Contents
Preface
Chapter One Introduction 1
1.1 Evolutionary History and Adaptations of Colobine Monkey 1
1.1.1 Evolutionary history of colobine monkeys 1
1.1.2 Adaptations of colobine monkeys 3
1.2 Classification of Asian Colobines 5
1.2.1 Brief review 5
1.2.2 Classification of Asian langur species 6
1.2.3 Classification of the francoisi group 7
1.3 Distribution of Asian Colobines 8
1.3.1 Brief review 8
1.3.2 Distribution of Trachypithecus species 10
1.3.3 Distribution of the limestone langurs 11
1.3.4 Distribution of T. francoisi 11
1.4 Ecological Studies of Trachypithecus Species 12
1.4.1 Ecological studies of the T. vetulus group 12
1.4.2 Ecological studies of the T. cristatus group 13
1.4.3 Ecological studies of the T. obscurus group 13
1.4.4 Ecological studies of the T. pileatus group 14
1.4.5 Ecological studies of the T. francoisi group15
1.5 Studies on Primate Behavioural Constraints 16
1.6 Conservation Issues 19
1.6.1 Brief review 19
1.6.2 Conservation of“l(fā)imestone langurs” 20
1.7 Purpose of This book 21
Chapter Two Study Area and Methods 23
2.1 Location of MNR and Human Pressure 23
2.1.1 MNR in general 23
2.1.2 Main study area 24
2.2 Climate and Vegetation Characteristic at MNR 25
2.2.1 Climate Characteristic 25
2.2.2 Vegetation 26
2.3 The Study Langur Groups 26
2.4 Methods 28
2.4.1 Climate 28
2.4.2 Methods on vegetation and phenology 28
2.4.3 Methods on behavioral data 31
2.4.4 Ranging behavior38
2.4.5 Data on local community 38
Chapter Three Climatic, Phenological Patterns, and Conservation 39
3.1 Climate and Seasonality 39
3.1.1 Temperature 39
3.1.2 Rainfall39
3.2 Plant Species Richness and Vegetation Composition 41
3.2.1 Species richness in the main study area41
3.2.2 Comparison between LYY and XGB43
3.3 Phenology of Food Species 45
3.3.1 Food species 45
3.3.2 Phenological pattern 46
3.4 Human Pressure in the Main Study Area 48
3.5 Discussion 49
3.5.1 Species richness and dominance 49
3.5.2 Seasonal variations of climate and phenological pattern 50
3.5.3 Human disturbance and its consequence for the langurs 53
3.6 Summary 55
Chapter Four Feeding Ecology 56
4.1 Introduction 56
4.2 Method Review 61
4.2.1 Data recording in the field 61
4.2.2 Data analysis 61
4.3 Results 63
4.3.1 Food species 63
4.3.2 Food items 70
4.3.3 Habitat effects on diet 76
4.3.4 Crop raiding 83
4.3.5 Rock lapping 84
4.3.6 Water drinking 85
4.4 Discussion 86
4.5 Summary 93
Chapter Five Range Use and Territoriality 95
5.1 Introduction 95
5.2 Review of field processes 100
5.3 Results 102
5.3.1 DTD 102
5.3.2 Home range 105
5.3.3 Range overlap and territoriality 111
5.3.4 Group leadership 115
5.3.5 Choice of sleeping site 117
5.4 Discussion 122
5.4.1 Daily travel distance 122
5.4.2 Range size 124
5.4.3 Range overlap & territoriality 125
5.4.4 Group leading 126
5.4.5 Choice of sleeping site 126
5.5 Summary 128
Chapter Six Activity Pattern and Time Budget 130
6.1 Introduction 130
6.2 Methods 134
6.2.1 Techniques review 134
6.2.2 Field procedure 136
6.2.3 Data pool and analysis 136
6.3 Results 137
6.3.1 Daily activity pattern 137
6.3.2 General time budget pattern and seasonal change 139
6.3.3 Habitat effects on time budget 144
6.3.4 Sex-age class differences 147
6.4 Discussion 149
6.4.1 Daily activity pattern 149
6.4.2 General time budget pattern and seasonal change 150
6.4.3 Habitat effect on time budget 152
6.4.4 Sex-age class differences 153
6.5 Summary 154
Chapter Seven Group Structure and Social Behavior 156
7.1 Introduction 156
7.1.1 Social organization 156
7.1.2 Dispersal 158
7.1.3 Group size 159
7.1.4 Sexual behavior, birth season and birth interval 160
7.1.5 Group takeover and infanticide 162
7.1.6 Social behavior 164
7.2 Methods 165
7.3 Results 166
7.3.1 Dynamic of target groups 166
7.3.2 Social structure and individual dispersal 168
7.3.3 Sexual behavior and birth 168
7.3.4 Group takeover and infanticide 172
7.3.5 Social behaviors 175
7.4 Discussion 176
7.4.1 Social structure and individual dispersal 176
7.4.2 Sexual behavior, breeding season and birth interval 179
7.4.3 Group takeover and infanticide 180
7.4.4 Social behavior 182
7.5 Summary 182
Chapter Eight Infant Behavioral Development and Pelage Coloration Change 184
8.1 Introduction 184
8.1.1 Dependent to independent feeding 184
8.1.2 Dependent to independent locomotion 185
8.1.3 Mother-infant physical proximity to socialization & allomothering 186
8.1.4 Coloration of natal pelage 187
8.2 Methods 189
8.3 Results 190
8.3.1 Dependent to independent feeding 190
8.3.2 Dependent to independent locomotion 192
8.3.3 Mother-infant physical proximity to socialization & allomothering 194
8.3.4 Coloration of natal pelage 195
8.4 Discussion 201
8.4.1 Dependent to independent feeding 201
8.4.2 Dependent to independent locomotion 202
8.4.3 Mother-infant physical proximity to socialization & allomothering 202
8.4.4 Coloration of natal pelage 204
8.5 Summary 205
Chapter Nine Conclusions 207
9.1 Socioecology of Franois’ Langur at MNR 207
9.1.1 Climatic and Phenology 207
9.1.2 Feeding ecology 207
9.1.3 Ranging behavior 208
9.1.4 Daytime activity time budget 209
9.1.5 Group structure and social behavior 210
9.2 Behavioral Flexibility in Response to Different Habitats 211
9.3 Infant Behavioral Development and Color Change of Natal Coat 213
9.3.1 Infant behavioral development 213
9.3.2 Color change of natal pelage 214
9.4 Threats to the Langurs at MNR and Conservation Recommendation 214
9.5 Final Conclusions 216
9.5.1 How does the food availability change throughout the year? 216
9.5.2 How does crop-raiding influence the langur? 216
9.5.3 Which area of MNR contains most langurs and what are the habitat characteristics of this area? 216
9.5.4 How does the social organization and behavior respond to ecological variables? 216
9.5.5 How does habitat destruction influence the survival of Franois’ langur? 217
References 218
AppendixⅠ List of woody plant species in the main study area at MNR 245
Appendix Ⅱ Transect sampling record in the main study area 252
Appendix Ⅲ Overall feeding records of food species at Mayanghe (Listed according to Hutchinson’s System) 258
Appendix Ⅳ Food species (as families) contribution to feeding record 266
Appendix Ⅴ Comparison of relative utilization intensity and annual feeding percentage of 74 common food species between LYY and XGB (Unit: %) 268
Appendix Ⅵ Comparison of feeding record seasonality of 74 common food species between LYY and XGB*(Unit:%) 272
Appendix Ⅶ Comparison of feeding item seasonality of 74 common food species between LYY and XGB 276
Plates 283
List of Tables
Table 1-1 Classification of Asian colobines 6
Table 2-1 Group size and dynamic of study langur groups at MNR 26
Table 2-2 Altmann’s recommendations for sampling methods (1974) 32
Table 3-1 Monthly and seasonal temperature (Mean±SE) in the main study area at MNR (Unit: ℃) 39
Table 3-2 Monthly and seasonal rainfall in the main study area at MNR(Unit: mm) 41
Table 3-3 List of 18 common families in the main study area at MNR (based on transect sampling) 42
Table 3-4 List of 15 common species in the main study area at MNR (Based on transect sampling,each accounted for over 1.5% of total individuals) 42
Table 3-5 Vegetation transect sampling comparison between LYY and XGB 43
Table 3-6 Comparison of top 15 common and dominant plant species between LYY and XGB 45
Table 3-7 Monthly availability of phenological items at MNR(Unit: %) 47
Table 3-8 Seasonal variation of the availability of phenological items at MNR (Based on Oneway ANOVA Test) 47
Table 3-9 Correlation between monthly temperature, rainfall and availability of phenological items at MNR (Based on Spearman Rank Correlation Coefficient) 48
Table 3-10 Comparison of phenological patterns between LYY and XGB (Based on Paired-Samples Test and Mann-Whitney U-test) 48
Table 3-11 Climate pattern, habitat type and tree species richness at different sites of Trachypithecues species 50
Table 4-1 Composition and contribution of food species of Franois’ langur at Mayanghe 64
Table 4-2 All the year round and seasonal food species composition and contribution at Mayanghe 65
Table 4-3 Seasonality of food species composition and contribution at Mayanghe 66
Table 4-4 Comparison of the top food 26 species in the term of feeding percentage, dominance index and selection index at MNR 67
Table 4-5 Correlation test among feeding percentage, dominance index, and selection index of food species at Mayanghe 68
Table 4-6 Annual diet of Franois’ langur and its seasonal change at MNR 70
Table 4-7 Oneway ANOVA test of seasonality of food items at MNR (Based on daily feeding record) 71
Table 4-8 Comparison of seasonal feeding record and availability of major food items at MNR (Based on seasonal data) 72
Table 4-9 Comparison of contribution of tree species and other species in the term of food
items at MNR (Contribution was measured as the percentage of the feeding record of each food item) 76
Table 4-10 Comparison of seasonal feeding contribution of the 74 common food species between LYY and XGB 77
Table 4-11 Comparison of food item contributions of the 74 common food species between LYY and XGB 78
Table 4-12 Comparison of food species composition and contribution between LYY and XGB 79
Table 4-13 Comparison of seasonality of food species composition and contribution between LYY and XGB 79
Table 4-14 Comparison of seasonality of staple food species between LYY and XGB 80
Table 4-15 Comparison of stable and seasonal food species Between LYY and XGB 80
Table 4-16 Comparison of annual diet and diet seasonality between LYY and XGB 81
Table 4-17 Oneway ANOVA test of langur diet seasonality at LYY and XGB 82
Table 4-18 Comparison of contributions of tree species and other food species to diet between LYY and XGB 82
Table 4-19 Oneway ANOVA and U-Test of seasonal feeding percentage of food items between LYY and XGB (Based on daily feeding percentage) 82
Table 4-20 Systematic records of rock lapping at Mayanghe 85
Table 4-21 Systematic records of water drinking at Mayanghe 86
Table 4-22 Feeding comparisons among Trachypithecus 86
Table 5-1 Comparison of daily travel distance and range size of Trachypithecus species 102
Table 5-2 Seasonality of daily travel distance and range size at Mayanghe 103
Table 5-3 Correlation between daily travel distance and intake of food items at Mayanghe (Pearson Correlation Rank based on seasonal & daily data) 103
Table 5-4 Comparison of daily travel distance between LYY and XGB 104
Table 5-5 Correlation between daily travel distance and intake of food items at LYY and XGB (Pearson correlation rank based on daily records) 104
Table 5-6 Range size, habitat type and use intensity of langur groups at Mayanghe 106
Table 5-7 Range use intensity of langur groups at Mayanghe 109
Table 5-8 Range overlap between adjacent langur groups at Mayanghe 112
Table 5-9 Group leading by different sex-age classes at Mayanghe 115
Table 5-10 Seasonal change of group leading by different sex-age classes at MNR 116
Table 5-11 Number of sleeping caves used by langur groups at Mayanghe 117
Table 5-12 Physical features influencing sleeping site selection of the langurs at Mayanghe 121
Table 6-1 Comparison of time-budget strategy to shortage of food supply among primate species 132
Table 6-2 Annual pattern of time-budget of Franois’ langur and its seasonality at MNR 140
Table 6-3 Correlation between time budget, rainfall, and temperature at MNR (Perason & Spearman rank) (Based on seasonal data) 141
Table 6-4 Correlation between time budget and temperature at MNR (Person & Spearman Rank) 141
Table 6-5 Correlation between time budget and food item intake at MNR (Spearman Rank) 142
Table 6-6 Tentative absolute time allocation pattern of Franois’ langur and seasonality at MNR* 143
Table 6-7 Time budget correlation between behavioral categories at MNR (Spearman Rank) 144
Table 6-8 Seasonality of annual time budget pattern in LYY and XGB 145
Table 6-9 Comparison of annual and seasonal time budget pattern between LYY and XGB145
Table 6-10 Comparison of time budget pattern between sex-age categories 147
Table 6-11 Comparison of time budget pattern between several Trachypithecus species 151
Table 7-1 Records of female solicitation and subsequent copulation at MNR 169
Table 7-2 Birth records and their distribution at MNR 171
Table 7-3 Birth records and intervals at MNR (Based on 11 birth records in LYY-G1 from 2001 to 2007) 172
Table 7-4 Social grooming and play distribution among sex-age categories at MNR (Based on a total of 1,809 grooming and 5082 play records) (Unit: %) 175
Table 7-5 Different roles of sex-age categories in social grooming at MNR (Based on 917 GM-groomer and 892 GMP-groomee records*) (Unit: %) 175
Table 7-6 Distribution of initiating and receiving grooming among sex-age categories at MNR (Based on 917 GM and 892 GMP records) 176
Table 8-1 Dependent and Independent behavior record of infants in LYY-G1 190
Table 8-2 Age changes in behavior of infants in LYY-G1 191
Table 9-1 Comparison of langurs in intact and in disturbed habitats 212