Ecology and Population Status of the Puku Antelope (Kobus vardonii LIVINGSTONE, 1857) in Zambia Dissertation zur Erlangung des Doktorgrades (Dr rer nat.) der Mathematisch-Naturwissenschaftlichen Fakultät der Rheinischen Friedrich-Wilhelms-Universität Bonn vorgelegt von Vera Rduch aus Kleve Bonn, September 2013 Angefertigt mit der Genehmigung der Mathematisch-Naturwissenschaftlichen Fakultät der Rheinischen Friedrich-Wilhelms-Universität Bonn Dissertation for the degree of Doctor of Science (Dr rer nat.) in Zoology at the Rheinische Friedrich-Wilhelms-Universität Bonn This thesis was compiled with the permission of the Faculty of Mathematics and Natural Sciences of the Rheinische Friedrich-Wilhelms-Universität at Bonn Gutachter: Prof Dr Wolfgang Böhme Gutachter: Prof Dr Horst Bleckmann Tag der Promotion: 16 Dezember 2013 Erscheinungsjahr: 2014 Erklärung Hiermit erkläre ich, dass ich diese Dissertation persönlich, selbstständig und unter Offenlegung der erhaltenen Hilfen angefertigt habe und dass diese Arbeit noch nicht anderweitig als Dissertation eingereicht wurde Stellen der Arbeit, die anderen Werken in Wortlaut oder Sinn entnommen wurden, sind unter Angabe der Quellen als Entlehnung kenntlich gemacht Vera Rduch Bonn, im September 2013 Declaration I declare that the research described within this thesis is my own work and that this thesis is my own composition I certify that this thesis has never been submitted for any other degree or professional qualification Vera Rduch Bonn, September 2013 Diese Dissertation wurde innerhalb des ZamBio-Projektes des Zoologischen Forschungsmuseum Alexander Koenig in Bonn in Zusammenarbeit mit der Zambian Wildlife Authority durchgeführt This thesis was conducted within the ZamBio Project of the Zoologisches Forschungsmuseum Alexander Koenig at Bonn in cooperation with the Zambian Wildlife Authority Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Contents Abstract: “Ecology and population status of the puku antelope (Kobus vardonii LIVINGSTONE, 1857) in Zambia” i Zusammenfassung: “Ökologie und Populationsstatus der Puku Antilope (Kobus vardonii LIVINGSTONE, 1857) in Sambia“ iii Résumé: “L’écologie et l’état des populations de l’antilope puku (Kobus vardonii LIVINGSTONE, 1857) en Zambie“ v General Introduction 1.1 1.2 1.3 General conception The puku – an outline The study region Population status, distribution patterns and habitat choice 16 2.1 2.2 2.2.1 2.2.2 2.2.3 2.2.4 2.3 2.3.1 2.3.2 2.3.3 2.3.4 2.3.5 2.3.6 2.4 2.4.1 2.4.2 2.4.3 2.4.4 2.4.5 2.4.6 2.4.7 2.4.8 Introduction 17 Material and Methods 22 Data collection on the transect line 22 Location and short description of the transect lines 25 Setting camera-traps 28 Data analyses 30 Results 36 Sightings on the transect lines 36 Habitat choice of antelopes 40 Population estimates with DISTANCE 46 Results of the camera-trap survey 53 Social organisation of the puku 55 Body condition of puku 62 Discussion 64 General remarks to line transects and DISTANCE analyses 64 Distribution patterns of puku in the dry season 66 The puku’s distribution in the late rainy season 75 The development of the puku’s population in Kasanka NP 75 The camera-trap survey 77 Spatial overlap between puku and other antelopes 80 Looking inside the puku’s populations 85 Conclusions 92 Diet and dietary overlap of puku and other bovids 95 3.1 3.2 3.2.1 3.2.2 3.2.3 3.2.4 3.2.5 Introduction 96 Material & Methods 100 Assessment and collection of grass species 100 The reference collection of grasses 102 Collection of faecal samples of bovid species 105 Analysis of faecal samples 108 Data analyses 109 Ecology and population status of the puku antelope (Kobus vardonii) in Zambia 3.3 3.3.1 3.3.2 3.3.3 3.3.4 3.3.5 3.4 3.4.1 3.4.2 3.4.3 3.4.4 3.4.5 3.4.6 Results 111 Grass species in the study regions 111 Diet of puku and other bovid species 112 Food preferences in Kasanka NP 122 Measurements of diet breadth 127 Evaluation of dietary overlap 129 Discussion 138 General remarks about the methods 138 General classification of the puku’s diet 139 Grasses in the puku’s diet 140 Variation in the puku’s diet 146 Dietary competition between bovid species 150 Conclusions 153 Predation on the puku antelope 156 4.1 4.2 4.2.1 4.2.2 4.2.3 4.3 4.3.1 4.3.2 4.4 4.4.1 4.4.2 4.4.3 4.4.4 Introduction 157 Methods 159 Searching for signs of carnivore species 159 Analysis of the faeces of carnivores 161 Data analyses 163 Results 165 Faeces and other signs of predators 165 Determination of prey in the faecal samples 169 Discussion 174 Assigning the faecal samples to predators 174 Occurrence of predators in the study regions 176 The puku as prey 178 Conclusion: Top down regulation of puku 187 Activity patterns of puku 189 5.1 5.2 5.3 5.4 Introduction 190 Methods 192 Results 195 Discussion 205 General Synthesis 210 References 217 List of Figures .230 List of Tables .233 Abbreviations .235 Acknowledgment 236 Appendix 241 Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Appendix Figure A 18: Distribution of Melinis nerviglumis along the transect lines in Kasanka NP Figure A 19: Distribution of Panicum spp along the transect lines in Kasanka NP 276 Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Appendix Figure A 20: Distribution of Setaria spp along the transect lines in Kasanka NP Figure A 21: Distribution of Sporobolus spp along the transect lines in Kasanka NP 277 Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Appendix Description of the “Study regions for Predators” in Kasanka NP A – grid B – grid C – grid D – grid E – grid F – grid Figure A 22: Examples of habitats covered by the grids Miombo woodland covered by grid (A), grid (C) and grid (F), types of grasslands covered by grid and and floodplain habitat covered by grid (D) 278 Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Appendix Grid (Figure A 22 A) is situated in the western part of Kasanka NP close to Luwombwa River and Luwombwa Lodge although the floodplain or the borders of the river are not covered It covers an area west from the transect line and comprises miombo woodland The rock appeared close to the surface as a least in some parts small clearings with rocky ground and few soil are present Some parts of the area were burnt, other parts already provided fresh leaves Grid (Figure A 22 B) is situated in the north of Chikufwe Plain It comprises a part of grassland, the area around the waterhole and miombo woodland adjacent to Chikufwe Plain The waterhole still provided water and thus green and fresh plants grew around it Contrastingly the miombo and other parts of the grasslands are either dry or burnt The miombo rather provides few understory Grid (Figure A 22 C and D) is situated at Kabwe Area It comprises small amounts of dambo and floodplain areas where the grass is either dry, burnt or moist and green The same applies for the miombo woodland included in this grid Some parts of the miombo here provide a very dense understory or grasslayer Grid (Figure A 22 E) at Fibwe Plain comprises different habitat types: grassland areas with rather long and dry grass, grassland with grased or burnt areas, moist grassland with green grass, dry miombo with few understory and moist woodland Grid (Figure A 22 F) near Wasa is almost entirely covered by miombo The aspect was either burnt or freshly sprouting leaves The understody was rather absent Further, some areas in the north of the grid are characterised by termitaria 279 Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Appendix A B C D E F Figure A 23: Aspects of scats from different predators in Zambia: a latrine site of spotted hyena (Crocuta crocuta) near Kaingu Lodge (A); scats from crocodile (Crocodylus niloticus) (B), caracal (Caracal caracal) (C), serval (Leptailurus serval) (D), African civet (Civettictis civetta) (E) and white-tailed mongoose (Ichneumia albicauda) (F) in Kasanka NP The adjacent scale gives cm 280 Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Appendix Table A 13: Ethogram: This ethogram to assess the behaviour of puku and other antelopes during this study is based on the ethogram I used to study behaviour patterns of puku in Luambe NP (RDUCH 2008) Grazing The animal stands with the head bowed onto the grasslayer If the animal moves some few steps with the head still bowed down, the activity is still comprised in ‘grazing’ Browsing The animals are standing with the head in trees or scrubs The head of the animal or branches or leaves of the tree or scrub are moving Moving Walking comprises all forms of movements exhibited by the animals indifferent if the movements are slow (walk) or fast (gallop) Jumps are included 281 Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Appendix Vigilant behaviour The animals show a face and a general body posture of alert staring attentively into a direction Vigilant behaviours can also occur while the animal is standing or lying down Reproductive and territorial behaviours Displaying behaviours and postures of males towards females and towards other males belong to this category, as well as agonistic interactions, as fights between males Behaviour patterns of females that make the male continue are included but indifferent and refusing behaviours of the females are not Mating is included Suckling The category suckling includes both the female that lactates its young as well as the young itself 282 Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Appendix Resting While resting, the animal shows a relax posture, a relaxed face and relaxed ears Ruminating as visible by moving jaws can occur Resting is subdivided into three categories Resting (standing): Resting can occur while standing Resting (lying, head held up): Resting can occur while lying on the ground Mostly the legs are folded under the body The belly and the legs can be the contact surface to the ground The head is raised Resting (lying, head on the ground): Resting can occur while lying on the ground The legs can be folded under the body or stretched towards the side Contact surface to the ground can be the belly and the legs or the side of the animals The head lies on the ground Behaviour of comfort The animal scratches itself with the help of teeth, hooves or horns, while standing or lying down Watching at the observer The animal shows vigilant behaviour while starring at the observer Other behaviour patterns These are behaviours not covered by the categories mentioned above Out of sight An animal was noted out of sight if it was around but only partly visible so that the behaviour could not be evaluated 283 Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Appendix A B Figure A 24: Amount of observations of puku (Kobus vardonii) at Kabwe Plain (Kasanka NP) during the survey for activity patterns In cool dry season 2009 (A) und in hot dry season 2010 (B) 284 Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Appendix A B Figure A 25: Amount of observations for puku (Kobus vardonii) (A) and Impala (Aepyceros melampus) (B) during the survey for activity patterns at Puku Pan 285 Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Appendix A B Figure A 26: Air temperature [°C] and relative air humidity [%] during the day during the different survey for activity patterns 286 Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Appendix Table A 14: Quality of the behaviour sampling along the transect lines: Number of individuals encountered and those assessed for behaviours on the transect line survey and the percentage amount of animals not assessed for behaviours antelope time of individuals individuals assessed individuals not assessed species the day observed for behaviours for behaviours [%] puku am 213 186 20.19 10 am 191 170 18.32 pm 190 165 22.5 pm 251 223 16.92 impala am 300 227 35.67 10 am 227 210 10.57 pm 357 291 20.17 pm 182 157 14.29 287 [...].. .Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Abstract: Ecology and population status of the puku antelope (Kobus vardonii LIVINGSTONE, 1857) in Zambia by Vera Rduch Dissertation for the degree of Doctor of Science (Dr rer nat.) in biology at the Rheinische Friedrich-Wilhelms-Universitọt Bonn Explaining patterns of animal distribution and abundance is... des populations, le choix de lhabitat, analyses microhistologiques, nourriture, predation, comportement vi Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Here I lay, looking at the graceful forms and motions of beautiful pokus, leches and other antelopes, DAVID LIVINGSTONE Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Ecology and population. .. impalas diet, the dietary overlap i Ecology and population status of the puku antelope (Kobus vardonii) in Zambia between the impala and the puku increased, while it was rather low in the cool dry season Thus, one reason for high population densities of the puku in the study regions might be a generally low dietary competition, especially in the dry season To assess the impact of predation on puku populations,... of the puku and the development of the populations can be evaluated in order to improve natural conservation measures 16 Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Chapter 2: Population status, distribution patterns and habitat choice 2.1 Introduction Knowledge of distribution and abundance, of the population density or the population size is virtually important and. .. (C); The transition zone between miombo and open habitats in Kasanka NP (D) and the floodplain along Kasanka River (E) in the cool dry season and a dambo in the early rainy season (F) Riverine vegetation along Kafue River near McBridess Camp (G) and near Puku Pan Lodge (H) in the cool dry season 15 Ecology and status of population of the puku antelope (Kobus vardonii) in Zambia Chapter 2: Population status, ... (HUFFMAN 2011) Within the tribe of the Reduncini, the puku is classified nowadays as a member of the genus Kobus Two subspecies, 6 Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Chapter 1: General Introduction K.v .vardonii and K.v.senganus, are mentioned by ANSELL (1960a, 1978) They are considered invalid by HUFFMAN (2011a), but recent findings suggest that puku in Tanzania... Highlight of the 11 Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Chapter 1: General Introduction NP, however, is the millions of straw coloured fruit bats (Eilodon helvum) that occupy the NP from November to December Ten species of antelopes occur in the park: the sitatunga (Tragelaphus spekii) population is the most visible and densest in Africa and the very abundant population. .. important to know how these numbers emerge Following this the general question is: What affects the abundance and distribution of the puku in selected areas of Zambia? 1 Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Chapter 1: General Introduction Three factors determine distribution patterns: the dispersal capacities of the species, the spatial distribution of favourable environmental... differences in the diets 3 Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Chapter 1: General Introduction Puku are supposed to be prey of different carnivore species But which predators do actually occur in the study regions and what about it the amount of the puku in their diet? What activity patterns do puku show? They are suggested to have specific times of day and night... of the puku in Zambia, the investigation of the pukus ecological niche, its role in the Zambian ecosystem is important and thus the common theme of this study The term niche used here conforms to ELTONs definition of the ecological niche which means the animals place in its community, its relation to food and enemies, and to some extent to other factors (ELTON 1927) The ecological niche is one of the ... LIVINGSTONE Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Ecology and population status of the puku antelope (Kobus vardonii) in Zambia Chapter 1: General Introduction... to an increasing amount of grasses in the impalas diet, the dietary overlap i Ecology and population status of the puku antelope (Kobus vardonii) in Zambia between the impala and the puku increased,... puku antelope (Kobus vardonii) in Zambia Abstract: Ecology and population status of the puku antelope (Kobus vardonii LIVINGSTONE, 1857) in Zambia by Vera Rduch Dissertation for the degree of Doctor