Seasonal variation in the ranging behavior of elephants in the Laikipia‐Samburu ecosystem

Abstract

African savanna elephants are a highly mobile species that ranges widely across the diversity of ecosystems they inhabit. In xeric environments, elephant movement patterns are largely dictated by the availability of water and suitable forage resources, which can drive strong seasonal changes in their movement behavior. In this study, we analyzed a unique movement dataset from 43 collared elephants, collected over a period of 10 years, to assess the degree to which seasonal changes influences home range size of elephants in the semi‐arid, Laikipia‐Samburu ecosystem of northern Kenya. Auto‐correlated Kernel Density Estimation (AKDE) was used to estimate elephants' seasonal home range size. For each individual elephant, we also calculated seasonal home range shifts, as the distance between wet season home range centroids and dry season home range centroids. Core areas (50% AKDE isopleths) of all individual elephants ranged from 3 to 1743 km² whereas total home range sizes (the 95% AKDE isopleths) ranged between 15 and 10,677 km². Core areas and home range sizes were 67% and 61% larger, respectively, during the wet season than during the dry season. On average, the core area centroids for all elephants were 17 km away from the nearest river (range 0.2–150.3 km). Females had their core areas closer to the river than males (13.5 vs. 27.5 km). Females differed from males in their response to seasonal variation. Specifically, females tended to occupy areas farther from the river during the wet season, while males occupied areas further from the river during the dry season. Our study highlights how elephants adjust their space use seasonally, which can be incorporated into conservation area planning in the face of increased uncertainty in rainfall patterns due to climate change.

Full text

Ecology and Evolution. 2024;14:e70198.  
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https://doi.org/10.1002/ece3.70198
www.ecolevol.org
Received:17March2024
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Revised:30July2024
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Accepted:1August2024
DOI: 10.1002/ece 3.70198
RESEARCH ARTICLE
Seasonal variation in the ranging behavior of elephants in the
Laikipia- Samburu ecosystem
Loise W. Kuria1 | Duncan M. Kimuyu1,2 | Mwangi J. Kinyanjui1 |
George Wittemyer3 | Festus W. Ihwagi3
ThisisanopenaccessarticleunderthetermsoftheCreativeCommonsAttributionLicense,whichpermitsuse,distributionandreproductioninanymedium,
providedtheoriginalworkisproperlycited.
©2024TheAuthor(s).Eco logy an d EvolutionpublishedbyJohnWiley&SonsLtd.
1SchoolofNaturalResourcesand
EnvironmentalStudies,Karatina
University,Karatina,Kenya
2MpalaResearchCenterandWildlife
Foundation,Nanyuki,Kenya
3SavetheElephants,Nairobi,Kenya
Correspondence
LoiseW.Kuria,SchoolofNatural
ResourcesandEnvironmentalStudies,
KaratinaUniversity,P.O.Box1957,10101
Karatina,Kenya.
Email:loisewangui3@gmail.com
Funding information
CenterforMountainandClimateChange;
SavetheElephants
Abstract
Africansavannaelephantsareahighlymobilespeciesthatrangeswidelyacrossthe
diversityofecosystemstheyinhabit.Inxericenvironments,elephantmovementpat-
ternsare largelydictatedbythe availability ofwaterandsuitableforageresources,
whichcan drivestrongseasonal changesin theirmovement behavior.Inthisstudy,
weanalyzedauniquemovementdatasetfrom43collaredelephants,collectedovera
periodof10 years,toassessthedegreetowhichseasonalchangesinfluenceshome
range size of elephants in the semi-arid, Laikipia-Samburu ecosystem of northern
Kenya.Auto-correlatedKernel Density Estimation(AKDE)wasused to estimateel-
ephants' seasonalhome rangesize.Foreachindividualelephant,we alsocalculated
seasonal home range shifts, asthe distancebetween wet season home range cen-
troidsanddryseasonhomerangecentroids.Coreareas(50%AKDEisopleths)ofall
individualelephantsrangedfrom3to1743 km2whereastotalhomerangesizes(the
95%AKDEisopleths)rangedbetween15and10,677 km2.Coreareasandhomerange
sizeswere67%and61%larger,respectively,duringthe wetseasonthanduringthe
dry season. On average, thecorearea centroids forallelephants were17 kmaway
fromthe nearest river (range0.2–150.3 km). Femaleshad theircore areascloserto
theriverthanmales(13.5vs.27.5 km).Femalesdifferedfrommalesintheirresponse
toseasonal variation. Specifically,females tended to occupyareas fartherfromthe
riverduringthewetseason,whilemalesoccupiedareasfurtherfromtheriverduring
thedryseason.Ourstudyhighlightshowelephantsadjusttheirspaceuseseasonally,
whichcan beincorporatedintoconservation areaplanningintheface of increased
uncertaintyinrainfallpatternsduetoclimatechange.
KEYWORDS
coreareas,habitatuse,homerangeshift,homerangesize,Loxodonta africana,mega-
herbivores
TAXONOMY CLASSIFICATION
Behaviouralecology

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1 | INTRODUCTIO N
Elephantsact as ecosystemengineersbyinfluencing the structure
and compl exity of the ha bitats in wh ich they occur (Ge bremeskel
Haile et al., 2019),resultinginamyriadofcascadingeffectsonother
trophiclevels(Calengeetal.,2002;Smallie&O'Connor,2000).The
scaleofthiseffectis relativelybroadbecauseelephantsaremega-
herbivoreswithlargehomerangesandhighmobility,allowingthem
to cover large distance s across the landsc ape as they search for
forage re sources (Bolla & H ovorka, 2012). As mi xed feeders wi th
diversediets,elephantscanexhibitgreatplasticityinrangingbehav-
ior(Bastille-Rousseau & Wittemyer,2019; Ortega & Eggert,2004;
Sukumar,2003).Suchchangesmaybedrivenbyseasonalvariation
intheavailabilityofforageandwaterresources(Roeveretal.,2012;
Sukumar,2003; Valls-Fox et al ., 2018; Witte myer et al., 2017 ) as
well as physio logical and beh avioral diffe rences among indi vidual
elephants(Fortinet al.,2022; Moss et al., 2 011). Homerangeslink
themovementofanimalstothedistributionoftheresourcesneces-
sary forsurvival andreproduction.Home range size, location,and
shapemaychangedependingonthestateoftheindividualandthe
conditionsoftheexternalenvironment (Börgeretal., 2006, 2008;
Kenwardetal.,2001).Thereisgreatinterestinexaminingspatialand
temporalvariationinelephant-rangingbehavior,butthelackoflong-
termlongitudinaldatasetsseriouslyconstrainssuchstudies.
Seasonal pulses in the availability and distribution of forage and
water can have enormous impacts on elephants' ranging behavior
(Bastille-Rousseauetal.,2020).Generally,forageandwatertendtobe
morewidelyavailableduringwetthandryseasons.Forexample,rain-
fallcreatestemporarypoolsof water that maybeutilizedbyanimals,
negatingtheneed to rely onmorepermanentwater sourcessuch as
riversandsprings.Similarly,anincreaseinperceivedforageavailabil-
ityduringthewetseasonmayallowanimalstoutilizesomeareasthat
aregenerallyavoided during the dryseason.Inresponsetoseasonal
fluctuations inresourceavailability,elephantsmay eithermigrateout
ofanareaorexhibitseasonalrangefidelity.Migration,definedasthe
repeatedseasonalmovementbetween two non-overlappingregions
(Dingle & Drake, 2007),allowselephants toescapesevereseasonal
declinein resources.Elephantsmayemployanextensivecontinuum
ofmovementbehaviorsthatincludesmigration,highlyvariablehome
ranges, or resident behavior (Bartlam-Brooks et al., 2011; Purdon
et al., 2018).Seasonalrangefidelit yoccurswhenanindividualchanges
thesizeofitsrangewhilemaintainingthecoreareaofhabitatuse,con-
sequentlypresenting relativelyhigh range fidelity but withachange
inthedegreeofrangeoverlap(Damuth,1981 ; Lindstedt et al., 1986).
Sitefidelityisattributabletopredictableaccesstoresources(Burton-
Robertsetal.,2022).
Maleandfemaleelephantsrangingbehaviorvariesacrossspace
andtime due to theirsocial organization(Fortinetal., 2022; Moss
et al., 2011;Wittemyer,Douglas-Hamilton,etal.,2005)aswellasthe
differenceinforagingstrategy(Duffyetal.,2011;Kiokoetal.,2020;
Lee et al., 2 011; Woo lley et al., 2009). Male e lephants may have
largerhomerangesthanfemalesastheydispersetounfamiliarhab-
itatstoseekfoodandmates(duToit&Moe,2014; Lee et al., 2011).
Unlike maleelephants,female African elephants live in matrilineal
families consisting of individuals of different ages. Family-ranging
behaviormaybeconstrainedbycalvesthatmaynotbeabletomove
fastandfarfromwatersources(Ngene,2010).Malee lepha nts ,how-
ever,moveandforagealoneorinbachelorherdswithoutcalvesthat
wouldlimittheirmovement(Ngene,2010).
ElephantsintheLaikipia-Samburuecosystemareknowntomove
overlarge areas over time because theecosystem is stronglysea-
sonal(Bastille-Rousseauetal.,2020;Wittemyeretal.,2008).Inlight
ofthis,therewasacriticalneedtodocumentandcompareelephant
movementsovertheselargeareasasonemightexpecttheirranging
behaviortovaryseasonally.Tounderstandvariationsindistribution
patternsandspaceuseofelephantsinthisresource-limitingregion,
rigorous fielddata modeled withecologically meaningfulpredictor
variablesisneeded. Understanding andpredicting patterns of ani-
malspaceuse isparticularlyimportant for heavilymanagedanimal
populationsandforspeciesthatmayseverelyaffectecologicalpro-
cesses(Reineckeetal.,2014).
Togainadeeperunderstandingofdriversofelephantspaceuse,
weanalyzedthemovementdatasetfrom43collaredelephants,col-
lectedover10 yearsintheLaikipia-Samburuecosysteminnorthern
Kenya.Ouranalysisaimedtoassess(i)thevariationinhomerange
sizesacrossseasonsand(ii)theextentofseasonalhomerangeshifts
ofelephantsinrelationtotheirproximitytotheriver.Wepredicted
thatelephantswouldhavealargerhomerangeduringthewetsea-
son to capi talize on disper sed resources. We exp ected that male
elephantswouldhavelarger homerangesizesthan femalesdueto
theirmovementinsearchofmatesanddispersedresources.Incon-
trast,becausefemaleelephantsliveinmatrilinealfamilieswithde-
pendent calves,theywould beconstrictedto areasnearriversand
watersourcesandexpectedtohavesmallerrangesthanmales.We
expectedelephantstomovefromonelocationtoanotherdepend-
ingonseasonalresources(primarywater)and,therefore,predicted
that the elephants' home range would shift to areas closer to the
EwasoNg'iroriver,whichisakeyperennialwaterresource(Barkham
&Rainy,1976)duringthedryseasonsintheecosystem.Wediscuss
theimplicationsofourresultsforelephantconservation.
2 | METHODS
2.1 | Study area
Thisstudywascondu cte di nt heLai kipia-Sambu ruecosy stemofKe ny a
using datafor 2001and 2021 (Figure 1).The ecosystem isbounded
by coordina tes 0.2° S to 1. 5° Na nd 36.2° E to 3 8° E. The ecos ystem
isdefined by the geographicextent ofthe Ewaso NyiroRiverand its
tributaries,encompassingapproximately36,790 km2(Thouless,1995)
and the his torical elepha nt migration range (G eorgiadis, 2011). Th e
study areais semi-arid withawide range of habitats linked withthe
elevation and climatic gradients that characterize the region: from
cool, wet highlands in t he south to hot, dr y lowlands in the nor th
(Georgiadis, 2011). The rainfall is highly variable and bimodal, with

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peaksinMayandNovemberandayearlyrangefrom<400 mminthe
northtoamaximumof600 mminthesouth(Barkham&Rainy,1976 ;
Ihwagi et al., 2012).Theterrainiscomprisedofexpansiveplainsinter-
rupted by rugged terrain and isolated hills. Wildlife shares the land-
scape freely with the predominantly pastoral communities (Ihwagi
et al., 2015).TheconfirmedLaikipia-Samburuelephantrangeencom-
passes six major land use types: community conservancies, private
ranches, communal pastoral areas, state-protected forest reserves,
settlementsmainlyundersedentarysubsistenceproduction,andthe
nationalreservesthatare either owned by individuals, government,
orcommunities.Theprivate,government,andcommunitylandscom-
prise30%,11%,and59%ofthelandscape,respectively.TheLaikipia–
Samburuelephant(Loxodonta africana)populati onisthesecon dlargest
inKenya,withapproximately7347individuals,primarilyrelyingonthe
rangeoutsideofgovernmentallyprotectedareas(Litorohetal.,2010).
2.2 | Data collection
2.2.1 | Elephantmovementbehavior
ExistingGPS-satellitedata from43 collared elephants(14malesand
29females)thatuse theLaikipia-Samburu ecosystem wereanalyzed
(Figure 2).MaleAfricanelephantsaresolitary,whereasfemaleAfrican
elephantsliveinmatrilinealfamiliesthatcanhaveasmanyas36mem-
bers(Wittemyer,2001;Wittemyer,Daballen,etal.,2005;Wittemyer,
Douglas-Hamilton, et al., 2005). Only one individual elephant was
tracked for family groups consisting of multiple females. Save the
ElephantscarriedoutelephantcollaringoperationswithKenyaWildlife
Serviceusingstandardoperatingprocedures.Thesecollarscamefrom
eitherAfricanWildlifeTrackingfromSouthAfrica,SavannahTracking
from Kenya,or Followit AB from Sweden. Costinfluences the collar
used, theenvironmentthey aredeployed in,and technical specifica-
tions.Thecollarsrecordedthelocationofeachelephantatasetof1-h
intervals.Elephanttrackingdatawereretrievedfromacentralizedda-
tabaseusingcustomizedsoftwarethatemploysadatafiltertoremove
erroneousGPSfixesbasedonamaximumrateoftravelof7 km/h(Wall
et al., 2014). The dataprojections were on the UniversalTransverse
Mercator (UTM) WGS-84 reference system zone 37 N. Data were
stored in ESRI Geodatabase (ArcGIS version 10). Data collected be-
tween2001and2021wereusedforthisstudy.All43elephantsthat
wereselectedforthisstudyhadbeentrackedatvariousintervalsdur-
ing this pe riod, but each e lephant had at le ast 1 year of continu ous
trackingdatawithatleast10 monthsoftrackingdatainanygivenyear.
Theaveragetrackingperiodforall43individualswas4.3 years(range
1–9)years,withthemosttrackedbetween2015and2018(Table S1).
FIGURE 1 ThelocationoftheLaikipia-SamburuecosysteminNorthernKenya.

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2.2.2 | Seasonsdatadelineation
Ecoscope(Copyright2022,WildlifeDynamics,h t t p s : / / e c o s c o p e . i o /  )
anopen-sourcePythonlibraryusedforenvironmentalandconser-
vation dataanalyseswasused to identifytransitions between wet
anddryseasons.Usingstd_ndvi_valsfunctionintheEcoscopetool,
weextractedstandardizedNDVI(NormalizedDifferenceVegetation
Index)valueswithinthestudyarea.NDVIvaluesrangebetween−1
and1;withvaluescloserto1representinghigherproductivity(i.e.,
wetseasons)andvaluescloserto−1representinglowerproductiv-
ity(i.e.,dryseasons).NDVIreflectsactualchangesinvegetationat-
tributes,therefore,itmaybeconsideredabetterproxyforseasonal
changes,thanrainfalldata.TheNDVIvalueswereextractedforthe
wholestudyperiod(2001–2021).Theval_cutsfunctionwasusedto
calculatetheseasonaltransitionpoint,whichisthepointwherethe
NDVIvalueschangefromincreasingtodecreasingorviceversa.The
seasonalwindowsfunctionwasthenusedtodeterminetheseasonal
time windows. The output was a data frame containing each sea-
son's start and end dates, along with a label foreach season.The
seasonaltimewindowsdataframewasexportedtoaCSVfile.The
season'sdatawereusedtodeterminewhetheranelephantlocation
fixwasduringthewetordryseasons,hencesplittingtheelephant
movementdata intotwo (Figure 3).Thiswasachievedbyassigning
anelephantlocationGPSfixtimetoeitherwetordrydependingon
the season.
2.3 | Data analyses
TheAutocorrelatedKernelDensityEstimation(AKDE)methodwas
usedtoestimatetheindividualelephants'corearea(areasbounded
by50%isopleths)andtotalhomerangesize(areasboundedby95%
isopleths)forevery wet anddry instance in a year.Weconsidered
anyperiod lasting morethan80 days of continuous wet or drype-
riodsasa‘significant’wetordryinstance.Wetseasonsinthearea
typicallylast80 days,andtheareaexperiencestwowetseasons in
ayear.Therefore,anaverageyearwouldhavetwowetandtwodry
instances. AKDE was considered appropriate because it was de-
signed tobestatisticallyefficient whendealing withthecomplexi-
tiesand biasesofmodern movementdata,suchasautocorrelation
andmissingdata (Silvaetal., 2022). Semi-variancefunctions (SVF)
werecalculated for eachelephantto assessforrangeresidencyin
alldatasets(Calabreseetal.,2016).Rangeresidencyisanassump-
tionoftheAKDEapproach and isnecessary forthemethodto de-
fineahome rangeaccurately.Theelephantsusedinthis studyare
residents of the Samburu Laikipia ecosystem. We calculated the
WeightedAKDEc(or wAKDEc)tocater forwhen the devicehad a
malfunction, leading toGPS fixesshiftingfrom one fixperhour to
onefixper‘n’hours.Weestimatedtheseasonalhomerangeshiftin
relationtotheirproximitytoriversbycalculatingthedisplacement
ofthecentroidsofindividualelephantcoreareasduringthewetand
dryseasons.WeusedArcGIS(Esri,2011)tocomputethecorearea
FIGURE 2 Rangeextentofthe43
elephants(eachrepresentedbyarandom
color)trackedintheLaikipia-Samburu
ecosystem.

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centroids,whichwere calculated using thecentreofgravity algo-
rithm (Ba rtlett e t al., 2016). Th e centroids' coor dinates were the n
extractedandusedtodeterminetheshiftintheindividualelephant
core areas a cross seasons. A rcGIS was also u sed to calculate t he
distancefromthecentroidstotheriver.Thisanalysis allowedusto
quantifyandcomparethemovementpatternsofelephantsbetween
thedryandwetseasons,providinginsightsintohowtheircoreareas
change in response to seasonal changes.
Generalized line ar mixed models (GLMMs) with Gamma error
struc ture and log link f unction were t hen used to test t he effect
ofseason and sex ofindividual elephants on boththe total home
rangeandcoreareas.AllourmodelsincludedelephantIDasaran-
domeffecttoaccountforrepeatedsamplingofindividuals'(multiple
tracki ng instances). To account for di fferences in sampling ef fort
(differences in thenumber of daysper tracking instance), we have
included sampling days per instance as an offset in our models.
Wefittedall ourmodelsusing the generalized linear mixed effect
model ‘glm er’ function of t he ‘lme4’ package ( Bates, 2010; B ates
et al., 2015). Generalized linearmodel(GLM) wasused to test the
effect of season and sex on seasonal home range shift in relation
to their pr oximity to river Ewaso N gi'ro. Similarly, to accoun t for
differencesinsamplingeffort,wehave includedsamplingdaysper
instanceasanoffsetin our models. Akaike's Information Criterion
(AIC) Weighting was used for model averaging and to evaluate
suppor t for competing ra nging behavior mo dels. Model sele ction
andaveragingwereperformedusingtheRpackageMuMInversion
4.2.0(Barton,2016).AllstatisticalanalyseswereperformedusingR
(RCoreTeam,2021),andalltestswerecarriedoutatasignificance
levelof0.05.
3 | RESULTS
Atotal of 771,919hourlyGPS fixes were collectedduringthe dry
seasonand990,772duringthewetseason.Thesedatarepresented
340individual seasonsacross2001–2021.Allindividualelephants'
coreareasrangedbetween3and1743 km2,whereasthetotalhome
range ra nged betwee n 15 and 10,677 km2. Base d on AICc value s,
thebestranking models included season only,and season and sex
asthebestpredictorsoftotalhomerangesize,aswellascoreareas
(Table 1).Weconsideredthelatermodel(includingseasonandsex)
becauseitalignsbetterwith theoreticalexpectations.Onaverage,
the core (50 % isopeth) home r ange size was 67% (101 km2) larger
during wetthan thedry season ( χ2 = 21.06,p < .001).Similarly,the
total home range size was 61% (521 km2) larger during wet than
thedryseason( χ2 = 15.45, p < .001).Sexdidnotinfluence the indi-
vidualelephant'scoreareaandhomerangesize(χ2 = 0.63,p = .429;
χ2 = 1.07,p = .300respectively,Figure 4).
FIGURE 3 Rangeextentofelephants
duringthewetanddryseasonsfromGPS
trackingdatacollectedfrom43individuals
intheLaikipia-Samburuecosystem.

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Onaverage,thecoreareacentroidsforallelephantswere17 km
awayfromthenearestriver(range0.2–150.3 km).Femaleshadtheir
coreareasclosertotheriverthanmales(13.5vs.27.5 km).Basedon
AICcvalues,thebestrankingmodelincludedseasonandsexasthe
bestpredictorofhomerangeshift(Table 1).Femalesdifferedfrom
males in their response to seasonal variation ( χ2 = 6.43, p = .011).
Females tended to occupyareasfarther from the riverduringthe
wetseason,whilemalesoccupiedareasfurtherfromtheriverduring
thedryseason(Figure 5).
4 | DISCUSSION
Understandingelephant rangingbehaviorinaridenvironmentscan
provideinsightintoecologicalconstraintsandrequirementsforthe
species.Wehadpredicted thatelephantswould havelarger home
rangesandcoreareasduringthewets eas ontha nduri ngthedrysea -
son,andthat maleswouldhaveoverall largerrangesthanfemales.
Wefoundevidenceinsupport of ourfirst prediction; but notthat
males have la rger home rang es than female s. Our predic tion that
elephantswouldshifttheirhomerangebasedonseasonswassup-
ported.Lastly,ourpredictionthat elephantswouldshifttheircore
areastooccupyareasproximatetotheriver(perennialwatersource)
duringthedryseasonwaspartiallysupported.
The differencesbetweenwet anddry season coreareas and
the home range size observedin thisstudycan be attributed to
increased dispersion of water and forage resources during the
wetseason, allowing elephants to utilize sectionsof the habitat
thatarenormallyavoidedduringthedryseason.Inourstudy,the
river Ewaso Ng 'iro is the only w ater source duri ng the dry se a-
son.However,numerousephemeralwaterpoolsbecomeavailable
duringthewetseason(Ihwagietal.,2010)resultinginhigh-quality
forageresourcesbecomingmoreavailableawayfromtheriver,im-
plyingt hatel ephant scantakelongforaysawayfromtheriver. Our
findingsconcurwith several other studies thathave shownthat
elephantstendtoreducetheirhomerangesduringthedryseason
byconcentrating their foraging activitiesin areasclose towater
(ChamaillÉ-Jammesetal.,2007;deBeeretal.,2006; Leggett , 2006;
Osborn&Parker,2003;Owen-Smith,2004;Redfernetal.,2003;
Smit et al. , 2007). The ef fect of seasona l shrinking and ex pan-
sionofelephanthomerange warrants subsequentinvestigation.
One possi bility is that re duction in hom e range size during th e
dryseasonmayconcentrateelephantbrowsingdamagetosmall-
isolated patches, significantly impacting vegetation. However,
TAB LE 1 Candidatemodelsofthetemporalvariationofthe
elephantrangingbehaviorinrelationtotheseasonintheLaikipia-
Samburuecosystem,Kenya,2001–2021.
Model AICc Delta
Totalhomerangesize Season 5 089.15 0
Sex + season 5090.14 0.99
Season * sex 5092.17 3 .027
15101 .97 12.819
Sex 5102.95 13.802
Coreareasize Season 3918.16 0
Sex + season 39 19. 59 1.433
Season * sex 3921.45 3.291
13936.21 18.055
Sex 39 3 7. 6 3 19. 476
Coreareashift Sex * season 2195.84 0
Sex 2197. 95 2.119
Sex + season 2200 4.168
12226.39 30.55
Season 2227.12 31.29
Note:Allmodelsincludedelephantidentificationasarandomfactorand
thenumberofdayspertrackinginstanceasanoffset.
FIGURE 4 Seasonaldifferencesin
predicted(a)totalhomerangeand(b)core
areasofelephantsinLaikipia-Samburu
ecosystem.Predictedhomerangevalues
areobtainedfromourbest-candidate
model.Errorbarsrepresentstandard
errors(±oneSE).

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such patch es may recover from h eavy browsin g once elephant s
disperseduringthewetseason.
Theobservedincreaseinhome rangeandcoreareasizeduring
thewetseasonwasconsistentforbothmaleandfemaleelephants.
These f indings concu r with an earlie r study that sh owed that the
home ran ge of sexually imm ature males, n on-musth , sexually ma-
turemales,andfemalesissimilar(Whitehouse&Schoeman,2003).
However,sexually active males have been reported to have more
extensivehomerangesthanfemales(Barnes,198 3; Leggett, 2006;
Poole&Moss,1989;Tayloretal.,2020;Viljoen,1989;Whitehouse&
Schoeman,2003).Inourstudy,wedidnotaccountforthereproduc-
tivestatusof individual elephants. However,we expect suchsimi-
laritiescould bemootedbecause theSamburu-Laikipia ecosystem
experiences longdry periods,and bothmales'and females' move-
mentsarelimitedtoareasnearwaterpointsduringthedryseason.
Thisstudyshowedthatonaverage,femalestendedtooccupy
areas further fromthe river during the wet season,while males
occupiedareasfurtherfromtheriverduringthedryseason.This
isbecauseduringthedryseasonstheresourcesavailable,thatis,
waterand food,become limited.Therefore, for the freeroaming
adultmaleelephantsmustmoveawayfromtheriverinsearchof
food. In t he wet season, ma le elephant s prefer to stay cl ose to
theriver.Thisisbecauseoftheavailabilitywaterandhighquality
forage near theriverduringthe wet season withthe consistent
needtodrinkwateraselephantrequiredrinkingwatereveryone
or2 days(Douglas-Hamilton,1973).Shannon concurred withthe
resultsasshenotedthatelephantsareattractedtohabitatsnear
rivers anddams,which notonly provide drinking water but also
anabundanceof forage(Shannonet al., 2006).Astudydone by
Leggettfoundsimilarresultswithdistinctseasonalmovementof
collared elephantsbetween theirwetand dryseasoncore areas
(Leggett, 2006). However, Viljoen reported that the elephants
of the northwestrestrict themselves to seasonalranges within
their individual home ranges, irrespective of higher rainfall or
riverfloodsinareasadjacenttotheirhomeranges(Viljoen,1989).
During thedryseason,female elephants are morelikely toshift
their core a rea to occupy area s near the river, wher e water re-
sources are moreavailable becausefemaleelephants'movement
isrestrictedbydependentcalves. Thisexplainswhyfemaleshad
their core areascloser to theriverthan males.This seasonal be-
haviorunderscorestheimportanceofriversandwatersourcesfor
the Samburu-Laikipia ecosystem elephants' sur vival and move-
mentpatterns.
TheAKDEhomerangesobservedforGPS-collaredelephants
intheLaikipia-Samburuecosystemrangedfrom22to10,677 km2.
Thehomerangesizesofthisstudyarecomparabletohomeranges
reportedinotherstudies(Table 2).Themaximumareaofthetotal
home ran ge for the female r eported in t his study is 10,677 k m2
while the t otal home rang e for males in this s tudy is 6921 km2.
The large home ranges are comparable to the desert-dwelling
elephan ts of Mali with an ap proximate home ra nge of 11,500–
23,980 km2 (Wall et al., 2013) as they had larger home ranges
thanpreviouslyreportedforsavannaelephants.Thelargerhome
ranges of elephants in northwest Namibia probably reflect the
typeandqualityofavailablevegetation(Viljoen,1989).Thelarge
homerange sizesreportedinthis studycould be because of the
unpredictableavailabilityofwater.Secondly,foodavailabilitymay
belimited,thusforcinganimalstomoveoverawiderareaand/or
th e s p atiald i s tributio n ofhab i t a tt y p e sofh i ghernu t r i t ionalq u a l it y
mayresultinawiderareabeing traversed. Thehomerangesizes
determinedbyradio-trackingcollarswereusuallylargerthanthe
onesrevealedbyvisualidentification(Leuthold,1977 ).Thesmall
homerange sizeofsomeelephants couldbe because ofphysical
barrie rs therefore con fining them to a sm aller range of habi tat.
Thehomerangesizesof14to52 km2fortheAfricanelephantat
LakeManyara NationalPark,Tanzania(Douglas-Hamilton,1972)
FIGURE 5 Seasonaldifferencesinpredictedcoreareashift
ofelephantsinLaikipia-Samburuecosystem.Predictedcorearea
valuesareobtainedfromourbestcandidatemodel.Errorbars
representstandarderrors(±oneSE).
TAB LE 2 Autocorrelatedkerneldensityestimation(AKDE)home
rangeestimatesofelephantsfromBukitTigapuluh,Khaudum,
Etosha,Zambezi,andKunene.
Region
Home range size
(km2)References
BukitTigapuluh,Indonesia 275–518 0 Moßbrucker
etal.(2016))
KhaudumNationalPark,
Namibia
3000–12,000 Benitez
etal.(2022)
EtoshaNationalPark,
Namibia
280–38,000 Benitez
etal.(2022)
Zambezi,Namibia 1400–25,000 Benitez
etal.(2022)
Kuneneregionof
northwesternNamibia
1500–37,000 Benitez
etal.(2022)

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couldbeimposedbythedensevegetationinthepark,likelypro-
vidingahighdensityofforageforelephants.
Sambur u-Laikipia e cosystem is amon g the many ecosys tems
worldwidethat are facing increased fragmentation. Factors con-
tributingtosuchfragmentationincludeintensiveagriculture,over-
grazing , conversion of large a reas to human sett lements, ill egal
logging,urbanization,etc.Suchfragmentationwilllikelyaffectthe
distributionpatterns ofelephantsandotherherbivoresin these
systems . The future clima tic projections s uggest an increa se in
droughtfrequencyandseverityintheregion(GebremeskelHaile
et al., 2019).Such changesmayhavesevere implicationsforele-
phantsaswellasotherbiodiversity.Elephants,amongotherspe-
cies,co ntra c tth eirran getor emainn earperm ane ntw ate rso urces.
Spendingmoretimeinthevicinityofwaterintensifiesthebrows-
ingpressure inthoseregionsandmaycauseremarkable changes
invegetationcoverandcomposition.IntheLaikipia-Samburueco-
system,EwasoNg'iroRivertendstoretainwaterfor aprolonged
perioddur ingdr ought ,at trac tinga nimal sf romdi fferentr egion sto
thearea.Conservationeffortsshouldfocusonmaintaininganet-
workof conservationareasthatislargeenoughtoaccommodate
seasonalexpansion inhomerangeor migrationofanimalswhile
stillmaintainingaccesstocriticalresourcessuchaswateranddry
seasonforaginghabitats.
In conclusion, the behavioral patterns of elephants in the
Samburu-Laikipia ecosystemare intricately tiedto seasonal varia-
tions.Thewetseasonpromptstheexpansionofhomerangeaswater
and food resources become more abundantacross the landscape.
Contrarytoour predictions,male elephantsdidnotexhibitsignifi-
cantly larger homerangesthan females, indicatingthat females as
well as males demonstrate the use of vast areas. The prolonged
dry pe riods in the La ikipia-Sambu ru ecosyste m facilitate el ephant
home rangeshifts acrossseasons. Thisraisesquestions about po-
tentialtrade-offsbetweenforagingnearwateringpointswithlower-
qualityresourcesandembarkingoncostlyjourneystodistantfood
patches.Given these challenges, theL aikipia-Samburuecosystem
necessitates tailored conservation strategies such as protection
andmanagementof movementcorridors,communityengagement
andeducation,habitatrestorationamongothersthatwouldensure
thelongevityofelephantpopulationswhilesafeguardinglandscape
connectivityand crucialmovementcorridors,addressingbothim-
mediateandlong-termthreats.
AUTHOR CONTRIBUTIONS
Loise W. Kuria: Conceptualization(equal);datacuration(lead);for-
mal analysis (lead); methodology (equal); writing – original draft
(lead); writing – review and editing (equal). Duncan M. Kimuyu:
Conceptualization (equal); data curation (equal); formal analysis
(equal); me thodolog y (equal); writi ng – review and edi ting (equal).
Mwangi J. Kinyanjui:Fundingacquisition(supporting);writing–re-
viewandediting(supporting).George Wittemyer:Writing–review
and editing (equal). Festus W. Ihwagi: Conceptualization (equal);
data curation (equal); formal analysis (equal); funding acquisition
(lead);methodology(equal);writing–reviewandediting(equal).
ACKNOWLEDGEMENTS
Wethankeveryonewhocontributed to this research paperinone
way or anoth er.Wi thout your sup port, this w ork would not have
been accomplished. Wethank Save the Elephants and the Center
for Mountain and Climate Change for providing the financialsup-
porttopursuethisstudy.Tomyfellowcolleagues,AlfredKibungei,
PaulMunene,andRobertAng'ilaIacknowledgeandappreciateyour
support.
CONFLICT OF INTEREST STATEMENT
Thecorrespondingauthorconfirmsonbehalfofallauthorsthatno
involvementsmightraisethequestionofbiasinworkreportedorin
theconclusions,implications,oropinionsstated.
DATA AVAIL AB I LI T Y STATE MEN T
Thedata onhome rangesize and shift acrossseasonsare publicly
available on the Dryad website (h t t p s : / / d o i . o r g / 1 0 . 5 0 6 1 / d r y a d . 
djh9w0w77). GPS positions of elephant distribution are ethically
restrictedasthelocationsmayjeopardizetheanti-poachingopera-
tionsand thusputelephants atgreaterrisk. The dataarehowever
available on written request from the Office of Senior Assistant
DirectorofKenyaWildlifeService.
ORCID
Loise W. Kuria https://orcid.org/0009-0000-4422-7029
George Wittemyer https://orcid.org/0000-0003-1640-5355
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SUPPORTING INFORMATION
Additional supporting information can be found online in the
SupportingInformationsectionattheendofthisarticle.
How to cite this article: Kuria,L.W.,Kimuyu,D.M.,
Kinyanjui,M.J.,Wittemyer,G.,&Ihwagi,F.W.(2024).
Seasonalvariationintherangingbehaviorofelephantsinthe
Laikipia-Samburuecosystem.Ecology and Evolution, 14,
e70198. https://doi.org/10.1002/ece3.70198