7
Research Article Impacts of Bush Encroachment on Wildlife Species Diversity, Composition, and Habitat Preference in Ol Pejeta Conservancy, Laikipia, Kenya Cyrus M. Kavwele, 1 Johnstone K. Kimanzi, 2 and Mwangi J. Kinyanjui 1 1 Department of Natural Resources, Karatina University, Karatina, Kenya 2 Department of Wildlife Management, University of Eldoret, Eldoret, Kenya Correspondence should be addressed to Cyrus M. Kavwele; [email protected] Received 23 August 2017; Accepted 5 November 2017; Published 6 December 2017 Academic Editor: Daniel I. Rubenstein Copyright © 2017 Cyrus M. Kavwele et al. is is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Savannah ecosystems are currently facing a biome shiſt that changes grasslands to woody dominated landscapes, attributable to habitat degradation. In Ol Pejeta Conservancy (OPC), Euclea divinorum, an unpalatable and invasive woody species, is expanding to former savannah ecosystems with potential effects on herbivores key resources, wildlife species diversity, composition, and habitat use. We investigated wildlife species diversity, composition, and habitat preference or avoidance by wildlife in the conservancy. Infrared camera traps were deployed at the centroids of 2 km by 2 km, 50 cm above ground surface for 14 days and nights with 9 camera traps in each habitat type. Shannon wiener index revealed that wildlife species diversity was highest in E. divinorum dominated habitats and lowest in open grassland. Hierarchical Cluster Analysis revealed level of similarity in wildlife species composition between E. divinorum and mixed bushland. Jacobs index revealed that E. divinorum and mixed bushland were avoided by all guilds; however E. divinorum was significantly avoided while A. drepanolobium and open grassland were both preferred by all guilds. However, A. drepanolobium dominated habitats were significantly preferred compared to open grasslands. e findings are useful in management of sustainable ecosystems. 1. Introduction Woody species encroachment has been described as gradual conversion of grasslands to woody and shrub dominated savannahs [1], a phenomenon that has been documented well in various parts of the world [2]. ese increasingly changing grasslands [3] form persistent patches which alter composition and structure of flora in savannah ecosystems [4]. Overgrazing by exerting pressure on grazing lawns and increase in precipitation have shown positive correlation with woody species encroachment [5], hence touted as potential driver of the vegetation shiſts. Further, soil conditions [6, 7] and fire suppression [8] among other perturbations are also attributable to this phenomenon. According to Ward [9] an increase in woody species can significantly affect livestock and wildlife conservation efforts, as such, further indirectly or directly affecting livelihoods of the local people [10]. Particular facets of economy likely to be affected include but not limited to ecotourism, a key source of revenue in these ecosystems due to poor visibility, hence affecting game viewing experience [11]. From another ecological position, according to Wigley et al. [11], increase in woody species translates to increase in fuel wood, timber, and fencing material among other as well as increase in resource for browsing herbivores and structural diversity for avian biodiversity. Even worse, encroachment in isolated ecosystems can result in decline and/or extinction of native species and can potentially affect species diversity, distribution, and abun- dance [12]. is also holds true according to Dalle et al. [13] where encroachment in savannah ecosystems poses a serious threat to ecosystems function especially tree-grass coex- istence. Grass-tree balance influences grassland/rangeland Hindawi International Journal of Ecology Volume 2017, Article ID 5620125, 6 pages https://doi.org/10.1155/2017/5620125

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Page 1: Impacts of Bush Encroachment on Wildlife Species Diversity ...downloads.hindawi.com/journals/ijecol/2017/5620125.pdf · Impacts of Bush Encroachment on Wildlife Species Diversity,

Research ArticleImpacts of Bush Encroachment on Wildlife SpeciesDiversity Composition and Habitat Preference in Ol PejetaConservancy Laikipia Kenya

Cyrus M Kavwele1 Johnstone K Kimanzi2 andMwangi J Kinyanjui1

1Department of Natural Resources Karatina University Karatina Kenya2Department of Wildlife Management University of Eldoret Eldoret Kenya

Correspondence should be addressed to Cyrus M Kavwele cyruskavweleyahoocom

Received 23 August 2017 Accepted 5 November 2017 Published 6 December 2017

Academic Editor Daniel I Rubenstein

Copyright copy 2017 Cyrus M Kavwele et al This is an open access article distributed under the Creative Commons AttributionLicense which permits unrestricted use distribution and reproduction in any medium provided the original work is properlycited

Savannah ecosystems are currently facing a biome shift that changes grasslands to woody dominated landscapes attributable tohabitat degradation InOl Pejeta Conservancy (OPC)Euclea divinorum an unpalatable and invasivewoody species is expanding toformer savannah ecosystems with potential effects on herbivores key resources wildlife species diversity composition and habitatuse We investigated wildlife species diversity composition and habitat preference or avoidance by wildlife in the conservancyInfrared camera traps were deployed at the centroids of 2 km by 2 km 50 cm above ground surface for 14 days and nights with9 camera traps in each habitat type Shannon wiener index revealed that wildlife species diversity was highest in E divinorumdominated habitats and lowest in open grassland Hierarchical Cluster Analysis revealed level of similarity in wildlife speciescomposition between E divinorum andmixed bushland Jacobs index revealed that E divinorum andmixed bushland were avoidedby all guilds however E divinorum was significantly avoided while A drepanolobium and open grassland were both preferred byall guilds However A drepanolobium dominated habitats were significantly preferred compared to open grasslands The findingsare useful in management of sustainable ecosystems

1 Introduction

Woody species encroachment has been described as gradualconversion of grasslands to woody and shrub dominatedsavannahs [1] a phenomenon that has been documentedwell in various parts of the world [2] These increasinglychanging grasslands [3] form persistent patches which altercomposition and structure of flora in savannah ecosystems[4] Overgrazing by exerting pressure on grazing lawns andincrease in precipitation have shown positive correlationwithwoody species encroachment [5] hence touted as potentialdriver of the vegetation shifts Further soil conditions [6 7]and fire suppression [8] among other perturbations are alsoattributable to this phenomenon

According to Ward [9] an increase in woody species cansignificantly affect livestock and wildlife conservation effortsas such further indirectly or directly affecting livelihoods

of the local people [10] Particular facets of economy likelyto be affected include but not limited to ecotourism a keysource of revenue in these ecosystems due to poor visibilityhence affecting game viewing experience [11] From anotherecological position according toWigley et al [11] increase inwoody species translates to increase in fuel wood timber andfencing material among other as well as increase in resourcefor browsing herbivores and structural diversity for avianbiodiversity

Even worse encroachment in isolated ecosystems canresult in decline andor extinction of native species and canpotentially affect species diversity distribution and abun-dance [12] This also holds true according to Dalle et al [13]where encroachment in savannah ecosystems poses a seriousthreat to ecosystems function especially tree-grass coex-istence Grass-tree balance influences grasslandrangeland

HindawiInternational Journal of EcologyVolume 2017 Article ID 5620125 6 pageshttpsdoiorg10115520175620125

2 International Journal of Ecology

25 0 25 5 75 10

Chimps_HouseMain_gatesMash

Major_facilitiesMajor_roadsEwaso_Nyiro_River

OPC_Boundary

Gate_5

Gate_3

Gate_2

Kamok workshop

Serena HotelsHeadquarters

Chimps Facility

Airstrip

(km)

N

Figure 1 Map of the study site

economic services biodiversity conservation and ecosystemfunction at local and landscape scales

In Ol Pejeta Conservancy (OPC) encroachment byunpalatable E divinorum encroachment towards Acaciadrepanolobium grasslands and other open bushland vegeta-tion cover types has become a concern for conservationistsKey resources for megafauna in these ecosystems espe-cially the critically endangered Eastern black rhino Dicerosbicornis International Union for Conservation of Nature(IUCN) red listing [14] and African elephants Loxodontaafricana vulnerable (IUCN) red listing [15] among otherherbivores can potentially be affected This encroachmentcan further reduce the available ranging lands and to someextent exterminate some of the wild flora and fauna Herewe investigated (a) wildlife species diversity and evenness inencroached and nonencroached habitats (b) wildlife speciescomposition in encroached and nonencroached habitats and(c) habitat preference or avoidance by various feeding guildswithin the conservancy The findings demonstrate impactsof bush encroachment on biodiversity hence beneficial toconservationists

2 Methods

21 Study Site This study was undertaken in Ol PejetaConservancy in Laikipia County Central Kenya as shown inFigure 1 The property which covers 90000 acres (360 km2)lies in between Mt Kenya and Aberdares (at 0∘ 72881015840N36∘423841015840E and 0∘ 86341015840N 37∘ 06051015840E) (0∘ 18311015840S 36∘465781015840E and 0∘ 570251015840S 37∘ 24921015840E) It has an averagealtitude of 1810m mean annual rainfall of 739mm andmean maximum and minimum temperatures of 28∘C and12∘C respectively It is a private owned conservancy chieflya sanctuary for black rhinos (Diceros bicornis) Howeverother wild animals are also conserved in the same propertycomposed of various feeding guilds such as mixed feederscarnivores grazers and browsers It is also a sanctuary forchimpanzee (Pan troglodytes) providing refuge site for seizedchimps from black markets It has one permanent riverflowing through the conservancy as well as various man-made water holes to supplement water scarcity in dry spells

Habitat cover types include grasslands Acacia drepanolo-bium A xanthophloea Euclea divinorum and mixed bush-lands The conservancy is surrounded by an electric fence

International Journal of Ecology 3

Adrepanolobium

Edivinorum

Mixedbushland

Opengrassland

Habitat types

0

05

1

15

2

25

0

01

02

03

04

05

06

Spec

ies e

venn

ess (

HS

)

Spec

ies d

iver

sity (H

)

Figure 2 Wildlife species diversity (left 119910 axis) and wildlife species evenness (right 119910 axis) in different habitats in OPC

with three ldquocorridorsrdquo to allow movement of wild animals inand out of OPC (but movement of rhino species is restricteddue to the risks involved) The conservancy is surroundedby agropastoral communities and towards the north by otheradjoining conservancies

22 Data Collection and Analysis The entire conservancywas overlaid on 2 times 2 km grid as shown in Figure 1 (map)and centroids of each grid were identified At every centroidmotion triggered infrared camera traps of Reconyx Rapid-fire RM45 model were deployed systematically taking intocognisance of animal trails to capture wildlife species withingiven cover types Camera traps were set to take photos withno delay and remain active for 24 hrs at 50 cm above theground surface either attached to a tree using elastic strings(cords) or housed in a metal cage in case of grassland areasThey remained in the field for 14 consecutive days and nightsthough they were checked on the 7th day for battery chargelevel remaining storage size in secure cards (SD) and anyother malfunctions occasioned by knocking of the traps byrhinos or elephants A total of 9 camera traps were deployedin each habitat type namely open grassland E divinorumA drepanolobium and mixed bushland Upon realising lowbattery charge or near filled up storage there were replace-ments of such to avoid loss of data due to nonperformingcamera traps Immediately after removal of camera trapsfrom the field all photographic data were downloaded andfurther sorted into species name size time and habitat typethe camera trap was deployed in Further duplicates wereremoved where thirty (30) minutes after the last photo wastaken was considered as another photographic event

After data cleaning species diversity and compositionusing Shannon Weiner index (where if index is 5 it implieshighest diversity and if it is 1 it implies lowest diversity)[16] and Hierarchical Cluster Analysis [17] respectively wereperformed in Paleontological Statistics Software Package forEducation and Data Analysis (PAST) software while habitatpreference or avoidance was performed using Jacobsrsquo index[18] Further Jacobsrsquo index values were tested for signifi-cance preference or avoidance (where +1 implies maximumpreference while minus1 implies maximum avoidance) using 119905-tests against means of zero (0) in R version 312 Prior to

these tests normal distribution and homogeneity of variancewere performed to ensure that conditions for parametric testswere not violated [19] Overlay of grids on the spatial extentof the conservancy was performed in Quantum GeographicInformation Systems (QGIS) platform version 282

3 Results

31 Wildlife Species Diversity and Evenness ShannonWeinerindex revealed highest wildlife species diversity in E divi-norum dominated habitats (1198671015840 = 2291) followed by Adrepanolobium habitats (1198671015840 = 2058) mixed bushland (1198671015840 =1728) and least wildlife species diversity in open grassland(1198671015840 = 1715) Wildlife species evenness (119867119878) was highest inarea under E divinorum (119867119878 = 05201) followed by mixedbushland (119867119878 = 03751) and then closely followed by Adrepanolobium (119867119878 = 03404) and finally open grassland(119867119878 = 02647) as shown in Figure 2

32 Wildlife Species Composition Hierarchical Cluster Anal-ysis (HCA) compared wildlife species composition (commonwildlife species) across the four habitat types namely E divi-norum A drepanolobium mixed bushland and open grass-land Results revealed that E divinorum and mixed bushlandhabitats shared 45 similarity in wildlife species composi-tion Additionally E divinorum habitat and mixed bushlandshared 39 similarity in wildlife species composition with Adrepanolobium dominated habitat On the other hand opengrassland habitat shared 27 similarity in wildlife speciescomposition with three habitat types namely E divinorumA drepanolobium and mixed bushland as shown in Figure 3

33 Habitat Preference or Avoidance by Various Feeding Guildsin OPC Jacobs index revealed that carnivores preferredA drepanolobium (119863 = 0469) mostly followed by opengrassland (119863 = 0327) and least preferred mixed bushland(119863 = 0066) but avoided E divinorum dominated areas (119863 =minus0698) (Figure 4(a)) Grazers showed great preference foropen grassland (119863 = 0773) and A drepanolobium (119863 =0040) but avoided E divinorum (119863 = minus0917) and mixedbushland (119863 = minus0192) habitats (Figure 4(b))

4 International Journal of Ecology

A d

rep

E d

ivin

orum

Mix

ed b

ushl

and

Ope

n gr

assla

nd

02

03

04

05

06

07

08

09

10

Sim

ilarit

y

Figure 3 Percentage similarity in wildlife species composition of 4 habitats in OPC

minus08

minus06

minus04

minus02

0

02

04

06

Jaco

bsrsquo i

ndex

for c

arni

vore

s

Adrepanolobium

Edivinorum

Mixed_bushland Opengrassland

Habitat types

(a)minus12

minus1minus08minus06minus04minus02

002040608

1

Adrepanolobium

Edivinorum

Mixed_bushland Opengrassland

Jaco

bs in

dex

for g

raze

rs

Habitat types

(b)

0

minus08

minus06

minus04

minus02

02

04

06

08

Jaco

bs in

dex

for b

row

sers

Habitat types

Adrepanolobium

Edivinorum

Mixed_bushland Opengrassland

(c)

Adrepanolobium

Edivinorum

Mixed_bushland Opengrassland

minus1minus08minus06minus04minus02

002040608

Jaco

bs in

dex

for m

ixed

feed

ers

Habitat types

(d)Figure 4 Habitat preference or avoidance (Jacobsrsquo index) for (a) carnivores (b) grazers (c) browsers and (d) mixed feeders in OPC

Browsers showed preference for A drepanolobium (119863 =0674) and mixed bushland habitat (119863 = 0175) but avoidedboth E divinorum (119863 = minus0673) and open grasslands (119863 =minus0116) dominated habitats (Figure 4(c)) However mixedfeeders preferred habitats dominated by open grassland (119863 =0688) and A drepanolobium (119863 = 0523) but avoided Edivinorum (119863 = minus0858) and mixed bushland (119863 = minus0420)

dominated habitats (Figure 4(d)) A total of thirty (30)wildlife species were recorded as per the table in the sup-plementary material showing detection and nondetection ofwildlife species in various habitat cover types

Mean preference or avoidance revealed that E divinorumand mixed bushland were avoided however E divinorumwas significantly avoided (119905

1= 2253 df = 3 and

International Journal of Ecology 5

minus05

05

Jaco

bsrsquo i

ndex

mea

ns

E div m_bushland OGA drepHabitat types

Figure 5 Jacobs index means for all guilds across the habitat types

119901 lt 001) compared with mixed bushland (1199051= 2353 df

= 3 and 119901 = 027) On the other hand A drepanolobiumand open grassland were both preferred by all feedingguilds however A drepanolobium dominated habitats weresignificantly preferred (119905

1= 2353 df = 3 and 119901 = 003)

compared to open grasslands (1199051= 2353 df = 3 and 119901 =

006) as shown in Figure 5

4 Discussions

Wildlife species diversity and evenness were highest in Edivinorum dominated areas compared to other habitat typesThis is attributable to existence of resource rich patchesthat are hardly accessed by most wildlife species As suchattraction of several wildlife species into this dense vegetation(E divinorum) contributed to higher diversity chiefly smallcarnivores Animals use cues such as landscape structureand vegetation structure like tree species phenology resourceavailability (probability of encounter quantity and quality)and predatorparasite risks among others in order to explorea given habitat type This implies that spatial and temporalhabitat heterogeneity drives faunal species diversity andcomposition from local to regional to global scales Henceaggregation of resources may create dynamic impacts wherethese small sitespatches become a focus for higher propor-tion of faunal diversity and composition [20] Accordingto Sirami et al [21] savannah ecosystems are diverse instructure and floral composition hence different guilds willuse different habitats based on their requirements

Animals have different preference to certain habitats [22]as a function of direct and indirect effects of prey availabilitydetectabilitycover and resource availability [23] Spatialheterogeneity of ecosystem is important for maintenance ofdiverse wild animal species and acts as buffer against changesin resources availability for wild animals in era of climatechange [24] On the other hand woody encroachment canpotentially change this heterogeneity and affect wide range ofwild animals

Similarity in wildlife species composition was highestbetween E divinorum and mixed bushland habitats Thesetwo habitat types have nearly homogeneous vegetation struc-ture (dense canopy) This phenomenon is an illumination tohigher percentage similarity in wildlife species compositionin both E divinorum and mixed bushland dominated areasdue to nearly similar structure of the vegetation (canopy anddense bushland) providing ideal habitats for wildlife specieswhich require such areas According to Dupuch et al [25]

prey and predator can explore freely across habitats thatdiffer in resource quantity quality and inherent habitat risksAs such predatorsrsquo movement and habitat selection are asa function of prey density competition level cover andinherent risks while prey habitat selection and movement arebalance of forage quality (benefits of foraging) and risk ofpredation [26]

On the other hand habitats dominated by A drepanolo-bium and open grassland were preferred by feeding guildsmajorly herbivores due to their associated lesser predationrisk This is chiefly influenced by presence of relatively openlandscapes allowing prey to detect predators from a distanceand hencemaintain safer flight distance (improved visibility)The phenomenon is consistent with other findings accordingto Dupuch et al [25] where prey habitat selection is largelyinfluenced by the need to minimise predation risks overforage quality As such these factors influence preferenceof habitats that offer significant benefits for most wildlifespecies

5 Conclusion and Recommendations

The study revealed that bush encroachment by E divinorumhad impact on species diversity composition and habitatselection In particular species diversity was highest in Edivinorum dominated areas and least in open grasslands apositive impact albeit long term impacts are unknown Like-wise percentage similarity in species composition was high-est between E divinorum and mixed bushland dominatedhabitats Further feeding guilds avoided areas dominatedby E divinorum but preferred A drepanolobium and opengrassland dominated habitats hence bush encroachmentby E divinorum affected habitat suitability thus it has anegative impact Based on this finding active managementof the encroaching species is recommended Further stud-ies on activity budget of wildlife species to ascertain howhabitat types are being used are key Additionally long termstudy to provide insights into the effect of seasonality onspecies diversity dynamics across various land cover typesin the conservancy is crucial This information will be vitalin designing habitat management protocols for sustainableecosystem management

Conflicts of Interest

The authors declare no conflicts of interest

Acknowledgments

This study was funded by Rufford Foundation United King-dom (UK) Grant no 19422-1 The authors are grateful toCarol Ngwrsquoeno and Bernard Chira from Ol Pejeta Conser-vancy (OPC) for allowing them to carry out research intheir property and Andrew Bradley and Dan Smith bothfrom Bangor University for their help in camera trap imageclassification

6 International Journal of Ecology

Supplementary Materials

Table 1 Wildlife species detection or nondetection across thevarious habitat cover types (Supplementary Materials)

References

[1] A T Hudak C A Wessman and T R Seastedt ldquoWoodyoverstorey effects on soil carbon and nitrogen pools in SouthAfrican savannardquo Austral Ecology vol 28 no 2 pp 173ndash1812003

[2] S Archer T W Boutton and K A Hibbard ldquoTrees in grass-lands biogeochemical consequences of woody plant expan-sionrdquo inGlobal Biogeochemical Cycles in the Climate System pp115ndash137 Academic Press San Diego 2001

[3] I A Dickie S A Schnitzer P B Reich and S E Hobbie ldquoIsoak establishment in old-fields and savanna openings contextdependentrdquo Journal of Ecology vol 95 no 2 pp 309ndash320 2007

[4] S R Wangen and C R Webster ldquoPotential for multiple lagphases during biotic invasions Reconstructing an invasion ofthe exotic tree Acer platanoidesrdquo Journal of Applied Ecology vol43 no 2 pp 258ndash268 2006

[5] D F Joubert A Rothauge andG N Smit ldquoA conceptual modelof vegetation dynamics in the semiarid Highland savanna ofNamibia with particular reference to bush thickening byAcaciamelliferardquo Journal of Arid Environments vol 72 no 12 pp 2201ndash2210 2008

[6] M Sankaran J Ratnam and N Hanan ldquoWoody cover inAfrican savannas The role of resources fire and herbivoryrdquoGlobal Ecology and Biogeography vol 17 no 2 pp 236ndash2452008

[7] G Oba E Post P O Syvertsen andN C Stenseth ldquoBush coverand range condition assessments in relation to landscape andgrazing in southern Ethiopiardquo Landscape Ecology vol 15 no 6pp 535ndash546 2000

[8] O W Van Auken ldquoShrub invasions of North American semi-arid grasslandsrdquo Annual Review of Ecology Evolution andSystematics vol 31 pp 197ndash215 2000

[9] D Ward ldquoDo we understand the causes of bush encroachmentinAfrican savannasrdquoAfrican Journal of RangeampForage Sciencevol 22 no 2 pp 101ndash105 2005

[10] A T Hudak and C A Wessman ldquoTextural analysis of highresolution imagery to quantify bush enroachment in MadikweGame Reserve South Africa 1955-1996rdquo International Journalof Remote Sensing vol 22 no 14 pp 2731ndash2740 2001

[11] B J Wigley W J Bond and M T Hoffman ldquoBush encroach-ment under three contrasting land-use practices in a mesicSouth African savannardquo African Journal of Ecology vol 47 no1 pp 62ndash70 2009

[12] D R Towns I A E Atkinson and C H Daugherty ldquoHave theharmful effects of introduced rats on islands been exaggeratedrdquoBiological Invasions vol 8 no 4 pp 863ndash891 2006

[13] G Dalle B LMaass and J Isselstein ldquoEncroachment of woodyplants and its impact on pastoral livestock production in theBorana lowlands southern Oromia Ethiopiardquo African Journalof Ecology vol 44 no 2 pp 237ndash246 2006

[14] R Emslie ldquoDiceros bicornis The IUCN Red List of ThreatenedSpecies 2012rdquo 2012 eT6557A16980917 Downloadedon 17April httpdxdoiorg102305IUCNUK2012RLTST6557A16980917en 2016

[15] J Blanc ldquoLoxodonta africanaThe IUCNRed List ofThreatenedSpecies 2008rdquo2008 eT12392A3339343 httpdxdoiorg102305IUCNUK2008RLTST12392A3339343enDownloadedon 17 April 2016

[16] C E Shannon and W Weaver The Mathematical Theory ofCommunication University of Illinois Press Chicago 1949

[17] H G Gauch Multivariate Analysis in Community EcologyCambridge University Press Cambridge UK 1982

[18] J Jacobs ldquoQuantitative measurement of food selection - Amodification of the forage ratio and Ivlevrsquos electivity indexrdquoOecologia vol 14 no 4 pp 413ndash417 1974

[19] F Palomares M Delibes E Revilla J Calzada and J MFedriani ldquoSpatial ecology of iberian lynx and abundance ofeuropean rabbits in southwestern spainrdquo Wildlife Monographsvol 148 pp 1ndash36 2001

[20] D J D Natusch J A Lyons G Brown and R Shine ldquoCom-munally nesting migratory birds create ecological hot-spotsin Tropical Australiardquo PLoS ONE vol 11 no 10 Article IDe0162651 2016

[21] C Sirami C Seymour GMidgley and P Barnard ldquoThe impactof shrub encroachment on savanna bird diversity from local toregional scalerdquo Diversity and Distributions vol 15 no 6 pp948ndash957 2009

[22] A R E Sinclair S Mduma and J S Brashares ldquoPatterns ofpredation in a diverse predator-prey systemrdquo Nature vol 425no 6955 pp 288ndash290 2003

[23] W J Ripple and R L Beschta ldquoWolves and the ecology of fearCan predation risk structure ecosystemsrdquo Bioscience vol 54no 8 pp 755ndash766 2004

[24] G Wang N T Hobbs R B Boone et al ldquoSpatial and temporalvariability modify density dependence in populations of largeherbivoresrdquo Ecology vol 87 no 1 pp 95ndash102 2006

[25] A Dupuch L M Dill and P Magnan ldquoTesting the effectsof resource distribution and inherent habitat riskiness onsimultaneous habitat selection by predators and preyrdquo AnimalBehaviour vol 78 no 3 pp 705ndash713 2009

[26] P A Abrams ldquoHabitat choice in predator-prey systems Spatialinstability due to interacting adaptive movementsrdquoThe Ameri-can Naturalist vol 169 no 5 pp 581ndash594 2007

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ClimatologyJournal of

Page 2: Impacts of Bush Encroachment on Wildlife Species Diversity ...downloads.hindawi.com/journals/ijecol/2017/5620125.pdf · Impacts of Bush Encroachment on Wildlife Species Diversity,

2 International Journal of Ecology

25 0 25 5 75 10

Chimps_HouseMain_gatesMash

Major_facilitiesMajor_roadsEwaso_Nyiro_River

OPC_Boundary

Gate_5

Gate_3

Gate_2

Kamok workshop

Serena HotelsHeadquarters

Chimps Facility

Airstrip

(km)

N

Figure 1 Map of the study site

economic services biodiversity conservation and ecosystemfunction at local and landscape scales

In Ol Pejeta Conservancy (OPC) encroachment byunpalatable E divinorum encroachment towards Acaciadrepanolobium grasslands and other open bushland vegeta-tion cover types has become a concern for conservationistsKey resources for megafauna in these ecosystems espe-cially the critically endangered Eastern black rhino Dicerosbicornis International Union for Conservation of Nature(IUCN) red listing [14] and African elephants Loxodontaafricana vulnerable (IUCN) red listing [15] among otherherbivores can potentially be affected This encroachmentcan further reduce the available ranging lands and to someextent exterminate some of the wild flora and fauna Herewe investigated (a) wildlife species diversity and evenness inencroached and nonencroached habitats (b) wildlife speciescomposition in encroached and nonencroached habitats and(c) habitat preference or avoidance by various feeding guildswithin the conservancy The findings demonstrate impactsof bush encroachment on biodiversity hence beneficial toconservationists

2 Methods

21 Study Site This study was undertaken in Ol PejetaConservancy in Laikipia County Central Kenya as shown inFigure 1 The property which covers 90000 acres (360 km2)lies in between Mt Kenya and Aberdares (at 0∘ 72881015840N36∘423841015840E and 0∘ 86341015840N 37∘ 06051015840E) (0∘ 18311015840S 36∘465781015840E and 0∘ 570251015840S 37∘ 24921015840E) It has an averagealtitude of 1810m mean annual rainfall of 739mm andmean maximum and minimum temperatures of 28∘C and12∘C respectively It is a private owned conservancy chieflya sanctuary for black rhinos (Diceros bicornis) Howeverother wild animals are also conserved in the same propertycomposed of various feeding guilds such as mixed feederscarnivores grazers and browsers It is also a sanctuary forchimpanzee (Pan troglodytes) providing refuge site for seizedchimps from black markets It has one permanent riverflowing through the conservancy as well as various man-made water holes to supplement water scarcity in dry spells

Habitat cover types include grasslands Acacia drepanolo-bium A xanthophloea Euclea divinorum and mixed bush-lands The conservancy is surrounded by an electric fence

International Journal of Ecology 3

Adrepanolobium

Edivinorum

Mixedbushland

Opengrassland

Habitat types

0

05

1

15

2

25

0

01

02

03

04

05

06

Spec

ies e

venn

ess (

HS

)

Spec

ies d

iver

sity (H

)

Figure 2 Wildlife species diversity (left 119910 axis) and wildlife species evenness (right 119910 axis) in different habitats in OPC

with three ldquocorridorsrdquo to allow movement of wild animals inand out of OPC (but movement of rhino species is restricteddue to the risks involved) The conservancy is surroundedby agropastoral communities and towards the north by otheradjoining conservancies

22 Data Collection and Analysis The entire conservancywas overlaid on 2 times 2 km grid as shown in Figure 1 (map)and centroids of each grid were identified At every centroidmotion triggered infrared camera traps of Reconyx Rapid-fire RM45 model were deployed systematically taking intocognisance of animal trails to capture wildlife species withingiven cover types Camera traps were set to take photos withno delay and remain active for 24 hrs at 50 cm above theground surface either attached to a tree using elastic strings(cords) or housed in a metal cage in case of grassland areasThey remained in the field for 14 consecutive days and nightsthough they were checked on the 7th day for battery chargelevel remaining storage size in secure cards (SD) and anyother malfunctions occasioned by knocking of the traps byrhinos or elephants A total of 9 camera traps were deployedin each habitat type namely open grassland E divinorumA drepanolobium and mixed bushland Upon realising lowbattery charge or near filled up storage there were replace-ments of such to avoid loss of data due to nonperformingcamera traps Immediately after removal of camera trapsfrom the field all photographic data were downloaded andfurther sorted into species name size time and habitat typethe camera trap was deployed in Further duplicates wereremoved where thirty (30) minutes after the last photo wastaken was considered as another photographic event

After data cleaning species diversity and compositionusing Shannon Weiner index (where if index is 5 it implieshighest diversity and if it is 1 it implies lowest diversity)[16] and Hierarchical Cluster Analysis [17] respectively wereperformed in Paleontological Statistics Software Package forEducation and Data Analysis (PAST) software while habitatpreference or avoidance was performed using Jacobsrsquo index[18] Further Jacobsrsquo index values were tested for signifi-cance preference or avoidance (where +1 implies maximumpreference while minus1 implies maximum avoidance) using 119905-tests against means of zero (0) in R version 312 Prior to

these tests normal distribution and homogeneity of variancewere performed to ensure that conditions for parametric testswere not violated [19] Overlay of grids on the spatial extentof the conservancy was performed in Quantum GeographicInformation Systems (QGIS) platform version 282

3 Results

31 Wildlife Species Diversity and Evenness ShannonWeinerindex revealed highest wildlife species diversity in E divi-norum dominated habitats (1198671015840 = 2291) followed by Adrepanolobium habitats (1198671015840 = 2058) mixed bushland (1198671015840 =1728) and least wildlife species diversity in open grassland(1198671015840 = 1715) Wildlife species evenness (119867119878) was highest inarea under E divinorum (119867119878 = 05201) followed by mixedbushland (119867119878 = 03751) and then closely followed by Adrepanolobium (119867119878 = 03404) and finally open grassland(119867119878 = 02647) as shown in Figure 2

32 Wildlife Species Composition Hierarchical Cluster Anal-ysis (HCA) compared wildlife species composition (commonwildlife species) across the four habitat types namely E divi-norum A drepanolobium mixed bushland and open grass-land Results revealed that E divinorum and mixed bushlandhabitats shared 45 similarity in wildlife species composi-tion Additionally E divinorum habitat and mixed bushlandshared 39 similarity in wildlife species composition with Adrepanolobium dominated habitat On the other hand opengrassland habitat shared 27 similarity in wildlife speciescomposition with three habitat types namely E divinorumA drepanolobium and mixed bushland as shown in Figure 3

33 Habitat Preference or Avoidance by Various Feeding Guildsin OPC Jacobs index revealed that carnivores preferredA drepanolobium (119863 = 0469) mostly followed by opengrassland (119863 = 0327) and least preferred mixed bushland(119863 = 0066) but avoided E divinorum dominated areas (119863 =minus0698) (Figure 4(a)) Grazers showed great preference foropen grassland (119863 = 0773) and A drepanolobium (119863 =0040) but avoided E divinorum (119863 = minus0917) and mixedbushland (119863 = minus0192) habitats (Figure 4(b))

4 International Journal of Ecology

A d

rep

E d

ivin

orum

Mix

ed b

ushl

and

Ope

n gr

assla

nd

02

03

04

05

06

07

08

09

10

Sim

ilarit

y

Figure 3 Percentage similarity in wildlife species composition of 4 habitats in OPC

minus08

minus06

minus04

minus02

0

02

04

06

Jaco

bsrsquo i

ndex

for c

arni

vore

s

Adrepanolobium

Edivinorum

Mixed_bushland Opengrassland

Habitat types

(a)minus12

minus1minus08minus06minus04minus02

002040608

1

Adrepanolobium

Edivinorum

Mixed_bushland Opengrassland

Jaco

bs in

dex

for g

raze

rs

Habitat types

(b)

0

minus08

minus06

minus04

minus02

02

04

06

08

Jaco

bs in

dex

for b

row

sers

Habitat types

Adrepanolobium

Edivinorum

Mixed_bushland Opengrassland

(c)

Adrepanolobium

Edivinorum

Mixed_bushland Opengrassland

minus1minus08minus06minus04minus02

002040608

Jaco

bs in

dex

for m

ixed

feed

ers

Habitat types

(d)Figure 4 Habitat preference or avoidance (Jacobsrsquo index) for (a) carnivores (b) grazers (c) browsers and (d) mixed feeders in OPC

Browsers showed preference for A drepanolobium (119863 =0674) and mixed bushland habitat (119863 = 0175) but avoidedboth E divinorum (119863 = minus0673) and open grasslands (119863 =minus0116) dominated habitats (Figure 4(c)) However mixedfeeders preferred habitats dominated by open grassland (119863 =0688) and A drepanolobium (119863 = 0523) but avoided Edivinorum (119863 = minus0858) and mixed bushland (119863 = minus0420)

dominated habitats (Figure 4(d)) A total of thirty (30)wildlife species were recorded as per the table in the sup-plementary material showing detection and nondetection ofwildlife species in various habitat cover types

Mean preference or avoidance revealed that E divinorumand mixed bushland were avoided however E divinorumwas significantly avoided (119905

1= 2253 df = 3 and

International Journal of Ecology 5

minus05

05

Jaco

bsrsquo i

ndex

mea

ns

E div m_bushland OGA drepHabitat types

Figure 5 Jacobs index means for all guilds across the habitat types

119901 lt 001) compared with mixed bushland (1199051= 2353 df

= 3 and 119901 = 027) On the other hand A drepanolobiumand open grassland were both preferred by all feedingguilds however A drepanolobium dominated habitats weresignificantly preferred (119905

1= 2353 df = 3 and 119901 = 003)

compared to open grasslands (1199051= 2353 df = 3 and 119901 =

006) as shown in Figure 5

4 Discussions

Wildlife species diversity and evenness were highest in Edivinorum dominated areas compared to other habitat typesThis is attributable to existence of resource rich patchesthat are hardly accessed by most wildlife species As suchattraction of several wildlife species into this dense vegetation(E divinorum) contributed to higher diversity chiefly smallcarnivores Animals use cues such as landscape structureand vegetation structure like tree species phenology resourceavailability (probability of encounter quantity and quality)and predatorparasite risks among others in order to explorea given habitat type This implies that spatial and temporalhabitat heterogeneity drives faunal species diversity andcomposition from local to regional to global scales Henceaggregation of resources may create dynamic impacts wherethese small sitespatches become a focus for higher propor-tion of faunal diversity and composition [20] Accordingto Sirami et al [21] savannah ecosystems are diverse instructure and floral composition hence different guilds willuse different habitats based on their requirements

Animals have different preference to certain habitats [22]as a function of direct and indirect effects of prey availabilitydetectabilitycover and resource availability [23] Spatialheterogeneity of ecosystem is important for maintenance ofdiverse wild animal species and acts as buffer against changesin resources availability for wild animals in era of climatechange [24] On the other hand woody encroachment canpotentially change this heterogeneity and affect wide range ofwild animals

Similarity in wildlife species composition was highestbetween E divinorum and mixed bushland habitats Thesetwo habitat types have nearly homogeneous vegetation struc-ture (dense canopy) This phenomenon is an illumination tohigher percentage similarity in wildlife species compositionin both E divinorum and mixed bushland dominated areasdue to nearly similar structure of the vegetation (canopy anddense bushland) providing ideal habitats for wildlife specieswhich require such areas According to Dupuch et al [25]

prey and predator can explore freely across habitats thatdiffer in resource quantity quality and inherent habitat risksAs such predatorsrsquo movement and habitat selection are asa function of prey density competition level cover andinherent risks while prey habitat selection and movement arebalance of forage quality (benefits of foraging) and risk ofpredation [26]

On the other hand habitats dominated by A drepanolo-bium and open grassland were preferred by feeding guildsmajorly herbivores due to their associated lesser predationrisk This is chiefly influenced by presence of relatively openlandscapes allowing prey to detect predators from a distanceand hencemaintain safer flight distance (improved visibility)The phenomenon is consistent with other findings accordingto Dupuch et al [25] where prey habitat selection is largelyinfluenced by the need to minimise predation risks overforage quality As such these factors influence preferenceof habitats that offer significant benefits for most wildlifespecies

5 Conclusion and Recommendations

The study revealed that bush encroachment by E divinorumhad impact on species diversity composition and habitatselection In particular species diversity was highest in Edivinorum dominated areas and least in open grasslands apositive impact albeit long term impacts are unknown Like-wise percentage similarity in species composition was high-est between E divinorum and mixed bushland dominatedhabitats Further feeding guilds avoided areas dominatedby E divinorum but preferred A drepanolobium and opengrassland dominated habitats hence bush encroachmentby E divinorum affected habitat suitability thus it has anegative impact Based on this finding active managementof the encroaching species is recommended Further stud-ies on activity budget of wildlife species to ascertain howhabitat types are being used are key Additionally long termstudy to provide insights into the effect of seasonality onspecies diversity dynamics across various land cover typesin the conservancy is crucial This information will be vitalin designing habitat management protocols for sustainableecosystem management

Conflicts of Interest

The authors declare no conflicts of interest

Acknowledgments

This study was funded by Rufford Foundation United King-dom (UK) Grant no 19422-1 The authors are grateful toCarol Ngwrsquoeno and Bernard Chira from Ol Pejeta Conser-vancy (OPC) for allowing them to carry out research intheir property and Andrew Bradley and Dan Smith bothfrom Bangor University for their help in camera trap imageclassification

6 International Journal of Ecology

Supplementary Materials

Table 1 Wildlife species detection or nondetection across thevarious habitat cover types (Supplementary Materials)

References

[1] A T Hudak C A Wessman and T R Seastedt ldquoWoodyoverstorey effects on soil carbon and nitrogen pools in SouthAfrican savannardquo Austral Ecology vol 28 no 2 pp 173ndash1812003

[2] S Archer T W Boutton and K A Hibbard ldquoTrees in grass-lands biogeochemical consequences of woody plant expan-sionrdquo inGlobal Biogeochemical Cycles in the Climate System pp115ndash137 Academic Press San Diego 2001

[3] I A Dickie S A Schnitzer P B Reich and S E Hobbie ldquoIsoak establishment in old-fields and savanna openings contextdependentrdquo Journal of Ecology vol 95 no 2 pp 309ndash320 2007

[4] S R Wangen and C R Webster ldquoPotential for multiple lagphases during biotic invasions Reconstructing an invasion ofthe exotic tree Acer platanoidesrdquo Journal of Applied Ecology vol43 no 2 pp 258ndash268 2006

[5] D F Joubert A Rothauge andG N Smit ldquoA conceptual modelof vegetation dynamics in the semiarid Highland savanna ofNamibia with particular reference to bush thickening byAcaciamelliferardquo Journal of Arid Environments vol 72 no 12 pp 2201ndash2210 2008

[6] M Sankaran J Ratnam and N Hanan ldquoWoody cover inAfrican savannas The role of resources fire and herbivoryrdquoGlobal Ecology and Biogeography vol 17 no 2 pp 236ndash2452008

[7] G Oba E Post P O Syvertsen andN C Stenseth ldquoBush coverand range condition assessments in relation to landscape andgrazing in southern Ethiopiardquo Landscape Ecology vol 15 no 6pp 535ndash546 2000

[8] O W Van Auken ldquoShrub invasions of North American semi-arid grasslandsrdquo Annual Review of Ecology Evolution andSystematics vol 31 pp 197ndash215 2000

[9] D Ward ldquoDo we understand the causes of bush encroachmentinAfrican savannasrdquoAfrican Journal of RangeampForage Sciencevol 22 no 2 pp 101ndash105 2005

[10] A T Hudak and C A Wessman ldquoTextural analysis of highresolution imagery to quantify bush enroachment in MadikweGame Reserve South Africa 1955-1996rdquo International Journalof Remote Sensing vol 22 no 14 pp 2731ndash2740 2001

[11] B J Wigley W J Bond and M T Hoffman ldquoBush encroach-ment under three contrasting land-use practices in a mesicSouth African savannardquo African Journal of Ecology vol 47 no1 pp 62ndash70 2009

[12] D R Towns I A E Atkinson and C H Daugherty ldquoHave theharmful effects of introduced rats on islands been exaggeratedrdquoBiological Invasions vol 8 no 4 pp 863ndash891 2006

[13] G Dalle B LMaass and J Isselstein ldquoEncroachment of woodyplants and its impact on pastoral livestock production in theBorana lowlands southern Oromia Ethiopiardquo African Journalof Ecology vol 44 no 2 pp 237ndash246 2006

[14] R Emslie ldquoDiceros bicornis The IUCN Red List of ThreatenedSpecies 2012rdquo 2012 eT6557A16980917 Downloadedon 17April httpdxdoiorg102305IUCNUK2012RLTST6557A16980917en 2016

[15] J Blanc ldquoLoxodonta africanaThe IUCNRed List ofThreatenedSpecies 2008rdquo2008 eT12392A3339343 httpdxdoiorg102305IUCNUK2008RLTST12392A3339343enDownloadedon 17 April 2016

[16] C E Shannon and W Weaver The Mathematical Theory ofCommunication University of Illinois Press Chicago 1949

[17] H G Gauch Multivariate Analysis in Community EcologyCambridge University Press Cambridge UK 1982

[18] J Jacobs ldquoQuantitative measurement of food selection - Amodification of the forage ratio and Ivlevrsquos electivity indexrdquoOecologia vol 14 no 4 pp 413ndash417 1974

[19] F Palomares M Delibes E Revilla J Calzada and J MFedriani ldquoSpatial ecology of iberian lynx and abundance ofeuropean rabbits in southwestern spainrdquo Wildlife Monographsvol 148 pp 1ndash36 2001

[20] D J D Natusch J A Lyons G Brown and R Shine ldquoCom-munally nesting migratory birds create ecological hot-spotsin Tropical Australiardquo PLoS ONE vol 11 no 10 Article IDe0162651 2016

[21] C Sirami C Seymour GMidgley and P Barnard ldquoThe impactof shrub encroachment on savanna bird diversity from local toregional scalerdquo Diversity and Distributions vol 15 no 6 pp948ndash957 2009

[22] A R E Sinclair S Mduma and J S Brashares ldquoPatterns ofpredation in a diverse predator-prey systemrdquo Nature vol 425no 6955 pp 288ndash290 2003

[23] W J Ripple and R L Beschta ldquoWolves and the ecology of fearCan predation risk structure ecosystemsrdquo Bioscience vol 54no 8 pp 755ndash766 2004

[24] G Wang N T Hobbs R B Boone et al ldquoSpatial and temporalvariability modify density dependence in populations of largeherbivoresrdquo Ecology vol 87 no 1 pp 95ndash102 2006

[25] A Dupuch L M Dill and P Magnan ldquoTesting the effectsof resource distribution and inherent habitat riskiness onsimultaneous habitat selection by predators and preyrdquo AnimalBehaviour vol 78 no 3 pp 705ndash713 2009

[26] P A Abrams ldquoHabitat choice in predator-prey systems Spatialinstability due to interacting adaptive movementsrdquoThe Ameri-can Naturalist vol 169 no 5 pp 581ndash594 2007

Submit your manuscripts athttpswwwhindawicom

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental and Public Health

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcosystemsJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

MeteorologyAdvances in

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental Chemistry

Atmospheric SciencesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Waste ManagementJournal of

Hindawi Publishing Corporation httpwwwhindawicom Volume 201

International Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Geological ResearchJournal of

EarthquakesJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

OceanographyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Journal of Computational Environmental SciencesHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ClimatologyJournal of

Page 3: Impacts of Bush Encroachment on Wildlife Species Diversity ...downloads.hindawi.com/journals/ijecol/2017/5620125.pdf · Impacts of Bush Encroachment on Wildlife Species Diversity,

International Journal of Ecology 3

Adrepanolobium

Edivinorum

Mixedbushland

Opengrassland

Habitat types

0

05

1

15

2

25

0

01

02

03

04

05

06

Spec

ies e

venn

ess (

HS

)

Spec

ies d

iver

sity (H

)

Figure 2 Wildlife species diversity (left 119910 axis) and wildlife species evenness (right 119910 axis) in different habitats in OPC

with three ldquocorridorsrdquo to allow movement of wild animals inand out of OPC (but movement of rhino species is restricteddue to the risks involved) The conservancy is surroundedby agropastoral communities and towards the north by otheradjoining conservancies

22 Data Collection and Analysis The entire conservancywas overlaid on 2 times 2 km grid as shown in Figure 1 (map)and centroids of each grid were identified At every centroidmotion triggered infrared camera traps of Reconyx Rapid-fire RM45 model were deployed systematically taking intocognisance of animal trails to capture wildlife species withingiven cover types Camera traps were set to take photos withno delay and remain active for 24 hrs at 50 cm above theground surface either attached to a tree using elastic strings(cords) or housed in a metal cage in case of grassland areasThey remained in the field for 14 consecutive days and nightsthough they were checked on the 7th day for battery chargelevel remaining storage size in secure cards (SD) and anyother malfunctions occasioned by knocking of the traps byrhinos or elephants A total of 9 camera traps were deployedin each habitat type namely open grassland E divinorumA drepanolobium and mixed bushland Upon realising lowbattery charge or near filled up storage there were replace-ments of such to avoid loss of data due to nonperformingcamera traps Immediately after removal of camera trapsfrom the field all photographic data were downloaded andfurther sorted into species name size time and habitat typethe camera trap was deployed in Further duplicates wereremoved where thirty (30) minutes after the last photo wastaken was considered as another photographic event

After data cleaning species diversity and compositionusing Shannon Weiner index (where if index is 5 it implieshighest diversity and if it is 1 it implies lowest diversity)[16] and Hierarchical Cluster Analysis [17] respectively wereperformed in Paleontological Statistics Software Package forEducation and Data Analysis (PAST) software while habitatpreference or avoidance was performed using Jacobsrsquo index[18] Further Jacobsrsquo index values were tested for signifi-cance preference or avoidance (where +1 implies maximumpreference while minus1 implies maximum avoidance) using 119905-tests against means of zero (0) in R version 312 Prior to

these tests normal distribution and homogeneity of variancewere performed to ensure that conditions for parametric testswere not violated [19] Overlay of grids on the spatial extentof the conservancy was performed in Quantum GeographicInformation Systems (QGIS) platform version 282

3 Results

31 Wildlife Species Diversity and Evenness ShannonWeinerindex revealed highest wildlife species diversity in E divi-norum dominated habitats (1198671015840 = 2291) followed by Adrepanolobium habitats (1198671015840 = 2058) mixed bushland (1198671015840 =1728) and least wildlife species diversity in open grassland(1198671015840 = 1715) Wildlife species evenness (119867119878) was highest inarea under E divinorum (119867119878 = 05201) followed by mixedbushland (119867119878 = 03751) and then closely followed by Adrepanolobium (119867119878 = 03404) and finally open grassland(119867119878 = 02647) as shown in Figure 2

32 Wildlife Species Composition Hierarchical Cluster Anal-ysis (HCA) compared wildlife species composition (commonwildlife species) across the four habitat types namely E divi-norum A drepanolobium mixed bushland and open grass-land Results revealed that E divinorum and mixed bushlandhabitats shared 45 similarity in wildlife species composi-tion Additionally E divinorum habitat and mixed bushlandshared 39 similarity in wildlife species composition with Adrepanolobium dominated habitat On the other hand opengrassland habitat shared 27 similarity in wildlife speciescomposition with three habitat types namely E divinorumA drepanolobium and mixed bushland as shown in Figure 3

33 Habitat Preference or Avoidance by Various Feeding Guildsin OPC Jacobs index revealed that carnivores preferredA drepanolobium (119863 = 0469) mostly followed by opengrassland (119863 = 0327) and least preferred mixed bushland(119863 = 0066) but avoided E divinorum dominated areas (119863 =minus0698) (Figure 4(a)) Grazers showed great preference foropen grassland (119863 = 0773) and A drepanolobium (119863 =0040) but avoided E divinorum (119863 = minus0917) and mixedbushland (119863 = minus0192) habitats (Figure 4(b))

4 International Journal of Ecology

A d

rep

E d

ivin

orum

Mix

ed b

ushl

and

Ope

n gr

assla

nd

02

03

04

05

06

07

08

09

10

Sim

ilarit

y

Figure 3 Percentage similarity in wildlife species composition of 4 habitats in OPC

minus08

minus06

minus04

minus02

0

02

04

06

Jaco

bsrsquo i

ndex

for c

arni

vore

s

Adrepanolobium

Edivinorum

Mixed_bushland Opengrassland

Habitat types

(a)minus12

minus1minus08minus06minus04minus02

002040608

1

Adrepanolobium

Edivinorum

Mixed_bushland Opengrassland

Jaco

bs in

dex

for g

raze

rs

Habitat types

(b)

0

minus08

minus06

minus04

minus02

02

04

06

08

Jaco

bs in

dex

for b

row

sers

Habitat types

Adrepanolobium

Edivinorum

Mixed_bushland Opengrassland

(c)

Adrepanolobium

Edivinorum

Mixed_bushland Opengrassland

minus1minus08minus06minus04minus02

002040608

Jaco

bs in

dex

for m

ixed

feed

ers

Habitat types

(d)Figure 4 Habitat preference or avoidance (Jacobsrsquo index) for (a) carnivores (b) grazers (c) browsers and (d) mixed feeders in OPC

Browsers showed preference for A drepanolobium (119863 =0674) and mixed bushland habitat (119863 = 0175) but avoidedboth E divinorum (119863 = minus0673) and open grasslands (119863 =minus0116) dominated habitats (Figure 4(c)) However mixedfeeders preferred habitats dominated by open grassland (119863 =0688) and A drepanolobium (119863 = 0523) but avoided Edivinorum (119863 = minus0858) and mixed bushland (119863 = minus0420)

dominated habitats (Figure 4(d)) A total of thirty (30)wildlife species were recorded as per the table in the sup-plementary material showing detection and nondetection ofwildlife species in various habitat cover types

Mean preference or avoidance revealed that E divinorumand mixed bushland were avoided however E divinorumwas significantly avoided (119905

1= 2253 df = 3 and

International Journal of Ecology 5

minus05

05

Jaco

bsrsquo i

ndex

mea

ns

E div m_bushland OGA drepHabitat types

Figure 5 Jacobs index means for all guilds across the habitat types

119901 lt 001) compared with mixed bushland (1199051= 2353 df

= 3 and 119901 = 027) On the other hand A drepanolobiumand open grassland were both preferred by all feedingguilds however A drepanolobium dominated habitats weresignificantly preferred (119905

1= 2353 df = 3 and 119901 = 003)

compared to open grasslands (1199051= 2353 df = 3 and 119901 =

006) as shown in Figure 5

4 Discussions

Wildlife species diversity and evenness were highest in Edivinorum dominated areas compared to other habitat typesThis is attributable to existence of resource rich patchesthat are hardly accessed by most wildlife species As suchattraction of several wildlife species into this dense vegetation(E divinorum) contributed to higher diversity chiefly smallcarnivores Animals use cues such as landscape structureand vegetation structure like tree species phenology resourceavailability (probability of encounter quantity and quality)and predatorparasite risks among others in order to explorea given habitat type This implies that spatial and temporalhabitat heterogeneity drives faunal species diversity andcomposition from local to regional to global scales Henceaggregation of resources may create dynamic impacts wherethese small sitespatches become a focus for higher propor-tion of faunal diversity and composition [20] Accordingto Sirami et al [21] savannah ecosystems are diverse instructure and floral composition hence different guilds willuse different habitats based on their requirements

Animals have different preference to certain habitats [22]as a function of direct and indirect effects of prey availabilitydetectabilitycover and resource availability [23] Spatialheterogeneity of ecosystem is important for maintenance ofdiverse wild animal species and acts as buffer against changesin resources availability for wild animals in era of climatechange [24] On the other hand woody encroachment canpotentially change this heterogeneity and affect wide range ofwild animals

Similarity in wildlife species composition was highestbetween E divinorum and mixed bushland habitats Thesetwo habitat types have nearly homogeneous vegetation struc-ture (dense canopy) This phenomenon is an illumination tohigher percentage similarity in wildlife species compositionin both E divinorum and mixed bushland dominated areasdue to nearly similar structure of the vegetation (canopy anddense bushland) providing ideal habitats for wildlife specieswhich require such areas According to Dupuch et al [25]

prey and predator can explore freely across habitats thatdiffer in resource quantity quality and inherent habitat risksAs such predatorsrsquo movement and habitat selection are asa function of prey density competition level cover andinherent risks while prey habitat selection and movement arebalance of forage quality (benefits of foraging) and risk ofpredation [26]

On the other hand habitats dominated by A drepanolo-bium and open grassland were preferred by feeding guildsmajorly herbivores due to their associated lesser predationrisk This is chiefly influenced by presence of relatively openlandscapes allowing prey to detect predators from a distanceand hencemaintain safer flight distance (improved visibility)The phenomenon is consistent with other findings accordingto Dupuch et al [25] where prey habitat selection is largelyinfluenced by the need to minimise predation risks overforage quality As such these factors influence preferenceof habitats that offer significant benefits for most wildlifespecies

5 Conclusion and Recommendations

The study revealed that bush encroachment by E divinorumhad impact on species diversity composition and habitatselection In particular species diversity was highest in Edivinorum dominated areas and least in open grasslands apositive impact albeit long term impacts are unknown Like-wise percentage similarity in species composition was high-est between E divinorum and mixed bushland dominatedhabitats Further feeding guilds avoided areas dominatedby E divinorum but preferred A drepanolobium and opengrassland dominated habitats hence bush encroachmentby E divinorum affected habitat suitability thus it has anegative impact Based on this finding active managementof the encroaching species is recommended Further stud-ies on activity budget of wildlife species to ascertain howhabitat types are being used are key Additionally long termstudy to provide insights into the effect of seasonality onspecies diversity dynamics across various land cover typesin the conservancy is crucial This information will be vitalin designing habitat management protocols for sustainableecosystem management

Conflicts of Interest

The authors declare no conflicts of interest

Acknowledgments

This study was funded by Rufford Foundation United King-dom (UK) Grant no 19422-1 The authors are grateful toCarol Ngwrsquoeno and Bernard Chira from Ol Pejeta Conser-vancy (OPC) for allowing them to carry out research intheir property and Andrew Bradley and Dan Smith bothfrom Bangor University for their help in camera trap imageclassification

6 International Journal of Ecology

Supplementary Materials

Table 1 Wildlife species detection or nondetection across thevarious habitat cover types (Supplementary Materials)

References

[1] A T Hudak C A Wessman and T R Seastedt ldquoWoodyoverstorey effects on soil carbon and nitrogen pools in SouthAfrican savannardquo Austral Ecology vol 28 no 2 pp 173ndash1812003

[2] S Archer T W Boutton and K A Hibbard ldquoTrees in grass-lands biogeochemical consequences of woody plant expan-sionrdquo inGlobal Biogeochemical Cycles in the Climate System pp115ndash137 Academic Press San Diego 2001

[3] I A Dickie S A Schnitzer P B Reich and S E Hobbie ldquoIsoak establishment in old-fields and savanna openings contextdependentrdquo Journal of Ecology vol 95 no 2 pp 309ndash320 2007

[4] S R Wangen and C R Webster ldquoPotential for multiple lagphases during biotic invasions Reconstructing an invasion ofthe exotic tree Acer platanoidesrdquo Journal of Applied Ecology vol43 no 2 pp 258ndash268 2006

[5] D F Joubert A Rothauge andG N Smit ldquoA conceptual modelof vegetation dynamics in the semiarid Highland savanna ofNamibia with particular reference to bush thickening byAcaciamelliferardquo Journal of Arid Environments vol 72 no 12 pp 2201ndash2210 2008

[6] M Sankaran J Ratnam and N Hanan ldquoWoody cover inAfrican savannas The role of resources fire and herbivoryrdquoGlobal Ecology and Biogeography vol 17 no 2 pp 236ndash2452008

[7] G Oba E Post P O Syvertsen andN C Stenseth ldquoBush coverand range condition assessments in relation to landscape andgrazing in southern Ethiopiardquo Landscape Ecology vol 15 no 6pp 535ndash546 2000

[8] O W Van Auken ldquoShrub invasions of North American semi-arid grasslandsrdquo Annual Review of Ecology Evolution andSystematics vol 31 pp 197ndash215 2000

[9] D Ward ldquoDo we understand the causes of bush encroachmentinAfrican savannasrdquoAfrican Journal of RangeampForage Sciencevol 22 no 2 pp 101ndash105 2005

[10] A T Hudak and C A Wessman ldquoTextural analysis of highresolution imagery to quantify bush enroachment in MadikweGame Reserve South Africa 1955-1996rdquo International Journalof Remote Sensing vol 22 no 14 pp 2731ndash2740 2001

[11] B J Wigley W J Bond and M T Hoffman ldquoBush encroach-ment under three contrasting land-use practices in a mesicSouth African savannardquo African Journal of Ecology vol 47 no1 pp 62ndash70 2009

[12] D R Towns I A E Atkinson and C H Daugherty ldquoHave theharmful effects of introduced rats on islands been exaggeratedrdquoBiological Invasions vol 8 no 4 pp 863ndash891 2006

[13] G Dalle B LMaass and J Isselstein ldquoEncroachment of woodyplants and its impact on pastoral livestock production in theBorana lowlands southern Oromia Ethiopiardquo African Journalof Ecology vol 44 no 2 pp 237ndash246 2006

[14] R Emslie ldquoDiceros bicornis The IUCN Red List of ThreatenedSpecies 2012rdquo 2012 eT6557A16980917 Downloadedon 17April httpdxdoiorg102305IUCNUK2012RLTST6557A16980917en 2016

[15] J Blanc ldquoLoxodonta africanaThe IUCNRed List ofThreatenedSpecies 2008rdquo2008 eT12392A3339343 httpdxdoiorg102305IUCNUK2008RLTST12392A3339343enDownloadedon 17 April 2016

[16] C E Shannon and W Weaver The Mathematical Theory ofCommunication University of Illinois Press Chicago 1949

[17] H G Gauch Multivariate Analysis in Community EcologyCambridge University Press Cambridge UK 1982

[18] J Jacobs ldquoQuantitative measurement of food selection - Amodification of the forage ratio and Ivlevrsquos electivity indexrdquoOecologia vol 14 no 4 pp 413ndash417 1974

[19] F Palomares M Delibes E Revilla J Calzada and J MFedriani ldquoSpatial ecology of iberian lynx and abundance ofeuropean rabbits in southwestern spainrdquo Wildlife Monographsvol 148 pp 1ndash36 2001

[20] D J D Natusch J A Lyons G Brown and R Shine ldquoCom-munally nesting migratory birds create ecological hot-spotsin Tropical Australiardquo PLoS ONE vol 11 no 10 Article IDe0162651 2016

[21] C Sirami C Seymour GMidgley and P Barnard ldquoThe impactof shrub encroachment on savanna bird diversity from local toregional scalerdquo Diversity and Distributions vol 15 no 6 pp948ndash957 2009

[22] A R E Sinclair S Mduma and J S Brashares ldquoPatterns ofpredation in a diverse predator-prey systemrdquo Nature vol 425no 6955 pp 288ndash290 2003

[23] W J Ripple and R L Beschta ldquoWolves and the ecology of fearCan predation risk structure ecosystemsrdquo Bioscience vol 54no 8 pp 755ndash766 2004

[24] G Wang N T Hobbs R B Boone et al ldquoSpatial and temporalvariability modify density dependence in populations of largeherbivoresrdquo Ecology vol 87 no 1 pp 95ndash102 2006

[25] A Dupuch L M Dill and P Magnan ldquoTesting the effectsof resource distribution and inherent habitat riskiness onsimultaneous habitat selection by predators and preyrdquo AnimalBehaviour vol 78 no 3 pp 705ndash713 2009

[26] P A Abrams ldquoHabitat choice in predator-prey systems Spatialinstability due to interacting adaptive movementsrdquoThe Ameri-can Naturalist vol 169 no 5 pp 581ndash594 2007

Submit your manuscripts athttpswwwhindawicom

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental and Public Health

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcosystemsJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

MeteorologyAdvances in

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental Chemistry

Atmospheric SciencesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Waste ManagementJournal of

Hindawi Publishing Corporation httpwwwhindawicom Volume 201

International Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Geological ResearchJournal of

EarthquakesJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

OceanographyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Journal of Computational Environmental SciencesHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ClimatologyJournal of

Page 4: Impacts of Bush Encroachment on Wildlife Species Diversity ...downloads.hindawi.com/journals/ijecol/2017/5620125.pdf · Impacts of Bush Encroachment on Wildlife Species Diversity,

4 International Journal of Ecology

A d

rep

E d

ivin

orum

Mix

ed b

ushl

and

Ope

n gr

assla

nd

02

03

04

05

06

07

08

09

10

Sim

ilarit

y

Figure 3 Percentage similarity in wildlife species composition of 4 habitats in OPC

minus08

minus06

minus04

minus02

0

02

04

06

Jaco

bsrsquo i

ndex

for c

arni

vore

s

Adrepanolobium

Edivinorum

Mixed_bushland Opengrassland

Habitat types

(a)minus12

minus1minus08minus06minus04minus02

002040608

1

Adrepanolobium

Edivinorum

Mixed_bushland Opengrassland

Jaco

bs in

dex

for g

raze

rs

Habitat types

(b)

0

minus08

minus06

minus04

minus02

02

04

06

08

Jaco

bs in

dex

for b

row

sers

Habitat types

Adrepanolobium

Edivinorum

Mixed_bushland Opengrassland

(c)

Adrepanolobium

Edivinorum

Mixed_bushland Opengrassland

minus1minus08minus06minus04minus02

002040608

Jaco

bs in

dex

for m

ixed

feed

ers

Habitat types

(d)Figure 4 Habitat preference or avoidance (Jacobsrsquo index) for (a) carnivores (b) grazers (c) browsers and (d) mixed feeders in OPC

Browsers showed preference for A drepanolobium (119863 =0674) and mixed bushland habitat (119863 = 0175) but avoidedboth E divinorum (119863 = minus0673) and open grasslands (119863 =minus0116) dominated habitats (Figure 4(c)) However mixedfeeders preferred habitats dominated by open grassland (119863 =0688) and A drepanolobium (119863 = 0523) but avoided Edivinorum (119863 = minus0858) and mixed bushland (119863 = minus0420)

dominated habitats (Figure 4(d)) A total of thirty (30)wildlife species were recorded as per the table in the sup-plementary material showing detection and nondetection ofwildlife species in various habitat cover types

Mean preference or avoidance revealed that E divinorumand mixed bushland were avoided however E divinorumwas significantly avoided (119905

1= 2253 df = 3 and

International Journal of Ecology 5

minus05

05

Jaco

bsrsquo i

ndex

mea

ns

E div m_bushland OGA drepHabitat types

Figure 5 Jacobs index means for all guilds across the habitat types

119901 lt 001) compared with mixed bushland (1199051= 2353 df

= 3 and 119901 = 027) On the other hand A drepanolobiumand open grassland were both preferred by all feedingguilds however A drepanolobium dominated habitats weresignificantly preferred (119905

1= 2353 df = 3 and 119901 = 003)

compared to open grasslands (1199051= 2353 df = 3 and 119901 =

006) as shown in Figure 5

4 Discussions

Wildlife species diversity and evenness were highest in Edivinorum dominated areas compared to other habitat typesThis is attributable to existence of resource rich patchesthat are hardly accessed by most wildlife species As suchattraction of several wildlife species into this dense vegetation(E divinorum) contributed to higher diversity chiefly smallcarnivores Animals use cues such as landscape structureand vegetation structure like tree species phenology resourceavailability (probability of encounter quantity and quality)and predatorparasite risks among others in order to explorea given habitat type This implies that spatial and temporalhabitat heterogeneity drives faunal species diversity andcomposition from local to regional to global scales Henceaggregation of resources may create dynamic impacts wherethese small sitespatches become a focus for higher propor-tion of faunal diversity and composition [20] Accordingto Sirami et al [21] savannah ecosystems are diverse instructure and floral composition hence different guilds willuse different habitats based on their requirements

Animals have different preference to certain habitats [22]as a function of direct and indirect effects of prey availabilitydetectabilitycover and resource availability [23] Spatialheterogeneity of ecosystem is important for maintenance ofdiverse wild animal species and acts as buffer against changesin resources availability for wild animals in era of climatechange [24] On the other hand woody encroachment canpotentially change this heterogeneity and affect wide range ofwild animals

Similarity in wildlife species composition was highestbetween E divinorum and mixed bushland habitats Thesetwo habitat types have nearly homogeneous vegetation struc-ture (dense canopy) This phenomenon is an illumination tohigher percentage similarity in wildlife species compositionin both E divinorum and mixed bushland dominated areasdue to nearly similar structure of the vegetation (canopy anddense bushland) providing ideal habitats for wildlife specieswhich require such areas According to Dupuch et al [25]

prey and predator can explore freely across habitats thatdiffer in resource quantity quality and inherent habitat risksAs such predatorsrsquo movement and habitat selection are asa function of prey density competition level cover andinherent risks while prey habitat selection and movement arebalance of forage quality (benefits of foraging) and risk ofpredation [26]

On the other hand habitats dominated by A drepanolo-bium and open grassland were preferred by feeding guildsmajorly herbivores due to their associated lesser predationrisk This is chiefly influenced by presence of relatively openlandscapes allowing prey to detect predators from a distanceand hencemaintain safer flight distance (improved visibility)The phenomenon is consistent with other findings accordingto Dupuch et al [25] where prey habitat selection is largelyinfluenced by the need to minimise predation risks overforage quality As such these factors influence preferenceof habitats that offer significant benefits for most wildlifespecies

5 Conclusion and Recommendations

The study revealed that bush encroachment by E divinorumhad impact on species diversity composition and habitatselection In particular species diversity was highest in Edivinorum dominated areas and least in open grasslands apositive impact albeit long term impacts are unknown Like-wise percentage similarity in species composition was high-est between E divinorum and mixed bushland dominatedhabitats Further feeding guilds avoided areas dominatedby E divinorum but preferred A drepanolobium and opengrassland dominated habitats hence bush encroachmentby E divinorum affected habitat suitability thus it has anegative impact Based on this finding active managementof the encroaching species is recommended Further stud-ies on activity budget of wildlife species to ascertain howhabitat types are being used are key Additionally long termstudy to provide insights into the effect of seasonality onspecies diversity dynamics across various land cover typesin the conservancy is crucial This information will be vitalin designing habitat management protocols for sustainableecosystem management

Conflicts of Interest

The authors declare no conflicts of interest

Acknowledgments

This study was funded by Rufford Foundation United King-dom (UK) Grant no 19422-1 The authors are grateful toCarol Ngwrsquoeno and Bernard Chira from Ol Pejeta Conser-vancy (OPC) for allowing them to carry out research intheir property and Andrew Bradley and Dan Smith bothfrom Bangor University for their help in camera trap imageclassification

6 International Journal of Ecology

Supplementary Materials

Table 1 Wildlife species detection or nondetection across thevarious habitat cover types (Supplementary Materials)

References

[1] A T Hudak C A Wessman and T R Seastedt ldquoWoodyoverstorey effects on soil carbon and nitrogen pools in SouthAfrican savannardquo Austral Ecology vol 28 no 2 pp 173ndash1812003

[2] S Archer T W Boutton and K A Hibbard ldquoTrees in grass-lands biogeochemical consequences of woody plant expan-sionrdquo inGlobal Biogeochemical Cycles in the Climate System pp115ndash137 Academic Press San Diego 2001

[3] I A Dickie S A Schnitzer P B Reich and S E Hobbie ldquoIsoak establishment in old-fields and savanna openings contextdependentrdquo Journal of Ecology vol 95 no 2 pp 309ndash320 2007

[4] S R Wangen and C R Webster ldquoPotential for multiple lagphases during biotic invasions Reconstructing an invasion ofthe exotic tree Acer platanoidesrdquo Journal of Applied Ecology vol43 no 2 pp 258ndash268 2006

[5] D F Joubert A Rothauge andG N Smit ldquoA conceptual modelof vegetation dynamics in the semiarid Highland savanna ofNamibia with particular reference to bush thickening byAcaciamelliferardquo Journal of Arid Environments vol 72 no 12 pp 2201ndash2210 2008

[6] M Sankaran J Ratnam and N Hanan ldquoWoody cover inAfrican savannas The role of resources fire and herbivoryrdquoGlobal Ecology and Biogeography vol 17 no 2 pp 236ndash2452008

[7] G Oba E Post P O Syvertsen andN C Stenseth ldquoBush coverand range condition assessments in relation to landscape andgrazing in southern Ethiopiardquo Landscape Ecology vol 15 no 6pp 535ndash546 2000

[8] O W Van Auken ldquoShrub invasions of North American semi-arid grasslandsrdquo Annual Review of Ecology Evolution andSystematics vol 31 pp 197ndash215 2000

[9] D Ward ldquoDo we understand the causes of bush encroachmentinAfrican savannasrdquoAfrican Journal of RangeampForage Sciencevol 22 no 2 pp 101ndash105 2005

[10] A T Hudak and C A Wessman ldquoTextural analysis of highresolution imagery to quantify bush enroachment in MadikweGame Reserve South Africa 1955-1996rdquo International Journalof Remote Sensing vol 22 no 14 pp 2731ndash2740 2001

[11] B J Wigley W J Bond and M T Hoffman ldquoBush encroach-ment under three contrasting land-use practices in a mesicSouth African savannardquo African Journal of Ecology vol 47 no1 pp 62ndash70 2009

[12] D R Towns I A E Atkinson and C H Daugherty ldquoHave theharmful effects of introduced rats on islands been exaggeratedrdquoBiological Invasions vol 8 no 4 pp 863ndash891 2006

[13] G Dalle B LMaass and J Isselstein ldquoEncroachment of woodyplants and its impact on pastoral livestock production in theBorana lowlands southern Oromia Ethiopiardquo African Journalof Ecology vol 44 no 2 pp 237ndash246 2006

[14] R Emslie ldquoDiceros bicornis The IUCN Red List of ThreatenedSpecies 2012rdquo 2012 eT6557A16980917 Downloadedon 17April httpdxdoiorg102305IUCNUK2012RLTST6557A16980917en 2016

[15] J Blanc ldquoLoxodonta africanaThe IUCNRed List ofThreatenedSpecies 2008rdquo2008 eT12392A3339343 httpdxdoiorg102305IUCNUK2008RLTST12392A3339343enDownloadedon 17 April 2016

[16] C E Shannon and W Weaver The Mathematical Theory ofCommunication University of Illinois Press Chicago 1949

[17] H G Gauch Multivariate Analysis in Community EcologyCambridge University Press Cambridge UK 1982

[18] J Jacobs ldquoQuantitative measurement of food selection - Amodification of the forage ratio and Ivlevrsquos electivity indexrdquoOecologia vol 14 no 4 pp 413ndash417 1974

[19] F Palomares M Delibes E Revilla J Calzada and J MFedriani ldquoSpatial ecology of iberian lynx and abundance ofeuropean rabbits in southwestern spainrdquo Wildlife Monographsvol 148 pp 1ndash36 2001

[20] D J D Natusch J A Lyons G Brown and R Shine ldquoCom-munally nesting migratory birds create ecological hot-spotsin Tropical Australiardquo PLoS ONE vol 11 no 10 Article IDe0162651 2016

[21] C Sirami C Seymour GMidgley and P Barnard ldquoThe impactof shrub encroachment on savanna bird diversity from local toregional scalerdquo Diversity and Distributions vol 15 no 6 pp948ndash957 2009

[22] A R E Sinclair S Mduma and J S Brashares ldquoPatterns ofpredation in a diverse predator-prey systemrdquo Nature vol 425no 6955 pp 288ndash290 2003

[23] W J Ripple and R L Beschta ldquoWolves and the ecology of fearCan predation risk structure ecosystemsrdquo Bioscience vol 54no 8 pp 755ndash766 2004

[24] G Wang N T Hobbs R B Boone et al ldquoSpatial and temporalvariability modify density dependence in populations of largeherbivoresrdquo Ecology vol 87 no 1 pp 95ndash102 2006

[25] A Dupuch L M Dill and P Magnan ldquoTesting the effectsof resource distribution and inherent habitat riskiness onsimultaneous habitat selection by predators and preyrdquo AnimalBehaviour vol 78 no 3 pp 705ndash713 2009

[26] P A Abrams ldquoHabitat choice in predator-prey systems Spatialinstability due to interacting adaptive movementsrdquoThe Ameri-can Naturalist vol 169 no 5 pp 581ndash594 2007

Submit your manuscripts athttpswwwhindawicom

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental and Public Health

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcosystemsJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

MeteorologyAdvances in

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental Chemistry

Atmospheric SciencesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Waste ManagementJournal of

Hindawi Publishing Corporation httpwwwhindawicom Volume 201

International Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Geological ResearchJournal of

EarthquakesJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

OceanographyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Journal of Computational Environmental SciencesHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ClimatologyJournal of

Page 5: Impacts of Bush Encroachment on Wildlife Species Diversity ...downloads.hindawi.com/journals/ijecol/2017/5620125.pdf · Impacts of Bush Encroachment on Wildlife Species Diversity,

International Journal of Ecology 5

minus05

05

Jaco

bsrsquo i

ndex

mea

ns

E div m_bushland OGA drepHabitat types

Figure 5 Jacobs index means for all guilds across the habitat types

119901 lt 001) compared with mixed bushland (1199051= 2353 df

= 3 and 119901 = 027) On the other hand A drepanolobiumand open grassland were both preferred by all feedingguilds however A drepanolobium dominated habitats weresignificantly preferred (119905

1= 2353 df = 3 and 119901 = 003)

compared to open grasslands (1199051= 2353 df = 3 and 119901 =

006) as shown in Figure 5

4 Discussions

Wildlife species diversity and evenness were highest in Edivinorum dominated areas compared to other habitat typesThis is attributable to existence of resource rich patchesthat are hardly accessed by most wildlife species As suchattraction of several wildlife species into this dense vegetation(E divinorum) contributed to higher diversity chiefly smallcarnivores Animals use cues such as landscape structureand vegetation structure like tree species phenology resourceavailability (probability of encounter quantity and quality)and predatorparasite risks among others in order to explorea given habitat type This implies that spatial and temporalhabitat heterogeneity drives faunal species diversity andcomposition from local to regional to global scales Henceaggregation of resources may create dynamic impacts wherethese small sitespatches become a focus for higher propor-tion of faunal diversity and composition [20] Accordingto Sirami et al [21] savannah ecosystems are diverse instructure and floral composition hence different guilds willuse different habitats based on their requirements

Animals have different preference to certain habitats [22]as a function of direct and indirect effects of prey availabilitydetectabilitycover and resource availability [23] Spatialheterogeneity of ecosystem is important for maintenance ofdiverse wild animal species and acts as buffer against changesin resources availability for wild animals in era of climatechange [24] On the other hand woody encroachment canpotentially change this heterogeneity and affect wide range ofwild animals

Similarity in wildlife species composition was highestbetween E divinorum and mixed bushland habitats Thesetwo habitat types have nearly homogeneous vegetation struc-ture (dense canopy) This phenomenon is an illumination tohigher percentage similarity in wildlife species compositionin both E divinorum and mixed bushland dominated areasdue to nearly similar structure of the vegetation (canopy anddense bushland) providing ideal habitats for wildlife specieswhich require such areas According to Dupuch et al [25]

prey and predator can explore freely across habitats thatdiffer in resource quantity quality and inherent habitat risksAs such predatorsrsquo movement and habitat selection are asa function of prey density competition level cover andinherent risks while prey habitat selection and movement arebalance of forage quality (benefits of foraging) and risk ofpredation [26]

On the other hand habitats dominated by A drepanolo-bium and open grassland were preferred by feeding guildsmajorly herbivores due to their associated lesser predationrisk This is chiefly influenced by presence of relatively openlandscapes allowing prey to detect predators from a distanceand hencemaintain safer flight distance (improved visibility)The phenomenon is consistent with other findings accordingto Dupuch et al [25] where prey habitat selection is largelyinfluenced by the need to minimise predation risks overforage quality As such these factors influence preferenceof habitats that offer significant benefits for most wildlifespecies

5 Conclusion and Recommendations

The study revealed that bush encroachment by E divinorumhad impact on species diversity composition and habitatselection In particular species diversity was highest in Edivinorum dominated areas and least in open grasslands apositive impact albeit long term impacts are unknown Like-wise percentage similarity in species composition was high-est between E divinorum and mixed bushland dominatedhabitats Further feeding guilds avoided areas dominatedby E divinorum but preferred A drepanolobium and opengrassland dominated habitats hence bush encroachmentby E divinorum affected habitat suitability thus it has anegative impact Based on this finding active managementof the encroaching species is recommended Further stud-ies on activity budget of wildlife species to ascertain howhabitat types are being used are key Additionally long termstudy to provide insights into the effect of seasonality onspecies diversity dynamics across various land cover typesin the conservancy is crucial This information will be vitalin designing habitat management protocols for sustainableecosystem management

Conflicts of Interest

The authors declare no conflicts of interest

Acknowledgments

This study was funded by Rufford Foundation United King-dom (UK) Grant no 19422-1 The authors are grateful toCarol Ngwrsquoeno and Bernard Chira from Ol Pejeta Conser-vancy (OPC) for allowing them to carry out research intheir property and Andrew Bradley and Dan Smith bothfrom Bangor University for their help in camera trap imageclassification

6 International Journal of Ecology

Supplementary Materials

Table 1 Wildlife species detection or nondetection across thevarious habitat cover types (Supplementary Materials)

References

[1] A T Hudak C A Wessman and T R Seastedt ldquoWoodyoverstorey effects on soil carbon and nitrogen pools in SouthAfrican savannardquo Austral Ecology vol 28 no 2 pp 173ndash1812003

[2] S Archer T W Boutton and K A Hibbard ldquoTrees in grass-lands biogeochemical consequences of woody plant expan-sionrdquo inGlobal Biogeochemical Cycles in the Climate System pp115ndash137 Academic Press San Diego 2001

[3] I A Dickie S A Schnitzer P B Reich and S E Hobbie ldquoIsoak establishment in old-fields and savanna openings contextdependentrdquo Journal of Ecology vol 95 no 2 pp 309ndash320 2007

[4] S R Wangen and C R Webster ldquoPotential for multiple lagphases during biotic invasions Reconstructing an invasion ofthe exotic tree Acer platanoidesrdquo Journal of Applied Ecology vol43 no 2 pp 258ndash268 2006

[5] D F Joubert A Rothauge andG N Smit ldquoA conceptual modelof vegetation dynamics in the semiarid Highland savanna ofNamibia with particular reference to bush thickening byAcaciamelliferardquo Journal of Arid Environments vol 72 no 12 pp 2201ndash2210 2008

[6] M Sankaran J Ratnam and N Hanan ldquoWoody cover inAfrican savannas The role of resources fire and herbivoryrdquoGlobal Ecology and Biogeography vol 17 no 2 pp 236ndash2452008

[7] G Oba E Post P O Syvertsen andN C Stenseth ldquoBush coverand range condition assessments in relation to landscape andgrazing in southern Ethiopiardquo Landscape Ecology vol 15 no 6pp 535ndash546 2000

[8] O W Van Auken ldquoShrub invasions of North American semi-arid grasslandsrdquo Annual Review of Ecology Evolution andSystematics vol 31 pp 197ndash215 2000

[9] D Ward ldquoDo we understand the causes of bush encroachmentinAfrican savannasrdquoAfrican Journal of RangeampForage Sciencevol 22 no 2 pp 101ndash105 2005

[10] A T Hudak and C A Wessman ldquoTextural analysis of highresolution imagery to quantify bush enroachment in MadikweGame Reserve South Africa 1955-1996rdquo International Journalof Remote Sensing vol 22 no 14 pp 2731ndash2740 2001

[11] B J Wigley W J Bond and M T Hoffman ldquoBush encroach-ment under three contrasting land-use practices in a mesicSouth African savannardquo African Journal of Ecology vol 47 no1 pp 62ndash70 2009

[12] D R Towns I A E Atkinson and C H Daugherty ldquoHave theharmful effects of introduced rats on islands been exaggeratedrdquoBiological Invasions vol 8 no 4 pp 863ndash891 2006

[13] G Dalle B LMaass and J Isselstein ldquoEncroachment of woodyplants and its impact on pastoral livestock production in theBorana lowlands southern Oromia Ethiopiardquo African Journalof Ecology vol 44 no 2 pp 237ndash246 2006

[14] R Emslie ldquoDiceros bicornis The IUCN Red List of ThreatenedSpecies 2012rdquo 2012 eT6557A16980917 Downloadedon 17April httpdxdoiorg102305IUCNUK2012RLTST6557A16980917en 2016

[15] J Blanc ldquoLoxodonta africanaThe IUCNRed List ofThreatenedSpecies 2008rdquo2008 eT12392A3339343 httpdxdoiorg102305IUCNUK2008RLTST12392A3339343enDownloadedon 17 April 2016

[16] C E Shannon and W Weaver The Mathematical Theory ofCommunication University of Illinois Press Chicago 1949

[17] H G Gauch Multivariate Analysis in Community EcologyCambridge University Press Cambridge UK 1982

[18] J Jacobs ldquoQuantitative measurement of food selection - Amodification of the forage ratio and Ivlevrsquos electivity indexrdquoOecologia vol 14 no 4 pp 413ndash417 1974

[19] F Palomares M Delibes E Revilla J Calzada and J MFedriani ldquoSpatial ecology of iberian lynx and abundance ofeuropean rabbits in southwestern spainrdquo Wildlife Monographsvol 148 pp 1ndash36 2001

[20] D J D Natusch J A Lyons G Brown and R Shine ldquoCom-munally nesting migratory birds create ecological hot-spotsin Tropical Australiardquo PLoS ONE vol 11 no 10 Article IDe0162651 2016

[21] C Sirami C Seymour GMidgley and P Barnard ldquoThe impactof shrub encroachment on savanna bird diversity from local toregional scalerdquo Diversity and Distributions vol 15 no 6 pp948ndash957 2009

[22] A R E Sinclair S Mduma and J S Brashares ldquoPatterns ofpredation in a diverse predator-prey systemrdquo Nature vol 425no 6955 pp 288ndash290 2003

[23] W J Ripple and R L Beschta ldquoWolves and the ecology of fearCan predation risk structure ecosystemsrdquo Bioscience vol 54no 8 pp 755ndash766 2004

[24] G Wang N T Hobbs R B Boone et al ldquoSpatial and temporalvariability modify density dependence in populations of largeherbivoresrdquo Ecology vol 87 no 1 pp 95ndash102 2006

[25] A Dupuch L M Dill and P Magnan ldquoTesting the effectsof resource distribution and inherent habitat riskiness onsimultaneous habitat selection by predators and preyrdquo AnimalBehaviour vol 78 no 3 pp 705ndash713 2009

[26] P A Abrams ldquoHabitat choice in predator-prey systems Spatialinstability due to interacting adaptive movementsrdquoThe Ameri-can Naturalist vol 169 no 5 pp 581ndash594 2007

Submit your manuscripts athttpswwwhindawicom

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental and Public Health

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcosystemsJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

MeteorologyAdvances in

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental Chemistry

Atmospheric SciencesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Waste ManagementJournal of

Hindawi Publishing Corporation httpwwwhindawicom Volume 201

International Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Geological ResearchJournal of

EarthquakesJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

OceanographyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Journal of Computational Environmental SciencesHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ClimatologyJournal of

Page 6: Impacts of Bush Encroachment on Wildlife Species Diversity ...downloads.hindawi.com/journals/ijecol/2017/5620125.pdf · Impacts of Bush Encroachment on Wildlife Species Diversity,

6 International Journal of Ecology

Supplementary Materials

Table 1 Wildlife species detection or nondetection across thevarious habitat cover types (Supplementary Materials)

References

[1] A T Hudak C A Wessman and T R Seastedt ldquoWoodyoverstorey effects on soil carbon and nitrogen pools in SouthAfrican savannardquo Austral Ecology vol 28 no 2 pp 173ndash1812003

[2] S Archer T W Boutton and K A Hibbard ldquoTrees in grass-lands biogeochemical consequences of woody plant expan-sionrdquo inGlobal Biogeochemical Cycles in the Climate System pp115ndash137 Academic Press San Diego 2001

[3] I A Dickie S A Schnitzer P B Reich and S E Hobbie ldquoIsoak establishment in old-fields and savanna openings contextdependentrdquo Journal of Ecology vol 95 no 2 pp 309ndash320 2007

[4] S R Wangen and C R Webster ldquoPotential for multiple lagphases during biotic invasions Reconstructing an invasion ofthe exotic tree Acer platanoidesrdquo Journal of Applied Ecology vol43 no 2 pp 258ndash268 2006

[5] D F Joubert A Rothauge andG N Smit ldquoA conceptual modelof vegetation dynamics in the semiarid Highland savanna ofNamibia with particular reference to bush thickening byAcaciamelliferardquo Journal of Arid Environments vol 72 no 12 pp 2201ndash2210 2008

[6] M Sankaran J Ratnam and N Hanan ldquoWoody cover inAfrican savannas The role of resources fire and herbivoryrdquoGlobal Ecology and Biogeography vol 17 no 2 pp 236ndash2452008

[7] G Oba E Post P O Syvertsen andN C Stenseth ldquoBush coverand range condition assessments in relation to landscape andgrazing in southern Ethiopiardquo Landscape Ecology vol 15 no 6pp 535ndash546 2000

[8] O W Van Auken ldquoShrub invasions of North American semi-arid grasslandsrdquo Annual Review of Ecology Evolution andSystematics vol 31 pp 197ndash215 2000

[9] D Ward ldquoDo we understand the causes of bush encroachmentinAfrican savannasrdquoAfrican Journal of RangeampForage Sciencevol 22 no 2 pp 101ndash105 2005

[10] A T Hudak and C A Wessman ldquoTextural analysis of highresolution imagery to quantify bush enroachment in MadikweGame Reserve South Africa 1955-1996rdquo International Journalof Remote Sensing vol 22 no 14 pp 2731ndash2740 2001

[11] B J Wigley W J Bond and M T Hoffman ldquoBush encroach-ment under three contrasting land-use practices in a mesicSouth African savannardquo African Journal of Ecology vol 47 no1 pp 62ndash70 2009

[12] D R Towns I A E Atkinson and C H Daugherty ldquoHave theharmful effects of introduced rats on islands been exaggeratedrdquoBiological Invasions vol 8 no 4 pp 863ndash891 2006

[13] G Dalle B LMaass and J Isselstein ldquoEncroachment of woodyplants and its impact on pastoral livestock production in theBorana lowlands southern Oromia Ethiopiardquo African Journalof Ecology vol 44 no 2 pp 237ndash246 2006

[14] R Emslie ldquoDiceros bicornis The IUCN Red List of ThreatenedSpecies 2012rdquo 2012 eT6557A16980917 Downloadedon 17April httpdxdoiorg102305IUCNUK2012RLTST6557A16980917en 2016

[15] J Blanc ldquoLoxodonta africanaThe IUCNRed List ofThreatenedSpecies 2008rdquo2008 eT12392A3339343 httpdxdoiorg102305IUCNUK2008RLTST12392A3339343enDownloadedon 17 April 2016

[16] C E Shannon and W Weaver The Mathematical Theory ofCommunication University of Illinois Press Chicago 1949

[17] H G Gauch Multivariate Analysis in Community EcologyCambridge University Press Cambridge UK 1982

[18] J Jacobs ldquoQuantitative measurement of food selection - Amodification of the forage ratio and Ivlevrsquos electivity indexrdquoOecologia vol 14 no 4 pp 413ndash417 1974

[19] F Palomares M Delibes E Revilla J Calzada and J MFedriani ldquoSpatial ecology of iberian lynx and abundance ofeuropean rabbits in southwestern spainrdquo Wildlife Monographsvol 148 pp 1ndash36 2001

[20] D J D Natusch J A Lyons G Brown and R Shine ldquoCom-munally nesting migratory birds create ecological hot-spotsin Tropical Australiardquo PLoS ONE vol 11 no 10 Article IDe0162651 2016

[21] C Sirami C Seymour GMidgley and P Barnard ldquoThe impactof shrub encroachment on savanna bird diversity from local toregional scalerdquo Diversity and Distributions vol 15 no 6 pp948ndash957 2009

[22] A R E Sinclair S Mduma and J S Brashares ldquoPatterns ofpredation in a diverse predator-prey systemrdquo Nature vol 425no 6955 pp 288ndash290 2003

[23] W J Ripple and R L Beschta ldquoWolves and the ecology of fearCan predation risk structure ecosystemsrdquo Bioscience vol 54no 8 pp 755ndash766 2004

[24] G Wang N T Hobbs R B Boone et al ldquoSpatial and temporalvariability modify density dependence in populations of largeherbivoresrdquo Ecology vol 87 no 1 pp 95ndash102 2006

[25] A Dupuch L M Dill and P Magnan ldquoTesting the effectsof resource distribution and inherent habitat riskiness onsimultaneous habitat selection by predators and preyrdquo AnimalBehaviour vol 78 no 3 pp 705ndash713 2009

[26] P A Abrams ldquoHabitat choice in predator-prey systems Spatialinstability due to interacting adaptive movementsrdquoThe Ameri-can Naturalist vol 169 no 5 pp 581ndash594 2007

Submit your manuscripts athttpswwwhindawicom

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental and Public Health

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcosystemsJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

MeteorologyAdvances in

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental Chemistry

Atmospheric SciencesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Waste ManagementJournal of

Hindawi Publishing Corporation httpwwwhindawicom Volume 201

International Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Geological ResearchJournal of

EarthquakesJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

OceanographyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Journal of Computational Environmental SciencesHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ClimatologyJournal of

Page 7: Impacts of Bush Encroachment on Wildlife Species Diversity ...downloads.hindawi.com/journals/ijecol/2017/5620125.pdf · Impacts of Bush Encroachment on Wildlife Species Diversity,

Submit your manuscripts athttpswwwhindawicom

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental and Public Health

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcosystemsJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

MeteorologyAdvances in

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Environmental Chemistry

Atmospheric SciencesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Waste ManagementJournal of

Hindawi Publishing Corporation httpwwwhindawicom Volume 201

International Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Geological ResearchJournal of

EarthquakesJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

OceanographyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Journal of Computational Environmental SciencesHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ClimatologyJournal of