.4.4 Modelling of Scenarios: Threats, Metapopulation and Conservation Strategies 6.4.4.1 Impact of Threats: Fire

6.4.4.2 Impact of Threats: Road-Kill

6.4.4.3 Impact of Threats: Infectious Disease (Leptospirosis)

6.4.4.4 Tapir Metapopulation Scenario

6.4.4.5 Corridors: A Potential Conservation Strategy for Tapirs?

Chapter 7

Conclusions & Conservation Recommendations

7.1 Viability of Lowland Tapir Populations in the Atlantic Forest of the Interior of the Pontal do Paranapanema Region

7.2 Conservation Recommendations: A First Step in Building a Regional Action Plan for the Conservation of Lowland Tapirs in the Pontal do Paranapanema Region

7.2.1 Action Planning for Species Conservation

7.2.2 Recommendations for the Conservation of Lowland Tapirs in the Atlantic Forests of the Pontal do Paranapanema Region 7.2.2.1 Future Research Priorities and Research Gaps

• Tapir Population Monitoring: Demography, Epidemiology, and Genetics

• Research on Reproductive and Mortality Parameters for Lowland Tapirs 7.2.2.2 Habitat Protection • Continuous and Effective Protection of Existing Protected Areas • Creation of Private Protected Reserves: Reservas Particulares do Patrimônio Natural (RPPN)

• Protection and Restoration of Riparian Habitats

7.2.2.3 Habitat Restoration and Re-Establishment of Landscape Connectivity • Establishment of Forested Buffer-Zones around Forests Fragments

• Establishment of Wildlife Corridors and Stepping-Stones

7.2.2.4 Neutralisation of Threats

7.2.2.5 Tapir Management: Translocations and Re-Introduction

7.3 Determinants for Long-Term Persistence of Animal Populations across Fragmented Landscapes

References

´

Appendix I. Activity 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008

Appendix II. Period Animal Date of Capture Age of Radio- ID Capture Site Sex Class Monitoring Tracking Note 1 Note 2 Joana Paulete Docinho Luizinho Chu-Chu Batista Xuxa Gringo Marinho Mikilique João Patrícia

Sem Nome James Bond Georgete Dedinho Esperta Baby Cicinho Chico Júlia Cidão Tonha Tina Mama Tunga Joares Diana Cris Gatinho Branquinha Volverina Regininha Fêmea 08JL Robinho

Appendix III. Year/ West Southeast Northwest Month Border Border Border Centre JOA PAU LUI CHU XUX JOA PAT DOC JAM GEO DED ESP BAB CHI JUL CID TON TIN MAM 1997 1998 (3) (3) 1999 (22)

2000 (36) (2) (33) 2001 (35) 2002 (2) 2003 (30) (25) (29) (4) 2004 (22) (12) (8) 2005 (28) (18) 2006 (27) (33)