Thevegetatio n ofManyar a Scale-dependent states and transitions in the African Rift Valley CENTRALE LAND B OUW CAT ALO GU S 0000 0801 1260 Promotoren: dr. H.H.T. Prins hoogleraar in het natuurbeheer ind etrope n en deecologi e van vertebraten dr. J. vanAnde l hoogleraar in de plantenoecologie Rijksuniversiteit Groningen »)fio.>V . '.: -' Paul EduardLot h Thevegetatio n ofManyar a Scale-dependent statesan dtransition s inth eAfrica n RiftValle y Proefschrift ter verkrijging van degraa dva n doctor opgeza g van derecto r magnificus van deWageninge n Universiteit, dr. CM. Karssen, inhe t openbaar te verdedigen opvrijda g 8oktobe r 1999 Doctoral thesis (1999); ISBN 90-5808-126-5 Also published in Tropical Resource Management PapersN o 28 (1999);ISS N 0926-9495 Wageningen University and Research Centre BIiiUOTKY^ lANDBCL^UKlV^JTI? f This study was financially supported by theNetherland s Foundation for the Advancement of Tropical Research (WOTRO),Projec t number W84-370 . Cover design: Paul Loth, Ernst van Cleef t)i\0%''ir • ::; 4 '' Stellingen behorende bij het proefschrift: 'The vegetation of Manyara: scale-dependent states andtransition s inth eAfrica n Rift Valley' 1. Deveelhei d aanvegetatievorme ndi ebeslote n ligti n deter m savanne reflecteert het dynamischekarakte r vansavannes . 2. Dehard e zaadhuidva nAcacia soorte n ismed eal saanpassin g aanvuu r tot standgekomen . 3. Intropisch e savannesbepaal t dewaterbeschikbaarhei d in debode m uiteindelijk hetvoorkome nva nbomen . Dit proefschrift Devegetati e van savannes bepaalt de samenstellingva n deherbivoren - gemeenschap, nietandersom . Van Wijngaarden, W. (1985) ITCPub l 4, Enschede Dit proefschrift 5. Opgrot e schaalverandere n vegetatiepatronen niet. Dit proefschrift 6. Het gewicht van groteherbivore n alsparamete r inmodelle n dient gestandariseerd teworden . 7. Groeionderzoek aantropisc hhou t gaathe tbest eme the tblot eoog . 8. Het leefmilieu in stedelijke gebieden wordt, als gevolg vanhe t gedoogbeleid, onvoldoende beschermd tegen het dagelijkse mestoverschotva nhonden . 9. Zolangbestuurder smogelij k belanghebbe nbi j deuitkoms t van wetenschappelijke publicaties, dienthe t auteursrecht bij de wetenschapper te blijven. NRC Handelsblad 28 augustus 1999 Wb 18, 3jun i 1999;W b 26,2 September 1999 10. Ooki n deacademisch e wereld gaattegenwoordi g winst voorwerk . 11. Alshe tbergafwaart s gaatword td emass akritisch . 12. Hetalar mo puniversiteitsgebouwe n dienteerde rte r zelfbescherming van degebruike r dante r beschermingva nhe tgebouw . 13. Tegen open deuren isgee n stelling tenemen . PaulE .Lot h Wageningen, 8oktobe r 1999 .voorMimi,Susanne en Gine Abstract Loth, P.E. (1999). The vegetation of Manyara: scale-dependent states and transitions in the African Rift Valley. Doctoral thesis (1999); ISBN 90-5808-126-5. Also published as Tropical Resource Management Papers No 28; ISSN 0926-9495, Wageningen University and Research Centre, TheNetherlands . This study focuses on scale dependency, both temporal and spatial, of vegetation changes. At different spatial levels, starting from the level of the individual plant, via patch or stand, plant community, and, finally, at the level of ecosystems, the changes in the vegetation of an East African savanna in the Rift Valley are related to the time span considered. The dominant treeAcacia tortilis inLak eManyar a National Park was taken as atypica l case inthi s respect. A. tortilis seed germination and seedling establishment is most successful on bare patches, inth e absence ofothe rvegetation . Forth e reconstruction oftre e establishment in the past, allometric tree growth was studied. Changes in floristic composition between the 1970s and the 1990s were predominantly caused by changes in abiotic factors. The influence of herbivores on the vegetation appears to be minimal on the long-term. The structure of the herbivore assemblage in Lake Manyara National Park, however, is more likely to be determined by changes in the vegetation. Although faunistic and floristic changes occur, these changes must be considered as fluctuations within the ecosystem, and do not lead to a change of the ecosystem itself. It is proposed that scale, both temporal and spatial, must be incorporated in state and transition models. Keywords :tempora l and spatial scale,vegetatio n dynamics, biotic and abiotic factors, Acacia tortilis Acknowledgements I gratefully acknowledge the permission to carry outresearc h in Tanzania granted to meb yth eGovernmen t ofth e United Republic ofTanzania . Thestud y would nothav e been possible without theassistanc e ofman y individuals andinstitutions . Inparticula r Ia mgratefu l toth eCommissio n for Science andTechnolog y (COSTECH),th eSerenget i Wildlife Research Institute (SWRl), and Tanzania National Parks (TANAPA) to allow me towor k in oneo fth e most beautiful areas of East Africa: Lake Manyara National Park. In particular I thank Dr C.A. Mlay and Dr George A. Sabuni of SWRIan d Messrs. B.C.Mwasaga , A.R.Kajuni , M.Melamari , andG . Bigurube ofTANAPA .I am thankful to the Chief Ecologists of TANAPA, Messrs. J. Magombe, A.R. Kajuni, and especially Mr. B.C.Mwasaga , for their kind assistance. Mr.J . Meijer andMaaik eva n Vliet of the Royal Netherlands Embassy have been very helpful in overcoming the initialhurdles . I thank the Netherlands Foundation for the Advancement of Tropical Research (WOTRO)fo r thefinancia l support ofth e study. Ireceive d additional support from the Tropical Nature Conservation and Vertebrate Ecology Group in the Department of Environmental Sciences. Ials othan k my supervisors. Prof. Dr.Herber t Prins andProf . Dr. Jelte van Andel who supervised me, both in the field and during the follow-up phase inth eNetherlands . Foradministrativ e matters Ireceive d full assistance from the Department inth e person ofGerd a Martin,fo rwhic hI than khe rver ymuch . My sincere thanks also to Dr W. Mziray, Director of the National Herbarium of Tanzania, to allow Mr. D.K. Sitoni, to participate in the study. Likewise 1 thank the Department ofWate r topermi t Messrs. P.Mchang a andM . Hieronimi to carry out the ground water survey. Ithan k Mr. E.Ntumbo , Principal ofth e Forest Training Institute Olmotonyi, for the use of equipment. I greatly appreciate the assistance I received from theVillag e Council ofRhoti a Village for theus eo fth e village scale. Ithan kth e International Institute for Aerospace Survey and Earth Sciences (ITC) for providing the opportunity to David Duli to carry out the vegetation survey. In particular Ia m grateful toJa nd eLeeu w ofIT Cfo rhi s enthusiasm andstimulatin g ideas. In the field I was fully supported by the management and staff of Lake Manyara National Park. Iwoul d like tothan k especially thePrincipa l Wardens ofth e Park,Mr . B. Kanza andMrs .M .Kibas a forthei r cooperation, andth e Park's Ecologists, Messrs. F. Leihora andF . Shilkiluwasha, theWarde n Community Conservation Service, Mr. Mtui and the Warden Tourism Mr. M. Mushi for their assistance and interest inth e research. AllRanger s inth e Park lent a hand whenever thenecessit y arose, for whichI thankthem . I am grateful for all the help I received from the following persons. Thad, David and Mike Peterson of DOROBOSafari s whoallowe d met ous ethei r equipment inth e field. Paul and Finny Strebel permitted me to cut trees on their farm and provided hospitality while I did the destructive sampling of trees. The 'Dodoma brothers', Giteh, Paul, Rajinder and Suleh Mandh, were always prepared to assist, and often 1 visited their hide-out. Friends, I thank you. I thank Mr. Ganji of Rift Wall Estate to allow me to carry out experiments on his farm, and for the hospitality I received. Likewise, I thank Paul vand e Vijver to let me work on Manyara Estate. I thank Mr. A.R. Chimbalambala, locally known as Mr. Karangi, Estate Manager of Manyara Estate for guiding me around the farm. Ithan k Kees Terhell and Patricia McCauley of WEGS Consultants for their assistance with the soil auguring project. Hans Baart of Multiflower is gratefully acknowledged for his participation in experiments. I thank Mr. Gordon Alexander of Gargill for providing me with material for experiments. I thank Gerard van 't Land of the Mbulu District Rural Development Programme and Wim van Kampen of the Royal Institute for the Tropics (KIT)fo r their assistance and fruitful discussions, andfo r theirhospitality . Paul Oliver andhi s office staff in Arusha providedth enecessar y links ofcommunication , for whichman ythanks . I wish to mention Joseph Masembe of Aerophoto Systems Ltd. in Nairobi for the aerial photographs made of the Park. I was impressed by the quality of the infra-red photographs thatwer e made in addition toth eblac k and white photography. I can only speak in praise about the quality of the work of Aerophoto Systems Ltd., but unfortunately the delivery cameuntimely . It is hard to express my gratitude to Professors Herbert Prins and Jelte van Andel, who provided conituous support in all stages of my study, not in the least during the writing ofthi s thesis. Iwa s fortunate that both Herbert andJelt e proved tob e hands-on supervisors, who came outt oth ebush . I also benefited from the visits of my colleagues Han Olff, Pieter Ketner, Sip van Wieren, and Arend Brunsting. I shared fruitful discussions and leisurely sundowners with Claudius van de Vijver and Margje Voeten, fellow-PhD students in nearby Tarangire National Park. Also with Peter Hetz and Marianne Kuitert I had in-depth exchanges ofthoughts , alternatedwit hfabulou s family excursionsint oth e Serengeti. The work in the field meant hard work for the crew who carried out the auguring programme. With respect I remember the long days of continuous auguring under the supervision of Peter Mchanga, with the assistance of Michael Hieronimi, Aminiel Gabriel, Shael Josephat and Mashaka Ngurala. Peter Mchanga and his colleague carried out the survey of the bore hole transects.