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Population Size and Genetic Diversity of Nigerian Lions (Panthera Leo)

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Population Size and Genetic Diversity of Nigerian Lions (Panthera Leo) POPULATION SIZE AND GENETIC DIVERSITY OF NIGERIAN LIONS (PANTHERA LEO) POPULATION SIZE AND GENETIC DIVERSITY OF NIGERIAN LIONS (PANTHERA LEO) Talatu Tende AKADEMISK AVHANDLING Som för av filosofie doktorsexamen vid naturvetenskapliga fakulteten, Lunds universitet, kommer att offentligen försvaras i Blå Hallen, Ekologihuset, Lund, Fredagen den 31 January 2014, kl.9:00. ACADEMIC DISSERTATION Presented in fulfilment of the requirements for the degree of Philosophie Doctor at the Faculty of Science, Lund University, to be defended publicly in the Blue Hall, Ecology Building, Sölvegatan 37, Lund, Sweden, Friday 31st January 2014, 9 AM. Fakultetsopponent: Göran Spong, Swedish University of Agricultural Sciences, Umeå POPULATION SIZE AND GENETIC DIVERSITY OF NIGERIAN LIONS (PANTHERA LEO) Talatu Tende A doctoral thesis at a university in Sweden is produced either as a monograph or as a collection of papers. In the latter case, the introductory part constitutes the formal thesis, which summa- rizes the accompanying papers. These have either already been published or are manuscripts at various stages (in press, submitted or in ms). Copyright © Talatu Tende Department of Biology | Lund University Cover art and intro chapter title art: Stina Andersson Layout and formatting: Katarina Eriksson Printed in Sweden at Media Tryck, Lund, 2013 ISBN: 978-91-7473-773-8 CONTENTS List of papers 8 List of contribution 8 Introduction 11 References 30 Popular Summary 37 Acknowledgments 39 Papers Paper I 45 Paper II 57 Paper III 73 Paper IV 85 Paper V 91 7 LIST OF PAPERS This thesis is based on the following papers which are referred to by their Roman numerals. I Ulf Ottosson, Talatu Tende, Christian Hjort & Bengt Hansson. Populations and geographical distribution of Lions (Panthera leo) in West Africa from ancient to present times, especially during the last 200 years. - Submitted II Talatu Tende, Bengt Hansson, Ulf Ottosson, Staffan Bensch. 2014. Evaluating preservation medium for the storage of DNA in the African lion (Panthera leo) faecal samples. - Current zoology in press III Talatu Tende, Staffan Bensch, Ulf Ottosson, Bengt Hansson. Dual phylogenetic origins of Nigerian lions (Panthera leo). - Submitted IV Talatu Tende, Ulf Ottosson, Bengt Hansson, Mikael Åkesson, Staffan Bensch. 2010. Population size of lions in Yankari Game Reserve as revealed by faecal DNA sampling. - African Journal of Ecology 48:949-952 V Talatu Tende, Bengt Hansson, Ulf Ottosson, Mikael Åkesson, Staffan Bensch. 2013. Individual Identification and genetic variation of lions Panthera( leo) from two protected areas in Nigeria. - Plos One in press Papers reprinted with permission from the publishers. LIST OF CONTRIBUTIONS Paper I. The idea was formulated by Ulf Ottosson (UO), Talatu Tende (TT) and Bengt Hansson (BH). UO and Christian Hjort (CH) analysed the data and wrote the manuscript with support from co-authors. Paper II. The study was planned by TT and UO. Sample collection was organised by TT and UO. TT analysed the data, and wrote the manuscript with support from other co-authors. Paper III. The idea was formulated by all co-authors. TT carried out the analyses with support from BH and Staffan Bensch (SB). TT wrote the manuscript with support from all co-authors. Paper IV. The study was conceived by TT and UO. Field data collection was organised by TT and UO. Laboratory work was performed by TT. Analyses were done by TT, BH and Mikael Åkesson (MÅ). TT wrote the manuscript with support from co- authors. Paper V. TT and UO conceived the study. Field data collection was organised by TT and UO. Laboratory work was performed by TT, MÅ & BH. All analyses were done by TT with assistance from BH and SB. TT wrote the manuscript with support from other co-authors. 8 POPULATION SIZE AND GENETIC DIVERSITY OF NIGERIAN LIONS (PANTHERA LEO) POPULATION SIZE AND GENETIC DIVERSITY OF NIGERIAN LIONS (PANTHERA LEO) INTRODUCTION 2006). If small isolated populations are vulnerable to extinctions, then large carnivores High growth rate in human populations, are especially prone to extinction due to agricultural developments and industriali- the aforementioned reasons (Woodroffe & zation are leading increasingly to biodiversity Grinsberg, 1998). Many studies have been loss (Ceballos & Ehrlich 2002, Estes et al. carried out on carnivores which revealed that 2011). These anthropogenic activities are most populations are now declining (Craigie responsible for the loss of natural habitats, et al. 2010) and even the remnant populations habitat fragmentation and isolation of natural exist in small, fragmented isolates (Riggio et al. 2012). For instance a study by Proctor et populations. This is driving many species to al. (2005) in the United States of America exist in only small-sized populations. Once a on grizzly bears (Ursus arctos ssp.) revealed continuous population has been fragmented a range contraction and that the remaining into smaller patches with reduction in populations exist in only four fragmented size, isolation from other populations isolates in vulnerably small number (≤100 with restriction in movement, random individuals). The Arabian leopards (Panthera demographic and genetic processes can lead pardus nimr) in Israel have also faced a severe the population rapidly towards extinction reduction in their population size due to (Wilcox & Murphy 1985, Nowell & Jackson anthropogenic activities (Perez et al. 2006). 1996, Turner 1996 in Callens et al. 2011). This population is predicted to be at the brink Large carnivores have suffered great of extinction. Also the wild Scandinavian wolf reductions in their populations and range (Canis lupus) experienced extinction during sizes over the past 50 years mostly due to the first half of the 20th century before re- anthropogenic activities (Nowell & Jackson establishment by immigrants from Finland/ 1996, Estes et al. 2011, Riggio et al. 2012). Russia in the beginning of 1980s (Liberg et Their requirements for large home range, al. 2005, Bensch et al. 2006). limited dispersal ability, low reproductive The lions were once known to roam most rates and subsequent low densities have parts of Africa, Southern Europe, the Middle made them exceptionally sensitive to habitat East and Asia (Bauer & Merwe 2004, Nowell changes (Proctor et al. 2005, Howe et al. & Jackson 1996). Today they are only found 11 Population size and Genetic Diversity of Nigerian Lions (Panthera leo) Population size and Genetic Diversity of Nigerian Lions (Panthera leo) in sub-Saharan Africa and at one locality close relatives i.e. inbreeding results in in India, where they are being increasingly recessive deleterious alleles coming into the restricted to supposedly protected areas limelight as homozygous in the offspring. and often in declining numbers (Nowell & This increases the chances of offspring being Jackson 1996, Chardonnet 2002, Bauer & affected by harmful effects, and indeed Merwe 2004, Wilson & Mittermeier 2009). A inbreeding is known to cause deleterious recent survey has shown that the African lions effects on all aspects of reproduction and are in rapid decline due to habitat loss, with survival (Frankham et al. 2002, Keller & a population decrease from around 100,000 Waller 2002, Bjorklund 2003). The negative animals just fifty years ago to as few as 32,000 consequences of inbreeding on fitness is called today (Packer et al. 2013). Alarmingly, lions inbreeding depression. It can be expressed as have been predicted to be at the verge of reduction in clutch size (in the case of birds), extinction in the near future, just 20-40 years fewer or smaller cubs, lower cub survival from now (Nowell & Jackson 1996, Bauer & (in the case of mammals), and physiological Merwe 2004, Riggio et al. 2012, Packer et al. defects as well as stillborn. Loss of genetic 2013). variation due to genetic drift and inbreeding In West Africa lions are found only in depression is assumed to be a serious problem protected areas such as national parks, game for the conservation of small populations reserves and zoological gardens. In Nigeria (Gilpin & Soule 1986). All species experience today only two protected areas still have wild environmental change whether it is climate lions; Yankari Game Reserve in Central North change, altered competition, new diseases East and Kainji-Lake National Park in the West. or new predators (Frankham & Kingslover, When the population of any species 2004). Genetic diversity is the raw material becomes reduced in size and gets isolated for needed in order to evolve the ability to cope a long period of time it brings about erosion with the aforementioned environmental of genetic diversity. This compromises the challenges. When the genetic diversity is high evolutionary potential of such population to it is more likely that some individuals within environmental changes (Falconer & Mackay a population will possess the alleles suited to 1996, Frankham 1995, 1996, Keller & cope with the environmental change. There Waller 2002). Also such small populations are are many studies on lions demonstrating often faced with a higher risk of extinction the harmful effects of inbreeding. During from environmental catastrophes (Saccheri the period of 2003 and 2008 in a zoological et al. 1998, Dunham et al. 1999, Bijlsma et garden in northern Italy, 19 white lions were al. 2000, Higgins & Lynch 2001) than large recorded to be born to parents that were interconnected populations (Lande 1993, half-sibs (Scaglione et al. 2010). Out of these Frankham 1996, Lacy 2000). The smaller
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