Tsetse Invasion As an Emerging Threat to Socioecological Resilience of Pastoral Communities in Karamoja, Uganda

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Article Tsetse Invasion as an Emerging Threat to Socioecological Resilience of Pastoral Communities in Karamoja, Uganda

Anthony Egeru 1,2,* , Joseph Opio 2, Aggrey Siya 3, Bernard Barasa 2 , John Paul Magaya 4 and Justine J. Namaalwa 2

1 Training and Community Development, Regional Universities Forum for Capacity Building in Agriculture (RUFORUM), Wandegeya-Kampala P.O. Box 16811, Uganda 2 Department of Environmental Management, College of Agricultural and Environmental Sciences, Makerere University, Kampala-Uganda P.O. Box 7062, Uganda; [email protected] (J.O.); [email protected] (B.B.); [email protected] (J.J.N.) 3 Department of Biosecurity, Ecosystems and Veterinary Public Health, College of Veterinary Medicine, Animal Resources and Biosecurity, Makerere University, Kampala-Uganda P.O. Box 7062, Uganda; [email protected] 4 Department of Geomatics and Land Management, College of Engineering Design Art and Technology, Makerere University, Kampala-Uganda P.O. Box 7062, Uganda; [email protected] * Correspondence: [email protected]

Received: 16 January 2020; Accepted: 15 February 2020; Published: 20 February 2020

Abstract: Over 70% of Uganda is infested by the tsetse fly, which has negative effects on human and livestock health. From colonial to post-independent Uganda, the Government of Uganda has worked to eradicate the tsetse menace. Despite these efforts, recent veterinary reports from the Karamoja sub-region have indicated widespread tsetse invasion. This study investigated the potential impact of tsetse invasion on the socioecological resilience of pastoral communities in the Karamoja sub-region. Results indicated that tsetse invasion is spreading from north to south of Karamoja. The tsetse transmission route emerging from southern Karamoja is perceived to be a continuation of the tsetse belt from West Pokot, Kenya. Cases of livestock deaths, livestock abortions, decreased milk yields, restricted access to prime grazing lands, heightened human-wildlife conflicts and disruption on crop cultivation have been reported. A computed socioecological resilience index in the study area was positive but low. Owing to the transboundary characteristics of tsetse invasions and sources and the associated documented effects, an urgent, strategic and system-wide intervention should be undertaken to control the tsetse invasion in this sub-region.

Keywords: Glossina spp.; health; sleeping sickness; tsetse invasion; Karamoja sub-region; Africa

1. Introduction Trypanosomiasis transmitted by tsetse flies (genus Glossina) continues to constitute a major factor limiting rural development in vast areas of tropical Africa and affects 37 countries in sub-Saharan Africa (Figure1). More than 10 million square kilometres of Africa, especially sub-Saharan Africa, are infested by tsetse [1]. Three types of tsetse flies are found in Africa: Palpalis in riverine habitats and Fusca in forests and isolated patches. In contrast, Morsitans is mainly found in savannahs dominated by woodlands and scattered thickets [2]. These haematophagous vectors are responsible for transmitting animal African trypanosomiasis (AAT) and human African trypanosomiasis (HAT), also known as livestock nagana and sleeping sickness, respectively [3].

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(Figure 2). The flagellate protozoa infect a wide range of wild and domestic animals as well as humans in sub-Saharan Africa [5]. Infected livestock and humans present diverse clinical manifestations. For example, persons infected with sleeping sickness may experience multi-organ dysfunction, high-grade fever, dizziness and constipation [6,7] . In the case of one Polish tourist who experienced a tsetse fly bite in Queen Elizabeth National Park in Uganda, a high-grade fever was observed. The fever was preceded by chills, bleeding from the gums and oral mucosa. Haemorrhage at the sites of venepuncture, numerous ecchymoses, fine-spotted skin rashes and tachycardia were observed. In addition, hepatosplenomegaly, dehydration, jaundice, dyspnoea, hypoxaemia, Sustainability 2019, 11, x FOR PEER REVIEW 2 of 25 generalised oedema and oliguria have been observed [6]. Additionally, livestock anaemia, (Figure 2). The flagellate protozoa infect a wide range of wild and domestic animals as well as lacrimation, Sustainabilityenlarged2020 lymph, 12, 1599 nodes, abortions and decreased fertility are signs 2associated of 26 with humans in sub-Saharan Africa [5]. Infected livestock and humans present diverse clinical infected livestock.manifestations. In other For example, cases, infectedpersons infected livestock with sleeping exhibited sickness a loss may of experience appetite, multi-organ poor body condition and productivitydysfunction,Both and HAThigh-grade early and mortality AAT fever, are dizziness caused [8,9]. by and flagellate constipation protozoa [6,7] transmitted. In the case byof one the tsetsePolish [tourist4]. The who tsetse experiencedprovides ana tsetse ambient fly environmentbite in Queen whereElizabeth the National parasites Park complete in Uganda, their unique a high-grade breeding fever life was cycle observed.(Figure2 The). The fever flagellate was preceded protozoa by infect chills, a wide bleedi rangeng from of wild the andgums domestic and oral animals mucosa. as Haemorrhage well as humans at inthe sub-Saharan sites of venepuncture, Africa [5]. Infectednumerous livestock ecchymoses, and humans fine-spotted present sk diversein rashes clinical and tachycardia manifestations. were For observed.example, In persons addition, infected hepatosplenomegaly, with sleeping sickness dehydration, may experience jaundice, multi-organ dyspnoea, dysfunction, hypoxaemia, high-grade generalisedfever, dizziness oedema and and constipation oliguria [have6,7]. Inbeen the caseobserved of one [6]. Polish Additionally, tourist who livestock experienced anaemia, a tsetse lacrimation,fly bite in Queenenlarged Elizabeth lymph Nationalnodes, abortions Park in Uganda,and decreased a high-grade fertility fever are wassigns observed. associated The with fever infectedwas preceded livestock. by In chills,other cases, bleeding infected from livestock the gums exhibited and oral a loss mucosa. of appetite, Haemorrhage poor body at condition the sites of andvenepuncture, productivity and numerous early mortality ecchymoses, [8,9]. fine-spotted skin rashes and tachycardia were observed. In addition, hepatosplenomegaly, dehydration, jaundice, dyspnoea, hypoxaemia, generalised oedema and oliguria have been observed [6]. Additionally, livestock anaemia, lacrimation, enlarged lymph nodes, abortions and decreased fertility are signs associated with infected livestock. In other cases, infected livestock exhibited a loss of appetite, poor body condition and productivity and early mortality [8,9].

Figure 1. Predicted distribution of tsetse fly in Africa, 2000 (Source: [10]).

Figure 1. Predicted distribution of tsetse ﬂy in Africa, 2000 (Source: [10]). Figure 1. Predicted distribution of tsetse fly in Africa, 2000 (Source: [10]).

Figure 2. Life cycle of trypanosomes [4].

Sustainability 2019, 11, x; doi: FOR PEER REVIEW www.mdpi.com/journal/sustainability Figure 2. Life cycle of trypanosomes [4].

Figure 2. Life cycle of trypanosomes [4].

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TheThe tsetsetsetse phenomenonphenomenon presentspresents aa highhigh diseasedisease burdenburden onon thethe humanhuman population.population. ThisThis highhigh incidenceincidence oftenoften leads leads to decreasedto decreased agricultural agricultural productivity productivity and livestock and livest deathsock [deaths5]. Sleeping [5]. Sleeping sickness issickness an important is an important public health public disease health in disease sub-Saharan in sub-Saharan Africa. On Africa. the basis On ofthe the basis HAT of relatedthe HAT mortality, related itmortality, has been it ranked has been ninth ranked out of ninth 25 human out of infectious 25 human parasitic infectious diseases parasitic in Africadiseases [4]. in Meanwhile, Africa [4]. livestockMeanwhile, nagana livestock remains nagana a major remains obstacle a major to livestock obstacle development to livestock in development several sub-Saharan in several African sub- countries.Saharan African It causes countries. the death It of causes an estimated the death three of million an estimated cattle annually. three million Furthermore, cattle annually. losses of approximatelyFurthermore, losses 26% inof dairyapproximately yield and 26% 50% in of dairy cattle yield herds and in high50% of potential cattle herds agricultural in high areaspotential are attributedagricultural to areas tsetse are infestation. attributed to These tsetse losses infestatio leadn. to These an estimated losses lead economic to an estimated loss of US$4.5economic billion loss annuallyof US$4.5 in billion Africa annually [11,12]. Inin theAfrica case [11,12]. of Uganda, In the approximately case of Uganda, 90% approximately of sleeping sickness 90% of casessleeping are causedsickness by cases tsetse areGlossina caused fuscipes by tsetse fuscipes Glossina[3] andfuscipesGlossina fuscipes f. fuscipes [3] andand GlossinaGlossina f. fuscipes palidipes andare Glossina known topalidipes occur inare most known of Ugandato occur (Figurein most3 of). Uganda The infection (Figure of 3). livestock The infection in cattle-rearing of livestock communities in cattle-rearing by naganacommunities has led by to nagana a livestock has led crisis to a arisinglivestock from crisis livestock arising from deaths livestock and sale deat restrictions.hs and sale This restrictions. pattern hasThis exacerbated pattern has foodexacerbated and nutrition food and insecurity nutrition [13 in,security14]. Trypanosomiasis [13,14]. Trypanosomiasis prevalence prevalence and resistance and areresistance predicted are to predicted become moreto become pronounced more pronounced and persistent and in persistent many African in many tsetse African belts, signifying tsetse belts, an aggravationsignifying an of aggravation the phenomenon of the [ 15phenomenon]. These patterns [15]. areThese attributed patterns to are changes attributed in land to usechanges and cover in land as welluse and as climate cover as variability well as climate and change variability [16,17 and]. Evidence change from[16,17]. within Evidence Uganda from show within treatment Uganda failure show ratestreatment are as failure high as rates 30% are in northernas high as Uganda 30% in [northern18]. Uganda [18].

FigureFigure 3.3. Historical distribution ofof thethe three majormajor tsetsetsetse speciesspecies inin Uganda,Uganda, 19821982 [[10].10]. MapsMaps havehave beenbeen recreatedrecreated byby thethe authorsauthors ofof thisthis article.article.

OverOver 70%70% (140,000(140,000 kmkm22)) of Uganda’s land area hashas beenbeen invadedinvaded byby tsetsetsetse flies,flies, butbut withwith varyingvarying levelslevels ofof prevalence prevalence [19 [19].]. Uganda Uganda has has approximately approximat elevenely eleven tsetse tsetse fly subspecies. fly subspecies. However, However, two species, two Glossinaspecies, f.Glossina fuscipes f.and fuscipesGlossina and palidipes, Glossinahave palidipes, received have significant received verification significant and verification documentation and (Figuredocumentation3). G. f.(Figure fuscipes 3). isG. thef. fuscipes most abundantis the most and abundant widespread and widespread tsetse species tsetse and species is found and inis approximatelyfound in approximately 70% of the 70% country’s of the totalcountry’s infested total area. infested Uganda area. is theUganda only countryis the only in thecountry region in that the hasregion endemic that has foci endemic of Trypanosoma foci of Trypanosoma brucei rhodesiense bruceiand rhodesienseT. b. gambiense and T. b.[20 gambiense]. Tsetse [20]. studies Tsetse in Uganda studies havein Uganda shown have that shown the country that the has country had three has had major three epidemics major epidemics of sleeping of sleeping sickness sickness since the since 1890s the (Figure1890s (Figure4). Over 4). the Over years, the the years, geographical the geographical extent of ex tsetsetent of prevalence tsetse prevalence and trypanosomosis and trypanosomosis in Uganda in hasUganda evolved. has Itevolved. has expanded It has expanded from the traditional from the trad concentrationitional concentration in the Buganda in the sub-region Buganda (1905–1920)sub-region to(1905–1920) the Busoga, to south-eastern the Busoga, andsouth-eastern mid-eastern and parts mid- ofeastern the Teso parts sub-region of the and Teso the sub-region northern and and West the Nilenorthern areas and of Uganda West Nile [3, 21areas,22]. of Uganda [3,21,22]. StudiesStudies inin partsparts ofof KenyaKenya andand UgandaUganda havehave revealedrevealed thatthat climate,climate, knowledgeknowledge ofof tsetsetsetse andand theirtheir controlcontrol methods,methods, culture, culture, farming farming practices practices and and demographic demographic factors factors have have a significant a significant influence influence on itson prevalenceits prevalence [23 ,[23,24].24]. The The occurrence occurrence belt belt observed observ ined easternin eastern and and mid-eastern mid-eastern Uganda Uganda now now appears appears to beto progressingbe progressing towards towards the Karamoja the Karamoja sub-region sub-region [19,22,25 [19,22,25].]. A spike A in thespike reports in the of tsetsereports prevalence of tsetse inprevalence the Karamoja in the sub-region Karamoja sub-region has been reported, has been particularlyreported, particularly in the districts in the ofdistricts Kaabong of Kaabong and Kotido. and InKotido. these In areas, these the areas, areas the that areas are that most are a ffmostected affected include include Sidok, Sidok, Lolelia, Lolelia, Kapedo Kapedo and Kathile. and Kathile. This outbreakThis outbreak in tsetse in tsetse represents represents a possible a possible invasion inva aftersion decades after decades of no reports of no reports of new of species new spreadspecies spread after the eradication campaigns of the 1960s [26]. Despite these observations, the extent to

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Sustainability 2019, 11, x FOR PEER REVIEW 4 of 25 after the eradication campaigns of the 1960s [26]. Despite these observations, the extent to which tsetse invasionwhich has tsetse impacted invasion the has socioecological impacted the resilience socioecological of pastoral resilience communities of pastoral in communities Karamoja remains in ambiguous.Karamoja The remains socioecological ambiguous. resilience The socioecological of ecosystems resilience depends of ecosystems on its ability depends to absorbon its ability change to and disturbance,absorb change without and shifting disturbance, to a new without regime, shifting governed to a new by aregime, diﬀerent governed set of processes by a different and structures.set of In thisprocesses case, resilience and structures. is the system’s In this case, ability resilience to resist is the transforming system’s ability into to a newresist system transforming state [ 27into]. Often,a transformationsnew system state and the[27]. associated Often, transformations threat of additional and the associated transformations threat of additional have critical transformations implications for both humanhave critical well-being implications and for resource both human management well-being [28 and]. resource management [28].

FigureFigure 4. Sleeping4. Sleeping sickness sickness epidemic epidemic cycl cycleses in in Uganda Uganda (Source: (Source: [22]). [22 ]).

SuccessfulSuccessful vector vector control control requires requires soundsound coordinationcoordination of of the the on on ground ground measures measures that thatare are foundedfounded on detailed on detailed knowledge knowledge of of the the vector’s vector’s distribution,distribution, movement movement patterns, patterns, connectivity connectivity across across socioecologicalsocioecological systems. systems. This This approachapproach requires requires a asound sound understanding understanding of the of spatial the spatial dynamics dynamics of of humanhuman and and animalanimal trypanosomiasis, trypanosomiasis, connectivity connectivity across across the landscape the landscape and good and coordination good coordination ability ability[3,29]. [3,29 A]. sub-regional A sub-regional study study in eastern in eastern and southern and southern Africa, Africa,[30] opined ref. [that30] changes opined thatin land-use changes in patterns and climate affected fly abundance and distribution. Furthermore, recent distribution land-use patterns and climate affected fly abundance and distribution. Furthermore, recent distribution patterns have become more confined to protected areas such as game parks and reserves. It has also patternsbeen have noted become that vector more distribution confined may to occur protected in limited areas biotopes, such assuch game as water parks points and and reserves. associated It has also beenareas notedfrequented that vectorby mammals. distribution Depending may on occur hydrographic in limited network biotopes, variations, such as waterdifferences points in and associateddistribution areas frequentedcould exist. For by example, mammals. a forest Depending region’s on vector hydrographic distribution network could be variations,uniquely different differences in distributionfrom that of could a savannah exist. For [31]. example, In this regard, a forest the region’s identification vector of distribution distributional could patterns be uniquely is critical di forfferent fromthe that execution of a savannah of control [31 measures]. In this [3] regard, and for the determining identification the challenge of distributional index that patterns reflects tsetse is critical for theabundance execution and of infection control measuresrates in an [ 3area] and [32]. for This determining identification the is challenge particularly index important that reflects in areas tsetse abundancethat are and experiencing infection recurrent rates in and an area sporadic [32]. tsetse This outbreaks, identification e.g., in is the particularly traditional important endemic areas in areas of south-eastern Uganda and the previously unaffected areas in northern Uganda. In these areas, that are experiencing recurrent and sporadic tsetse outbreaks, e.g., in the traditional endemic areas of HAT and AAT remain a major health threat to both humans and livestock [5,33]. south-eastern Uganda and the previously unaffected areas in northern Uganda. In these areas, HAT Vectors and vector-borne disease dynamics continue to change with changes in land use because and AATthey affect remain host-parasite a major health dynamics threat at tothe both landscape humans level. and These livestock dramatic [5 ,changes33]. have subsequently Vectorsled to a andloss vector-borneof resilience (social disease and dynamics ecological) continue that has to traditionally change with underpinned changes in landthe adaptive use because they acapacityffect host-parasite within dryland dynamics environments at the landscape and soci level.eties These[17,34]. dramatic This effect changes arises have because subsequently the led tosocioecological a loss of resilience system (social connections and ecological) within the that dryl hasands traditionally include the underpinnedintricate interactions the adaptive among capacitythe withinenvironment, dryland environments vectors, their relationships and societies with [17 huma,34].ns This and etheffect transmission arises because of bacteria, the socioecological viruses or systemprotozoa connections [35,36]. within Consequently, the drylands numerous include stud theies intricate on tsetse interactions distribution, among prevalence the environment, and vectors,trypanosome their relationships infection risk with have humans been conducted and the transmission [3,30]. However, of bacteria, there is a viruses paucity or of protozoainformation [35 ,36].

Consequently,Sustainability 2019 numerous, 11, x; doi: studiesFOR PEER onREVIEW tsetse distribution, prevalence ww andw.mdpi.com/journal/sustainability trypanosome infection risk have been conducted [3,30]. However, there is a paucity of information on the linkage between tsetse invasion and the socioecological resilience of communities. In this study, we assessed the Sustainability 2020, 12, 1599 5 of 26

Sustainability 2019, 11, x FOR PEER REVIEW 5 of 25 potentialon the impactlinkage ofbetween tsetse invasiontsetse invasion on the and socioecological the socioecological resilience resilience of pastoral of communities. communities In inthis the Karamojastudy, we sub-region. assessed the This potential approach impact was of taken tsetse because invasion tsetse on the distribution socioecological and prevalenceresilience of represent pastoral i) a socioecologicalcommunities in challengethe Karamoja that sub-region. requires interdisciplinary This approach was perspectives taken because for ittsetse to be distribution addressed andand ii) haveprevalence the potential represent to cause i) a changessocioecological in the areaschallenge where that the requires species interdisciplinary occurs, including perspectives transformations for it of theto socioecological be addressed and systems ii) have in responsethe potential to induced to cause perturbations. changes in the areas where the species occurs, including transformations of the socioecological systems in response to induced perturbations. 2. The Study Area 2. The study area The Karamoja sub-region is located between latitudes 2◦30’N and 4◦15’N and longitudes 33◦30’E and 34◦The35’E Karamoja in north-eastern sub-region Uganda is located (Figure between5). latitudes Rainfall 2°30’N in the sub-regionand 4°15’N and is highly longitudes variable 33°30’E with intermittentand 34°35’E drought in north-eastern episodes. Uganda Droughts (Figure in the 5). sub-region Rainfall in havethe sub-region been recorded is highly since variable the 1920s with and haveintermittent debilitating drought effects, episodes. including Droughts severe in livestock the sub-region mortality have [ 37been]. Therecorded region’s since temperatures the 1920s and are relativelyhave debilitating high with effects, high evapotranspiration including severe livestock levels. Karamojamortality is[37]. mostly The region’s covered temperatures by open savannah are grasslands,relatively high woodlands, with high thickets evapotranspiration and shrublands levels.[ 38Karamoja]. The is vegetation mostly covered in Karamoja by open savannah consists of Acacia–Combretum–Terminaliagrasslands, woodlands, thicketsspecies and associations,shrublands [38]. with The a grassvegetation layer in of KaramojaHyparrhenia, consists Setaria, of Acacia– Themeda, ChrysopogonCombretum–Terminaliaand Sporobolus speciesspp. associations, In this sub-region, with a grass five landlayer coverof Hyparrhenia, classes have Setaria, been Themeda, identified: subsistenceChrysopogon farmlands, and Sporobolus woodlands, spp. In grasslands, this sub-region thickets, five and land shrublands cover classe ands bushlands. have been Anidentified: increased emergencesubsistence of bushlands,farmlands, woodlands, especially ingrasslands, northern thic andkets southern and shrublands Karamoja, and has bushlands. been recorded An increased [38]. emergence of bushlands, especially in northern and southern Karamoja, has been recorded [38].

Sustainability 2019, 11, x; doi: FOR PEERFigure REVIEW 5. Location of the Karamoja sub-region. www.mdpi.com/journal/sustainability

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Livestock is an important mainstay of the Karamoja sub-region. The 2008 national livestock census estimated livestock populations in the sub-region as follows: 2,253,960 cattle, 2,025,293 goats, 1,685,500 sheep, 960 donkeys and 32,870 camels. Cattle are traditionally the most important livestock and are a source of wealth and status. However, cattle ownership and trade have declined in recent years due to disease, raiding and the transition to cropping. On average, 44% of families in Karamoja own some livestock [39]. The Uganda National Household Census of 2014 estimated the total population of the Karamoja sub-region to be close to one million people, with one of the highest population growth rates (Amudat 4.3%; Kotido 3.3%; Nakapiripirit 4.6%) in Uganda [40]. The sub-region is a major biodiversity conservation area, with 12% and 41% of the land area dedicated to forest reserves and wildlife conservation areas, respectively [41]. These areas are also important migratory zones for wild mammals, especially buﬀalos and elephants. These large mammals migrate southwards from Kidepo Valley National Park to other rangelands in central and southern Karamoja. The seasonal movements of these wildlife species have been attributed to inﬂuence tsetse distribution in the sub-region.

3. Research Methodology and Methods

3.1. Tsetse Entomological Survey A tsetse survey in the month of June-July 2016, was undertaken to provide baseline information on the Glossina morsitans distribution in northern Karamoja (Kotido and Kaabong districts). The two districts were selected because invasion reports by the District Veterinary Officers in the sub-region indicated that the two districts were most affected. Ten by ten kilometre (10 10 km) grids were × overlain in the districts of Kaabong and Kotido. The grids were used to ensure unbiased sampling during the tsetse survey and survey team assignment. Using the grids as operational units, five teams were allocated to the survey grids, and each grid had a unique identifier code as shown (Figure6) for ease of georeferencing and geotagging. Each team was led by an entomologist and field assistants. Sampling within the grids was based on the heterogeneity of habitats. Biconical tsetse traps were used in the survey (Figure7). The traps were odour-baited and set at a height of 30 cm above the ground. A total of 444 georeferenced traps were established. The trapping covered all the sub-counties in the study sites. However, trapping coverage depended on the probability of registering a catch as directed by local guides. Trapping at each site lasted 72 h in accordance with the established protocol [42]. Collections were made at the end of the 72-h period. At each collection, the parameters recorded in the predesigned entomological survey sheet included the trap code, eastings (X), northings (Y), elevation (Z), vegetation type at the trap site, start date and time, end date and time, species trapped, number of females or males and flies of unidentified sex and number of other biting insects. Caught tsetse flies were identified to the species level using the Food and Agriculture Organisation (FAO of the United Nations Training Manual for Tsetse Control Personnel [43]. Fly trap density (FTD), a measure of tsetse prevalence, was obtained by dividing the total tsetse catch per trap by the number of three trap days (72 h). Sustainability 2020, 12, 1599 7 of 26 Sustainability 2019, 11, x FOR PEER REVIEW 7 of 25

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Figure 6.6. A 10 × 1010 km km grid grid network network guide for team and trap deployment. Figure 6. A 10 × 10× km grid network guide for team and trap deployment.

Figure 7.7. A biconical trap at one of the ﬁeldfield sites.

3.2. Participatory assessment Assessment of of tsetse Tsetse prevalence Prevalence in in Karamoja Karamoja Participatory assessmentFigure involving 7. A biconical thethe use trap of focusfo cusat one group of discussionsthe field sites. (FGDs) and key informant interviews was was undertaken undertaken in in five ﬁve (Kaabong, (Kaabong, Kotido, Kotido, Nakapiripirit, Nakapiripirit, Amudat Amudat and Napak) and Napak) of the ofseven the sevendistricts districts in the Karamoja in the Karamoja sub-region. sub-region. The FGDs The were FGDs conducted were conducted in Sidok in (Figure Sidok 8), (Figure Lolelia,8), Lolelia,Loyoro, 3.2. Participatory assessment of tsetse prevalence in Karamoja Loyoro,Usake (Kaabong Usake (Kaabong district), district),Rengen (Kotido Rengen district), (Kotido Lorengdwat district), Lorengdwat (Nakapiripirit) (Nakapiripirit) and Karita and (Amudat Karita Participatory(Amudatdistrict). district).Additionally, assessment Additionally, key involving informants key informants the were use of wereinterviewed focus interviewed group from discussions from the theAmudat, Amudat, (FGDs) Napak, Napak, and Moroto, Moroto, key informant Nakapiripirit, Kotido and Kaabong districts. Though considered a low-risk location, Usake was interviewsNakapiripirit, was undertaken Kotido and in five Kaabon (Kaabong,g districts. Kotido, Though Nakapiripirit, considered a low-risk Amudat location, and Napak) Usake wasof the seven included because it is a major grazing area inin thethe drydry season. It is also a relatively high-altitude area districtsand in the aa cattlecattle Karamoja tradingtrading corridor corridorsub-region. with with South South The Sudan FGDsSudan Turkana Turkanawere conducted (Kenya). (Kenya). Furthermore, Furthermore, in Sidok it (Figure isit ais key a key exit8), exit routeLolelia, route for Loyoro, Usake (Kaabongcattlefor cattle raided raided district), by the by Turkanathe Rengen Turkana entering (Kotido entering Kenya district),Kenya and Toposaand Lorengdwat Toposa of South of South Sudan (Nakapiripirit) Sudan (Figure (Figure9). Participants 9). and Participants Karita in the (Amudat district).FGDsin Additionally,the andFGDs the and key the informant key key informantinformants interviewees interviewees were asked interviewed we tore deliberateasked to deliberatefrom on the historicalthe on Amudat,the patternshistorical ofNapak, patterns tsetse in Moroto, Nakapiripirit,theof tsetse sub-region. Kotidoin the Theysub-region. and had Kaabon to They relate ghad these districts. to trends relate totheseThough key politicaltren dsconsidered to events key political in the a sub-regionlow-risk events in andthelocation, sub-region in Uganda. Usake was This approach was meant to enable the research team to better position the event to relative historical includedand because in Uganda. it is This a major approach grazing was meant area into enablethe dr they season. research Itteam is also to better a relatively position the high-altitude event to area time.relative Participants historical alsotime. provided Participants information also provi aboutded the information perceived eaboutﬀects the of tsetse perceived on household effects of welfare tsetse and a cattleon household trading welfarecorridor and with on cultural South and Sudan social Turkana relations. (Kenya).The FGD participants Furthermore, consisted it is ofa elderskey exit route for cattleand raided Karachunas by the-herders Turkana whose entering knowledge Kenya and folklore and Toposa is well developed. of South KeySudan informant (Figure interviews 9). Participants in the FGDswere undertakenand the key with informant the District intervieweesGovernment Lead weers,re askedVeterinary to deliberateOfficers and ontwo the resident historical senior patterns of tsetse in the sub-region. They had to relate these trends to key political events in the sub-region and in Uganda.Sustainability This 2019, 11approach, x; doi: FOR PEERwas REVIEW meant to enable the research team www.mdpi.com/journal/sustainability to better position the event to relative historical time. Participants also provided information about the perceived effects of tsetse on household welfare and on cultural and social relations. The FGD participants consisted of elders and Karachunas-herders whose knowledge and folklore is well developed. Key informant interviews were undertaken with the District Government Leaders, Veterinary Officers and two resident senior

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Sustainability 2020, 12, 1599 8 of 26 and on cultural and social relations. The FGD participants consisted of elders and Karachunas-herders whose knowledge and folklore is well developed. Key informant interviews were undertaken with the SustainabilityDistrict Government 2019, 11, x FOR Leaders, PEER REVIEW Veterinary Oﬃcers and two resident senior researchers in the sub-region.8 of 25 Information gathered from the key informants was critical in triangulating the FGDs and cross-sectional researchers in the sub-region. Information gathered from the key informants was critical in researcherssurvey data. in the sub-region. Information gathered from the key informants was critical in triangulating the FGDs and cross-sectional survey data.

Figure 8 8.. FocusFocus group group discussions discussions (FGD) (FGD) in in Sidok sub-county.

Figure 9. A research team at Usake with the herd of rustled cattle recovered by the Uganda People’s Figure 9.9. A research team at Usake with the herd of rustled cattle recovered by the Uganda People’s Defense Forces (UPDF) from the Toposa of South Sudan. Defense Forces (UPDF) from the Toposa ofof SouthSouth Sudan.Sudan.

3.3. Cross-sectional Cross-Sectional survey Survey A householdhousehold cross-sectional cross-sectional survey survey was conductedwas conducted using semi-structuredusing semi-struc questionnairestured questionnaires involving 152involving randomly 152 selectedrandomly households selected households in Kaabong in and Kaab Kotidoong and districts. Kotido Respondents districts. Respondents provided information provided informationon the perceived on the eff ectsperceived of tsetse effects at the of household tsetse at the and household community and levels. community During thelevels. cross-sectional During the cross-sectionalsurvey, guided interviewssurvey, guided were usedinterviews in the administrationwere used in the of the administration semi-structured of the questionnaires. semi-structured This questionnaires.approach was preferred This approach because was of highpreferred illiteracy because rates of in high the area,illiteracy which rates were in estimatedthe area, which at 88% were [44]. estimatedEnumerators at 88% that administered[44]. Enumerators the semi-structured that administered questionnaires the semi-structured were selected questionnaires from within were the selectedregion. They from werewithin required the region. to have They an were education required qualification to have an of education the Uganda qualification Advanced of Certificate the Uganda of AdvancedEducation (HighCertificate School of Education Certificate). (High They School were knowledgeable Certificate). They in thewere local knowledgeable dialect used byin the local dialectcommunities used by in the Kotido local (Jie) communities and Kaabong in Kotido (Dodoth). (Jie)The and enumerators Kaabong (Dodoth). were trained The enumerators in the instrument were trainedadministration in the instrument process, procedures administ andration purpose. process, A procedures pre-test of the and semi-structured purpose. A pre-test questionnaires of the semi- was structuredundertaken questionnaires as part of the training was undertaken process. Afteras part the of pre-test, the training questions process. were After fine-tuned the pre-test, for consistency questions wereand accuracy. fine-tuned The for survey consistency ran for and a period accuracy. of two The weeks. survey ran for a period of two weeks.

4. Research analysis implementation

4.1. Tsetse prevalence in Karamoja

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4. Research Analysis Implementation

4.1. Tsetse Prevalence in Karamoja Fly trap density (FTD), a measure of tsetse prevalence, was obtained by dividing the total tsetse catch per trap by the number collected over the three trap days (72 h). To determine mean differences in the FTDs, a least significant difference (LSD) procedure using Duncan’s multiple range test for mean comparisons was used. Prior to running an LSD, analysis of variance (ANOVA) was performed to determine whether there were significant differences in FTDs among the sites. FGD data were thematically analysed. Emerging themes were grouped into thematic clusters to provide a logical result. The thematic analysis of participatory assessments was used because of its ability to provide critical insights among local pastoral communities [45,46]. Furthermore, the cross-sectional survey data were descriptively analysed, generating frequencies and means to describe the effects of tsetse at the household level.

4.2. Perceived Effect of Tsetse Invasion on Socioecological Resilience in Karamoja Resilience, a multifaceted term, has evolved over the last four decades in its meaning and application. It has subsequently risen in prominence in various disciplines. In the 1970s, as ecologists were grappling with explaining nonlinear dimensions within ecological systems, defined resilience from an ecological theory perspective [47]. It was also considered to be the ability of a system to absorb changes or shocks from a disturbance while maintaining its internal relations [48]. Since then, resilience has come to be used in a number of ways, often as an evocative metaphor in sustainability discourse. Second, resilience is a measurable quantity in a given socioecological system [49]. Increasingly, the concept has been recognised within the context of uncertainties and rapid changes that affect rural areas. Ecological, economic and social systems often become increasingly entangled, with the interactions among these systems increasing in scale and intensity [50]. Further, it has been noted that socioecological resilience is the capacity to adapt or transform in the face of change in socioecological systems [51]. Four critical issues are identifiable within the socioecological resilience realm: i) socioecological resilience relies on assumptions of nonlinearity in dynamics of change; ii) existence of cross-scale coordination; iii) adaptive management/co-management and learning; and iv) transformability into new pathways [52,53]. The measurement of resilience is a complex process, partly because long periods of observation may be required. However, in most cases, these data requirements are generally lacking, especially in rural sociological systems. Furthermore, considering historical contingency and path dependency contexts within complex ecological systems, it may be problematic to make generalisations about causal relationships [54]. Taking these perspectives into account, it becomes apparent that resilience is an emergent property of complex systems rather than a directly measurable characteristic [55]. As such, surrogates of socioecological systems are utilised to measure resilience. Such surrogates are those that are deemed to have the potential to enhance resilience and adaptive capacity [56]. Three components are important surrogates often measured to derive the resilience of communities: absorptive, adaptive and transformative capacities. Absorptive capacity refers to the various coping strategies used by individuals and/or households to moderate or buffer the impacts of shocks and stresses on their livelihoods and basic needs. They enable individuals and households to avoid suffering permanent and negative impacts on longer-term well-being [57,58]. Absorptive capacity is identified from a range of household capitals such as number of livestock owned [59]. Adaptive capacity refers to the ability of a household or community to adjust to changing social, economic and environmental conditions, including climate variability and extremes. It enables households and/or communities to moderate potential damages, take advantage of opportunities, or cope with the consequences and shocks [60]. It describes the proactive responses taken by individuals, households and communities in response to extreme events, e.g., tsetse invasion in this study. Conversely, transformative capacity of communities refers to the ability to create a new system when ecological, economic or social structures Sustainability 2020, 12, 1599 10 of 26 make the existing system untenable [61]. It reflects deliberate actions of the people in a given area. These actions could be autonomous as a result of spontaneous events. However, these actions could be unintentional, arising from forced transitions imposed from outside the system as a result of a perturbation. To establish the relationship, adaptive, absorptive and transformative indices were individually computed following the approach by [62]. As such, the adaptive capacity index was computed as:

ADCI = ADCn 0 + ADC 1 + ADCm 2 + ADC 3 (1) × l × × h × where,

ADCI = Adaptive capacity index

ADCn = Frequency of households that rate adaptation capacity as having no importance ADCl = Frequency of households that rate adaptation capacity as having low importance ADCm = Frequency of households that rate adaptation strategy as having moderate importance ADCh = Frequency of households that rate adaptation strategy as having high importance.

The relative importance of absorption capacity on socioecological resilience was calculated based on the following index formula:

ADCI = ABCn 0 + ABC 1 + ABCm 2 + ABC 3 (2) × l × × h × where,

ABCI = Absorptive capacity index

ABCn = Frequency of households that rate absorption capacity as having no importance ABCl = Frequency of households that rate absorption capacity as having low importance ABCm = Frequency of households that rate absorption capacity as having moderate importance ABCh = Frequency of households that rate absorption capacity as having high importance.

The relative importance of the transformative capacity on socioecological resilience was calculated based on the following index formula:

TRCI = TRCn 0 + TRC 1 + TRCm 2 + TRC 3 (3) × l × × h × where,

TRCI = Transformative capacity index

TRCn = Frequency of households that rate transformative capacity as having no importance TRCl = Frequency of households that rate transformative capacity as having low importance TRCm = Frequency of households that rate transformative capacity as having moderate importance TRCh = Frequency of households that rate transformative capacity as having high importance.

To obtain the resilience index, principal components using principal component analysis (PCA) and the varimax rotation method were applied to the factors in the absorptive capacity, adaptive capacity and transformative capacity. Factor loadings with a principal component >0.5 were identified and used in the computation of the resilience index. Accordingly, the principal components used were as follows: diversified economic activities and livestock off-take for absorptive capacity; access to livestock drugs, deployment of paravets-community animal health workers, access to information and early warming for adaptive capacity; and tsetse control infrastructure, access to extension services, level of sedentary households and urbanization for transformative capacity. These components formed the latent variables used to compute the resilience index; this formed the first stage. In the second Sustainability 2019, 11, x FOR PEER REVIEW 11 of 25

The results indicated that Kaabong district had a higher prevalence of males and females as well as a higher derived fly trap density (FTD) of tsetse populations than did Kotido district (Table 1). The Sustainability 2020, 12, 1599 11 of 26 distribution of females was found to be higher than that of males in both districts. In these districts, tsetse prevalence wasstage, higher the resilience in index the was areas derived of using Kare a factornga, analysis Loyoro, of the interacting Nakitoit, components Lokori, estimated Kmolicher, Longaro in the ﬁrst stage, and Locherep in Kaabong district (Figure 10); theseX areas are adjacent to Kidepo Valley National Park. Ri = wjFj (4) j in which the resilience index is a weighted sum of the factors (F ) generated by the scoring method, and Table 1. Geographical comparison of the distribution ofj tsetse in Kaabong and Kotido districts. the weights (wj) are the proportions of variance explained by each factor. In this study, the resilience index was computed from the principal component analysis factor loadings for component 1 that Altitude showed close similarity output Kaabong to the anticipated hypothesised Kotido outcomes. SE P-value

Parameter5. Results Tsetse Males 20.698a 1.033b 4.12903 0.0101 Tsetse Females5.1. Tsetse Spatial Distribution 23.893 and Prevalencea 3.1b 4.13741 0.0008 Tsetse Total The results indicated that 44.59 Kaabonga district had a higher 4.13b prevalence of males and 8.13339 females as well 0.0026 as a higher derived ﬂy trap density (FTD) of tsetse populations than did Kotido district (Table1). The a b Tsetse FTDdistribution of females was 14.864 found to be higher than that1.378 of males in both districts. 2.71113 In these districts, 0.0026 Means withtsetse prevalencedifferent was supers higher incripts the areas within of Karenga, a row Loyoro, are Nakitoit,significantl Lokori,y Kmolicher, different Longaro at p≥0.05. and Locherep in Kaabong district (Figure 10); these areas are adjacent to Kidepo Valley National Park.

Figure 10. Spatial distribution of tsetse ﬂies in northern Karamoja (Source: primary data). Figure 10. Spatial distribution of tsetse flies in northern Karamoja (Source: primary data).

5.2. Historical patterns and dispersal routes of tsetse in the Karamoja sub-region Through the participatory assessment undertaken in the districts of Kaabong, Kotido, Moroto, Amudat, Napak and Nakapiripirit, a historical account of tsetse in the sub-region was provided. Participants observed that the current invasion of tsetse was traceable to a period between 2003 and 2007 (Table 2). This period is in tandem with the renewed forceful disarmament exercise in the sub- region. The tsetse prevalence is accelerating within two wildlife migratory routes (eastern and western routes). The northern Karamoja to southern Karamoja apex route is aligned with the buffalo and elephant seasonal migrations that occur because the animals are in search of water and pasture. The eastern route commences from Kidepo Valley National Park through Pire-Nawuntos-Kalopeto- Kangaleta-Logum-Tukutan-Lokasirim to Remarim. Upon reaching Remarim, this route subdivides into two minor routes: one that enters Loyoro-Kotein to Sare and another that crosses into Kenya (Figure 11). The elephants and buffalos spend, on average, approximately three months at the water points at Loyoro, Kotein and Sare. These are the major convergence points. The second major migratory-dispersal route commences from Kidepo Valley National Park and runs through Nataba Lokure-Lofa-Omodoch-Lolelia-Lokapir. Upon reaching Lolelia, two minor routes emerge: one

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Sustainability 2020, 12, 1599 12 of 26

Table 1. Geographical comparison of the distribution of tsetse in Kaabong and Kotido districts.

Altitude Kaabong Kotido SE P-value Parameter Tsetse Males 20.698 a 1.033 b 4.12903 0.0101 Tsetse Females 23.893 a 3.1 b 4.13741 0.0008 Tsetse Total 44.59 a 4.13 b 8.13339 0.0026 Tsetse FTD 14.864 a 1.378 b 2.71113 0.0026 Means with different superscripts within a row are significantly different at p 0.05. ≥

5.2. Historical Patterns and Dispersal Routes of Tsetse in the Karamoja Sub-Region Through the participatory assessment undertaken in the districts of Kaabong, Kotido, Moroto, Amudat, Napak and Nakapiripirit, a historical account of tsetse in the sub-region was provided. Participants observed that the current invasion of tsetse was traceable to a period between 2003 and 2007 (Table2). This period is in tandem with the renewed forceful disarmament exercise in the sub-region. The tsetse prevalence is accelerating within two wildlife migratory routes (eastern and western routes). The northern Karamoja to southern Karamoja apex route is aligned with the buffalo and elephant seasonal migrations that occur because the animals are in search of water and pasture. The eastern route commences from Kidepo Valley National Park through Pire-Nawuntos-Kalopeto-Kangaleta-Logum-Tukutan-Lokasirim to Remarim. Upon reaching Remarim, this route subdivides into two minor routes: one that enters Loyoro-Kotein to Sare and another that crosses into Kenya (Figure 11). The elephants and buffalos spend, on average, approximately three months at the water points at Loyoro, Kotein and Sare. These are the major convergence points. The second major migratory-dispersal route commences from Kidepo Valley National Park and runs through Nataba Lokure-Lofa-Omodoch-Lolelia-Lokapir. Upon reaching Lolelia, two minor routes emerge: one towards the areas of Leterua-Naperetom-Sidok and Lopoet and one to Lolelia through Lomodoch-Kamoringaetyang-Sangar-Kotor-Kapeta-Lobanya-Kaicheri-Abim and Morulem (Figure 11). Key informants noted that tsetses on this western route have a risk of merging with those observed in parts of the Teso, Lango and Acholi sub-regions (Figure 12). Both the eastern and western routes open towards central and southern Karamoja. Presently, prevalence is largely concentrated in northern Karamoja in the districts of Kaabong and Kotido, with limited reports in the Amudat district in southern Karamoja. Within the two transmission routes in northern Karamoja, five tsetse fly concentration zones were identified in the communities as follows: i) Loyoro zone (Losululut, Aterak, Sar, Tapajei, Natelo, Musorod, Loglech, Loyile, Lobuneit, Bwangakou, Loumo, Maechit, Timira and Nakutan), ii) Regen zone (Lobel, Kalokitido, Morunyang, Waliwal Valley dam, Katukenyang Valley dam, Makal, Kanakori, Kanachom and Kaleta), iii) Kacheri zone (Lolelia, Kalingalem, Kayirang and Kapethinyang), iv) Usake-Kameon zone (Naminyit, Nawurat, Puta, Adumakuj, Lotila and Pire) and v) Lolelia zone (Kamoni, Narogole, Kumet, Kekuruk, Kamerisogol, Kamugemuge, Kalamaikol, Kotor, Lochokoi, Kaka, Lomodoch and Lochwai). Participants in the Napak and Nakapiripirit districts pointed to the areas of Narisai and sometimes to the bushes of Lotome and Narentogo in Nakapiripirit district. These areas are on the western migratory route, in which the northern Karamoja and central-to-southern Karamoja routes are connected through the Pian-Upe and Bokora wildlife reserves. In addition, some tsetse catches were reported in the Amudat district in southern Karamoja. Participatory assessment in Amudat indicated that the prevalence was higher in the areas of the Karita and Losidok sub-counties and the Katabok area towards the foot slopes of Mt. Kadam. Participants noted that heightened prevalence was observed at a time when the Pokot kinsmen from West Pokot (Kenya) crossed over to Uganda during the period of dry season grazing. Sustainability 2020, 12, 1599 13 of 26

Sustainability 2019,Sustainability 11, x FOR PEER 2019 REVIEW,Sustainability 11, x FOR PEER 2019 REVIEW, 11Sustainability, x FOR PEER 2019 REVIEW, 11, x FOR PEER REVIEW 14 of 25 14 of 25 14 of 25 14 of 25 Table 2. Historical timeline of events and tsetse prevalence in the Karamoja sub-region. Table 2. HistoricalTable timeline 2. Historical of eventsTable timeline and 2. Historical tsetse of events prevalenceTable timeline and 2. tsetse Historical inof theevents prevalence Karamoja timelineand tsetse insub-region. ofthe prevalenceevents Karamoja and intsetsesub-region. the Karamojaprevalence sub-region. in the Karamoja sub-region. Colonial Period through 1950s Post-Colonial Period 1962–Early1970s 1970s–Early 2000s Mid-2000s to Post–Disarmament Period

Post-colonial governmentPost-colonial governmentPost-colonial governmentPost-colonial governmentIn the mid-2000s In the mid-2000sIn the mid-2000s In the mid-2000s Post-colonial government recognised the tsetse recognised the tsetserecognised menace the tsetserecognised menace the tsetserecognised menace the tsetseapproximately menace 2003,approximately a 2003,approximatelyIn thea mid-2000s 2003,approximately approximately a 2003, 2003, a a disarmament menace in the sub-region. A tsetse control officer Ayela, a re-known tsetse eradication in the sub-region.in Athe tsetse sub-region. in Athe tsetse sub-region. inA thetsetse sub-region. disarmament A tsetse exercisedisarmament by the exercisedisarmament exerciseby the exercise by thedisarmament Government by the exercise of Uganda by the was initiated. called Ayela had been posted to Karamoja who officer, had been killed in the early 1970s. control officer calledcontrol Ayela officer calledcontrol Ayela officer calledcontrol Ayela officer Government called Ayela of UgandaGovernment was of UgandaGovernmentFirst, was thereof UgandaGovernment was a was peaceful of disarmamentUganda was involving Colonial administration was established established a camp in Kotido and parts of Kaabong The Karamojong were now armed, and had been postedhad to Karamoja been posted had to Karamoja been posted tohad Karamoja been posted initiated. to Karamoja First, thereinitiated. was aFirst, thereinitiated. wasvoluntary First,a thereinitiated. declaration was a First, and there return was of a guns to the in Karamoja with several effects closer to Kidepo Valley. ControlAyela, mechanisms, a re-knownAyela, tsetse a there-known rapidAyela, proliferationtsetse a re-known ofAyela, light tsetse weapons a re-known tsetse who establishedwho a camp established in who a camp established in awho camp established in apeaceful camp in disarmament peaceful disarmamentpeacefulUganda disarmament People’speaceful Defense disarmament Forces (UPDF). Failure to including the well-established including a gazette of communitieseradication offi in onecer,eradication had been offi becamecer,eradication had morebeen offi prominentcer,eradication had been in the officer, had been Kotido and partsKotido of Kaabong and parts Kotido of Kaabong and parts ofKotido Kaabong and parts ofinvolving Kaabong voluntaryinvolving voluntaryinvolvingachieve successvoluntaryinvolving and continued voluntary counter raids, forced subdivision of cultural groupings in location/camps and mobilisingkilled able-bodied in the early menkilled 1970s. to in the sub-region.earlykilled 1970s. in Governancethe early killed1970s. systems in the early 1970s. closer to Kidepocloser Valley. to Kidepocloser Valley. to Kidepo Valley.closer to Kidepodeclaration Valley. and returndeclaration of and declarationreturn theof Government and returndeclaration of of Uganda and return to launch of a forceful Karamoja.Colonial Tsetse administration wasColonial recognised was administration as aColonialcut was andadministration burnColonial all bushes was administration and trees.The Karamojong Much was vegetation The were Karamojong now collapsedThe were Karamojong now with regards wereThe to Karamojong now public were now Control mechanisms,Control mechanisms,Control mechanisms,Control mechanisms,guns to the Ugandaguns People’s to the Uganda guns People’s todisarmament the Uganda guns People’s exercise. to the Uganda Upon the People’s disarmament, the problemestablished in the region in andKaramojaestablished a control camp in Karamojaestablishedwas cut,in Karamoja transformingestablished woodlandsin Karamojaarmed, into and open the rapidarmed, andauthority the rapidarmed, and administration. and the rapidarmed, Intense and the rapid including a gazetteincluding of a gazetteincluding of a gazetteincluding of a gazetteDefense of Forces Defense(UPDF). Forces Defense(UPDF).UPDF Forces then (UPDF). providedDefense Forces security (UPDF). to communities and was established at Lolelia. A camp was withgrasslands. several effects Evidence of these remnant tree trunks livestock rustlingproliferation and wildlifeof light poaching with several effectswith several effects with several effectsproliferation of lightproliferation of light proliferation of lightherders. This marked the return of normalcy and also located on the fertile lands of communitiescan be observed in communitiesone at the Kidepo in Valley.communitiesone In addition, in onecommunities becameFailure in prominent.one to achieveFailure Wildlife success to were achieveFailure success to achieveFailure success to achieve success including the well-including the well-including the well-including the weaponswell- becameweapons more becameweapons more became moreweapons became more‘peace’ in the region. The wildlife that had hitherto Lokapir and Naperetom. This group location/campsherbicides that andlocation/camps killed everything andlocation/camps including bees andlocation/camps confinedand intocontinued and the Kidepo counterand Nationalcontinued raids, Park, counterand continued raids, counterand continued raids, counter raids, established subdivisionestablished of subdivisionestablished of subdivisionestablished of subdivisionprominent of in theprominent sub-region. in theprominent sub-region. in the sub-region.prominent in the sub-region.been confined to Kidepo Valley National Park slowly used to kill wildlife, including elephants,mobilising(non-selective) able-bodiedmobilising were men usedable-bodied to mobilising in the eradication men able-bodied to mobilising efforts. men which toable-bodied forced became menthe more Government to likeforced a refuge the of centre.Governmentforced the of Governmentforced of the Government of cultural groupingscultural in groupingscultural in groupingscultural in groupingsGovernance in systemsGovernance systemsGovernance systemsGovernance systemsstarted to return to graze in the outskirts of the as trophies. This was perhaps the worstcut andAt burn the fall all ofcutbushes the and Obote andburn Iall government cutbushes and andburn in all the bushes earlycut and and burnThis all periodUganda bushes also to and representedlaunch Uganda a forceful a to period launch Uganda of a forceful to launch Ugandaa forceful to launch a forceful Karamoja. TsetseKaramoja. was TsetseKaramoja. was Tsetse Karamoja.was Tsetsecollapsed was with regardscollapsed to with regardscollapsed to with regardscollapsed to with regardsNational to Park, especially during the dry season. With time for wildlife and livestock. Cattletrees.1970s, Much Karamojong vegetationtrees. Much acquiredwas vegetationtrees. guns afterMuch was overrunning vegetationtrees. wasMuch obscurity vegetationdisarmament of the was Karamoja exercise.disarmament sub-regionUpon exercise.disarmament Upon exercise.disarmament Upon exercise. Upon recognised as a problemrecognised in as a recognisedproblem in as a problemrecognised in as a problempublic inauthority publicand authoritypublic and authority andpublic authoritythe and return of the wildlife, the reinvasion of tsetse was were herded and confined to the eastcut, transformingthe Morotocut, woodlands barracks. transforming Light cut, weaponswoodlands transforming made entrycut,woodlands transforming fromthe governmentwoodlands disarmament, investment,the the disarmament, UPDF and the the disarmament, UPDF thethe disarmament,UPDF the UPDF the region and athe control region and athe control region and a controlthe region and aadministration. control Intenseadministration. Intenseadministration. Intenseadministration. Intensefirst reported in Kaabong and Kotido. As the wildlife of Karamoja. intointo open the life grasslands. of theinto pastoral open grasslands. communityinto open in thisgrasslands. region.into openKaramoja grasslands.thenwas provided viewed securitythen as a problemprovided to to securitythen provided to securitythen providedto security to camp was establishedcamp wasat establishedcamp wasat establishedcamp at was establishedlivestock at rustlinglivestock and rustlinglivestock and rustling livestockand rustling andcontinues to routinely return to the grazing and EvidenceAyela of these wasEvidence killedremnant by of Amin’s theseEvidence remnant soldiers, of and these thisEvidence remnant the of neighbouringthesecommunities remnant communities andcommunities herders. in Teso, andcommunities herders. and communitiesherders. and herders. Lolelia. A campLolelia. was also A campLolelia. was also A camp wasLolelia. also A campwildlife was also poaching wildlife became poaching wildlife became poaching wildlifebecame poaching waterholesbecame outside the park, the prevalence of tsetse is tree trunksmarked can thebetree endobserved trunks of the can at tsetse betree observed eradication trunks can at ebefforts treeobserved intrunks at can beThisLango, observed marked Sebei at the andThis return Acholi. marked of theThis return marked of the returnThis marked of the return of located on the fertilelocated lands on the fertilelocated lands on the fertilelocated lands on the fertileprominent. lands Wildlifeprominent. were Wildlifeprominent. were Wildlifeprominent. were Wildlife were also intensifying. the Kidepo Valley.the InKidepothe region. Valley.the InKidepo Valley.the In Kidepo normalcyValley. In and ‘peace’normalcy in the and ‘peace’normalcy in the and ‘peace’normalcy in the and ‘peace’ in the of Lokapir and Naperetom.of Lokapir and Naperetom.of Lokapir and Naperetom.of Lokapir and Naperetom.confined into theconfined Kidepo into theconfined Kidepo into the Kidepoconfined into the Kidepo addition, herbicidesaddition, that killed herbicides addition, that killed herbicides addition, that killed herbicides region. that The killed wildlife region. that The had wildlife region. that The had wildlife region. that hadThe wildlife that had This group usedThis to kill group usedThis to kill group used toThis kill group usedNational to kill Park, whichNational Park, whichNational Park, whichNational Park, which everything includingeverything bees includingeverything bees includingeverything bees includinghitherto bees been confinedhitherto to been confinedhitherto to been confinedhitherto to been confined to wildlife, includingwildlife, includingwildlife, includingwildlife, includingbecame more likebecame a refuge more likebecame a refuge more likebecame a refuge more like a refuge (non-selective) were(non-selective) used in were(non-selective) used in were(non-selective) used in Kidepo were used Valley in NationalKidepo ValleyPark NationalKidepo Valley Park NationalKidepo Park Valley National Park elephants, as trophies.elephants, This as trophies.elephants, This as trophies.elephants, This as trophies.centre. This This periodcentre. also This periodcentre. also This periodcentre. also This period also the eradication efforts.the eradication At the efforts.the eradication At the efforts.the eradication At the efforts.slowly Atstarted the toslowly return startedto toslowly return startedto to returnslowly to started to return to was perhaps the wasworst perhaps time thewas worst perhaps time the worstwas perhapstime the worstrepresented time a periodrepresented of a periodrepresented of a periodrepresented of a period of fall of the Obotefall I government of the Obote fall I government of the Obote Ifall government of the Obote graze I government in the outskirts graze ofin thethe outskirtsgraze ofin thethe outskirtsgraze of inthe the outskirts of the for wildlife and forlivestock. wildlife and forlivestock. wildlife and livestock.for wildlife andobscurity livestock. of the obscurityKaramoja of the obscurityKaramoja of the Karamojaobscurity of the Karamoja in the early 1970s,in the early 1970s,in the early 1970s,in the earlyNational 1970s, Park, especiallyNational Park, especiallyNational Park, especiallyNational Park, especially Cattle were herdedCattle and were herdedCattle and were herdedCattle and were herdedsub-region and fromsub-region government from sub-region government from governmentsub-region from government Karamojong acquiredKaramojong guns acquiredKaramojong guns acquiredKaramojong guns acquiredduring the guns dry season.during Withthe dry season.during Withthe dry season.during With the dry season. With confined to the eastconfined of to the eastconfined of to the eastconfined of to theinvestment, east of and investment,Karamoja and investment,Karamoja and Karamojainvestment, and Karamoja after overrunningafter the overrunningMoroto after the overrunning Moroto afterthe Moroto overrunning the thereturn Moroto of the the wildlife, return the of thethe wildlife, return theof the wildlife,the return the of the wildlife, the Karamoja. Karamoja. Karamoja. Karamoja.was viewed as awas problem viewed to as awas problem viewed to as a problemwas viewed to as a problem to barracks. Light weaponsbarracks. made Light weaponsbarracks. madeLight weaponsbarracks. made Light reinvasionweapons made of tsetse reinvasion was first of tsetsereinvasion was first of tsetsereinvasion was first of tsetse was first the neighbouringthe neighbouringthe neighbouring the neighbouring entry into the lifeentry of the into the lifeentry of the into the life entryof the into the reportedlife of the in Kaabongreported and in Kaabongreported and in Kaabongreported and in Kaabong and communities in communitiesTeso, in communitiesTeso, in Teso,communities in Teso, pastoral communitypastoral in this community pastoral in this community pastoral in this community Kotido. in this As the wildlifeKotido. As the wildlifeKotido. As the wildlifeKotido. As the wildlife Lango, Sebei andLango, Acholi. Sebei andLango, Acholi. Sebei and Lango,Acholi. Sebei and Acholi. region. Ayela wasregion. killed Ayela by wasregion. killed Ayela by wasregion. killed byAyela continueswas killed to by routinely continues return to routinelycontinues return to routinelycontinues return to routinely return Amin’s soldiers,Amin’s and this soldiers, Amin’s and this soldiers, andAmin’s this soldiers, andto the this grazing andto the grazing andto the grazing andto the grazing and marked the end markedof the tsetse the end markedof the tsetse the end ofmarked the tsetse the endwaterholes of the tsetse outside waterholes the park, outside waterholes the park, outside waterholes the park, outside the park, eradication effortseradication in the effortseradication in the effortseradication in the effortsthe prevalence in the ofthe tsetse prevalence is ofthe tsetse prevalence is of thetsetse prevalence is of tsetse is region. region. region. region. also intensifyingalso. intensifyingalso. intensifying. also intensifying. Participants attributedParticipants tsetse attributedParticipants invasion tsetse inattributed theParticipantsinvasion sub-region tsetse in attributed the invasionto sub-regionseveral tsetsein factors.the toinvasion sub-regionseveral The in pastoralfactors. theto severalsub-region The pastoralfactors. to severalThe pastoral factors. The pastoral communities acknowledgedcommunities acknowledged thatcommunities the tsetse acknowledged eradicationthatcommunities the tsetse efforts eradicationthat acknowledged of the the tsetse 1960s efforts eradication andthat of 1970s, thethe tsetse1960s efforts especially eradicationand of 1970s,the those 1960s especially efforts and of1970s, thethose 1960s especially and 1970s, those especially those championed bychampioned Ayela, were bychampioned effective Ayela, werein byreducing effective championedAyela, the werein tsetsereducing byeffective Ayela, populations the inwere tsetsereducing effectivein populations the the sub-region. intsetse reducing in populations the They thesub-region. tsetse in thepopulations They sub-region. in theThey sub-region. They observed that theobserved Republic that of theobserved Sudan Republic (now that of South the Sudanobserved Republic Su (nowdan) that of didSouth Sudanthe not RepublicSu undertake (nowdan) didSouth of Sudannotany Su undertakeactiondan) (now didtowards South notany undertake Suaction tsetsedan) towards did any not action tsetseundertake towards any tsetse action towards tsetse eradication in itseradication area of control, in itseradication area which of control,left in itstheeradication area problemwhich of control, left onlyin the its half problemwhicharea solved. of left control, only the Participants halfproblem which solved. leftonly noted Participantsthe half problemthat solved. the noted only Participants half that solved. the noted Participants that the noted that the source of the currentsource invasion of the currentsource appears invasion of tothe have current appearssource a transboundary invasion of to the have current appears a transboundary character invasion to have emerging aappears transboundary character from to have emerging South a character transboundary Sudan from emerging South character Sudan from South emerging Sudan from South Sudan and perhaps fromand perhapswestern fromKenyaand perhapswestern (based fromKenyaonand the western perhaps (basedcatches Kenya on fromreported the western(basedcatches in Amudaton Kenyareported the catches (baseddistrict). in Amudat reportedon They the district).catchesalso in Amudat reportedThey district).also in Amudat They alsodistrict). They also attributed the prevalenceattributed the to the prevalenceattributed disarmament the to the prevalence attributedexerci disarmamentse (removing to the the prevalence exerci disarmament AK47se (removing to guns) the exerci disarmament that AK47se was (removing guns) undertaken exerci that AK47 wasse in (removing guns) undertaken that AK47 was in undertaken guns) that wasin undertaken in the sub-region thefrom sub-region approximatelythe from sub-region approximately 2003, which fromthe played sub-region approximately 2003, awhich significant from played 2003, approximately role awhich significant in releasing played 2003, role athe significantwhich in buffalos releasing played androle the a in significant buffalos releasing and rolethe buffalosin releasing and the buffalos and elephants fromelephants the Kidepo from Valleyelephants the Kidepo National from Valley elephantsthePark Kidepo Nationalto graze from Valley Parkfreely the NationalKidepoto in graze other Valley Parkfreely parts to National inin graze otherthe sub-region.freely partsPark intoin othergrazethe sub-region. partsfreely in in the other sub-region. parts in the sub-region. Participants indicatedParticipants that indicatedpreviously,Participants that when indicatedpreviously,Participants the communitiesthat when previously,indicated the werecommunities that when full previously,y th armed,e werecommunities elephantswhenfully armed,th ewere andcommunities elephantsfully armed, wereand elephants fully armed, and elephants and buffalos were containedbuffalos were in the containedbuffalos park for were insafety, the containedbuffalos park as the for communitiesinwere safety, the contained park as the for killed communities safety, in the them. aspark the They forkilled communities safety,also them. noted as Theythe killedthat communities also them. noted They that killed also them.noted Theythat also noted that the creation ofthe valley creation dams of andthe valley creationpermanent dams of and valleywaterthe permanent creation podamsints and ofin water valleyareas permanent po thatdamsints originally in andwater areas permanent po thatdidints originallynot in waterareashave such thatpodidints originallynot in have areas suchdid that not originally have such did not have such

Sustainability 2019,Sustainability 11, x; doi: FOR 2019 PEER,Sustainability 11, REVIEWx; doi: FOR 2019 PEER, 11Sustainability, x;REVIEW doi: FOR 2019 PEER, 11 REVIEW, x; doi: FOR www. PEERmdpi.com/journal REVIEW www. /sustainabilitymdpi.com/journal www. mdpi.com/journal/sustainability www. /sustainabilitymdpi.com/journal /sustainability

Sustainability 2019, 11, x FOR PEER REVIEW 12 of 25 towards the areas of Leterua-Naperetom-Sidok and Lopoet and one to Lolelia through Lomodoch- Kamoringaetyang-Sangar-Kotor-Kapeta-Lobanya-Kaicheri-Abim and Morulem (Figure 11). Key informants noted that tsetses on this western route have a risk of merging with those observed in parts of the Teso, Lango and Acholi sub-regions (Figure 12). Both the eastern and western routes open towards central and southern Karamoja. Presently, prevalence is largely concentrated in northern

KaramojaSustainability in the2020 districts, 12, 1599 of Kaabong and Kotido, with limited reports in the Amudat 14district of 26 in southern Karamoja.

FigureFigure 11. Tsetse 11. Tsetse dispersal dispersal routes routes in Karamoja in Karamoja (Source: (Source: participatory participatory assessment assessment data). data). The Thereddish- brownreddish-brown arrows denote arrows tsetse denote emergi tsetseng emerging from Kidepo from Kidepo Valley Valley National National Park Park in innorthern northern Karamoja. Karamoja. The purpleThe arrows purple denote arrows the denote tsetse the perceived tsetse perceived to be to coming be coming from from West West Pokot Pokot in in Kenya. Kenya.

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Sustainability 2020, 12, 1599 15 of 26 Sustainability 2019, 11, x FOR PEER REVIEW 13 of 25

FigureFigure 12. 12. PredictionPrediction of of tsetse prevalenceprevalence for for the the three thr majoree major species species in Uganda in Uganda in 2011 in (Source: 2011 (Source: [63]). [63]). Participants attributed tsetse invasion in the sub-region to several factors. The pastoral communities acknowledged that the tsetse eradication efforts of the 1960s and 1970s, especially Within the two transmission routes in northern Karamoja, five tsetse fly concentration zones those championed by Ayela, were effective in reducing the tsetse populations in the sub-region. They wereobserved identified that in thethe Republic communities of Sudan as follows: (now South i) Loyoro Sudan) zone did not (Losululut, undertake Aterak, any action Sar, towards Tapajei, tsetse Natelo, Musorod,eradication Loglech, in its Loyile, area of Lobuneit, control, which Bwangakou, left the problem Loumo, only Maechit, half solved. Timira Participants and Nakutan), noted ii) that Regen zonethe (Lobel, source Kalokitido, of the current Morunyang, invasion appears Waliwal to have Valley a transboundary dam, Katukenyang character emerging Valley from dam, South Makal, Kanakori,Sudan andKanachom perhaps fromand western Kaleta), Kenya iii) (based Kacheri on the catcheszone reported(Lolelia, in Kalingalem, Amudat district). Kayirang They also and Kapethinyang),attributed the iv) prevalence Usake-Kameon to the disarmament zone (Naminyit, exercise Nawurat, (removing Puta, AK47 Adumakuj, guns) that Lotila was undertaken and Pire) and v) Loleliain the sub-regionzone (Kamoni, from approximately Narogole, Kumet, 2003, which Kekuruk, played Kamerisogol, a significant role Kamugemuge, in releasing the Kalamaikol, buffalos Kotor,and Lochokoi, elephants fromKaka, the Lomodoch Kidepo Valley and National Lochwai). Park Participants to graze freely in in otherthe Napak parts in and the sub-region.Nakapiripirit districtsParticipants pointed indicated to the areas that previously, of Narisai when and thesome communitiestimes to the were bushes fully armed, of Lotome elephants and and Narentogo buffalos in were contained in the park for safety, as the communities killed them. They also noted that the creation Nakapiripirit district. These areas are on the western migratory route, in which the northern of valley dams and permanent water points in areas that originally did not have such waterholes Karamoja and central-to-southern Karamoja routes are connected through the Pian-Upe and Bokora attracted elephants and buffalos into such areas. Accordingly, the migration and dispersion of buffalos wildlifeand reserves. elephants In have addition, widened some in the tsetse region, catches and theirwere residence reported time in the in particularAmudat district locations in where southern Karamoja.permanent Participatory water sources assessment exist has in increased. Amudat Consequently, indicated that there the isprevalence increased breeding,was higher dispersion in the areas of theand Karita distribution and Losidok in non-traditional sub-counties tsetse and host the locations Katabok in area the sub-region. towards the foot slopes of Mt. Kadam. Participants noted that heightened prevalence was observed at a time when the Pokot kinsmen from West Pokot (Kenya) crossed over to Uganda during the period of dry season grazing.

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Sustainability 2020, 12, 1599 16 of 26

5.3. Perceived Socioecological Effects of Tsetse in Karamoja Participants observed that herder movements to and from grazing grounds had become limited. They observed that grazing grounds with good pastures, such as Hyparrhenia newtonii (Hack.) (Emaa), Sporobolus pyramidalis Beauv (Ngajien), Eragrostis pilosa (L.) Beauv (Ngiletio) and Bracharia brizantha (Hochst.) Stapf (Elet), had a high tsetse prevalence. In this case, the herders had to realign their grazing movements by reducing residence time in the grazing grounds as was historically done. This approach creates a higher grazing pressure in areas with low-quality pastures. Participants observed that tsetse prevalence had significant effects on kinsmen from Turkana in particular, as it restricted their movement to the grazing lands in Uganda. They also observed that livestock diseases, particularly livestock nagana and other tick-borne diseases, became prevalent mainly in the Kaabong, Kotido and Amudat districts. These diseases have had negative effects on livestock health in the area. Participants indicated that because of their livestock’s inability to graze well, cattle milk yield became limited, livestock body conditions decreased and some livestock died over time. Furthermore, they observed that livestock abortions among cattle, donkeys, sheep and goats become more common than before. Participants from Lolelia and Sidok also noted that tsetse flies were most prevalent in the ‘fertile’ agricultural lands. This prevalence affected women’s active participation in crop production because women avoided staying in the gardens due to tsetse bites on them and on children. As a result, overall crop output, especially sorghum, a key food security crop in the sub-region, was affected. Despite the observed variability in livestock numbers, the results of the cross-sectional survey revealed that there was a visible perceived effect of tsetse flies on owned and sold livestock (Table3). Approximately 94.7% of the respondents indicated that their livestock numbers had decreased due to tsetse invasion. Meanwhile, 76.3% of the respondents indicated that their crop production activities in the fertile agricultural lands had been affected. Respondents participated in diverse and multiple livelihood activities: 55.3% relied on farming, 48.7% on charcoal burning, 9.2% on beekeeping, 11.8% on the sale of livestock and livestock products and 2.6% on gold mining.

Table 3. Livestock owned and livestock sold.

Livestock Owned Livestock Sold Livestock Affected by Tsetse Not Affected by Tsetse Affected by Tsetse Not Affected by Tsetse (N = 76) (N = 75) (N = 76) (N = 75) Cattle 5.8 5.5 5.8 7.7 1.3 1.9 1.1 1.6 ± ± ± ± Sheep 20.6 84.5 6.8 12.7 2.4 4.5 1.1 2.4 ± ± ± ± Donkeys 0.8 1.9 0.7 1.3 0.1 0.5 0.1 0.9 ± ± ± ± Goats 11.3 21.3 8.3 9.6 2.7 3.4 1.7 2.5 ± ± ± ±

In terms of the effect of tsetse invasion on socioecological resilience, 57% of respondents indicated that they had the capacity to adapt to tsetse invasion, 54% indicated that the area had the requisite transformative capacity and 35% indicated that there was a mechanism to absorb any shocks due to tsetse invasion. The results of the absorptive capacities showed that diversified economic activities were ranked first. This result constituted the most important absorption strategies to tsetse prevalence among respondents. Access to financial services was ranked second, followed by livestock off-take, bush burning and seeking loan facilities (Table4). The results of the adaptive capacities showed that diversified access to livestock drugs was ranked high and was most important among respondents’ adaptive capacities to tsetse prevalence. This was followed by the deployment of paravets, access to information and early warning systems, diversification of livestock and temporary migration (Table5). Sustainability 2020, 12, 1599 17 of 26

Table 4. Ranked order of the absorptive capacity to tsetse invasion.

Absorptive Capacity High Medium Low No ABCI Rank Diversified economic activities 14 54 7 1 157 1 Access to financial services 13 29 18 16 115 2 Livestock off-take 2 35 26 13 102 3.5 Bush burning 11 23 23 18 102 3.5 Seeking loan facility 4 35 17 20 99 5

Table 5. Ranked order of the adaptive capacity to tsetse invasion.

Adaptive Capacity High Medium Low No ADCI Rank Access to drugs 59 13 4 0 207 1 Deployment of paravets (CAHWs) 52 20 4 0 200 2 Access to information and early 25 36 12 3 159 3 warning systems Diversiﬁcation of livestock 17 42 11 6 146 4 Temporary migration 2 11 36 27 64 5

For transformative capacity, tsetse control infrastructure was ranked high in response to tsetse prevalence. This was followed by access to extension services (entomologists), level of sedentary population and urbanization (Table6). When all these capacities were combined, the factor analysis results revealed that the absorptive capacity of the respondents was 15.0%, the adaptive capacity was 22.1% and the transformative capacity was 24.5% (Table6). The results of the principal component analysis (PCA) were more stable in component 1. These factor loadings were used in the computation of the resilience index (Table7). Accordingly, the computed socioecological resilience index of the study area was 20.5% (Table8).

Table 6. Ranked order of the transformative capacity to tsetse invasion.

Transformative Capacity High Medium Low No TRCI Rank Tsetse control infrastructure 38 29 8 1 180 1 Access to extension services 34 25 14 3 166 2 (entomologists) Sedentary level 6 39 27 4 123 3 Urbanization 1 11 35 29 60 4

Table 7. Principal component analysis factor loadings.

Factor Comp1 Comp2 Comp3 Comp4 Comp5 Comp6 Unexplained Livestock off-take 0.1338 0.2675 0.2449 0.4227 0.3244 0.2339 0.2871 − − − Access to financial services 0.3083 0.3402 0.3808 0.2425 0.1411 0.2371 − Seeking a loan facility 0.3816 0.3184 0.3436 0.0556 0.1755 0.1939 − − Diversified economic activities 0.3189 0.3586 0.1308 0.1810 0.1907 0.2019 0.2768 − − Bush burning 0.3650 0.1085 0.1384 0.3779 0.4266 − Diversification of livestock 0.2767 0.1351 0.2258 0.4052 0.5365 0.2222 − Temporary migration 0.0929 0.1782 0.4880 0.2928 0.2090 0.2488 0.3173 − − Deployment of paravets (CAHWs) * 0.3971 0.2630 0.1318 0.1775 0.3587 − Access to information and early 0.3327 0.3271 0.1804 0.2127 0.1220 0.0741 0.3053 warning systems − − Access to drugs 0.1950 0.3140 0.1980 0.2766 0.2799 0.4527 0.249 − − − Tsetse control infrastructure 0.2034 0.4264 0.2428 0.2544 0.2356 0.2063 0.2206 − − Urbanization 0.1928 0.1585 0.2920 0.4505 0.1497 0.1398 0.3682 − − − − Sedentary level 0.1662 0.2266 0.2594 0.1437 0.6075 0.2235 0.2373 − − Access to central extension services 0.0939 0.5299 0.2290 0.1693 0.1096 0.0854 0.2581 (entomologists) − − * Community Animal Health Workers (CAHWs). Sustainability 2020, 12, 1599 18 of 26

Table 8. Resilience index.

25th 50th 75th Std. Dev. Variable Mean Min Max Percentile Percentile Percentile (Mean) Resilience Index 1.0802 2.2285 3.0526 2.052675 1.304063 0.5905 4.8386 − Rescaled Resilience Index 0.3077 0.5192 0.6710 0.4868532 0.2401987 0 1

6. Discussion

6.1. Tsetse Prevalence and Distribution Patterns The results showed a high tsetse prevalence in close proximity to the protected areas (Kidepo Valley National Game Park). This pattern could be attributed to the fact that these areas have remained undisturbed habitats for wildlife, including the buffalos and elephants that act as primary hosts of tsetse. Second, Kidepo Valley National Park is connected to the Kidepo Game Reserve in South Sudan, which is nearly four times larger. The Kidepo Game Reserve in South Sudan is believed to have continued to serve a breeding ground for tsetse in the region. No major traceable eradication intervention action was found to have been implemented in this area. Wildlife acts as hosts and influences the survival of tsetse due to a constant supply of food (i.e., blood). Furthermore, game parks act as breeding grounds [30,64]. Thus, the observed prevalence in proximity to the National Park should not be surprising. Current evidence indicates tsetse dispersal from the north to central southern Karamoja following wildlife migratory routes. Additionally, there is a dispersal route emerging from southern Karamoja towards central Karamoja. This particular route is linked to the seasonal migrations of Pokot from West Pokot in Kenya, as animals search for pasture and water during the dry season. Tsetse prevalence in western Kenya, including in Turkana and West Pokot that neighbour Karamoja, and whose people routinely migrate to Karamoja in search of water and pasture, have previously been documented [65]. This belt from southern Karamoja thus appears to be an expansion of the western Kenya tsetse belt that has previously been reported, with an expanding tsetse belt in the region bordering Uganda [23,24,66]. The dispersal of tsetse southwards is particular concern, as there is potential for this tsetse population to interact with other tsetse species (Figure 12) in eastern and northern Uganda [63]. If this happens, the tsetse belt in the country will become more complex with unprecedented effects. However, the current dispersal southwards is linked to the migration of wildlife in search of water and pasture, particularly during the dry season. Seasonally distributed tsetse patterns associated with livestock and wildlife migration patterns have previously been documented in the Maasai Steppe of Tanzania, with pronounced abundance during the dry season [67]. During the participatory assessments, participants noted that the construction of permanent waterholes, such as those at Loyoro, Kotein and Sare, was further complicating the migration patterns of wildlife in the sub-region. These artificial waterholes enabled the elephants and buffalos to spend more time grazing outside the park than traditionally known. These waterholes were introduced as a governmental response to frequent pastoral movements, restricted movements and conflict over water resources during the dry seasons [68]. Regardless of why they were developed, it is important to note that the extent of tsetse distribution in the sub-region can no longer be ignored. It is clear that tsetse has had a negative effect by affecting various aspects of life, culture and relations, production and economy in the sub-region. It has also affected relations with kinsmen and women in neighbouring Kenya, particularly in the Turkana and Pokot pastoral communities. By affecting livestock, tsetse is generating a livestock health crisis that has cumulative effects on socioeconomic and cultural well-being. In a review of five African countries, Meyer et al. observed that tsetse affected farmers’ livelihoods and welfare and food security and posed a threat to public health [69]. As indicated by participants in the assessments, livestock deaths caused by livestock nagana have become widespread, particularly in the Kaabong and Kotido districts. These observations corroborate earlier reports of tsetse infestation in Kaabong and Kotido districts [26]. Further, by affecting herder movements and reducing milk yields, tsetse invasion is likely to worsen the food Sustainability 2020, 12, 1599 19 of 26 security situation in the sub-region, particularly during the dry season. This situation could be more acute in Turkana, Kenya, whose food security situation is often more acute than that of Karamoja [45]. Both areas are renowned food insecurity hotspots in Kenya and Uganda [70,71]. A particular disruption to food security in the sub-region includes its effect on women, who are the major cultivators of food in the sub-region. Tsetse invasion on grazing grounds, perceived as having good pasture species, and the limit on adequate grazing cycles in these locations could trigger an ecological imbalance leading to bush encroachment that will affect the long-term availability of pasture feed resources. According to [38], bush encroachment in Karamoja is increasing. It has previously been observed that when bush encroachment occurs, high-quality grass species often disappear, leaving behind hardy and less palatable grasses. Further, bush encroachment could occur due to the removal of keystone species and/or the removal of grasses that often help with maintaining the balance of tree-grass interactions [72,73].

6.2. Socioecological Effects of Tsetse Invasion in Karamoja The resilience results from the study area have revealed a generally positive resilience estimated at 20.5%. This positive resilience outcome is unique in that it is a departure from some analyses of resilience that have shown a negative resilience pattern among pastoral communities [74]. A positive resilience pattern observed in the study area could be attributed to the deployment of paravets, access to extension services, diversified economic activities including crop production, livestock off-take, access to veterinary drugs and access to information including early warning systems. In the pastoral and agro-pastoral Somali, Afar and Oromia areas of Ethiopia, households with better access to veterinary services achieve better resilience. Community animal health workers (CAHWs) are critical players facilitating the delivery of much-needed livestock health services, including advisory services. These services help pastoral and agro-pastoral households maintain healthy herds [75]. Furthermore, institutional setup, including improving irrigation facilities and skills and expanding the participation of pastoral households in irrigation farming, increases pastoral resilience [76]. Additionally, creating opportunities for non-farming income and improving access to credit, integration into markets, health and veterinary services expands pastoral resilience. These practices, in addition to ethno-veterinary practices, positively improve household resilience to shocks [77]. The components as discussed by Fent et al. and Muricho et al. fall within the broad spectra of absorptive, adaptive and transformative capacities necessary to influence resilience [76,77]. However, it is also critical to note that there is a considerable narrative on the disruptiveness that vectors cause in pastoral and agro-pastoral communities. These vectors and other associated conditions predispose such communities to further weakened resilience. As seen in the resilience index results, the overall minimum index indicated a negative resilience situation. This is a revelation that there are households whose resilience has generally been affected by the tsetse invasion. Poor communities are often not resilient in the face of different hardships and shocks that erode their adaptive capacity [78]. This pattern has particularly been observed among many pastoral groups in East Africa, such as Boarana [79] and Afar of Ethiopia [80]. To this end, most of the food insecurity in pastoral communities is a result of weak resilience to shocks [81]. Vectors such as tsetse have also been recognised as causing instability within complex adaptive systems. Thus, they affect the resilience of such systems by interfering with their adaptive cycle [17]. For example, in parts of northern Uganda, the spread of sleeping sickness associated with tsetse is linked to restocking efforts after decades of military conflict [82,83]. Due to the agistments associated with pastoral communities, including Karamoja, they may be gifting their kinsmen with already infected livestock. Such practices lead to heterogeneous risks of transmission in both space and time. It is thus not surprising to observe limited socioecological resilience although the overall resilience index was positive [84]. Sustainability 2020, 12, 1599 20 of 26

6.3. Implications of the Assessment on Socioecological Practices of Tsetse Control This study assessed tsetse prevalence and its effect on socioecological resilience in the pastoral Karamoja sub-region. We found complex and multiple dimensions of socioecological interactions in the study region. These interactions included: i) livestock-wildlife interaction; ii) wildlife-human-livestock interaction; iii) agricultural land expansion into wildlife grazing locations; iv) cross-country human-livestock-wildlife movements; v) intensified water resource development for livestock production development without due consideration for wildlife grazing and water needs; and vi) seasonally oriented wildlife-livestock movements taking advantage of heterogeneously distributed water and pasture resources in space and time. These observations highlight the need for the application of transdisciplinary, interdisciplinary and system dimensions in planning, implementing and managing tsetse control interventions in the area. The application of multi-disciplinary approaches for addressing socioecological challenges is required [85]. By adopting transdisciplinary, interdisciplinary and system dimensions, we believe that the actors in the intervention sphere will be better poised to co-create integrative solutions. Collaborative co-creation of solutions brings stakeholders together to share knowledge and form networks that are vital in guiding intervention strategies and practices that increase socioecological resilience [86]. Furthermore, through these approaches, practitioners are in a better position to engage policymakers, decision leaders and local communities to adopt scale- and transboundary-level interventions [87,88]. This process is particularly important because resilience and sustainable development are possible only within a context where the processes affecting and the processes affected by the health of ecosystems and human societies are holistically considered [87,89]. In this regard, it will not be necessary to separate planning for wildlife grazing and watering needs from that of cattle. Further, the tsetse eradication effort in cattle populations was separated from that of wildlife in national parks and game reserves. Yet, these areas are believed to be source points. Accordingly, integrative and cross-sector approaches to tsetse control are critical [90]. The tsetse distribution in Karamoja bears a spatiotemporal distribution characteristic. Both entomological surveys and participatory assessments revealed a higher prevalence in northern Karamoja than in central and southern Karamoja. Of particular concern is the spatial distribution and spread southwards and the emerging south-to-west Karamoja pattern. These patterns call for the establishment of a permanent, robust and routine tsetse monitoring system in Karamoja. This robust system must be inclusive, taking into consideration local knowledge and information systems. It is also critical to develop approaches that can enable the timely transfer of tsetse-based information for prevalence and risk quantification. This speed is important in facilitating timely response [91]. Actors in tsetse control must pay close attention to seasonal dimensions because dispersal is associated with seasonal migrations of hosts, especially wildlife. High fly abundance and mobility during dry seasons have been reported elsewhere in Tanzania and South Sudan [92–94]. Considering the flux in wildlife and livestock-cattle mobility into Uganda during the dry season, it is vital that monitoring be conducted to ascertain tsetse dynamics associated with this mobility. Tsetse is a vector of transboundary importance whose control calls for coordinated multinational actions [95,96]. Temporal migratory route monitoring could perhaps aid in providing critical information regarding the progression of tsetse abundance and therefore help with the identification of risk hotspots, particularly because these routes are transformation foci.

7. Conclusions Tsetse invasion in the Karamoja sub-region is an important entomological and epidemiological concern. Although the socioecological resilience in the study is positive, the threat of tsetse in the sub-region is growing. A higher prevalence is observed in two districts in northern Karamoja that are in close proximity to Kidepo Valley National Park, and there were some reported in southern Karamoja. This result raises an urgent need for immediate intervention before a wider and total sprawl of tsetse occurs in the sub-region. Intervention is required to control the spread, particularly to minimise the Sustainability 2020, 12, 1599 21 of 26 potential of the tsetse belts from greater northern and eastern Uganda (Teso sub-region) merging with that from western Kenya. Tsetse control approaches that need to be implemented should be holistic and recognise the complex and multiple dimensions of socioecological interactions in the study region. Furthermore, given the perceived transboundary nature of tsetse prevalence and distribution in Karamoja, coordinated multination intervention is required. It is only through these dimensions that resilience in the sub-region can be sustained and further increased.

Author Contributions: Cconceptualization, A.E., J.O., and B.B.; methodology, A.E., J.O., B.B. and J.J.N.; software, J.O. and J.P.M.; validation, A.E., J.O., J.P.M. and B.B.; formal analysis, A.E., J.O. and J.P.M.; investigation, A.E. and J.O.; resources, A.E.; data curation, J.O., J.P.M. and A.S.; writing—original draft preparation, A.E.; writing—review and editing, A.E., J.O., B.B., and A.S.; visualization, A.E., J.O., B.B. and A.S.; supervision, J.J.N.; project administration, A.E.; funding acquisition, A.E. and J.J.N. All authors have read and agreed to the published version of the manuscript. Funding: This research was funded by the Carnegie Corporation of New York through the Nurturing Emerging Research Leaders Project at Makerere University (Post Doc 2017). The research also received tsetse data from the Ministry of Agriculture Animal Industry and Fisheries (MAAIF) through the tsetse surveillance programme, and the publication of this article was funded by the Regional Universities from for Capacity Building in Agriculture (RUFORUM) through the Transforming African Agricultural Universities to meaningfully contribute to Africa’s growth and development (TAGDev) supported by the Mastercard Foundation. Conﬂicts of Interest: The authors declare no conﬂict of interest.

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