Ecology of three apex predators in Majete Wildlife Reserve, Malawi

232 Pages·2017·5.52 MB·English

Ecology of three apex predators in
Majete Wildlife Reserve, Malawi
By
Willem Daniel Briers-Louw
Thesis presented in partial fulfilment of the requirements for the Degree of Master of Science,
Department of Conservation Ecology & Entomology, Stellenbosch University
Supervisor: Dr Alison J. Leslie
Faculty of AgriSciences
Department of Conservation Ecology & Entomology
December 2017
Stellenbosch University https://scholar.sun.ac.za
Declaration
By submitting this thesis electronically, I declare that the entirety of the work contained therein is my
own, original work, that I am the sole author thereof (save to the extent explicitly otherwise stated),
that reproduction and publication thereof by Stellenbosch University will not infringe any third party
rights and that I have not previously in its entirety or in part submitted it for obtaining any
qualification.
Willem Daniel Briers-Louw
December 2017
Copyright © 2017 Stellenbosch University
All rights reserved
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Abstract
Apex carnivores play an important role in the ecosystem by regulating prey via predation.
Anthropogenic influences have resulted in rapid range and population reductions of large carnivores
across the African continent. These carnivores are often reintroduced into protected areas to
compensate for human-induced losses, restore ecosystem functioning and promote eco-tourism.
Majete Wildlife Reserve in Malawi is a prime example, as human persecution resulted in the
extirpation of large carnivores, with the exception of a small spotted hyena (Crocuta crocuta; hereafter
hyena) population. As from 2003, African Parks attempted to rectify this problem by restoring and
developing the reserve. Between 2011 and 2012, three lions (Panthera leo) and six leopards (Panthera
pardus) were reintroduced. The aim of this study was to describe the ecology of the apex predators
and to determine whether the felid reintroduction was successful or not.
Lion and leopard movements and home ranges were determined using GPS collars. The reintroduction
of felids was considered successful. This was based on: (1) reduced post-release movements; (2) lack
of homing tendencies; (3) breeding success; and (4) population persistence. Mean home ranges of lion
(380.45 ± 117.70 km2 [SE]) and leopard (495.08 ± 80.99 km2), were the largest on record for any
reintroduced felid in Africa, which was likely due to a low competitor density. Thus, we expect home
range sizes to decrease with an increase in conspecific density.
Population abundances and densities were estimated with the use of camera traps. The known lion
population increased to eleven individuals in five years, while the leopard population was estimated
at 11 (range = 9–17). This indicates population persistence and growth. Both founder populations were
small and require additional translocations to maintain genetic diversity. Hyena density (2.62
hyenas/100 km2) and clan size (5.33 ± 0.67) were the lowest estimates in any woodland habitat and
comparable to arid areas. This may be explained by decades of direct persecution and poaching of
their prey, or a naturally low density.
Predator diets were described and compared by means of scat analysis. Lion and hyena exhibited a
high dietary overlap of medium to large herbivores. Using Jacobs’ preference index, both species
preferred warthog (Phacocoerus africanus) and waterbuck (Kobus ellipsiprymnus). Hyenas selected a
broader range of prey, likely reducing competition with lions (which almost exclusively selected only
four species). In contrast, leopards occupied a lower dietary niche, which consisted mainly of small-to
medium-sized ungulates. These findings indicate that the three apex predators use resource
partitioning to reduce competition.
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This study suggests that reintroduction is a viable tool for re-populating large carnivores in protected
areas in Malawi. The current predator population appeared to have a minimal impact on prey
populations due to their small population size. We recommend long-term monitoring of predator-
prey dynamics as the predator populations increase to prevent major ecological imbalances. Finally,
we encourage management to focus energy and resources on the formation of a managed carnivore
metapopulation to establish a genetically viable carnivore population within Malawi.

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Opsomming
Top roofdiere speel ‘n belangrike rol in die regulering van ekosisteme deur predasie. Menslike
invloede het gelei tot ‘n vinnige afname in karnivoor populasiegetalle en habitatverliese in Afrika.
Hierdie karnivore word dikwels hervestig in bewaarde areas om te kompenseer vir menslik ge-
induseerde verliese, funksionering van ekosisteme te herstel en eko-tourisme te bevorder.
Majete Wildreservaat is ‘n prima voorbeeld waar menslike vervolging groot karnivore uitgeroei het,
behalwe vir die gevlekte hiëna (Crocuta crocuta; hierna hiëna) populasie. Vanaf 2003 het African Parks
gepoog om hierdie probleem reg te stel deur restourasie en ontwikkeling van die reservaat. Tussen
2011 en 2012 was drie leeus (Panthera leo) en ses luiperds (Panthera pardus) hervestig. Die doel van
hierdie studie was om die ekologie van die top roofdiere te beskryf en te bepaal of die hervestiging
van leeus en luiperds suksesvol was.
Bewegings en gebiede van leeus en luiperds is bepaal met hulp van GPS-halsbande. Die hervestiging
van hierdie spesies was beskou as suksesvol. Na vrystelling was daar: (1) ‘n afname in beweging; (2)
geen behoefte om huiswaarts (herkoms) te keer nie; (3) sukses met teling; en (4) populasie oorlewing.
Die gemiddelde gebied van leeus (380.45 ± 117.70 km2)[SE]) en luiperds (495.08 ± 80.99 km2) was die
grootste op rekord vir enige hervestigde karnivoor in Afrika, moontlik as gevolg van lae kompetisie
digtheid. Dus verwag ons ‘n verkleining van gebiede namate die kompetisie digtheid toeneem.
Populasiegetalle- en digthede is geskat met behulp van kamerastrikke. In vyf jaar het die erkende leeu
populasie tot elf individue vermeerder, terwyl die luiperd populasie op 11 (omvang = 9–17) staan, wat
populasie oorlewing en groei aandui. Beide stigter populasies was klein, daarom is addisionele
translokasie nodig om genetiese diversiteit te behou. Hiëna digtheid (2.62 hiënas/100 km2) en
stamgroep grootte (5.33 ± 0.67) was die kleinste in soortgelyke habitat. Hierdie is moontlik as gevolg
van direkte vervolging en stroping van hulle prooi oor dekades of ‘n natuurlike lae digtheid.
Roofdierdiëte is beskryf en vergelyk deur misanalise. Leeus en hiënas se diëte van medium tot groot
prooi het tot ‘n groot mate oorvleuel. Gebaseer op Jacobs’ indeks, het beide spesies vlakvark
(Phacocoerus africanus) en waterbok (Kobus ellipsiprymnus) verkies. Hiënas het egter ‘n wye
verskeidenheid prooi geëet, moontlik weens mededinging met leeus (wat amper uitsluitlik net vier
spesies geëet het). In kontras met leeus en hiënas, het luiperds ‘n kenmerkende dieet, wat hoofsaaklik
uit klein tot medium grootte prooi bestaan. Hierdie resultate bewys dat die drie karnivore kompetitise
met mekaar verminder deur verskillende diëte te volg.
Hierdie studie dui daarop dat hervestiging van groot karnivore in bewaarde areas in Malawi ‘n vatbare
tegniek is. Tot dusver het die roofdierpopulasie ‘n klein impak op die prooipopulasies gehad weens
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die klein roofdier populasiegetalle. Ons beveel aan dat roofdier-prooi dinamika gemonitor word soos
wat die roofdiere toeneem om ekologiese wanbalanse te vermy. Ten slotte, moedig ons die bestuur
van African Parks aan om ‘n karnivoor meta-populasie in Malawi te stig, wat ‘n genetiese vatbare
populasie kan volhou binne die landsgrense.

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Acknowledgements
Thank you to African Parks (Majete) Pty Ltd. for allowing me to witness and be part of the amazing
success story that is Majete Wildlife Reserve. Thank you to everyone who made this study possible,
particularly those who were involved in the reintroduction and collaring of the lions and leopards.
Thank you to Majete management and the entire game capture team, for allowing me to be part of
the lion re-collaring, this was truly a once-in-a-lifetime experience. Next, I would like to thank Craig
Hay and Gervaz Tamala for their supervision and willingness to assist me with my research. To the rest
of the management team, thank you for making me feel part of the Majete family. Thank you to the
law enforcement team, especially Tizola Moyo, Martin Awazi and John Jiya for organising scouts,
providing valuable information and allowing access to reports. To all the scouts who assisted me in
the field I am greatly appreciative. Your knowledge about Majete really aided my project and helped
me learn so much about the bush. I would also like to thank all the stores-men and mechanics at the
workshop, particularly Africa’s best bush-mechanic, Isaac Mlilo, who kept our vehicles going despite
numerous setbacks and was always there when we needed him.
Thank you to Dr Alison Leslie who provided me the opportunity to experience the “warmest” heart of
Africa. Majete has become my home and I will never forget the stories and friends that have been
made along this incredible journey. I am overjoyed to have been part of this amazing team and to have
experienced this with you. I would like to thank all the Earthwatch volunteers for their contribution.
Thank you to Prof Arthur Rodgers for your correspondence and help with operating Home Range Tools
in ArcGIS. Thank you to Dr Matt Hayward for providing equations for predator carrying capacities and
Prof Martin Kidd for help with statistics. I would also like to thank Dr Dan Parker and Rhodes University
for providing access to their reference of animal hair cross-sections. A special thanks to Prof
Antoinette Malan for her generosity with laboratory equipment which proved invaluable for the scat
analysis.
Thank you to the Hay family for your support throughout my stay in Majete and being my role-models.
To all my fellow researchers, Anel Olivier, Claire Gordon, Frances Forrer and Kayla Geenen, thank you
for all your help and for the good times we shared in Majete. Thank you to my colleague and friend,
Charli de Vos, for assisting me with my challenging fieldwork. I am so privileged to have had the
opportunity to work with you as we battled through some difficult times together, but more
importantly, experienced the utter joy and satisfaction of life in the African bush. Thank you to my
family for allowing me to pursue my dream of wildlife conservation and providing financial support. I
am forever grateful. Finally, I thank my King for showing me the breath-taking beauty of His creation.

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Contents
Declaration ................................................................................................................................... i
Abstract ....................................................................................................................................... ii
Opsomming ................................................................................................................................ iv
Acknowledgements..................................................................................................................... vi
Contents .................................................................................................................................... vii
List of Tables ............................................................................................................................... xi
List of Figures ............................................................................................................................ xiv
List of Appendices ..................................................................................................................... xvi
Chapter 1 - Introduction ............................................................................................................... 1
1.1 Role of apex carnivores in ecosystems ......................................................................................... 1
1.2 Conservation status of apex carnivores ........................................................................................ 2
1.2.1 Lion (Panthera leo; Linnaeus, 1758)....................................................................................... 3
1.2.2 Leopard (Panthera pardus; Linnaeus, 1758) .......................................................................... 6
1.2.3 Spotted hyena (Crocuta crocuta; Erxleben, 1777) ................................................................. 9
1.3 Conservation status of apex carnivores in Malawi ..................................................................... 12
1.4 The importance of reintroduction .............................................................................................. 14
1.5 Aim and objectives ...................................................................................................................... 16
1.5.1 Aim ....................................................................................................................................... 16
1.5.2 Objectives ............................................................................................................................. 16
1.6 Thesis outline .............................................................................................................................. 17
1.7 References .................................................................................................................................. 18
Chapter 2 - Study area and reintroduction of study species ......................................................... 34
2.1 Study area ................................................................................................................................... 34
2.1.1 Location and history ............................................................................................................. 34
2.1.2 Climate ................................................................................................................................. 35
2.1.3 Topography and altitude ...................................................................................................... 35
2.1.4 Geology and soils ................................................................................................................. 35
2.1.5 Watercourses ....................................................................................................................... 36
2.1.6 Vegetation ............................................................................................................................ 36
2.2 Large carnivore reintroduction into MWR .................................................................................. 39
2.2.1 Pre-release management ..................................................................................................... 41
2.4 References .................................................................................................................................. 42
2.5 Appendices .................................................................................................................................. 44
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Chapter 3 - Early post-release monitoring and post-release breeding of two reintroduced felids in
Majete Wildlife Reserve, Malawi ................................................................................................ 45
3.1 Abstract ....................................................................................................................................... 45
3.2 Introduction ................................................................................................................................ 46
3.3 Methods ...................................................................................................................................... 48
3.3.1 Study site .............................................................................................................................. 48
3.3.2 Methods and materials ........................................................................................................ 49
3.3.3 Data analysis ........................................................................................................................ 51
3.4 Results ......................................................................................................................................... 53
3.4 Discussion .................................................................................................................................... 62
3.6 Conclusions ................................................................................................................................. 68
3.7 Acknowledgements ..................................................................................................................... 68
3.8 References .................................................................................................................................. 69
3.9 Appendices .................................................................................................................................. 75
Chapter 4 - Home range and habitat selection of reintroduced lion (Panthera leo) and leopard
(Panthera pardus) in Majete Wildlife Reserve, Malawi ................................................................ 79
4.1 Abstract ....................................................................................................................................... 79
4.2 Introduction ................................................................................................................................ 80
4.3 Methods ...................................................................................................................................... 82
4.3.1 Study site .............................................................................................................................. 82
4.3.2 Immobilisation and collaring ................................................................................................ 83
4.3.3 Statistical analysis ................................................................................................................ 84
4.4 Results ......................................................................................................................................... 87
4.5 Discussion .................................................................................................................................. 103
4.6 Conclusion ................................................................................................................................. 106
4.7 Acknowledgements ................................................................................................................... 107
4.8 References ................................................................................................................................ 107
Chapter 5 - Dietary ecology of three apex predators in Majete Wildlife Reserve, Malawi ........... 116
5.1 Abstract ..................................................................................................................................... 116
5.2 Introduction .............................................................................................................................. 117
5.3 Methods .................................................................................................................................... 118
5.3.1 Study area .......................................................................................................................... 118
5.3.2 Data collection ................................................................................................................... 119
3.3.3 Data analysis ...................................................................................................................... 122
3.4 Results ....................................................................................................................................... 124
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3.5 Discussion .................................................................................................................................. 137
5.5 Conclusion ................................................................................................................................. 143
5.6 Acknowledgements ................................................................................................................... 143
5.7 References ................................................................................................................................ 144
5.7 Appendices ................................................................................................................................ 153
Chapter 6 - Population dynamics and carrying capacity of two reintroduced felids and a resident
hyenid in Majete Wildlife Reserve, Malawi ............................................................................... 156
6.1 Abstract ..................................................................................................................................... 156
6.2 Introduction .............................................................................................................................. 157
6.3 Methods .................................................................................................................................... 159
6.3.1 Study site ............................................................................................................................ 159
6.3.2 Fieldwork ............................................................................................................................ 159
6.3.3 Statistical analysis .............................................................................................................. 161
6.4 Results ....................................................................................................................................... 164
6.5 Discussion .................................................................................................................................. 170
6.6 Conclusion ................................................................................................................................. 174
6.7 Acknowledgements ................................................................................................................... 174
6.8 References ................................................................................................................................ 174
6.9 Appendices ................................................................................................................................ 185
Chapter 7 - Research findings and implications for African Parks Majete management .............. 191
7.1 Overview ................................................................................................................................... 191
7.2 Research findings ...................................................................................................................... 192
7.2.1 Reintroduction success – importance of the pre- and post-release phase ....................... 192
7.2.2 A few big cats in a small reserve – insights from home range and habitat use analyses .. 192
7.2.3 Population persistence of apex predators – a look at population growth and density .... 193
7.2.4 Predator-prey interactions – are prey limited by predators? ............................................ 194
7.2.5 Competition for resources – evidence of dietary separation ............................................ 195
7.3 Management implications ........................................................................................................ 196
7.3.1 Managing large carnivores in MWR ................................................................................... 196
7.3.2 Conservation status of lion in Malawi................................................................................ 199
7.3.3 The managed metapopulation approach for carnivores in Malawi .................................. 200
7.2.4 Which cats go where? – considering the reintroduction of cheetah into MWR ............... 202
7.4 Conclusion ................................................................................................................................. 204
7.5 References ................................................................................................................................ 204
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