How elephants utilize a miombo-wetland ecoystem

How elephants utilize a miombo-wetland ecosystem in Ugalla landscape, Western

Elikana Kalumanga

Department of Physical Geography Stockholm University 2015

1 Elikana Kalumanga

© Elikana Kalumanga ISSN 1653-7211 ISBN 978-91-7649-161-4

Paper I © Elikana Kalumanga Paper II © Elikana Kalumanga Paper III © Elikana Kalumanga Paper IV © Elikana Kalumanga

Cover: Elephants drinking water from Ugalla River in Ugalla Game Reserve Photo by Elikana Kalumanga Publisher: Holmbergs, Malmö 2015 Distributor: Department of Physical Geography, Stockholm University

2 How elephants utilize a miombo-wetland ecoystem

Doctoral dissertation 2015 Elikana Kalumanga Department of Physical Geography Stockholm University

Abstract

African elephants are ‘keystone’ species with respect to biodiversity conservation in since they maintain habitats that support several animal communities by changing vegetation structure through foraging and by dispersing seeds between landscapes. Elephants are also ‘flagship’ species because, given their impressive size, they can make people sympathetic and stimulate local and international concerns for their protection. Economically, elephants contribute to national revenues as tourists are willing to pay to watch them. Despite all these factors, little is known however about elephant movement and how they utilize resources, especially in miombo-wetland ecosystems. This thesis investigates how elephants utilize resources in a miombo-wetland ecosystem in the Ugalla landscape of Western Tanzania over different protected areas containing different resource users. Using Global Positioning System (GPS) collars fitted to six elephants, it was observed that some elephant families are not confined in one protected area in the Ugalla landscape. Rather, they moved readily between different protected areas. Elephant movements were restricted to areas near the rivers, especially the Ugalla River, during the dry season and were dispersed widely during the wet season. As they move, elephants in the miombo woodlands of Ugalla selected the most abundant woody plants for browsing. Common to many woody plants, the browsed plants were short of mineral nutrients (e.g., sodium, calcium). Elephants obtained additional minerals by eating soils from certain termite mounds. Soils from termite mounds are richer in mineral elements (e.g., sodium, calcium, iron) compared to soils from the surrounding flood plain or compared to the browsed plants. However, the recorded termite mounds from which elephants eat soils were not evenly distributed in the landscape but confined mainly to the flood plains in the Ugalla Game Reserve. The Ugalla River, which is the main source of water for the elephants and other animals and also supports fishing activities by the local people in Ugalla during the dry seasons, is infested by the water hyacinth (Eichhornia crassipes). Such infestation potentially limits access to these precious surface water supplies. In addition at the regional level, the Ugalla River is among the major rivers that flow into the which is shared by the countries of Tanzania, , Democratic Republic of Congo and Zambia. Thus, the spread of water hyacinth if left unchecked threatens to impact Lake Tanganyika, affecting many countries and ecosystem services. This thesis highlights that sustainable conservation of biodiversity in different protected areas in the Ugalla landscape requires an integrated management approach that will embrace conservation of different interrelated landscape resources required by both wildlife and the rural poor populations for their livelihoods. Regular coordinated wildlife anti-poaching patrols should be initiated across the entire Ugalla landscape because the elephants, among other wildlife, utilize different protected areas in Ugalla. Local communities should also be engaged in conservation initiatives (e.g., controlling the spread of the water hyacinth) as these directly impact local livelihoods.

Keywords: Biodiversity, Browsing, Elephants, Forage, GPS collars, Mineral elements, Miombo woodlands, Protected areas, Termite mounds, Ugalla, Water hyacinth, Woody plants abundance

3 Elikana Kalumanga Sammanfattning

Den afrikanska elefanten är en så kallad nyckelart för biologisk mångfald på många platser i Afrika. Den påverkar sin livsmiljö genom att ändra vegetationen när den söker efter föda, och genom att sprida frön, vilket i sin tur påverkar många andra djur. Elefanter har också ett stort symbolvärde när bevarandestrategier diskuteras. Folks sympati och oro för elefanternas framtid och skydd finns både på lokal och internationell nivå. I ett land som Tanzania är turism en av de stora inkomstkällorna för utländskt kapital och safariturismen står för 80 % av inkomsterna. Många turister vill se just elefanter i sin naturliga miljö och de utgör därför en stor inkomstkälla för safariturismen. Elefanter är inte bara viktiga för den biologiska mångfalden utan för hela Tanzania. Afrikas elefanter har minskat kraftigt i antal under de senaste 200 åren. I början av 1800-talet fanns det cirka 24 miljoner elefanter. På grund av jakten på elfenben minskade antalet till 4 miljoner i slutet av 1800-talet. Trots stora insatser för att bevara elefanterna under 1900-talet finns det idag bara cirka 500 000 kvar i Afrika. Antalet fortsätter att minska på grund av tjuvjakt och krympande områden och svårare att röra sig mellan områdena. Ett stort djur som elefanter behöver stora områden att söka föda på men idag utgör förutom tjuvjakt också ett mer intensifierat jordbruk och ett ökat befolkningstryck ett hot mot de kvarvarande elefanterna. Tanzania har avsatt 25 % av sin landyta för olika naturskyddsområden, men trots det är utrymmet för elefanter begränsat. Även om elefanterna har ett stort symbolvärde vet vi idag lite om hur de rör sig i landskapet och vilka resurser de utnyttjar. Det mesta av den ekologiska forskningen i Tanzania har gjorts på savannen där kunskapen från andra typer av ekosystem är begränsad. I den här avhandlingen studeras hur elefanter rör sig och utnyttjar olika naturresurser i ett miombo-våtmarkssystem i Ugalla i västra Tanzania. Miombo-skog är ett ekosystem som kan ses som en övergång mellan den öppna savannen och tät regnskog. Oftast fäller träden bladen under torrperioder och skogsbränder är en naturlig del av ekosystemets störningsregim. Ugalla är en del av Malagarasi-Muyovozi Ramsarområdet vilket är ett stort våtmarkskomplex med många unika växt- och djurarter. Ramsarkonventionen är en global naturvårdskonvention om att bevara våtmarker och vattenmiljöer och nyttja dem på ett hållbart sätt. Våtmarkerna dräneras ut i Tanganyika-sjön. Ugalla är 5000 km2 stort men omges av andra skyddade områden som tillsammans omfattar mer än 13 000 km2. Idag bor det inga människor i Ugalla då det finns tsetse-flugor som sprider sömnsjuka. Ugalla är ett unikt reservat eftersom lokalbefolkningen tillåts fiska och bedriva biodling i området under torrperioden. Jaktbolag bedriver licensjakt på framförallt krokodil och flodhäst. Det finns idag cirka 4000 elefanter i Ugalla. Området patrulleras av beväpnande vakter för att skydda djuren från tjuvjakt. Trots detta tycks tjuvjakten öka i området på grund av en ökad mängd vapen, lättare att transportera ut tjuvgodset samt en stor efterfrågan på elfenben och elefanthår från Asien. Avhandlingen kartlägger hur elefanterna rör sig i landskapet, vad de äter och hur de får i sig nödvändiga mineral. Sex elefanter från olika grupper sövdes och förseddes med GPS-kragar under två år, vilket gav en position för varje elefant fyra gånger om dagen. Detta är den första studien i Tanzania från ett miombo-landskap som har använt GPS-märkning. Resultaten visar att vissa elefanter inte är begränsade till de skyddade områdena nära floderna utan rör sig över stora delar av landskapen runt omkring Ugalla. Under torrperioden håller sig grupperna nära floderna medan de skingras över stora områden under regnperioden då stora delar är översvämmade och det finns gott om vatten i landskapet. De rör sig i områden med de vanligaste trädarterna för att äta löv, grenar, bark och rötter. Elefanter anses vara generalister i sitt val av föda, men resultaten här visar på att de kan vara selektiva i vilka växter de äter. Gemensamt för de växter som elefanter äter är att de är fattiga på mineral och vissa näringsämnen (t.ex. natrium och kalcium). Elefanter får i sig mineral genom att äta jord från vissa termitstackar. Jorden i dessa termitstackar är rikare på mineralämnen (t.ex., natrium, kalcium, järn) jämfört med jorden från den omgivande översvämningsslätten. De termitstackar som elefanterna äter av är begränsade till vissa platser i landskapet varför det är viktigt att de inte blir förstörda och att området intill lämnas i fred för att inte störa elefanterna. Ugalla-floden utgör den huvudsakliga vattenkällan för elefanter och andra djur och det bedrivs fiske där av lokalbefolkningen under den torra årstiden. På senare tid har den invasiva växten vattenhyacint (Eichhornia crassipes) etablerats i floderna i området. Växten kommer ursprungligen från Sydamerika och den breder ut sig i täta tjocka mattor på vissa ställen av floden. På lokal nivå utgör växten ett hot då den begränsar den öppna vattenspegeln vilket lett till fler flodhästattacker, med dödlig utgång, på fiskare då utrymmet blir begränsat. När den öppna vattenspegeln försvinner

4 How elephants utilize a miombo-wetland ecoystem skapar det också problem och stora kostnader för jaktbolagen, vilket i sin tur kan skapa stort inkomstbortfall i framtiden. Djur får svårt att dricka och fiska i områden med täta bestånd av vattenhyacinter, och de kan också sprida växten vidare. Om vattenhyacinterna skulle sprida sig till Tanganyikasjön blir det ett internationellt problem eftersom fyra länder är beroende av sjön (Tanzania, Burundi, Demokratiska republiken Kongo och Zambia). Om vattenhyacinten invaderar Tanganyikasjön kan det få stora direkta konsekvenser för befolkningen och djur som lever i och av sjön. Denna avhandling visar att för att skapa ett hållbart bevarande av biologisk mångfald krävs integrerade styrmetoder som omfattar bevarandet av olika resurser som krävs för både djurliv och den fattiga landsbygdsbefolkningen. Regelbundna samordnade anti-tjuvjaktpatruller bör initieras över hela Ugalla och dess omgivande områden eftersom elefanterna, bland andra vilda djur, inte bara använder de områden som ligger närmast floden. Ramsarkonventionen rekommenderar ett lokalt engagemang och därför bör den lokala befolkningen engageras för att kontrollera spridningen av vattenhyacint eftersom det har en direkt påverkan på den lokala näringen. Olika områden och naturresurser faller under olika förvaltningar och ministerier, skog, turism, jordbruk, jakt osv men för att bevara den biologiska mångfalden och samtidigt människors möjlighet att få inkomster på ett hållbart sätt krävs en integrerad syn på Ugalla och dess tillgångar.

Nyckelord: Abundans, Biodiversitet, Elefanter, Födosök, GPS övervakning, Mineraler, Miombo, Näringsinnehåll, Skyddade områden, Termit, Ugalla, Vattenhyacint, Vegetation

5 Elikana Kalumanga

To my late father Mzee Ephraim Elisha Kalumanga

6 How elephants utilize a miombo-wetland ecoystem

How elephants utilize a miombo-wetland ecosystem in Ugalla landscape, Western Tanzania

Elikana Kalumanga

Department of Physical Geography, Stockholm University, Sweden

Thesis content

This doctoral thesis consists of a summary and four papers. The papers are referred to as Papers 1 to IV in the summary text:.

List of papers

Paper I: Kalumanga, E., Nicholson, K.L., Mpanduji, D.G., Cousins, S.A.O. Home ranges and utilization distributions of African elephants in Ugalla landscape, Western Tanzania. Manuscript to be submitted to the journal of Conservation Biology

Paper II: Elikana Kalumanga & Sara A. O. Cousins. Woody plant abundance drives the browsing patterns of elephants in the Ugalla landscape, of Tanzania Manuscript.

Paper III: Elikana Kalumanga, Donald G. Mpanduji, and Sara A. O. Cousins. Geophagic termite mounds as one of the resources for African elephants in Ugalla Game Reserve, Western Tanzania. Manuscript resubmitted after revision to the African Journal of Ecology.

Paper IV: Elikana Kalumanga. Why Integrated Natural Resource Management in Malagarasi- Muyovozi Ramsar Site, Tanzania? Insights from Ugalla Game Reserve. Manuscript resubmitted after revision to the Marine and Freshwater Research: Ramsar Special Issue.

Co-authorship

I have been responsible for the elephant collaring, under the guidance of the co-authors DGM and SAOC for paper I. I am responsible for the data analysis and lead the writing while KLN helped with the data analysis tools and interpretation of the results for paper I. In paper II and III, I designed the study together with SAOC and I collected all the data in the field, sampled leaves and soils, and did the statistical analyses and lead the writing. DGM contributed in the study design for paper III, and commented on the manuscripts.

7 Elikana Kalumanga

8 How elephants utilize a miombo-wetland ecoystem

biodiversity is limiting the mismatch between Introduction the scale of management and the scale(s) of the ecological processes being managed (Cumming et al., The sustainable development of our societies and our 2006; Andrade & Rhodes, 2012; Maciejewski et al., general wellbeing are linked to the existence, survival 2015). The mismatch problem is more pronounced and welfare of the many other species with which in areas where wildlife traverse in different areas or we share the planet Earth (Millennium Ecosystem where different stakeholders with different goals are Assessment 2005; Grant et al., 2008; Naeem et al., involved (e.g. Metcalfe & Kepe, 2008). In Tanzania 2009; Turner et al., 2014; WWF 2014). Biodiversity for example, more than 25% of the nation is set aside (i.e. the variety of genes, species, communities, for conservation within different protected areas ecosystems and ecological processes) provides various (Table 1). Still, despite these vast protected areas, the ecosystem goods and services essential to economic country is facing a number of conservation challenges prosperity and even survival. This is especially true in ranging from wildlife poaching (e.g. Jambiya et al., developing countries (Primack, 1993; Vining, 2003; 2007; Caro, 2008; Rija, 2011; Martin & Caro, 2012; Millennium Ecosystem Assessment 2005; Turner et EIA, 2014), loss of wildlife migratory corridors al., 2014). However, around the globe, biodiversity (Newmark, 1996; Jones et al., 2009; Kideghesho and its benefits to humanity are disappearing at an et al., 2013), encroachment into protected areas by alarming rate (Myers, 1988; da Fosenca et al., 2005; farming, settlements among other land use activities Sachs et al., 2009; Mittermeier et al., 2011). Threats (Madulu, 2001; Mangora, 2005; Sinclair et al., 2008), facing both terrestrial and aquatic environments and infestation of invasive species (Kideghesho et al., (and the species that occupy them) are increasing 2013). As a result of such mixed issues and interests, and becoming more persistent (Turner et al., 2014). it is increasingly challenging to sustain wide-ranging To a large extent, such loss of ecosystem services species that require large space and that sometimes translates to direct impact on the livelihood of rural come into conflict with people. An exemplar of this poor people. This is because, compared to many mismatch problem within conservation, particularly societies globally, it is the livelihoods of the rural at the regional level, would be elephants throughout poor that rely on ecosystem goods and services from its range countries in Africa (Lee & Graham, 2006; biodiversity (i.e. their ‘biological supermarkets’). Van Aarde et al., 2006; Metcalfe & Kepe, 2008; Any loss of biodiversity and changes in ecosystem Delsink et al., 2013). goods and services makes the rural poor more vulnerable (Madulu, 2007). The loss of biodiversity can Why studying African elephants also have impacts on national and regional economies. (Loxodonta africana Blumenbach 1797) For instance, the safari and hunting industry in Africa depends on the existence of biodiversity. In many Elephants are the largest terrestrial mammals on countries (e.g. Tanzania), tourism is one of the largest Earth. Compared to other wild mammals, elephants sources of foreign currency with wildlife-based are ‘keystone’ species because they play important tourism often accounting for more than 80% of all ecological, social and economic roles in the landscape. tourism (Mariki et al., 2011). The global community Elephants are known to play an important ecological has become increasingly aware of the importance role in savanna and forest ecosystems. They help to of biodiversity as it tries, to some extent, to develop maintain suitable habitats and subsequent linkages conservation plans to diminish losses (Grant et al., in the food-web by greatly affecting the structure of 2008; WWF, 2014). However, the world is big and vegetation that supports several animal communities resources are limited, thus conservation cannot be (Stephenson, 2004; Batweteera et al., 2007; Blake done everywhere equally or in the same way (da et al., 2009; CITES 2010; Campos-Arceiz & Blake, Fosenca et al., 2005). Biodiversity conservation 2011). Due to their popularity and appeal to an must be coordinated within certain geographical international audience, and probably also because areas or for some target species or habitats. Further, of their impressive size, elephants are ‘flagship’ there are needs to develop focused actions at both species and have an ability to stimulate the public, local and regional scales with the ultimate goal of governments and other institutions to catalyze achieving concrete, measurable and time-bound landscape-scale conservation goals (Barua et al., biodiversity conservation outcomes (da Fosenca et 2010). Economically, elephants are there for a huge al., 2005; Redford et al., 2011a; 2011b). Altogether, asset for local, national and international economies determining where and how to design conservation to (Håkansson, 2004; Coutu, 2011; Delsink et al., 2013). secure biodiversity is a daunting task. They have become an important source of revenue Globally and across different regions, many areas through tourism as many people are prepared to pay are completely set aside for protection; however, this large sums of money either to watch and photograph is not entirely necessary for biodiversity conservation elephants in the wild or, unfortunately, to hunt them (Thirgood et al., 2004). Well-managed, protected for sport. areas have been seen as the most effective tools to For several centuries, the elephant population in protect biodiversity at the site level (da Fosenca et Africa has been in decline due to hunting (mainly for al., 2005; Turner et al., 2014; Maciejewski et al., ivory) (Håkansson, 2004; Burn et al., 2011; Coutu, 2015). The challenge faced by societies in managing 2011; Maisels et al., 2013; Wittemyer et al., 2014),

9 Elikana Kalumanga culling programs (Rodgers & Lobo, 1980), and loss of cessation of slavery and restrictions in large-scale habitats (Jones et al., 2009; WWF, 2010). According ivory exploitation and colonial hunting meant that the to Håkansson (2004), killing of elephants for ivory in elephant population was able to increase after 1900 Africa can be traced back for more than 4000 years, (Rodgers & Lobo, 1980). Despite these restrictions, when ivory was a highly valued commodity in Egypt. the numbers of elephants still declined from 3-5 Massive killing of African elephants happened million in the 1930s-1940s to some 1.3 million in during the era of slavery and ivory trade before the 1979 primarily due to poaching (ATE, 2010; WWF, 1890s (Rodgers & Lobo, 1980; Häkansson, 2004; 2010; Coutu, 2011). Coutu, 2011). Between 1800 and 1900, the elephant By 1989, the estimated number of African elephants population in Africa is estimated to have declined from was about 632000 (WWF, 2010). Ten years later, 24 million to 4 million (Coutu, 2011). Between 1840 there were possibly fewer than 500000 individuals and 1875, the demand for ivory from the East African (Blanc et al., 2007, WWF, 2010). Thus, despite elephants to Britain alone reached approximately active conservation measures, more than 50% of from 200 tons to over 800 tons, which could roughly African elephants were killed (mainly for ivory) by be equivalent to the killing of between 4000 to 17000 poaching and culling programs between 1979 and elephants per year (Beachey, 1967). The boom in 1989 (Rodgers & Lobo, 1980). Recently, (especially ivory trade did not last beyond 1900, however, as from 2011), elephant poaching has been occurring at elephants became increasingly scarce and colonial the highest rates since the 1980s (TRAFFIC, 2012; governments attempted to restrict elephant hunting UNEP, 2012). Today, elephants are found in only 38 by setting quotas, by charging license fees and setting countries south of the Sahara; i.e. African Elephant aside game reserves (Rodgers & Lobo, 1980). The Range States1 (Blanc, 2008). Due to killing for ivory,

Table 1: Categories of protected areas in Tanzania Protection Resource use types Governing Law (s) Land category category 1. National Parks Highest level of resource protection. The National Parks Act, 2002 Reserved Conservation through non-consumptive (URT, 2002a) land utilization (e.g. Photographic safaris) 2. Game Reserves Both consumptive and non-consumptive The Wildlife Conservation Act, Reserved utilization through pre-set hunting quotas, 2009, (WCA-2009) which land except in the Ugalla Game Reserve where has repealed and replaced the some restrictions have not been applied to Wildlife Conservation Act, 1974 persons whose ordinary places of residence (URT, 2009) were within the Ugalla Game Reserve (WCA, 1974 Section 7 (1). Wagalla in Ugalla Game Reserve are legally permitted to conduct fishing and beekeeping. 3. Game Con- Both consumptive and non-consumptive The Wildlife Conservation Act, Reserved trolled Areas utilization. For instance, in MMRS, activities 2009 (URT, 2009). Contrary to land such as crop farming, livestock keeping, the Wildlife Conservation Act, fishing, beekeeping, licensed hunting had been 1974, the WCA-2009 prohibits done in GCAs (Yanda et al. 2002; URT 2007) residence, livestock keeping and cultivation. 4. Forest Reserves Only licensed activities (e.g. Timber The Forest Act 2002 (URT, Reserved harvesting) 2002b) land 5. Nature Reserve Another high level of resource protection. The Forest Act 2002 (URT, Reserved Main legal activity is wildlife-based tourism 2002b) land 6. Wildlife Man- Fishing, Beekeeping and licensed hunting The Wildlife Conservation Act, Village land agement Areas (URT, 2003) 1974 (Wildlife Conservation Act, 2009) (URT, 2009) 7. Ngorongoro Multiple land use area; Conservation, Tourism The Wildlife Conservation Act, Reserved Conservation and Pastoralism 2009 (URT, 2009) land Area 8. Open areas Adjacent to other protected areas; acting as The Wildlife Conservation Act, across buffer zones with restriction to different land 2009 (URT, 2009) Village, uses except licenses hunting General, Reserved lands

10 How elephants utilize a miombo-wetland ecoystem among other factors, elephants became extinct in Elephant population trend in Tanzania 1913 in Gambia, in the 1920s in Swaziland, in the 1970s in Burundi and in the 1980s in Mauritania Until the beginning of the colonial era (1896), (Blanc, 2008). In the 1980-1990s elephants were elephants were found in most landscapes in Tanzania reintroduced in Swaziland (Blanc, 2008). Besides (Rodgers & Lobo, 1980). During German rule in poaching for ivory, African elephants have presently Tanganyika (Tanzania mainland), some selected less suitable space than ever before (Stephenson, men were employed to shoot elephants in Southern 2004). There is continuous decline in the extent Tanganyika to supply elephant meat to German and quality of elephant habitat as their range is commissariat during the First World War (WW1) fragmented and made discontinuous due to various (Spinage, 2012). However, as Rodgers & Lobo, anthropogenic activities (TAWIRI, 2010). In total, (1980), puts it, ‘the elephant killing for meat and fat the elephant range extent in Africa is reported to have could not have significantly affected the population declined from 7.3 million km2 in 1979 to 5.9 million of elephants in these areas because the elephant km2 in 1987 and further to 5.7 million km2 in 1998 population was said to probably increase at the rate (WWF, 2010). Of the remaining range, almost 80% of 3,000 per year in 1932’. The main reason for this is is located outside protected areas (WWF, 2010), as likely that elephants had few restrictions to their range elephants often occupy ranges extending into both as people were sparsely inhibiting most of country. protected and unprotected areas (Mpanduji, 2004; For instance, until the 1930s villages in the Southern Blanc et al., 2007; Blanc, 2008). Tanzania had on average 3-10 huts separated at about The existing African elephants are capable to adapt a distance of 27 kilometers (Rodgers & Lobo, 1980). to various environments. They can be considered as After the Germans left Tanganyika, the British a generalist species surviving in deserts (e.g. Etosha Government (from late 1910s) established an elephant National Park in Namib, Gourma Sahel region in control scheme to reduce the human-elephant Mali), savanna, woodlands and wooded grasslands in conflicts. During the implementation of the scheme, East and Southern Africa, equatorial forests (Congo elephants were killed in some parts of Tanzania (by and Gabon), agricultural landscapes (e.g. Sebugwe then Tanganyika) wherever they came into conflict region in Zimbabwe), bushlands (e.g. Tsavo National with people (e.g. during crop raiding). The first Park in Kenya), swamps, as well as on high-altitude elephant control scheme took place in 1922 (South mountains (e.g. Kibale in Uganda, Kilimanjaro in Mahenge scheme) followed by five other schemes Tanzania and Aberdares range in Kenya) (Kikoti, by the year 1933 in the regions of Iringa, Tukuyu, 2002; de Beer et al., 2006; Rode et al., 2006; Blanc Kilwa (considering both hippos and elephants), et al., 2007; Blanc 2008; Ngene et al., 2008; Canney South Mahenge, Rufiji (considering both hippos and & Ganame, 2012; Granados et al., 2012; Vanleeuwe elephants) and Dar es Salaam District (Rodgers & et al., 2014). Currently, sustainable management and Lobo, 1980). The approval of large-scale elephant conservation of African elephant populations, both control schemes in Tanganyika (Tanzania mainland) within and around protected areas, is among the major in 1933 killed more elephants in subsequent years conservation challenges in Africa (Thouless, 1994; as compared to the similar schemes started in Hoare, 2000; Galanti et al., 2006). Despite the wealth 1922 (Table 2). As a result, the elephant control of studies on elephants, particularly from savannah schemes produced considerable quantities of ivory, ecosystems, there is still a lack of information on how the revenues of which accrued to the government elephants make use of their specific habitats across (Rodgers & Lobo, 1980). Until the early 1970s, spatial and temporal scales. In most cases, habitats regardless of legal and illegal elephant killings, used by elephants are described at broad scales and Tanzania had still one of the largest populations of generalized via a three scale measure: (1) the known elephants on the continent, second to the Democratic range, (2) the possible range and (3) the doubtful Republic of Congo (Spinage, 2012). However, range (Blanc et al., 2007). Thus, detailed ecological during the late 1970s the elephant control scheme in information to inform elephant conservation and Tanzania was ended bringing about conditions where management are missing throughout its range (Poole, the commercial ivory poaching increased. Large 1996; Blanc 2008) and particularly outside savannah scale commercial poaching of elephants for ivory ecosystems. was rampant during much of the 1980s, reaching its peak between 1986 and 1989 due to increased ivory demand in Asia and other markets (Siege & Baldus, 1 African Elephant Range States are: Angola, Benin, 2000). To combat elephant poaching, the Tanzania Botswana, Burkina Faso, Cameroon, Central African government initiated the Operation Uhai in 1989. Republic, Chad, Republic of Congo, The Democratic This was an anti-poaching campaign involving huge Republic of Congo, Ivory Coast, Equatorial Guinea, government resources to overcome illegal hunting. Eritrea, Ethiopia, Gabon, Ghana, Guinea, Guinea- For the first time, Tanzania’s armed forces were Bissau, Kenya, Liberia, Malawi, Mali, Mozambique, deployed around the country to help in combating the Namibia, Niger, Nigeria, Rwanda, Senegal, Sierra illegal killing of elephants (Siege & Baldus, 2000). Leone, Somalia, South Africa, Sudan, South Sudan, After the Operation Uhai in 1989 most elephant Swaziland, United Republic of Tanzania, Togo, population in Tanzania started to increase (Figure 1). Uganda, Zambia and Zimbabwe. However, since late 2000s the elephant population

11 Elikana Kalumanga Table 2: Number of elephants shot in Tanzania on control schemes from 1922 to 1975 (adapted from Rodgers and Lobo).

Year Total Year Total Year Total 1922 218 1941 1067 1960 2,497 1923 590 1942 1067 1961 3,171 1924 422 1943 1067 1962 3,250 1925 394 1944 1067 1963 3,247 1926 576 1945 1493 1964 2,993 1927 1,027 1946 2,272 1965 3,105 1928 450 1947 1,956 1966 3,430 1929 639 1948 1,651 1967 3,506 1930 619 1949 2,098 1968 3,580 1931 687 1950 2,075 1969 3,765 1932 604 1951 1,837 1970 2,886 1933 1,908 1952 2,420 1971 3,525 1934 2,716 1953 2,516 1972 2,835 1935 2,694 1954 2,283 1973 3,472 1936 2,674 1955 1,923 1974 1,957 (Southern Tanzania) 1937 1,481 1956 2,137 1975 1,965 (Southern Tanzania) 1938 1,653 1957 2,040 1939 1,067 1958 2,599 1940 1,067 1959 2,594 1922-1975 106,832 in Tanzania has started to decrease. For instance, the securing extended wildlife habitats and helping the largest elephant population in Tanzania is in the Selous local people to retain usage rights over the wildlife Game Reserve. This population was estimated to be resources (Wilfred, 2010). Almost all WMAs are between 50000 and 70000 individuals in 2006 which connected to other protected areas. As WMAs are has since declined to less than 40000 individuals in a relatively new category of protected areas in the 2009. The sharp drop in elephant population in Selous country, information on how wildlife species (e.g. Game Reserve was a result of increasing poaching elephants) utilize the WMAs in relation to other in Tanzania. Another sharp decrease in elephant protected areas is needed to ensure their management population was noted in Ugalla Game Reserve in success for both people and wildlife. western Tanzania where elephant numbers declined from 4000 individuals in 2006 to approximately 1000 individuals in 2009 (TAWIRI, 2010). Besides poaching, the elephant range in Tanzania has also Objectives and scope decreased from almost 460000 km2 in 1998 (49% of country area) to approximately 370000 km2 (39%) in 2009 (TAWIRI, 2010). The areas that the elephants The overall objective of this thesis is to increase our use to disperse between different protected areas, so understanding on biodiversity conservation in Africa, called elephant corridors, have also become more especially in areas set aside for wildlife protection. and more encroached by various human activities This is achieved by studying the spatial and temporal (Figure 2). Therefore, to sustain the remaining use of habitats by African elephants in a miombo- elephant populations in Tanzania, evidence-based wetland ecosystem in Western Tanzania. Specifically data on the elephants themselves (population) and the study intended to i) determine where and when their habitats (key resources) are needed to inform elephants utilize the landscapes surrounding Ugalla; focused conservation efforts in different parts of the ii) identify the spatial patterns of important food country. Such data are crucial because elephants in sources for the elephants; iii) analyse why elephants Tanzania utilize landscapes managed by different select some woody plants for browsing and some stakeholders with potentially different interests and/ termite mounds to eat soils (i.e. geophagy); and or needs. Specifically to address these concerns, finally iv) to determine how an infestation and spread the government of Tanzania introduced Wildlife of water hyacinths in Ugalla River might affect, not Management Areas (WMAs) as a new category of only elephants and wildlife in the Ugalla landscape, protected areas in the country in the early 1990s. but also human fishing access in the Ugalla River Different from other protected areas, WMAs are during the dry season. established on village lands with the intention of This kind of information (i.e., the what and how of

12 How elephants utilize a miombo-wetland ecoystem

Tanzania elephant population trend 1989 - 2009 150,000

125,000

100,000

75,000 Population size Population

50,000

25,000

0 1989 1994 1999 2004 2009 Year

Figure 1: Tanzania’s elephant population trend 1989-2009 (Adapted from TAWIRI, 2010) elephant-habitat interaction) is important to the local mm/year (Kashaigili & Majaliwa, 2010). The mean managers of Ugalla as it can enable them to develop maximum and minimum temperature during the dry and improve their day-to-day wildlife conservation and wet seasons ranges between 28-30oC and 15- activities. A better understanding of processes that 21oC, respectively (Wilfred, 2012). impact on biodiversity issues is also important Ugalla is covered by Miombo woodlands with for the Ministry of Natural Resource and Tourism some open flood-plains near the rivers that run (MNRT) representing the custodians of natural through the landscape (Kalumanga et al., unpublished resource management in the Tanzania. In Tanzania, 3). The Miombo ecosystem is the most widespread it is the MNRT that is responsible for enforcement woodland ecosystem in Africa bisecting the continent of conservation laws and for overseeing the directly south of the Congo Basin and East African implementation of various international conservation savannahs. It covers an area between 2.5 to 4 million treaties in the country, such as the Convention on km2 depending on the exact definition of Miombo Biodiversity Conservation (CBD) and the Ramsar considered (Campbell et al., 1996; Malmer, 2007; Convention on Wetlands. It is the MNRT that is Ribeiro et al., 2008; Munishi et al., 2011; Dewees directly responsible for establishment of wildlife et al., 2011). The dominating species in Miombo management areas (WMAs) in Tanzania. woodlands are characterized by the dominance of some genera such as Brachystegia, Julbernadia and Isoberlinia which seldom occur outside the Miombo ecosystem (Campbell et al., 1996; Frost, 1996; Study area Ribeiro et al., 2008; Munishi et al., 2011; Dewees et al., 2011). Location, climate and vegetation types Management and drainage The data for this thesis was collected from the Ugalla Ugalla is part of the Malagarasi-Muyovozi Ramsar landscape situated in Western Tanzania between o o Site (MMRS) combining different protected areas latitude 5-6 south and longitude 31-33 east. Here, the managed by different agencies under the Ministry Ugalla landscape is the region comprised of the Ugalla of Natural Resources and Tourism, the district Game Reserve, Luganzo Game Controlled Area authorities and the villagers. These areas include (GCA), Msima GCA, Ipole Wildlife Management forest reserves, game reserves, game controlled areas Area (WMA), Uyumbu WMA and North Ugalla and and wildlife management areas (Figure 3; Table 1). Walla River Forest Reserves (Figure 3). The climate At the regional level, MMRS constitutes more than of the region is tropical in nature defined by a wet 30% of the Lake Tanganyika drainage basin. Lake season (November to May) and a dry season (June to Tanganyika, the second deepest lake in the world, October). Ugalla receives bimodal rains; short rains is a trans-boundary lake shared by four countries in November-December and a main, longer rainy namely; Tanzania, Democratic Republic of Congo, season in April-May with an average of 800–1000 Burundi and Zambia. In relation to other water bodies

13 Elikana Kalumanga

Figure 2: Map of current elephant distribution and corridors in Tanzania. (Sources: Jones et al., 2009). globally, Lake Tanganyika contains almost 17% of Game Reserve between 1925 and 1927. Some of the world’s available surface freshwater, harbors the villages in MMRS (e.g. Maboha, Itebula, Mtego over 1500 species, out of which approximately 600 wa noti and Ilalanguru) are purely fishing villages are endemic species to Tanganyika, and supports one where most people work with catching, processing or of the largest fisheries on the African continent (LTA trading fish (Yanda et al., 2002; Yomba 2006; Salum Secretariat, 2012). Most of the rivers forming the Lake 2007). Similar to fishing, some villages in MMRS Tanganyika drainage basin on the Tanzanian side are (e.g. Itebula) are still known as places for traditional found within the MMRS (i.e. Moyowosi, Kigosi, beekeepers (Yanda et al., 2002). It is estimated that one Gombe, Malagarasi, Wala, Shama and Ugalla). household in Itebula village can have between 50-200 Even the Malagarasi River that is located primarily hives (Yanda et al., 2002). Even after the resettlement on Burundian land drains into the Lake Tanganyika from the Ugalla Game Reserve in 1954 necessary as through MMRS (Figure 3). Therefore, at the regional a consequence of infestation of sleeping sickness, level, MMRS plays an important role in sustaining the British colonial government legally permitted Lake Tanganyika and its associated ecosystem Wagalla people to conduct fishing and beekeeping goods and services supporting an estimated human in the reserve (Figure 4). After independence in the population of between 12.5 and 13.0 million (LTA year 1961, the Tanzanian government also retained Secretariat, 2012). the permission for fishing and beekeeping by the Wagalla in Ugalla Game Reserve. Today, about 1000 Livelihood activities people are permitted annually into the Ugalla Game Reserve for fishing and beekeeping. The fishing For ages, the Malagarasi-Muyovozi wetlands and and beekeeping in Ugalla Game Reserve and other their surrounding Miombo woodlands have been Miombo areas within MMRS is done at both for supporting the survival of people and wildlife (Figure subsistence and commercial profit (Figure 4). 4). Fisher (2002) gives a detailed account on how Outside Ugalla, livestock keeping has been the Wagalla people in MMRS have depended on increasing as a means of livelihood (John et al. 2012) fishing and beekeeping for their livelihood before as the wetlands and Miombo woodlands can support and after their resettlements in the current Ugalla much livestock with widespread pasture lands and

14 How elephants utilize a miombo-wetland ecoystem

Figure 3: The Map showing different protected areas in Malagarasi-Muyovozi Ramsar site (MMRS), the protected areas adjacent the MMRS, location of the villages in MMRS, and the sites already infested by water hyacinth in Ugalla River within the Ugalla Game Reserve, Western Tanzania. An inset map shows the location of MMRS in relation to Lake Tanganyika in Tanzania plenty of water. Due to the extreme and persistent Wildlife management droughts in other regions of Tanzania, people and their livestock have been migrating from the more- The wetlands and Miombo woodlands in the Ugalla drought devastated regions to the wetland areas of landscape (and the entire MMRS) are habitats for MMRS and other wetlands in Tanzania (Kangalawe a number of wildlife species. They also provide & Liwenga, 2005; Majule, 2007; Hamis et al. 2012). successful breeding sites for the two endangered bird This increases the strain placed on the region’s species: the stork (Balaeniceps rex) and ecosystem and resources. the Wattled Crane (Grus carunculatus) (John et al. 2012). The global population of both the and wattled cranes is small at less than 8000 birds

15 Elikana Kalumanga

A B

C D

E F

Figure 4: Photographs taken in Ugalla Game Reserve during field works. A)( A traditional beehives used by beekeepers (B). (C) Some fishermen with a local canoe setting their fishing nets in Ugalla River. (D) The author with fisher men in one of the fishing camps in Ugalla Game Reserve. (E) small hut used by Wagalla people (F) to perform their rituals at the beginning of the fishing season (July each year). Photos credit: Nyanda Lupondije, 2012-2013).

16 How elephants utilize a miombo-wetland ecoystem for each species (Birdlife International, 2014). In the local fishermen making it troublesome for both Tanzania, the numbers are in the low hundreds (John people and animals. et al., 2012). The wetlands in MMRS are also the main habitat for the near-threatened and wetland- specialist antelope (Tragelaphus spekei). Paper I These animals face population decreases globally Kalumanga, E., Nicholson, K.L., Mpanduji, D.G., (IUCN, 2008). MMRS also serves as habitat to a Cousins, S.A.O. Home ranges and utilization number of many other wild mammals beside African distributions of African elephants in Ugalla landscape, elephants (Loxodonta africana). For example, Western Tanzania. Manuscript to be submitted to the African Wild Dogs (Lycaon pictus), which along journal of Conservation Biology with the African elephant is considered to be one of the ‘vanishing species’ in East Africa (AWF, 2010), can be found within Ugalla and the MMRS. Due to Little is known about the basic ecology of most large the large number of big mammals (and crocodiles) and medium-sized mammals in Miombo ecosystems in the MMRS, these protected areas and the Ugalla in general and particularly in Western Tanzania (Caro, region are internationally known for wildlife hunting 1999; Waltert et al., 2008). To determine the size and tourism (e.g. Ugalla Game Reserve, Moyowosi- locations of elephants in Ugalla landscape, we used Kigosi Game Reserves, Luganzo and Gombe Game satellite Global Positioning System (GPS) receiver Controlled Areas as well as Uyumbu and Ipole collars (from African Wildlife Tracking, Pretoria, Wildlife Management Areas). This provides sources South Africa) fitted on six elephants (1 adult bull, 1 of employment and an influx of foreign currency to juvenile bull and 4 matriarchs). By using GPS collars, the region (Wilfred, 2012). we obtained location points four times a day for the six elephants for the duration between 16-29 months depending on the batteries life-span in the collars. We used the location points from each elephant to Methods analyse their movement pattern and determined the location and sizes of their home ranges in both wet To obtain a general view on how elephants are and dry seasons, and in relation to different protected distributed in the landscape a baseline survey areas in Ugalla landscape. was carried out in Ugalla by recording dung-pile densities. A scale was developed and implemented Paper II to identify high, medium and low dung density Elikana Kalumanga & Sara A. O. Cousins. areas. The initial idea was to use this dung density Woody plant abundance drives the browsing metric to estimate number of elephants within the patterns of elephants in the Ugalla landscape, of area (Appendix 1). Based on the initial dung density Tanzania Manuscript. surveys, observational transects were identified and set up to estimate how elephants utilized resources Elephants are widely known to be generalist feeders, in some parts of the region. These transects but they are still selective when they forage by allowed, for example, mapping of the types and selecting either different plant parts or types of plants abundance of woody plant species in different parts to feed upon (Jachmann & Bell, 1985; Guy, 1989; of Ugalla, identification and recording of elephant Jachmann, 1989; Sukumar, 1990; Osborn, 2002; browsing signs, and mapping of the distribution of Holdo, 2003; Kerley & Landman, 2006). So far, little termite mounds eaten by elephants. The aim was to is known about the specific factors that influence understand elephant browsing patterns and determine elephant forage preferences and their selections of why some woody plants are selected for browsing and specific plant species (Guy, 1989; Holdo, 2003; Staub some are not (paper 2), as well as to assess why soils et al., 2013). To be able to understand the types of from some termite mounds are eaten compared to woody plants browsed by elephants and try to link soil from other sites (paper 3). The results from these browsing to movement patterns, we mapped the two studies would be able to inform key habitat areas woody vegetation in 1860 plots along 62 transects in utilized by the elephants in Ugalla landscape. Besides the Ugalla region of Tanzania in order to assess the assessing areas that are important for elephant diet, types and abundance of woody plant species, their six elephants were collared with Global Positioning height, signs of elephant browsing and leaf nutrient System (GPS) collars to analyse the location and content. Leaf samples were collected for nutrient size of the elephant home ranges in Ugalla landscape analysis from 50 woody tree species comprised of (paper 1). Finally, to estimate the potential for current 14 highly elephant-browsed tree species, 18 less and future limitations to elephant-accessible water, elephant-browsed trees and 18 non-preferred tree the extent of water hyacinth infestation and spread species. The grouping of tree species was mainly in Ugalla River within the Ugalla Game Reserve was based on the abundance-browsing percentages with mapped (paper 4). Water hyacinth infestation makes highly browsed tree species (browsing ratio >10%), some areas of the Ugalla River unsuitable not only for less browsed tree species (browsing ratio between 2 much of the wildlife during the dry season (especially to 10%), and the non-preferred tree species (browsing the water dependant wildlife) but also for fishing by ratio < 2%) defined explicitly. 17 Elikana Kalumanga Paper III covering the entire water surface with impenetrable Elikana Kalumanga, Donald G. Mpanduji, and Sara mats (Figure 5). During the flight, the start and stop A. O. Cousins. Geophagic termite mounds as one of location of each block (section of river classified the resources for African elephants in Ugalla Game under the three-point scale) were recorded to Reserve, Western Tanzania. Manuscript resubmitted allow the mapping of the entire river into the three after revision to the African Journal of Ecology. different classes. These locations, defining transitions between hyacinth coverage classes were brought During the baseline survey it was noticed that into a geographical information system and used elephants eat soil from some specific termite mounds to delineate the length of the river covered by each in Ugalla Game Reserve. Soil from termite mounds block. This allowed for a quantification of the extent can provide an important part of the elephant diet and of water hyacinth infestation indicative of decreased it might be crucial to protect these termite mounds access to surface water in the region. from some anthropogenic activities that may destroy them (such as the construction of roads or air strips). While walking along transects in the Ugalla region all the termite mounds encountered were recorded Results and Discussion and categorized as to if they had signs of soil eating by elephants or not. To determine why soils from Location and size of elephant home ranges in some termite mounds were eaten by the elephants, Ugalla landscape (Paper I) we sampled soils from 10 termite mounds with signs of geophagy (soil eating) and from seven termite The analysis of the locations obtained from the GPS mounds without signs of soil eating. These samples collars showed clearly that the location and size of were analysed for nutrient/elemental composition. To the home ranges of individual elephants collared in have a wider comparison of the soils nutrient status Ugalla landscape differed between the individual we also sampled soil from 13 sites in the flood plain elephants and between wet and dry seasons. For at least 30 m away from termite mounds utilized by instance, among the four matriarchs, the locations the elephants. A total of 30 soil samples (250 g each), of the home ranges for the two matriarchs (Johari which were air-dried for a few days in the field and then and Ugalla) were situated mostly in Ugalla Game transported for analysis at the Agri-Lab in Uppsala, Reserve. They had occasionally visited the North Sweden. Clay and silt contents was determined Ugalla Forest Reserve and Msima Game Controlled using the sieving and sedimentation method (SS-ISO Area. On the other hand, the matriarch named 11277:2009, IDT-Swedish Standard) and different Francisca utilized the eastern side of the Ugalla mineral element concentrations were analysed using landscape (i.e., Uyumbu Wildlife Management Area the AL-Method as described by Balsberg-Påhlsson and eastern part of the Ugalla Game Reserve) while (1990). the elephant named Sara utilized the western part of the Ugalla landscape (Ugalla Game Reserve and the Msima Game Controlled Area). Within two years, Paper IV the adult bull named Kalumanga utilized almost the Elikana Kalumanga. Why Integrated Natural entire Ugalla landscape, whereas the location of the Resource Management in Malagarasi-Muyovozi home range for the juvenile bull Ruben was more Ramsar Site, Tanzania? Insights from Ugalla Game or less in the same places as the matriarch named Reserve. Manuscript resubmitted after revision to the Ugalla. During the dry seasons most of the collared Marine and Freshwater Research: Ramsar Special elephants were concentrated along the Ugalla River Issue. within the Ugalla Game Reserve. The Ugalla River is the only source of water during the dry seasons The Ugalla River, which is the main source of water in Ugalla landscape (especially between June to supporting wildlife and livelihood activities such November). During the wet seasons, elephant were as fishing, is becoming infested by water hyacinth able to move away from the Ugalla River into the (Eichhornia crassipes). Water hyacinths originate Miombo woodlands. This was because water is from the Amazon in South America and were brought readily available at many places in the landscape (e.g. to Africa as an ornamental species by the gardeners seasonal swamps) during the wet season. in late 1890s (El-Morsy, 2004). Their appearance This is the first study to track elephants in Western in Tanzania was noted for the first time in 1950s Tanzania using GPS collars. The results show that (). To understand the extent of the elephants are concentrated to the Ugalla Game water hyacinth infestation and spread in the Ugalla Reserve. This is probably not only because of the River, an aerial survey was conducted in August 2012 water availability in Ugalla River during the dry using a microlight aircraft along 87.6 kilometers of seasons but also because it is a relatively secure the Ugalla River in Ugalla Game Reserve. The water habitat compared to other adjacent protected areas. hyacinth infestation was recorded in different river There are regular anti-poaching patrols in the Ugalla ‘blocks’ using a three–point scale: i) almost absent Game Reserve compared to the adjacent protected ii) partially present, but not completely covering areas. Anti-poaching patrols are only rarely carried the water surface and iii) completely present and out in the forest reserves or any significant parts of the 18 How elephants utilize a miombo-wetland ecoystem

Msima Game Controlled Areas. This demonstrates situated in Miombo woodlands. The results from soil that well-managed wildlife management areas analysis show that, soils eaten by the elephants had (WMAs) in Tanzania can help to protect habitats for higher concentration of mineral elements compared wide-ranging species like elephants. These WMAs to the soils sampled in the surrounding flood plain or need to span multiple management/protected regions. termite mounds without signs of geophagy. Almost For example, as aforementioned, the home range of all the minerals we analysed, which included Mn, Fe, the matriarch named Francisca was situated between Cu, Zn, Na, Mg, P, S, K, Ca and Cr, were higher in the Ugalla Game Reserve and the Uyumbu Wildlife soils eaten by the elephants as compared to soils not Management indicating utilization of both regions. utilized by the elephants. Therefore, to rescue the remaining elephants (or Soil eating behaviour in both animals and human better secure their populations) in Ugalla landscape, beings (i.e. geophagy) has been reported since 1398, regular anti-poaching activities should be extended but a conclusive biological explanation is still lacking beyond Ugalla Game Reserve into the other adjacent (Mahaney et al., 1999; Ketch et al., 2001; Dudley et protected areas. al., 2012). There are four main hypotheses of why animals eat soils. From zoo-pharmacognosy (i.e. How woody plant abundance drives the study of self-medication in animals), geophagy the browsing patterns of elephants in is hypothesised to be therapeutic. The therapeutic the Ugalla landscape? (Paper 2) hypothesis is supported by the premise that geophagic soils used by elephants and other mammals contain A total of 129 different woody plant species a high clay content known to alleviate gastro- belonging to 35 families were recorded along the intestinal distress and upsets. Another hypothesis 62 transects. The Fabaceae family dominated the is that geophagy is a food additive behaviour woody plant community with 31 species, followed because typical herbivore diet (tree and grasses) by the Combrataceae (12 species) and the Rubiaceae has inadequate mineral concentrations (especially (12 species). Only half of the species had signs Na+ concentration) to meet the dietary demand of browsing by elephants. Two abundant woody of almost all the herbivores. Geophagic soils are plant species were found to be highly browsed in also known to detoxify high concentration of plant Ugalla landscape (i.e. Antidesma vernosum and secondary compounds (especially alkaloids) found Hymenocardia acida) that are not known to be in tropical forest trees, thus countering the effects highly browsed from other elephant browsing studies of lactic acidosis (Houston et al., 2001; Klaus et al, across the countries covered by Miombo woodlands 1998). Another hypothesis on geophagy is related to (Jachmann & Bell, 1985; Guy, 1989; Jachmann, 1989; buffering of gastric pH (Dudley et al., 2012; Mills et Osborn, 2002; Holdo, 2003; Osborn, 2004; Kerley & al., 2009). Landman, 2006). From the mineral and fiber analysis The results in our study are not conclusive but of woody plant species in Ugalla landscape it can be point towards a theory that elephants eat soil in seen that woody plant selection for browsing by the the Ugalla landscape because of the high mineral elephants is partly driven by the plant abundance and contents in the soils from the geophagic termite availability (which is measured by the height of the mounds. Plants browsed by elephants are short of trees). The leaf contents did not seem to be a major minerals to meet elephant dietary demand; thus, they factor driving browsing patterns of the elephants require the termite mounds to obtain different types of in Ugalla. Most of the highly-browsed plants were minerals missed in plants. Identifying such linkages abundant as compared to the lesser browsed trees. between the physical environment and species is This indicates that although elephants are widely central to successful management. In Tanzania, known to be generalist feeders, they selectively wildlife management is still passive; i.e. ‘Mother forage from types of plants as well as the parts of the Nature taking its own course’. There is no guaranteed plant they feed upon. In some woody plant species, provisioning of key resources such as water and elephants utilize only leaves, fruits, roots, and barks. artificial salt sources to different wildlife species. Thus, woody plant abundance could be a major factor Therefore, sites where animals are able to obtain their driving the browsing patterns of elephants in Miombo diet naturally (e.g. the geophagic termite mounds in woodlands of Ugalla landscape. Ugalla) should be protected from any destruction, for instance during the development of infrastructure Why elephants eat soils from some termite such as roads, airstrips, and camps. Inclusion of such mounds in Ugalla Game Reserve? (Paper 3) resources in the definition of habitat areas could lead to a more robust and richer conservation strategy. In total, we recorded 535 termite mounds along 62 transects in Ugalla Game Reserve; 465 termite Why integrated Natural Resource mounds perpendicular to the rivers (87%) and 70 Management in MMRS? (Paper 4) termite mounds (13%) in Miombo woodlands. Out of 465 termite mounds that we recorded perpendicular Out of the 87.6 kilometers of the Ugalla River in Ugalla to the rivers, 18 termite mounds had soil eating Game Reserve, 13 kilometers (15%) were completely signs by the elephants. No sign of soil eating by covered by water hyacinths, 17 kilometers (20%) the elephants was recorded in the termite mounds partially covered and 57 kilometers (65%) had no or

19 Elikana Kalumanga A B

C D

Figure 5: Photos showing different blocks covered by water hyacinth in Ugalla River within Ugalla Game Reserve. The first two photos (A) and (B) shows the water hyacinth in different river blocks with aquatic vegetation (A) and rocks (B). (C) Shows four hippopotamus with restricted fully utilization of some parts of the Ugalla River infested with the water hyacinth , and (D), shows a segment of the Ugalla where a fishing camp was closed due to water hyacinth infestation. Photos credit: Sara Cousins, 2014 A( ) and Elikana Kalumanga, 2014 (B, C, D). very few water hyacinths. Almost all the river blocks and animals. Equally important also is a reminder that where impenetrable mats of water hyacinth covered the Ugalla River in Ugalla Game Reserve (and also the entire water surface were found in places where large parts of the Ugalla landscape) constitutes parts the Ugalla River either meanders (short distances) of the Malagarasi-Muyovozi Ramsar Site (MMRS; or where there were other aquatic vegetation and/ Figure 3). When the Parliament of Tanzania ratified or rocks (Figure 5 A & B). Fundamentally, these the Ramsar Convention in February 1999 as a Ramsar are the regions of the river where the flow of the Contracting Party, or a Member State, Tanzania water is slow. Water hyacinth was also recorded in committed herself to implementing the Ramsar two oxbow lakes that are seasonally flooded and Convention by, among other things: i) designating adjacent to the Ugalla River channel. Where there suitable wetlands for the List of Wetlands of are thick mats of water hyacinth fishing cannot be International Importance (“Ramsar List”); ii) ensuring carried out and such sites also cannot be fully utilized effective management of designated Ramsar Sites and by wildlife (for example elephants) for drinking iii) working towards the wise use of all their wetlands water. This is especially critical for water-habitat through national land-use planning, appropriate dependent mammals such as hippopotamus (Figure policies and legislation, management actions, and 5 C). As a result, both fishermen and wildlife (e.g. public education (Ramsar Convention on Wetlands hippopotamus and crocodiles), are restricted to share 1971). At the heart of the Ramsar Convention is the some blocks of the Ugalla River, which are not full ‘Ramsar Wise Use Principle’, which is defined as infested with water hyacinth resulting into human- the maintenance of a wetland’s ecological character. wildlife conflicts and sometimes with deadly results. This should be achieved through the implementation The take home message from this survey is that the of ecosystem management approaches within the more parts of the Ugalla Riveris covered by water context of sustainable development. The ecological hyacinth, the less parts that are accessible for people character, as defined in the Ramsar Convention,

20 How elephants utilize a miombo-wetland ecoystem is the combination of the ecosystem components, crocodiles) depends on water in the rivers (e.g. Walla processes and benefits/services that characterise the and Ugalla Rivers). Across the globe, protection of wetland at a given point in time (Ramsar Convention numerous forest resources, hydrological catchments, on Wetlands1971). In other words, the infestation and and various ecosystems have benefited from the fauna spread of water hyacinth in Ugalla River is already residing in them. Primarily, this is because animals affecting the ecological character of the MMRS. The draw more conservation attention and sympathy thick mats of the water hyacinth are affecting fishing at local and international levels than trees. Such activities done by the local communities in MMRS benefits can clearly flow both ways since the linkages (Figure 5 D). Furthermore, the MMRS constitute among resources and their inherent connections to 30% of the Lake Tanganyika catchments (URT animals and people makes it such that impacts on 2007; Kashaigili & Majaliwa 2010). Spread of the one component can directly and/or indirectly impact water hyacinth beyond the Ugalla River in Ugalla the other components across the landscape mosaic.. Game Reserve might end up in the trans-boundary Therefore, the main conclusions from this thesis are: Lake Tanganyika, thus affecting more communities • To conserve the biodiversity in protected areas and countries. In the guidelines for management in Tanzania requires management systems that planning for Ramsar Sites and other wetlands, it is are able to adjust and integrate all the resources insisted that ‘legitimate stakeholders, particularly found in the area (e.g. water, forests and animals) local communities and indigenous people, should across various spatial and temporal scales. be strongly encouraged to take an active role in planning and in the joint management of the sites’ • Elephants utilize different parts of the Ugalla (Ramsar Convention on Wetlands 1971). landscape during the wet seasons and have Thus, the infestation and spread of water hyacinth restricted movement patterns during the dry in Ugalla River requires urgent attention. The seasons where they remain closer to the Ugalla eradication of water hyacinths is necessary because River. Ugalla River, apart from being used by elephants • Elephants cannot be considered true generalist and different wildlife species during the dry seasons feeders, because they are selective for the forages and as part of their habitats, is a societal asset. The to feed upon. fishing done by the local people depends onthe health of the Ugalla River. Further, the spread of • Soils from geophagic termite mounds are one the water hyacinth further down the Ugalla River of the important resources eaten by elephants might also affect not only fishing activities in the (among other mammals). Geophagic termite shallow lakes in MMRS (figure 3) but also the trans- mounds, thus, should be protected to avoid boundary Lake Tanganyika in general. To achieve damage from the development of infrastructure the eradication of the water hyacinth and reduce this such as roads. threat requires an integrated management approach • There is a need for coordinated anti-poaching not possible through sector-based management plans. patrols throughout the Ugalla landscape that This is because it requires engagement of different goes beyond the Ugalla Game Reserve to other stakeholders and organizations at both the catchment adjacent protected areas since these patrols and basin scales. influence elephant movement. • As recommended in the Ramsar Convention and under the supervision of the local managers, local Conclusions people should be engaged in addressing some of the problems occurring in the Ramsar site such controlling the spread of the water hyacinth in Resources managed in different protected areas Ugalla River. within the Ugalla landscape are interrelated. The water (in different rivers), the forests (in different • Wildlife management areas, if appropriately forest reserves), and the wild animals (in various managed and selected, can secure habitats for the game reserves, wildlife management areas and game wide-ranging species such as elephants. controlled areas) are intimately linked to one another. This last point is central as it gets at the heart For instance, the survival of all wild animals in Ugalla, of the scale mismatch problem prevalent within among other factors, depends on the Ugalla River conservation management. By understanding (especially during the dry season). Further, people also the linkages between the distribution of physical depend on and interact with these various linkages resources and the movement/distribution of wildlife, and resources. For example, fishing and trophy it may be possible to manage landscapes in a way hunting in Ugalla landscape are also done during the to both secure biodiversity and support livelihood dry season. The location of all the fishing and hunting development. camps (and even the hunting of hippopotamus and

21 Elikana Kalumanga Karin Holmgren (the program coordinator): asante Acknowledgement sana. A special thanks to my family and clan. Words This is one of the hardest sections to write in my cannot express how grateful I am for all of the thesis. There is no space to list the names of the sacrifices that you’ve made on my behalf. Many many people both in Sweden and Tanzania that have things have happened which needed my personal extended help and support to me in pursuit of my attention, but you stood in my place. A good example PhD. I will try to list a few names and some groups, is when our father passed away at my place in Dar es but to all of you, please accept my thanks and Salaam and I was in Sweden pursuing this PhD. You appreciations expressed in Kiswahili; Asanteni sana! demonstrated how a family and a clan should always I would like to give thanks to the Almighty God for stand. I really owe you much. I have nothing to pay keeping me alive up to this point. This journey wasn’t you back, but asanteni sana! It is your prayers for me easy for me. I place on record my sincere thank you that sustained me thus far. Thank you to my wife, to Prof. Sara Cousins who was my main advisor for Ruth, because you have suffered the consequences of sharing expertise, for sincere and valuable guidance my choice to pursue my PhD abroad. It was hard for and for encouragement throughout my PhD studies. you to stay alone with our sons Ngasa and Ngusa. Your repeated comments on my manuscripts of ‘you But you managed through this hard way. Thank you are getting there’ and support in the field gave me for understanding and hope time will heal your pains. the energy and power to keep on going. Asante sana! Many thanks to you, Mama, for the support you have To Prof. Gregory Donald Mpanduji, Dr. Steve Lyon been always giving us since our childhood until we (my co-advisor) and Dr. Jan Plue, your assistance were able to stand on our own. To Mr. and Mrs. Joel and comments greatly improved my manuscripts Ngusa, thank you for always standing with me. I and kept me going. Thank you so much. I really would also like to thank all of my friends in Tanzania appreciate! Since I arrived in Sweden, I have and Sweden who supported me and gave me the received moral and academic support from different incentive to strive towards my goal. Asanteni sana. people at the Department of Physical Geography. You are all wonderful people; keep on that good spirit! Members in the former Landscape Ecology Group (LEG) and current Biogeography and Biomatics Financial support Group have been of great help to me wherever I got stuck academically. Asanteni sana! Also, thank you, This PhD project has been supported by the Swedish Helle for your help in the last minutes of compiling International Development Cooperation Agency my thesis. (SIDA) through the Integrated Natural Resources Davis and Catarina Dirisha: I owe you much. Management Program executed between the Your moral and material support during all the times University of Dar es Salaam (Institute of Resource I spent in Sweden is highly appreciated. I have really Assessment) and Stockholm University (Department enjoyed your company and staying at your place of Physical Geography). every time you escape the winter by going to Dar es Salaam. To Simon Mwansasu, Rene Mbanguka and all members of the Association of Tanzanian Students in Sweden, thank you for a good company References and I know our journey is still continuing. Asanteni sana! African Wildlife Foundation- AWF (2010). African Wild Dogs. Back to Tanzania, I really appreciate the support I http://www.awf.org/content/wildlife/detail/africanwilddog downloaded on 21th April 2014 received from the Institute of Resource Assessment, Amboseli Trust for elephants-ATE (2010). Elephants and CITES: University of Dar es Salaam and from the Ministry the Facts, http://www.elephanttrust.org/node/618 downloaded of Natural Resources and Tourism through the on 26 September 2012 Andrade,G. S. M., & Rhodes, J. R. (2012). Protected areas Tanzania Wildlife Research Institute and the Wildlife and local communities: an inevitable partnership toward Division. My field work demanded more funds than successful conservation strategies? Ecology and what could be provided by the main SIDA program at Society 17(4): 14. http://dx.doi.org/10.5751/ES-05216-170414. the University of Dar es Salaam. Many thanks to the Babweteera, F., Savill, P. & Brown, N. (2007) Balanites Project Manager of the Ugalla Game Reserve, Mr. wilsoniana: Regeneration with and without elephants. Japhary Lyimo and the staff of the Tanzania Game Biological conservation 134(1), 40-47. Balsberg-Påhlsson, A. M. (1990). Handledning i kemiska Trackers Safaris and the Friedkin Conservation Fund, metoder vid växtekologiska arbeten. Meddelanden från whose material support greatly supplemented the Växtekologiska avdelningen. Nr. 52. (In Swedish.) funds obtained from the University of Dar es Salaam. Lunds Universitet, Lund, Sweden. Barua, M., Tamuly, J. & Ahmed, R.A. (2010). Mutiny or Clear My appreciation also goes to the rangers in Ugalla Sailing? Examining the Role of the Asian Elephant as a Game Reserve who supported me during transect Flagship Species. Human Dimensions of Wildlife 15 (2), 145- walks and elephant collaring. I also acknowledge 160. Beachey, R. (1967). The East African ivory trade in the nineteenth the financial support I received from our program at century. Journal of African History 8: 269–90. the Department of Physical Geography to facilitate BirdLife International (2014). ‘Important Bird Areas factsheet: the removal of elephant collars in Ugalla. To Prof. Moyowosi - Kigosi Game Reserves’. Available at http://www.

22 How elephants utilize a miombo-wetland ecoystem

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stages shown in Table 3. In the present study, only Appendix 1: Estimating ele- 15 dungs survived beyond 24 hours. Other dungs phant dung decay rate: the ex- were destroyed by dung beetles immediately after they were dropped on ground. All the surviving perience from Ugalla landscape dungs had more seeds and palm roughages (Figure 6). Thus the use of dung densities to estimate Initially, one of the specific objectives in this PhD number of elephants within a region proved to be study was to estimate the number of elephants in futile. Major factors known to affect dung decay rate Ugalla landscape by; i) recording dung piles per include rainfall (White, 1995; Barnes et al., 1997; km2, ii) determing the defaecation rate of elephants; Olivieret et al., 2009), the amount of fruits in the iii) determining the mean dung decay rate; and iv) diet, type of habitats, the microclimate where the combining the three estimates to get an estimate dung is deposited, the abundance and diversity of of numbers/density of elephants per km2 (Barnes, decomposers (particularly fungi and dung bettles) 1996; Barnes et al., 1997; Demeke & Bekele, 2000; (Breuer & Hockemba, 2007), foraging activity Mubalama, 2000; Mubalama & Sikubwabo, 2002; of small mammals and birds, sun exposure and Vanleeuwe & Probert, 2014). minimum temperatures in the days after deposition, In October 2011 to November 2011, 150 fresh (Mubalama and Sikubwabo, 2002). Similar elephant dungs were identified and monitored to studies should be aware of the field challenges and establish the dung decay rate (as a parameter in the understand the various factors known to affect dung model to refer dung density to elephant numbers). decay rates in different sites and different seasons. Dung decay rate is the time taken for the dung or dung piles to gradually decay into the different

A B

C D

Figure 6: Some elephant dung monitored to estimate the dung decay rate in Ugalla landscape, Western Tanzania. Survived dungs had more seeds (A) and palm roughages and corns (B, C, D). Photo credit: Elikana Kalumanga (2011-2012).

26 How elephants utilize a miombo-wetland ecoystem

Table 3: Dung-piles classification according to their shape (Adopted from Barnes & Jensen, 1987)

Dung category Description A Boli intact, very fresh, moist, with odour B Boli intact, fresh but dry, no odour C Some of the boli have disintegrated; others are still recognizable as boli D All boli completely disintegrated; dung-pile now forms and amorphous flat mass E Decayed to the stage where it would be unlikely to be detected at a range of two metres in the undergrowth

Mubalama, L. (2000). Population and Distribution of Elephants References: Appendix 1 (Loxodonta africana africana) in the Central Sector of the Virunga National Park, Eastern DRC. Pachyderm 28, 44-55. Mubalama, L. & Sikubwabo, C. (2002). Rate of decay of Barnes, R.F. W. (1996). Estimating forest elephant abundance elephant dung in the central sector of Parc National des by dung count. In ‘Studying elephant’. (Ed K. Kangwana.) Virunga, Democratic Republic of Congo. Pachyderm pp. 38-48. (African Wildlife Foundation, Nairobi, Kenya.) 33:43–49. Barnes, R.F.W. & Jensen, K.L. (1987). How to count elephants Olivier, P.I, Ferreira, S.M. & van Aarde, R.J. (2009). Dung in forests. IUCN, African Elephant and Rhino Specialist survey bias and elephant population estimates in southern Group. Technical Bulletin 1, 1-6. Mozambique. African Journal of Ecology 47:202 Barnes, R.F.W., Asamoah-Boateng, B., Naada Majam, J. & 2013. Agyei-Ohemeng, J. (1997). Rainfall and the population Vanleeuwe, H. & Probert, J. (2014). Decay rate of elephant dynamics of elephant dung piles in the forests of dung in Coukouati-Douli National Park, Republic of Congo. southern Ghana. African Journal of Ecology 35:39–52. Pachyderm 55, 89-91. Breuer, T. & Hockemba, M.N. (2007). Forest elephant dung White, L.J.T. (1995). Factors affecting the duration of elephant decay in Ndoki Forest, northern Congo. Pachyderm 43:43– dung piles in rain forest in the Lope Reserve, Gabon. African 51. Journal of Ecology 33, 142–150. Demeke, Y. & Bekele, A. (2000). Study on the Elephant of Mago National Park, Ethiopia. Pachyderm 28, 32-43.

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