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The Interspecific Relationships of Black Rhinoceros (Diceros Bicornis) in Hluhluwe-Imfolozi Park

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The Interspecific Relationships of Black Rhinoceros (Diceros Bicornis) in Hluhluwe-Imfolozi Park The interspecific relationships of black rhinoceros (Diceros bicornis) in Hluhluwe-iMfolozi Park Roan David Plotz B.Sc. (ConsBiolEcol) (Hons1); GradDipEd (Sec) A thesis submitted to Victoria University of Wellington in fulfilment of the requirement for the degree of Doctor of Philosophy in Ecology and Biodiversity 2014 1 2 “To Ryker, may the wild places of this world long remain protected to captivate and inspire you” Black rhino near the Black iMfolozi River in Hluhluwe-iMfolozi Park, Zululand, South Africa (Photograph by Dale Morris). “We learn more by looking for the answer to a question and not finding it than we do from learning the answer itself.” Lloyd Alexander 3 4 ABSTRACT As habitat loss, predators (human and non-human) and disease epidemics threaten species worldwide, protected sanctuaries have become vital to species conservation. Hluhluwe-iMfolozi Park (HiP) in South Africa is at the centre of one of the world’s greatest conservation success stories. The formal proclamation of HiP in 1895 prevented the extinction of the south-central black rhino (Diceros bicornis minor) population. In recent times HiP has been a strategic source population for the D. b. minor range expansion program, facilitating an 18-fold population increase across southern Africa. However, HiP’s own black rhino population appears to be in decline. Evidence for decline is most often attributed to overpopulation and poor habitat quality that is driving apparently significant increases in the average home range sizes, poor growth rates (i.e., low calf recruitment) and poor body condition of black rhino. Other factors such as non-human calf predation and parasitism have also been raised as potential causes of decline but remain untested. HiP does have some of the highest densities of lion (Panthera leo) and spotted hyena (Crocuta crocuta). HiP’s black rhino population also suffers from remarkably severe chronic haemorrhaging lesions caused by a filarial parasite (Stephanofilaria dinniki). Empirical evidence if or indeed why the HiP black rhino population might be in decline is lacking. Investigating this population’s true status and any potential causes of an apparent decline is urgently needed. This thesis therefore aimed to test three hypotheses for poor performance that included: (1) investigations of the average black rhino home range size, (2) confirmation of black rhino calf predation and (3) the relationship between filarial lesions and black rhino body condition. I inserted horn-implant VHF radio transmitters into 14 adult (i.e., >5 years) female black rhino in HiP and regularly monitored them on-foot over a three-year period. I found that average home range estimates (9.77 km2) were not significantly dissimilar to estimates using a similar technique obtained forty years prior (i.e., 7.5 km2). I also established the first confirmed link between predation attempts and tail amputation during a lion attack on a black rhino calf. Black rhino body condition, while significantly 5 inversely and temporally correlated to lesion severity, did not appear to be driven by lesion severity itself and highlights the need for further research. An additional research focus for my thesis developed while in the field. I regularly witnessed red-billed oxpeckers (Buphagus erythrorynchus) feeding at black rhino filarial lesions while also alarm calling to alert them to my presence. Studies have found it difficult to empirically show how oxpecker-host interactions have net positive benefits that make it a mutualism. Thus, two chapters were designed to determine if red-billed oxpeckers were predominately mutualistic or parasitic when visiting black rhino. Determining this depended on whether I could identify net positive benefits or net costs to black rhino. Oxpeckers provide rhino with two possible benefits i.e., benefit 1 is cleaning ectoparasites and benefit 2 is increasing vigilance, and one cost i.e., lesion parasitism. More than 50 hours of behavioural observations established that oxpeckers favoured haemorrhaging filarial lesions over sites of tick attachment on black rhino. Moreover, black rhino appeared to be completely tolerant of oxpeckers that fed at lesions. To test whether oxpeckers increased rhino’s anti-predator vigilance, I conducted 84 human approach trials towards black rhino both with and without oxpeckers present. Results showed that rhino were immediately responsive to oxpecker alarm calls and benefitted from more than a two-fold increase in human detection rate and detection distance. Rhino predominately orientated to face towards their sensory blind spot (i.e., downwind) after an oxpecker alarm call. The traditional name (Askari wa kifaru) of the red-billed oxpecker, which translates as the rhino’s guard, appears to be validated. However, future research will need to confirm whether black rhino’s tolerance of parasitic oxpeckers is directly related to vigilance benefits. In summary, black rhino managers in HiP can be confident that the average home range sizes have not increased significantly. Further, predation of calves might be a greater problem than previously realised and requires further investigation. Monitoring changes in the filarial lesion severity of black rhino might be a useful tool for detecting impending changes in a rhino’s condition. Finally, black rhino are clearly eavesdropping and benefitting from oxpecker alarm calls – a co-evolution that has implications for conserving oxpecker populations as well. 6 ACKNOWLEDGEMENTS I am most grateful to my supervisor Dr. Wayne Linklater for being so helpful and encouraging throughout. I could not have asked for someone that was more supportive of all that I was trying to achieve and for that I am forever thankful. Dr. Pete Ritchie, Victoria University of Wellington, was also a wonderfully supportive secondary supervisor to turn to – thank-you! Completing my and I presume all PhDs requires people that are at various times willing to provide assistance that is above and beyond a level I could ever have hoped for starting out. For this I especially thank Bom Ndwandwe for his invaluable efforts in the field. We spent countless days trekking out on foot together while dodging very large mammals, climbing trees and rushing to get back to our vehicle before darkness set in. My time with him afforded me opportunities to learn to read different animal behaviours, spoor and other bush survival techniques that kept me safe. The experience of being fully immersed into a vast untouched wilderness that is Zululand’s Hluhluwe-iMfolozi Park is a rare opportunity that will stay with me forever! Special thanks go to Craig Reid, Sue van Rensburg and Dave Druce from Hluhluwe-iMfolozi Park Management and Research for their support. I am also very thankful for the support and information provided from ongoing personal communications with Dave Cooper, Head Veterinarian Ezemvelo KwaZulu-Natal Wildlife (EKZNW) Game Capture, Jacque Flammand, Black Rhino Range Expansion Project and Dennis Kelly, Section Ranger Mkhuzi Game Reserve, EKZNW. Professor Graham Kerley, Nelson Mandela Metropolitan University, Dr. Doug Armstrong, Massey University, and Benjamin Pocock, Wellington Zoo, are also thanked for their assistance. Remote living in a single room hut in Masinda with no electricity and opportunity to move after dark made it essential to find opportunities to socialise outside the Park and for that opportunity I owe a huge gratitude to Chris Kelly and Simon Morgan at Mkhuze Game Reserve. They were not just good blokes to hang out with but highly experienced in the field with all things black rhino. I am thankful for the logistical support of Masinda and Mbhuzane Section Rangers San- Marie Ras and Emile Smidt. 7 I am very thankful to EKZNW, who approved my research (permit number: ZC/101/01). Regularly finding the rare beast that is the black rhino required horn- implant radio-transmitters to be inserted. I am grateful for the logistical expertise (i.e., helicopters, 4-WDs, people and rhino sedatives) of the amazing EKZNW Game Capture Team that made this possible. I especially thank Dave Cooper, Jeff Cooke, Paul Jennings and Quinton Rochat for both their professional expertise but also ongoing patience.The research was made possible through granted funds from the Australian Geographic Society, Rufford Small Grants (ref: 15.10.07), U.S. Fish and Wildlife Service administered Rhino and Tiger Conservation Act of 1994 (grant agreement numbers 98210-6-G102, 98218-8-G690) and Victoria University of Wellington Summer Scholarship Scheme grant. I also thank Kees Rookmaaker and the world’s largest rhino information website for access to references (Rhino Resource Center: http://www.rhinoresourcecenter.com). Although sporadic, my time as a student in the beautiful weather battered city of Wellington, New Zealand, was a pleasure. I thank everyone at the School of Biological Sciences at Victoria University of Wellington who played a part in making it such a colourful experience: Monica Awasthy, Beaux Berkeley, Ilse Corkery, Alex Dixson, Megan English, Elizabeth Heeg, Mary Murray, Gesine Pufal, Patricia Stein, Sandra Taylor and Gaius Wilson. I especially thank fellow VUW students Barnaby Dixson, Andrew Stringer and Leighton Thomas for being good friends in Wellington. Fellow black rhino PhD students Andrew Stringer and Rosalynn Anderson-Lederer are thanked for their field assistance, but also for being an ear for ongoing black rhino discussions. I thank Paul Marsden for running the SBS indoor Soccer team, providing me with a much needed physical outlet. Finally, I thank my family and in particular my parents Linda and Tony Stumbles and my wife Liana Cahill for everything but also for putting up with my long absences and providing the emotional and often financial support that a PhD conducted over three continents requires – I can never repay your kindness. Thanks also to Professor Des Cahill and Maria Cahill for their support, it is gratefully appreciated. Kelly and Mary-Jane Gate are thanked for looking after Liana at the Bethesda Hospital in Ubombo, where she volunteered for 6-months at a time while I pursued rhino.
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