Arboreality Increases Reptile Community Resistance to Disturbance from Livestock Grazing

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This is the Accepted Version of a paper published in the Journal of Applied Ecology:

Neilly, Heather, Nordberg, Eric J., VanDerWal, Jeremy, and Schwarzkopf, Lin (2018) Arboreality increases reptile community resistance to disturbance from livestock grazing. Journal of Applied Ecology, 55 (2). pp. 786-799.

http://dx.doi.org/10.1111/1365-2664.12982

This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. 10.1111/1365-2664.12982 doi: lead article this todifferences as version between this andthe ofPleasecite Version Record. been and throughtypesetting, proofreadingwhich may thecopyediting, process, pagination This article undergone has for and buthas accepted been not review publication fullpeer ABSTRACT title: Running Articletype : Research Article MS. HEATHERNEILLY (Orcid : ID 0000-0002-2006-2271) Accepted Article disturbance to resistance community reptile increases Arboreality Co 1. llege of ScienceJames and Cook llege of Engineering, University, Townsville, Australia

H. Neilly*, E. Nordberg, J. VanDerWal, and L. Schwarzkopf dwelling fauna has been examined, but there are few studies of arboreal fauna arboreal of studies few are there but examined, been has fauna dwelling ground- of grazing to response the Similarly, known. poorly are habitat arboreal habitat ground-level alters directly grazing livestock Domestic c the be may as them, avoiding or level ground at changes structural the from but species fauna vertebrate of decline the in implicated been has grazing Globally,

s wt abra seis Hr w eaie roel n tretil habitat terrestrial and arboreal examine we Here species. arboreal with ase Arboreal reptiletolivestock resistanceArboreal grazing oe pce apa resista appear species some *Corresponding author: *Corresponding from livestock grazing from livestock t te fet o gaig ete benefiting either grazing, of effects the to [email protected] nt

and arboreal microhabitats. and We showed reptiles Arboreal resi groups. functional or species certain conserving to relating goals include objectives management if particularly management, grazing livestock of impact the to responses to positively responded lizards featuressuch microhabitat treeas hollows. Arboreal areas. grazed heavily in reduced greatly were which structures, ground complex with associated positively were reptiles Terrestrial grazing. heavy by affected negatively were reptiles terrestrial microhabitat and landscape-scale variables. to relation in modelled were responses Species analyses. species-level individual and group functional community, used we assemblages, reptileterrestrial and arboreal of response grazing the compare a at treatments, We conducted arboreal and terrestrial reptile surveys reptile terrestrial and arboreal conducted We Arboreal reptile species were resistant to the impact of grazing, whereas whereas grazing, of impact the to resistant were species reptile Arboreal arb whether determine to grazing, to responses community reptile and responses ytei ad applications and Synthesis stance in a landscape that is grazed, but where trees have not been cleared. cleared. been not have trees where but grazed, is that landscape a in stance highlight the importance of retaining trees in rangelands for both terrestrial both for rangelands in trees retaining of importance the highlight oreal reptile speciesreptileareorealmoreresista 19 ya eprmna gaig ra i nrhr Australia. northern in trial grazing experimental -year . Arboreal and terrestrial reptiles have different different have reptiles terrestrial and Arboreal . nt terrestrial species.reptilethan . This has implications for rangeland for implications has This on four different grazing different four To This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. (Read 2002; Read& Cunninghamgrazing Germano, 2010; & Saslaw Rathbun 2012) p reptilesgrazing, showtheir and some dueAustralia responseNorth America, to no to grazing ordetectableresponse represented to no anincreasebe inby abundance, livestock (Carpentergrazing benegatively impacted etal. (i.e.by) not resistance grazing.aof degree Here, suggests to community,a lack or Within fauna a of the ofincreaserspecies, decreaserpresence species, inabundance increase (‘increasers’), havea negativeeffect(‘decreasers’) t a effect impactscan positive species, Grazing have 2016). them Schwarzkopf on causing (Neilly, ability access & avoidpredation, Vanderwal on their to andfood, thermoregulate vertebrate affect the structure live environments,fauna that in grazed potentially impacting Alterations in habitat exactlyhabitat.however, unclear effects grazing arboreal how is, It leaflitter(Eldridge,& James 2011; Val and debris Dorrough& RamseyBrown, 2011). vegetation,woodysuchas structures ground-level Grazinghabitat 1988). alters Lauenroth (Milchunas, onthe grazing muchthat by species& nicheshow that relies are Sala affected (Donald,abundance& 2001)HeathGreen and 2004) domesticby livestockGrazing across occurs Introduction reptiles, rangelands, Keywords: Accepted heavy promoted by theground higheropen reference and habitats for temperatures Article and is implicated as a contributing factor inandasimplicatedcontributing isofvertebrate a decline the factor species

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response mechanisms, agriculture, arborealresponse mechanisms, , habitat fragmentationhabitat 25% landEarth’s of the surfac . Thewillextentdependimpactofgrazing on ‘resi , herpetofauna, off-reserve conservation, herpetofauna, off-reserve stance ’ means the ability to toleratetheto means ability 2001 , e (Asner e or havenoeffect. or ) . Resistance Resistance . . et al. ’ richness In arid In

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This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. low branches saplings (Fleischnerconsuming or trampling 1994), compaction soil livestock impacts. However,affectground-based indirectlythrough treesmay obvious, the compared on grazing to effect habitatreceived The of arboreal has less attention other resources. forand competition food reduced whereTheymay benefit layer. also habitat other species have declined due to u Lindenmayer loose Therefore, and & they bark (Gibbons be 2002). may branches, hollows,treeupper-strata generally cracksuse suchfissuresin microhabitats dead as and are Arborealare abundant reptiles addressedin reptiles. explicitly re b towhichdegree The ‘arboreality’ influence niches by areas these usingother strata,avoid habitat examplearborealfor graz species those toresistant Bull Bylo,(Pettigrew Overall, & 2014; & Koper Molloy 2014). etal Kutt 1990; species(Currygroupmay level,grazingfunctional & Hackeror heavy facilitate predation (Martin Ata diversity, cleared providedtrees not 2007; & McIntyre &are Lusk 2013). Koper grazingby communities orincrease inremaineven showLikewise, bird can an unaffected naffected by ground-level disturbances because they spend little time in ground-levelaltereddisturbances spendthe they becauselittle naffected by timein bird where exploredforaging een birds, for height used has been topredict species Acceptedyet buthas tograzingto be sponse & (Martin Possingham & 2005; Kutt Martin 2010), Article as grazed by domestic livestock (Woinarski & Ash 2002; Knox, Cree & grazedand as Seddonby domestic & 2012) 2002;livestock(Woinarski Knox,Ash Cree ing either benefit directly or indirectly from the structural changesstructural ing indirectlyeither directly atbenefit or level, or ground thefrom or . 2013; Piana & Piana 2013; Marsden 2014) consuming palatable shrubs (Jones 1981). Furthermore, soilFurthermore,compaction(Jones 1981). consuming palatableshrubs s a speciesa s or affect predator avoidanceaffect strategies ’ response to grazing response has to impacts, in (P disturbed areas, includingdisturbed areas, itt et al. 1998), breakingetal. itt 1998), .

This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. Acceptedgrazing.effectsresistanceexhibittheof to to grazing of levels litterground-dwelling (e.g., skinksheterogeneity individualspecies. and predictgroups We reptilesrespond terrestrial aim arborealmoreWe impacted than be would habitat. hypothesising groundhabitat and on treatmentsthat grazing ground-level arboreal habitat, mammalianand fauna avian tomoreopposed vagile grazing atand,thisoperatesscalesappropriate trial,as importantly, size abundant tothe of woodlandsavannaAustralian, tropical cattlestrategies grazing long-running,at different a grazing experimental trialina northern four weinvestigate Here response the ofarboreal reptileand communities terrestrial to Article 2007). McIntyre (Gibbonsarboreal removingfauna impacting by habitat grassclearingdirectlygrowth,withtopromoteis livestock inTree used conjunction grazing managementsuch landas clearing. treerespond grazing-related techniques, also may to species Arboreal Blesky1997). todesertificationBlumenthal leading & (Fleischner1994; begins communityincludingstructure, tochangetrees, and absorption vegetation the 2000).waterin decrease A nutrient water(Yates, and loss evaporative Norton& Hobbs grasses, Valone& ground,lack(Castellano Bare 2007).created systems of a by herbaceous foliage, erosion, ultimatelyand aleads to excess run-off, in water infiltrationdecrease root to or even leaf litter, can leadleaf or can soiltoincreasedeven litter, temperatures, resultingin high versely,predict we . Con ed

to the grazing treatments as a community, as grazing a as functional the treatments . Thediverse andis reptile at thislocationcommunity ed ed that arborealmorelikely may reptile speciesbe that that those species that relied on ground-level that relied species on those that ) w to identifyand arboreal to how ould likelyrespond highernegatively to ould . Wemeasured the effectofdifferent ed & Lindenmayer 2000; Martin &Lindenmayer 2000; Martin

This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. Speciesand vegetationtype,adjacentsitestreatment were atleastapart. 400m a Within vegetationfrom 100m boundaries 200m and wateringcattlefrom points. sitesreflectsThewere communities their located relativeareaat least within eachpaddock. TheSilver-leafin Ironbark. eight ofsitesdifferentnumber inBox and vegetationthe River 16inReid Additionally, were treatments. sites located in differentvegetation types; siteswere located with1-ha sixestablished, in each Twenty-four ofthe grazing sites four R t ( grass species and hasof Currantanpatchily distributedunderstorey community Bush melanophloia RiverReid Box( types: O’Reagain, 2009; Bushell& Holme cattlestrategies production, prof on grazing determineFisheriestoeffectand Department Queensland the ofAgriculture ofseveral WGT The wassummerestablished a andseason in 1997with wet the winter dryby season. Averageannual rainfall extensivestudyatsurrounded by the grazing. cattleis 643mm, site TheQueensland, station Charters1040hatrialand is Towers, Australia. cattle near Grazingcommercial a wasTrialconducted study (WGT), located Wambiana aton theThis trialWambianagrazing Materials andmethods Carissa ovata herefore each treatment paddock was replicated twice (Tablepaddock1).each was treatment replicatedhereforetwice Acceptedsurveys eptile Article ; s ; ). Eight paddocks were randomly assigned one of four grazing treatments, paddocks randomlywere assigned ).Eight offour one

ee Kutt et al. 2012al. vegetationfordescriptions).vegetation et Kutt Each ee community Eucalyptus brownii)Eucalyptus

s 2011). s two vegetationWGT Thedominant 2011). consistsof and Silver-leaf Ironbark ( Ironbark Silver-leaf and itability,conditionland and (O’Reagain ptus Eucaly etal.

This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. c met variation- 1100Due in checked trappingand h). to eachthen shelters morning (0700 prior ACBsset allowing nights). were surveys, trap dayto the animalsto utilizea theup time andlizards in Aprilnocturnalarboreal of2304 both October(totaldiurnal and only of 2015 of Nordbergwere tomonitor(ACBs;Schwarzkopfpopulations boards used cover2015) & Insitesurveys addition, (5.3 spotlight per between man-hours) 2014 2015. and searchedarboreal reptiles.We trees 16 conducted observers ground, for the and bushes where spotlightingconductedtwice session, was ateach trapping nocturnal per site, Timed Arborealsurvey reptile atthe andsite ofcapture measured then released identified,weighed, animalsweretwice Captured checked daily. removed traps, from and opened eachPitfall offence. funneland3 ofthe werefor10 endsnights traps at the x(18cm x placedside18cm and x two 79cm), either 6 traps length); funnelofthe drift fence in comprised:a and arraysite,at 10mtrap x30cm each wasset-up diameter4 pitfall buckets At dry October(endand2015, April season). and wetof (end in 2014 ofthe season) the surveys conducted were occurred Thesein Four to assess the terrestrialreptilecommunity. surveyTerrestrialreptile treatments.among compare andprimary aim our wassite to detectability ommunity analysis, but was used individualspecies foranalyses. used analysis, but ommunity was Accepted Articletervals hods and survey dates, capture data of captureof anddata hods survey dates, in a ‘T’ina configuration, intersected fenceby drift wa s assumeds tobeequal standardised as survey were methods used at each

Cryptoblepharusaustralis, .

(one 20mlength 10mone and was excluded from excluded was 24 arboreal This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. Pairwise madecharacteristics.were comparisons Tukey using variab of andcover mean on thestructural vegetation grazing treatment effectstype the of analyzedhabitat t featureswereStructural using Habitat characteristics residuals. lsmeans pairwise modelusing optimal werethe comparisons made theTukey terms the of in testin InformationAkaike Criterion (AICc) using the Where relevant,2016). optimalwere models determinedby W DataAnalysis includingcanopy(Tablesite, numberofthe and cover,2). dead hollows trees werecharacteristics Arborealhabitat throughoutmeasured. in overstory measured trees diameteridentifiedwere (DBH)at theirbreast transects and height and was c such asground,bare litter, features groundgrass cover, etc.)leaf (e.g., cover were site surveys. At each complexity Structural of featuresw microhabitat surveysMicro-habitat ategorized along each transect (Table 2) alongeach ategorized transect

Acceptedcomplementarya univariatemultivariateanalysesusedR (R and Core rangeof e Team, in Article les inarborealles strata valuesmeanhabitatfor lower the (ground as level) well as

(Lenth 2016) and thefinal(Lenth byand modelswere examining validated thedeviance 2016)

, 3 x 100 m parallel transects, 50m apart,were mparallel x established transects, 50m 3 100

. Alleither treesonside ofthe metre withinone dredge as wo measured each the reptilemeasured during four of -way analysis ofvariance toinvestigate-way analysis function in comparing modelsbased onthe ’s tests. M uMIn

(Bartoń 2015), (Bartoń 2015), . Terrestrial This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. Acceptedco-efficientsGLMmodel a from standardised the responsewecommunity to plotted grazing reptileTovisualisecommunitiesmodelled overall separately. weretheterrestrial Arborealas explanatory variables.and and(and the interactions) year their season treatment,binomial type, negative vegetation weredistribution applied,with grazing mixed speciesthatsites. doeswereThis presentof infunction allowat leastfive not abundance for among communitiesvariance the tothe overall modeleach relativecontribution toindicate speciesfor their in thecalculated were Univariate statisticsandtest p-values can forbe function used testing. hypothesis GLMsMultiple analyses. fitted variables are tomany simultaneouslyand an mvabund generalised linear using the (GLM), modelfunction multivariateextension ofa werea explored using and Arboreal compositionalterrestrialcommunity differences Articlecompositioncommunity Reptile inincluding model.them before the as( random effects season in relationgrazing richness treatments to vegetationand (fixedtypeeffects), year with and species toexamineabundancewere overdispersion) and (accounting for used distribution species.Generalisedlinearwith abinomial terrestrial negative (GLMM) mixedmodels arboreal andwere for session collated (n=96) trapping species fora and richness Abundance abundanceReptile and richness effects models and so each year was analysed separately. Multivariate GLMsMultivariateeffects with anda models yearso each wasseparately.analysed (Wang et al. 2012). et lme4

Thisanalysismultivariate an alternativeis todistance-based

; Bates . We constructed a site- Wea constructed etal . 2015 ) . Variables were. Variables explored collinearity for by -species table populated by thetable-species by populated anova.manyglm manyglm in in

This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. Acceptedarborealonly connectivityand two characteristicsCanopy habitat Conversely, of# (% 10 cattlelargelowby leadwith complexity structurallitter areas to groundbareand consumptionthereduced leafH, inand greatly areasIn ofgrasses highstocking with rates. coarseleaflitter,and woody (Fig. Grass 1). grass height, cover, alltreatment were debris eightofSixgrazing terrestrial reptiles. featureswerehabitat significantly affected by available terrestrialahadtreatment to Grazing effect on major structuralcomplexitythe CharacteristicsMicro-habitat Results wadistributionbinomial negative year seasonasand using randomGLMMs effects ( graz Articlegro groups functional analysed terrestrialand two arboreal We terrestrial variables(Table microhabitat 2). mo wereofthe analyse usedwith a binomial to responses GLMs, negative distribution, the Individualand species functional group responses Wilson(2015). followed LASOO penalties with toa create st abundant arboreal and terrestrial species, and and abundant st arboreal terrestrial species, and up: arboreal geckos (see Appendix S4 in Supporting Information). Their to responses in SupportingAppendixS4 Information). (see arboreal up: geckos ing variables)analyseding were treatmentvegetationwithand (landscape-scale type

: diurnal litter skinks and terrestrial geckos, andskinksdiurnal arboreallitterand one terrestrial geckos, functional

‘heat s appliedwhere appropriate. - map’ (Brown et 2014) et al. (Brown

lme4 fu nctional groups,to nctional relevantthe ; Bates2015)etal. . Reptile speciesReptiletaxon . A poissonA . or om

y This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. mo optimal Modelabundance richnessmodelthe for coefficients model. null wasreptile the statisticallywas not it distinguishablefrom The optimal forterrestrialreptile modelsignificant. richnessincluded vegetation, although secondvariables, instatisticallythe two model,included best these although wasnot 3 supportedFig. higher(Table 3, in Handthiscase, abundances V vegetation type,however, significantlyreptile Arboreal abundancebyaffected was also grazing treatmentand Ironbark.than the terrestrial higher reptile abundance t significantlysitesHIronbark in the lower Overall, had terrestrialreptileall abundance than inbutH, wereterrestrial differenthigher reptile not each thanfrom other. abundance allR andhad (TableFig.vegetation3, V and M, significantly treatment grazing 3a). type optimalreptileTheGLMMabundance were included forterrestrial species observed. arboreal624of8 ACB reptiles trap 288 nights, 57.6ofspotlighting and Over observed. hours Ove abundanceand Reptile species richness S1, Table Appendix 1). (Fig.> grazing 30DBH)cm2, Trees were treatments significantly differentamong the he other grazing treatment and vegetation type combinations. The BoxvegetationheotherThe grazing had typetreatment vegetationand type combinations. Acceptedb) Article dels are included(Table1,2). dels in Appendix S2 . r 3840 pitfall and 5760 funnel trap nights, 684 terrestrial reptiles of 18 species were terrestrialfunnel 684reptilespitfalland trapof18 5760 3840 nights, r Furthermore,wasreptile abundance inhigher Ironbark in Box than

the nullthe(∆AICc=0.12). modelarboreal best The . The interactionThe of This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. bydriven strongly significant wasa effect inofseason 2014 ofvegetationand in2015, type In arboreal the reptile abundantcommunity in community this there most species subset. castlenaui dubia arborealwere geckosspecies includedhouse inFour the ( community analysis: was(Appendix S3,detecting differences Figure difficult so 1) treatments among allw abundance species of session, ofall abundance species,end exceptatthe in ofthe season2015. this dry During trapping wascommunitycommunities, with mostdifferent fromIronbark lower thea other 2015. 2014from Onlyto andgrazing interaction changed vegetation however individualspecies of the driving the speciesby same ( driven the individual (Tableresponsesseasonalgrazing The interaction were4). vegetation betweenand an In was more reptilecommunity grazing negative (Figure 4). response than the of arboreal the gecko( ( nobbi active, striped surface diurnal skink ( skinks dwelling, diurnal werespecieslitter- includedsmall, inthree theEight terrestrial community analysis: Community Reptile Diplodactylusconspiculatus

Accepted Articleand season 2014 2015, terrestrial reptileand explainedcommunity by composition was ) ), , ale-headed snakes ale-headed

Lucasiumsteindachneri and a groupa of ground-dwelling nocturnal,and p ),

and

easterngeckos spiny-tailed Carlia munda, Menetia greyii greyii CarliaMenetia munda, H. binoei

( ) Hoplocephalusbitorquatus ,

By ). Overall, the response of terrestrialresponse reptilethe ). Overall, community to noe’s gecko noe’s consistently influenced this interactionthis influenced consistently term as

( Ctenotus robustusCtenotus much lower than at any other time in any other year, atany much lowerthan other timeinany other year, C.munda, robustus, taeniopleuraC. greyii,M. M.

(

Strophuruswilliamsi ( Heternotiabinoei

geckos, including the fat-tailed geckofat-tailed geckos,the including and ) , ), northern ), a diurnaldragona Morethiataeniopleura ), and the box-patterned and ), ) .

Gehyra dubia Gehyra velvetgeckos

( Diporiphora . Gehyra . The H The was

( ) Oedura ,

larger, larger, the the ), This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. by and influenced fine-woody debris associatedandwith grass cover, gecko whereas positively terrestrial ground wasabundance associatnegativelymicrohabitat Litter featuresskinks various were (Table5). d whereasH,terrestrial geckos skinkstypically litterwere showed Overall, abundantinless evident for H were Mlowest and in in abundance 5, in(Tablespecies,and groups geckoand litterfunctional the oftheterrestrial skink four five S4)analysed effectwere significant ofgrazing was separatelya There Appendix (Tableon5). (see terrestrial terrestrialfunctionalspeciesmost andgroups five the The abundant Individualspecies and groupfunctional responses o surveysthewerewellas limitation as that inandtrappingmethoddifference (ACBs), effort C G.dubia abundance.beto theirinfluencedrawndue tothe not relativelyoverwhelming of lowDue G.dubia ry ifferent ingrazingifferent responses to differentvegetation types speciesis conducted examined however this nly in individually 2015,

AcceptedFig. 5a-g Article ptoblepharus australis ptoblepharus , Fitted 4). for value plots (Table individual species communitymore may individual than analysis analysis be appropriate ). H. binoei

Carlia munda (Fig 5b), where abundance was higher in Box in H, M and R inbut V wasBox highernot in (Fig R H,Mwhere abundance 5b), and was not included in the arboreal community analysisarboreal includedwasin not toa due the (Fig. 5a)

and C. . M. M. taeniopleu Theis grazinginteraction between vegetation and ovata H. bitorquatus,O. castlenauiandS.williamsi cover

ra (Fig. 5c) both had highest had (Fig.both 5c) . Individual species responded responded Individualto species .

ed with bare could could . . This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. (i.e.,Hvs. highestcontrastM) treatments theamong largest differences b Inandlarge, overstorycanopy more H, V trees.had lower than significantly connectivity wer treatments the only weConversely, differences inarborealfound featuresamonghabitat grazing the in thisstudy. ground-dwelling herpetofaunaimplications thefor (Dorrough et al. conditions other imp an significantly Inwas wasground-levelaltered, H,microhabitat habitat not arboreal affected. featuresWhile habitat Discussion cavities.positively to and number the and with of treeshollows respondedcomplexgeckos negativelytosmall Arboreal more trees bark). ( BarkwereIndex the positively associated(they moreand commononDBHwith treeswith 5 (trees Ironbark. in moreabundantwere Handwas innosignificant There V. effect ofgrazing abundant on mostandand least M R inabundanttograzing vegetation(Fig. and6a),were and responded arborealseparately Fig. two analysed species(Table Onlybe could 6a-c). 5, oth cases, ifthe havegrazing cases, drivingweremechanism, expect oth thenwewould

Acceptedvegetationofgreat structural d structuralwascomplexity reduced. Diverse is habitat Article ortance to ground-dwelling reptile communities as they create a mosaic anda of thermal reptile create theyortance communitiesas toground-dwelling – 10 cm 10 DBH) whereas e in terms of canopy connectivity and number of Trees > 30 cm DBH. R had had canopyofTrees > number 30DBH.cmand R connectivity in termsof at ground level were significantlyground level modified C.australis 2012 Gehyradubia ). The simplified ground habitat found in H had majorhad in).H simplifiedTheground habitat found was negatively associated with trees 10 associatednegatively was with trees was negatively associated with small associated treeswith negatively was by C.australis grazi . Further, younger Further, ng 5 Gehyradubia treatment,the – . 10cm DBH) Both species Both ed tosee the –

20 cm

This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. thefrom directimpactsThis ofgrazing, negative buffered microhabitatsuch as loss. stockinginincreasing abundance increasing rate with resi re many ground-dwelling where treatments,including heavily grazing the stocked paddocks, most arborealabundantThe reptiles, grazing.by (positively) abundancethe ofindividualspeciestreatment, grazing (namely tofourwas limited species reptilespeciesabundant arboreal muchless were than overallreptilesabundance were atsites, our patterns arboreal driven by a livestock resistantonly reptiles impactsheavy not of Arboreal were to the but had grazing, grazingArboreal to response reptile minimal. is overstory treeson impact arborealand habitat features atake longmay timetorespondgrazing to disturbance,after 19yearsbut WGT, ofthe the in and soildecreasedaswater resultcompaction, infiltration, a offuture, long-term the recruitmentindifference classesof suggesting no treatments,grazing amongthesize treeswere different not n apparent preference for H and V. However, while there was a diverse assemblage whilethere preference However, ofapparentwas a diverse Hand for n V. creased soil temperature (Yates, Norton & Hobbs 2000; Castellano & & & Hobbs2000;Castellano Trees Valone Norton creased 2007). soil temperature(Yates, Acceptedsufferptiles Article stant to the effects of heavy grazing, whereas effectsstant to the ofheavy grazing, ed . Cryptoblepharusaustralis . It is possible that changesis toarboreal that willhabitatIt possible beidentified in . While community composition was communitycompositionstrongly not by While affected G. dubia did not respond to grazing, and respondwas,therefore not did tograzing, and and G.dubia C. australis . Both specieswereBoth apparently G.dubia showed an increaser pattern, an showed , flourish , , G. dubiaG. so our community analysis our so

ed ) G.dubia in all all in was affected wasaffected of the . Most Most This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. ho wildlife,thattrees arethe presentsupport of wideespecially witha oldvariety trees woodlandsWGTWhileopen-canopy thelocation. such have as naturally sparse cover,tree overstory old, trees.Additionally, fireis regularly not tosuppressused woody growthat this and hasmany WGT therefore thegrazing, has notclearedwithin been years 100the last (Pringleoccupancyarborealother oflivestock Unlikeareasforlizards increase used 2008). (W acc (Manning,Cunningham world & Lindenmayer the distributionandwillhavethe of trees major impactsbiodiversity on acrossregions vast of loss As stylesleadingkeystonestructures, management grazing are major declines.to tree etal Pringle Fischer (2009) suggestthat structurecurrent(Ogada 2008). arboreal 2008; etal. can with interact large browsing nativegrazing by herbivores, soil long-term arboreal fauna(Gibbonsto & Lindenmayer 2017).2002; etal. Indirectly, Parsons imp indirect and depending onthefire, case other bethe extent clearing, not elsewhere, of tree arborealcommunitywas not the negatively Here, but may this heavy impacted by grazing, (Smart, & 2003). Whiting (low Twine disturbance) comparedrangeland toadjacentprotected rangelands(highcommunal disturbance area) wereseverallizard arborealin moresimilar species a study, supports abundant where

Accepted llows and & Lindenmayer bark 2012).flaking Bryant, &(Gibbons 2002; Dundas Fleming Article h umulation of course woody debris, arecourse animaldiverseof prime umulation communities for debris, habitat woody acts on arborealon acts habitats compaction iles & Grubaugh 1996). Even damaged trees increase structural complexity, and can Grubaugh increase treesstructural& complexity, damaged Even iles 1996). and , may suppress new tree growth (Fischer, Lindenmayer & Cowling 2004) and 2004) Lindenmayer& new growthCowling suppress (Fischer, tree . Treeclearing, grazing,often with isassociated majorthreata 2006) . Both dead and living trees, andlivingtrees, and the. Both dead resulting in changes to This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. effects. GLMMs, whereweusing year benefit species seasonthe andhad of treating as random community Theown. their is analysis greatly complemented assessment the ofindividualby d responses,speciescommunity atThea group compositionaland individual functional level. most grazingin clearly seenoverall abundance, increasing pressure. Thisrelationship was arboreal reptilesUnlike reptiles, generally terrestrial a association had negative with T (Scheffers 2014). resilience et al. the lizards offrogs and and to extreme events climatic (Martinresponse livestock& effectively Possingham used topredict2005)to bird grazing gradient’ ‘arboreality i morediversearboreal a community, t applied a In wehave binary ouranalyses, reptiles. arboreality ofnotion (either arboreal or strata, lower Hwere usingand By (on not V. stratasmall the shrubs) presentlower in or treesand either resistant effects appear to the ofgrazing. BothspeciesLindenmayer Fitzgerald al. atnight2002; & of trees(Gibbons et 2010). trunks bitorquatus werevegetation not linkedto type. Both grazing and every treatment arb model couldonly We errestrial), but in reality, arboreal species use vertical habitat strataerrestrial),verticalhabitat inIn reality, to but differentextents.species arboreal use errestrial reptile responseerrestrialgrazingreptile to Accepteddifficult interpretation making ofcomplexonifferences weresubtle, results these and Article oreal reptiles on the WGT. For example,orealonthereptiles For WGT. use loose bark and loosebark hollows anddiurnaland use as refugia on forage and branches the williamsi S. and use this as a predictor of resistance to disturbance. This has disturbance.been to Thishas predictor resistance use as thisand of a G. dubia may be less tolerantbe may impactsgrazing tothe of than other arboreal and and C.australis tmay be beneficialalong an toclassify species Strophuruswilliamsi O.castelnaui individually, but made observations of othermadeindividually, observationsofbut and and H. were generally found inthegenerally were found bitorquatus O . castelnaui were found in found were and and H. This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. increaserspecies, hasoftenassemblage a proportion ofhigher locations,At grazing.other reptile the susceptible terrestrial to particularly morearidareas, terrestrialThe reptile assemblageby diurnalwas dominated abundantskinks at locationthis arboreal terrestrialfauna. the and supports furtherimportance features the of retaining treesin grazed environments, forboth importanceThe& Cowling structure oftree-provided Lindenmayer toground2004). arelogs, which ofarboreal fallen structurehabitatand actually a function (Fischer, woodyleafas debris, reptiles litter often dwelling respondto suchcharacteristics habitat Ground-vegetationsurrounding community. tograzing response greatlyis influenced by the 2002), speciesin suggesting areasAsh further of this (Woinarskiabundant heavy& grazing wa gecko responseclearlyterrestrial influencedabundant the The of terrestrialgeckosby most is h likelyofin refugesgrazedand toa reductionthermal atlocations, other duelevel ground negatively inaccordance to responded grazing, response with heavy the of taeniopleura mo 2007). James 2003;2002; Kutt &Woinarski Australiangrazing other & Ash(Woinarski withsimilar studies terrestrialreptile assemblages group,supportingskinks results a performed littertypicaltheAs species, as decreaser from Accepted Article(Hacking,& eavilygrazed areas SchwarzkopfAbom 2014; & Schwarzkopf Abom 2016). s significantly less abundant inabundant less significantly s stly showedstly sameresponsethe grazing.negative increased to H. binoei. were both least abundant in leastheavilyabundantin the sites. stocked both were While seemingly unaffected by grazing in the Box vegetation type, seeminglyunaffected in grazing by While Boxvegetation the type, H Ironbark. In other vegetation types, Ironbark.other vegetationIn types,

The litter skinks that were analysedlitterThe separatelywere skinks that that prefer more open, less thatprefer moreopen, Carliamunda Ctenotusrobustus H.binoei Ctenotus sp. Ctenotus and and is more more is . binoei H M. M. also in in

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This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. (Neilly, & Vanderwal respondoftengroups into disturbancesdifferent opposite)ways (even Speciesand abundancefunctional richness. suchand measures as overall biodiversity just on separately, individual composition and species ratherin more detail, than focusing illustrateresults Our the importance ofexamining and arboreal terrestrial community tograzing. responses individual species behind mechanisms betterA understanding(butinofthedetail examined see etal.Rosi al. 2009; Villar 2013).et aresuggested or mechanisms assumed, very and havebeen few experimentally tested or Pafilis2014).2013; 2012;PettigrewBull responseal.grazing& Seddonet Most Cree & environmentsgrazed& Hackerpredator-prey (e.g. dynamicsCurry changed in Knox, 1990; Reptiles can be abundanceitby reptile andaffected would these. be beneficial totest arborealItlikelyareis theremechanismstoberesistant.complex indirect species driving we complexity, but determinedhave thenot mechanism arboreal allowing or groups t forthermoregulationrequirements,habitat responses interpretationof Our reptile c we lownumbers however,due to capture ground habitatsandtemperatures, open higher One agamid species Saslaw2012). found during study, the Rathbun 2002; & (Read Read groundenvironments 2010; & Cunningham complex Germano, he negative response of many species grazing response henegative ofmany to by loss driven is a ofmicrohabitat Accepted Articlegrazing its detect abundancetreatments. differencesnot in among ould ’ responses itmakepossiblemay to species predict to grazing to knowledgelimited ouris by ofspecies’ , predator avoidancewesuggestand predatorHere, food. D. nobbiD. , would likely prefer would This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. 2006). Therefore,universalone Fischer & Lindenmayer Vickery 2003; & Wilson Manning, increasesofbiodiversity landscape speciesretention and scale richnesstrees (Benton, (Fischer,Itspecies2004).acceptedwidely & CowlingLindenmayer is terrestrial that the indirectly species,extensive but providearboreal for forharbor microhabitats habitat 2016(Neilly,Schwarzkopf rangelands & globally Vanderwal in Australianarebirds,torangelands, found howeverand reptiles, arboreal unique mammals rangelands may similarcommunities Diverse to arboreal from biomes. be reptile relevant extent ofvertical(Asner etal. strata vary in their 2004) aglobe,the systems diversityAcross grazing and support systemsbiomes ofthese livestock 2016).Schwarzkopf & Vanderwal (Neilly, acalls nuancedapproachstrategies for recommendationsto management co arboreal geckosconserving wouldbe different compared toa strategygrazing ataimed grazing devising management strategies attributesofthe influence understanding that suchresistance,as arboreality, can whenhelp frameworksrisk-based rangelandtoguide (Chambersetal. Anmanagement 2017). successfully tobuildused ofplantanimalresistance and Measures resilience and have been Managementimplications appropriately. thuscommunity2016), responseSchwarzkopf importance wehighlight the data ofanalysing

Accepted varied litterresponse ofvertebratesdiurnal nserving The todifferentskinks.grazing Article . Incase,this recommendedthe grazing strategy for . Therefore, our findings willmoreTherefore, ourbe findings ). Furthermore, trees Furthermore,only trees not This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. Accepted files:WambianaData Reptile Community http://dx.doi.org/ availablefrom the Data Repository.DigitalDryad DOI: Data Accessibility gaveand approvalfinal publication. drafts for data, analysed collectedand and conceivedandJV LS EN, EN HN, the designedideasmethodology;and HN Author Article many assistantsO’Reagain,the field P. BushellJ. andand QLD DAF. Lyons thanksto the familytheir for and ‘Wambiana’ support hospitality. of also thankWe projectFunding forthis LS throughwasto awarded Meat Livestock and ManyAustralia. Acknowledgments animal where productionand conservation co-exist. biodiversity can where stocking rangelands ratesare trees moderate,are retained potentially are and areas Unlike moreintenseagricultural(e.g.woody cropping), standing land-uses treesand debris. practicetomanagementor retain arborealincrease terrestrial and be toretain fauna may s’ Contributions ledthe writingmanuscript.ofthe All authorscontributed critically to the HN 10.5061/dryad.fn02v

(Neillyet al.,

2017)

This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. AcceptedvehiclessheepSome Bury, R.B.and Busack, grazing & S.D., (1974) effects on of off-road (2012)Fleming,P.A. G.L.,S.J. Dundas, Bryant, Treehollows& are ofconservation H.Gibb,Cassis, The G.& (2014) fourth- Warton,D.I.,A.M., Brown, Andrew, N.R., Binns, M., G.W.,Brown, Dorrough, (2011)LandscapeRamsey, D.S.L. localJ.W. & and influenceson (2003)FarmlandWilson, J.D. T.G.,biodiversity: Vickery,J.A. is Benton, habitat & Article D., Maechler, M., B.,Bates, Bolker, & multiK. (2015)MuMIn:Bartoń, G.P., Elmore,A.J., Asner, Martin,L.P., R.E.Olander, (2004)systems,GrazingA.T. Harris, & R.& Schwarzkopf, responses Abom, ofreptile tofireinassemblages L. (2016)Short-term References importance for a near-threatened pythonspecies. importance fora near-threatened the environment. with solution corner lme4. using lizard populations in the Mojave desert. populationsin thelizard Mojave Landscapeand Planning Urban woodlands ofsouthernofreptile Australia. patterns occurrencein grazed temperate key?heterogeneity the Resources responses, globalchange.ecosystem and weedyand native tropical savannah. , 29 JournalStatisticalSoftware of , 261 – using using predictivetomodels howunderstand species traits interact – 299. MethodsinEcologyand Evolution TrendsinEcology& Evolution -model inference, R package version 1.15.1-model inference, version R package

Walker, , 103 , 277, GlobalEcology and Conservation S. (2015)S. Fitting linearmodels mixed-effects BiologicalConservation – , AnnualEnvironment and of Review 288. 67 , 1, –

48. JournalZoology of

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6 , 179 , , , 286 – 6 183. , 58- , ,81 66 – 92.

This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. AcceptedMcintyre,J., Dorrough, S.,Barrett, J., G.,Stol,A. Brown,G.& (2012) Differential R.E.Green,Donald, P.F., (2001) AgriculturalM.F. intensification& Heath, and collapseofthe Hacker, & R.B.Can P.J. Curry, pastoral (1990) grazing satisfymanagement endorsed ArticleWuenschel, (2017) A. Pyke,C.S.,J.D., J.C., Boyd, Chambers, Maestas, D.A., & Pellant, M. & (2007)Livestock,M.J., Valone,T.J. compactionwater Castellano, and soil infiltrationrate: S., B.,Walker,Carpenter, Abel,Anderies, & J.M. FrommetaphorN. (2001) measurement: to i intensity N.Koper,Bylo, L.N., K.A.(2014)Grazing& Molloy, Environments a Evaluating potential desertification recovery mechanism. resilience ofwhattowhat? Management inmixed-grass habitat prairies.use badger American 582. agriculture. reptile andto clearing: plant, responses bird ofplants, responses birds reptilesand tograzing fertilizermanagement, and tree Sciences populations.Europe’s farmland bird Management inobjectives conservation aridWestern Australia? 164. Greaterand ecosystems Sage-grouse. resilienceand Using conseptsresistancemanage to persistent threatstosagebrush

, 268

71 ,25 , , 67 30 , 97, – , 247, , 295, 29. – 108.

– – 254. 320. Ecosystems

Proceedings of the Royal Society B: BiologicalB: RoyalProceedingsthe Society of , RangelandEcology& Management 4 , 765-781.

JournalEnvironmental of nfluences ground squirrelgroundandnfluences RangelandEcology& AustralEcology JournalArid of , 37 , 70 , 569, , 149- – This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. AcceptedJ. C.D., Landsberg, Morton,S.R. watering & James, (1999)Provision of pointsin the Hacking,R.Abom,& Schwarzkopf,J., L. (2014)Why avoidweeds?lizards do & (2002)Tree P., Lindenmayer,D. Gibbons, hollowswildlifeconservation and inAustralia. Rathbun, D.J., Germano, ofgrazing G.B.Effects invasiveon Saslaw,L.R.and grasses & (2012) B. Lazell, Fitzgerald, M., R.(2010) Ecology& Shine, conservationthe of and Pale-headed ArticleStott,J., Forrester, R.I. Fischer, Zerger,J., (2009) ReversingG., Warren, A., a Sherren,K. & Lindenmayer,J., Fischer, (2004)challengeA. TheCowling, D.B. &multipleof managing J.D.J., Eldridge, Val, & James,A.I. effects (2011)Abiotic predominate underprolonged Snake ( Snake Sciences of Academy regenerationtree crisisinanendangered ecoregion. Invasions 87 arid Australian revieweffectszone: a on of biota. Collingwood, AU.CSIRO publishing, vertebratesCalifornia. desert in Applied Ecology atmultiplescales: species reptiles in an Australianlandscape. grazing Ecology disturbance: livestock-induced Livestock-induced inwoodlands. disturbance – 121. Hoplocephalusbitorquatus , 36 ,

16 , 367 : 935-947. : , – 41 377. , 106 , 32,

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– , Elapidae). Elapidae). , 10391. The JournalThe ManagementWildlife of AustralianZoologist Journalarid of environments Proceedings of National the , 35 , 283, , Biological Journal of 76 – ,670 290. Austral –

, 682. 41 , This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. AcceptedWoinarski, J.C.Z.A.S. & effects TheofKutt, (2007) and grazing fireonvegetationand the change. vegetation & BirdA.S. foraging Martin, Kutt, T.G. (2010) height predicts speciesbird response towoody A.S.,Kutt, Vanderduys, E.P. & O’Reagain, P. (2012) Spatial and temporal effects grazing of A.S., Vanderduys, J.J., Kutt, E.P.,Perry, Bateman, Perkins,J.E., G.C., Kemp, B.L.,Kanowski,J. Articleindirectgrazingof introduced effects (2012)by C.D., Cree,P.J. &Seddon, A. Knox, andDirect onstandgrazing Belsky, Joy & Blumenthal,(1997)Effectsand A. livestock D.M. dynamics of K.B. (1981)Effects onLizardJones, ofGrazing Abundance and Arizona.Diversity in Western ofvertebratesC.D. (2003)Response James, tofencelinecontrasts in grazing intensity in soils in West.in forestsupland soils of interior the SouthwesternNaturalist JournalTropical of Ecology Australia. inassemblagea woodlandvertebrate savanna in tropicalnorth-eastern Journal management andrainfall on vertebrate the fauna of a tropical savanna. afterlivestockgrazing. years cessation 5 of changeR. (2013) Signals & Jensen, of woodlandinsavanna tropicalvertebrate fauna 46 m semi- ammals on a native, arborealg native, ona ammals , 145, ar , – idwoodlands of Australia. eastern 34 152. , 173 Biodiversity andBiodiversity Conservation

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182. , 26 , 23 , 107 , 95 – ec 115. – ko( 106. Naultinus gemmeusNaultinus , AustralEcology 19 ConservationBiology WildlifeResearch , 2247 – 2262. , ). 28 , JournalHerpetology of ,137 39 , , The Rangeland ,386. 11 – 151. , 315, – 327.

, This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. AcceptedNeilly O.E. Milchunas, Sala, D.G., & Lauenroth, generalizedW.K. (1988)A model effectsof the of T.G. & Martin, Possingham, H.P.(2005) Predictingimpactlivestock the of grazing onbirds livestockclearingonand grazing tree ofT.G.birds & Martin, McIntyre,Impacts of (2007) S. ArticleJ. D.B. Fischer,& Lindenmayer,(2006)Scattered A.D., keystoneManning, are trees Cunningham,A.D., Manning, R.B.,Bringing Lindenmayer, D.B. &(2013) theforward benefits southwesternsurvival nest in songbird Koper, andJ.S. &(2013)GrazingLusk, N. R.V. (2016)Least Lenth, of coarse woody inof debris ecosystem recovery grazing under differentlevels of and Software structures - Implications conservation.- structures for vegetation density. , H., Vanderwal, J. & Schwarzkopf, L. (2016) Balancing and Balancing H.,biodiversity food , Vanderwal,J. L. (2016) & Schwarzkopf, production: betterproduction: A understandingofwildliferesponse will tograzing inform off- 436. conservation reserve on rangelands. Naturalist largeby herbivorescommunitygrazing ongrassland structure. Applied Ecology height data:using foraging Bird foraging impactheight predicts ofgrazing. w Saskatchewan. oodland and riparian and habitats. oodland

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This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. Accepted Vanderduys, S.A., Kutt,Q., Parsons, Perry, E.P., J.J. L. (2017)& Schwarzkopf, Exploring Anastasiou,I.,K. &Pafilis, P., E.D.Valakos, (2013)GrazingSagonas, goatsonislands by Bushell,P., O’Reagain, J. B.(2011)Managing & Holmes, rainfallfor variability: long ArticleBushell,P., A. O’Reagain, Holloway, J., C.&Reid, (2009) Managing rainfallfor variability: Gadd,D.L., Ogada, M.E., R.S., Young, Ostfeld, T.P. (2008)F. Impacts& Keesing, oflarge coverartificial bark boards: Using to L. (2015)Arboreal & Schwarzkopf, E.J. Nordberg, Neilly

H, sample crypticarboreallizards. sample AppliedEcology communityresistance reptile RangelandJournal betweenvertebrate relationships native diversity landgrazing and condition. islands. populations on the affects populationsofanendemic lizard:Mediterranean Grazing affects lizard AnimalProduction Science of profitability strategiesdifferent inaAustraliangrazing northern tropicalsavanna. ProductionScience ofgrazing effect strategycattle on productionin a dry savanna. tropical 387 abundancediversityon andin savanna. an herbivores bird African Nordberg - 397.

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Oecologia Journal Animal

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156 , This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. AcceptedCona,S., I.,Puig, M. I.,Videla,Rosi, M. Mendez,& Roig, F., (2009)E., G. V. a of Diet fossorial grazingcattle animalsonanferalCunningham, J.L. &and Relative impactsR.Read, (2010) of J.L. (2002)Experimental Read, trialofAustralian arid reptilesearly zone as warning Team(2016)A R: R Core environmentand language computing.forstatistical RFoundation ArticleR.M.Pringle, (2008)Elephants agents as creationofhabitat small for vertebrates atthe B.M. D.A Wikeem, Quinton, P.L., R.F.,Youwe, & Newman, M.D., Pitt, R.P.& Marsden,Piana, S.J. Impacts ofcattle (2014) ongrazing structureand forest raptor C.M.Bull, M.capture Pettigrew, & (2014)Prey by behaviour pygmy bluetonguewithlizards for Statistical Vienna, URL Austria.https://www.R-project.org/.Computing, for scale. patch RangeManagement index pressure to predictdamageofregeneratingcattle seedlings.tree 279. Centralin grazing the Monte(Argentina). availability, and(Octodontidae), food changesrelated tocattlerodent above-ground 314 arid reptileAustralian zone smalland mammal assemblage. ofovergrazing indicators cattle. by 559 a withinprotecteddistribution neotropical area. grazing. simulated – –

324. 572. Ecology New ZealandEcology of Journal , ,

89 51 , 26 , 417 – 33. – 422.

Australecology JournalArid Environmentsof Biodiversity andBiodiversity Conservation , , 27 38 ,55 ,45 – – 66. 52. AustralEcology .

(1998)grazinga Using , Journal of 73 , 273 , , 35 , , – 23

, This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. AcceptedAsh,J.C.Z.Responses & Woinarski, ofA.J. (2002) topastoralism,vertebrates military land (2015)S. Wilson, woodyGrubaugh,J.W. (1996)Whiles, ImportanceM.R., &ofcoarse tosouthern debris Article Y., U.,Naumann, Wang, & Warton,S. Wright,D.I. mvabund:(2012). model anR package for X.,Redpath,S. Lambin, Villar, N., Evans,(2013)evidence Experimental D., Pakeman, R.& R.,Whiting, Smart, W. (2005)Lizards M.J., landscapes:& Twine, andintegrating surveysfield Evans,(2014)T.A.Microhabitats B.R., Scheffers, Edwards,D.P., Williams,A., Diesmos, & S.E. Oecologica livestock that intensitygrazing affects activitygeneralistthe ofa predator. London, UK. London, version3.6.11. package of analysis based reptilesselected savannaSouthin a in Africa. disturbanceinterviews onlizard assemblagesand and toassess impactofhuman the 311 landscapeand in anuse Australian position savanna.tropical SE and Coarse Biodiversity in Woody SouthernDebris Forests.General TechnicalReport In:McMinn, J.W., herpetofauna,(Eds.), forest D.A. 94-100. Crossley Jr., p. animal’sreduce exposuretoclimateextremes. -94. USDAForest -94. Service. – 323. , 49 Afield reptilesQueensland.guide the of to , 12 , –

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, , 471, 122 20 Acta Acta , 495 , , 23 , - 474,R 474,R , – 27 – 31. 503. ,

- This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. Accepted(p<0.05)are differences reported. and significant Tukey werehoc tests toexamineused effectthe ofeachlevel factor =Post NB negative poisson, GLM binomial).All models a negative use binomial distribution. associationpositive‘ a and (GLM).(GLMM)generalisedlinearand generalised mixedmodelslinear models 5: Table contribution. species analysis,firstcommunity multivariategiven, forthethen broken and individualdown by andinteractions) for2014 values 2015. p Thevariablesin ofthe optimalarethe model terrestrial)and (arboreal grazing and vegetation treatment, seasonand type (andtheir 4: Table (p<0.05)are differences reported. Tukey werehoc tests Post examineused effect to the of each levelsignificantfactor and Articleby(GLMM)withgeneralized negative described a distribution. binomial linear mixed model as (fixedyeareffects) and and(random vegetationtype season effects) treatment 3: Table 2 Table 1 Table R.JC. J., Norton, D.A. Hobbs, Yates, & restoration. mi : Measured micro-habitat characteristics with a description of methodology.:micro-habitat a Measured of descriptionwith characteristics :grazingThetreatmentsof Wambiana the Trial. Grazing The response of reptile species and functional groups to habitat variables usinggroups species Theandfunctional tohabitatreptile response variables of relationshipThethe betweenanalysis reptileManyGLM showing assemblages The relationship between reptile abundance and speciesrichness reptile relationship abundancegrazing Theand between croclimate in fragmented woodlands in south-western Australia: implicationsAustralia: inin south-western for croclimate fragmentedwoodlands AustralEcology, - ’,a association. negative GLMM distributions indicated(P are =

25 . (2000)Grazing cover, effects on plant soil and , 36 – 47.

‘+’ indicates‘+’

This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. Accepted group functional modelsand 6: Figure and group species functional models. 5: Figure areassociations negative red. in strongerin green association,are The the and the square, brighterthe positive associations which variation donot Terms in any speciesexplain set LASOO model. response are tozero. generalised a standardisedfrom usingmodelvisualised the linearmodel- co-efficients 4 Figure ArticleGrazing~ + Abundance Vegetation. a) TerrestrialAbundanceGLMMs for: Reptile ~ Grazing Vegetation,Reptile+ and Arboreal b) 3: Figure test; 0.05).α posthoc = letters indicatedaresignificant presented, difference by different 2 Figure test; 0.05).α posthoc = letters (2-way ANOVA; Tukey indicatedaresignificant presented, difference by different 1 Figure : The response of arboreal and terrestrial reptile communities to grazing, as responseTheofarboreal : and communities terrestrial grazing, reptile to MeanSEtermsofarborealOnlymeasures : ±a habitatwith characteristics. termswithMeanSEOnly a ofmeasures : ± characteristics.terrestrial habitat Fittedvaluesarboreal with reptile forthe speciesintervalsconfidence optimal 95% Fittedvalues withreptile forthe terrestrial intervals confidence optimal 95% Theofthevalues fittedwithconfidence meannegative 95% binomial intervals

(2 -way ANOVA; Tukey This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. Accepted Heavy Variable Article(R)spelling wet Rotational Moderate treatment Grazing 1 Table - season

(H)

(V)

(M)

availability; 3 availability; withremaining accordance feed ofseason end wet the the in Stocking AE per wetthe season;during 7 ona rotation grazing) basis of ⅓ paddockand the (no spelled at Stocking per C S Description per AE per S tocking rate atrate tocking L the tocking attwice tocking LTCC;the 4 arrying arrying animal ( equivalent

C rate adjustedrate annually at apacity (LTCC) apacity

50% above50% the LTCC

- 12 12 ha per AE ong AE ; 8 ; - 10 10 ha T )

erm - 10 ha - 6ha variability can againstSpelling buffer rainfall maintains riskofoverMinimize the Reasoning especially duringyears wet especially Potentiallyhighprofitability, years wet allowsheavierstocking rates during over of whichminimize availability, riskthe rate Stocking feed to match - grazing during drybut years,

landcondition

- grazing,

This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. Accepted hollows (%) Tree (%) connectivity Canopy Index(1 Bark (m) tree nearest to Distance (%) Cover Canopy Trees Arboreal Articlefeatures Other cover Vegetative cover Ground Terrestrial characteristics Habitat 2 Table

The The lizards sheltering and flaky very represents know no little often and flakingbark, from 1 ranging roughness/flakiness ofbark Anindex (m) Distance cover canopy Estimated (m) height (cm) category and height (DBH) breast at diameter fell1 that tree Any burrows. and mounds, termite including transect along the noted 100m were features ground Other wasrecorded vegetativecover 100m Along the recorded was (CWD) (>10cmdiameter) woody debris coarse (FWD), (<10cmdiameter) woodydebris fine (mm), depth leaf and litter leaf(LL) litter rock, ground(BG), A Description ground The

tape measure was laid on the grounda was the on laid measure tape percentage of overstory trees sampled that that sampled trees ofoverstory percentage percentage of overstory trees sampled that that sampled trees ofoverstory percentage .

and converted into a percentage. intoa converted and

between adjacent trees adjacent between

transect, transect, .

- m either side of the 100m transect was identified and measuredfor and identified was ofthe 100mtransect side meither via

loose bark with ample refuge microhabitats available for available microhabitats with refuge bark ample loose the amount of theamount

spherical spherical

and converted into a percentage. intoa converted and

>2 m >2 tall n densitometer.

refuge options for sheltering lizards; 3 3 lizards; sheltering for options refuge grass long the 100m the long

. (and (and

had had had grass height grass

hollows or cavities visible from the from cavities visible or hollows branches or canopy overlapping - 3; 1 representing little or no or little representing 1 3; transect

)

. The amount The of . , shrub and other other and shrub ,

bare bare .

Accepted This article isprotected rights by All copyright. reserved. Richness Arboreal Reptile Abundance Arboreal Reptile Richness Terrestrial Reptile ArticleAbundance Terrestrial Reptile Variable Response 3 Table

Grazing Vegetation ~1 Grazing *Vegetation Vegetation Grazing Vegetation + Grazing ~1 ~Vegetation Grazing Grazing*Vegetation Grazing Vegetation + Model

(null model) (null model)

df 7 5 4 11 5 8 7 4 5 7 11 8

Log Likelihood Log ------99.406 100.042 100.222 213.398 220.585 214.477 166.359 169.737 168.561 265.634 259.424 262.837

AIC 214.1 210.8 208.9 451.9 451.8 446.6 348.0 347.9 347.8 546.5 544.0 543.3

∆ 5.2 1.87 0 5.33 5.23 0 0.20 0.12 0 3.21 0.66 0

AICc

0.050 0.263 0.669 0.060 0.063 0.856 0.228 0.237 0.252 0.105 0.374 0.521 weight AICc

Not significant Not significant As above Ironbark>Box Vegetation Variable>Moderate Heavy>Moderate Grazing Not significant As above Ironbark>Heavy Rotational Ironbark ModerateIronbark>HeavyIronbark IronbarkVariableBox> Heavy Box>Rotational HeavyIronbark IronbarkModerateBox> Heavy ModerateBox> HeavyBox Grazing *Vegetation Box>Ironbark Vegetation Variable>Heavy Rotational>Heavy Moderate>Heavy Grazing Posttest hoc

This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. Community Reptile Arboreal Community Reptile Terrestrial 4 Table 2015 2014 2015 2014

AcceptedGrazing*Vegetation Grazing*Vegetation Article Optimal Model Optimal model Optimal

Vegetation Season Season Season

Community Community Community (P value) (P (P value) (P <0.01 <0.01 <0.01 <0.01 <0.01 0.02

Carlia munda Carlia

<0.01 0.31 0.06 0.38 Gehyra dubia Gehyra

<0.01 0.01

Diplodactylus Diplodactylus conspiculatus

0.58 0.91 0.31 0.52

Hoplocephalus bitorquatus Hoplocephalus Menetia Menetia greyii 0.01 0.65 0.06 0.05 Individual Species Contributions (P value) (P Contributions Species Individual Individual Species contribution (P value) (P contribution Species Individual

0.113 1.00 Ctenotus Ctenotus robustus

<0.01

0.04 0.24 0.01

Diporiphora Diporiphora

nobbi 0.34 0.27 0.06 0.05

Oedura castlenaui Oedura Heteronotia Heteronotia binoei <0.01 0.19 0.14 0.07 0.68 1.00

steindachnerri

Lucasium Lucasium 0.25 0.15 0.14 0.79

Strophurus williamsi Strophurus

taeniopleura 0.23 1.00 Morethia Morethia <0.01 0.60 0.06 0.54

This article is protected by copyright. All rights reserved. This article isprotected rights by All copyright. reserved. AcceptedArborealgeckos australis Cryptoblepharus dubia Gehyra Species Arbroeal Geckos Terrestrial ArticleLitter greyii Menetia robustus Ctenotus taeniopleura Morethia binoei Heternotia munda Carlia Species Terrestrial 5 Table skinks

Grazing*Vegetation+(1|Year)+(1|Season) GLMM Model: Full LandscapeScale Grazing*Vegetation+(1|Year)+(1|Season) GLMM Model: Full LandscapeScale Grazing Vegetation Vegetation Grazing Grazing*Vegetation Grazing Vegetation Vegetation *Grazing Vegetation Grazing Vegetation Grazing* Grazing Vegetation Termsoptimal in model

Dist NB NB NB NB NB NB NB P P P

<0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 value 0.08 0.01 0.02 0.03 P

Post Hoc

Not significant Not

VB>HB MB>MI MB>HI MB>HI MB>HI RB>HI RB>HI RB>HI RB>HI VI>MI VI>MI VI>HI VI>HI VI>HI M>H M>H M>H V> H>M V>M H>M V>H V>H B>I I>B I>B I>B M

FWD (+) Grass(+) BG( TM(+) Grass(+) Grass(+) BG( Carissaovata (+) Carissaovata (+) FWD (+) BG( Carissaovata ( BG+CanopyCover LL<5mm+LL>5mm+TM + +CWD ovata +Carissa + GrassGrass+ + height GLM Model: Full MicrohabitatScale Hollows (+) DBH( Trees 5 (+) bark indexMean DBH( Trees 10 DBH( Trees 5 DBH 20 Trees 10 DBH+Trees 5 Trees dead+Trees <5cm +CanopyConnectivity + BarkIndex+Hollows MeanDist.NearTree+Mean GLM Model: Full MicrohabitatScale Carissaovata (+) optimal modeloptimal -

30cmDBH+Trees >30cm Terms in - - - ) ) ) +CanopyCover

- - - - -

) ) ) 10cm 10cm

- -

20cm 20cmTrees DBH+

- - 10 cm 10 DBH+ )

P valueP

<0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 0.11 0.02 0.02 0.06 0.02 0.05 0.01 FWD

Figure 1

B B

A A B

A A

B A A

AB A B A

B A A

B A

A B

B A

Veg A A

A A

etation type Grazing type treatmentetation

A B

BC A B

A A a a B

a a BC A B

A A B

AC A

B b

A A B

B A

AB AB B

Mean no. trees 1020cm DBH +/ SE Mean no. dead trees +/ SE A

Figure3 Accepted This article isprotected rights by All copyright. reserved. Article a) Terrestrial ReptileAbundance Terrestrial a)

b) Arboreal

Re ptile Abundanceptile

Accepted This article isprotected rights by All copyright. reserved. Article 4 Figure

d )

Accepted This article isprotected rights by All copyright. reserved. Article Figure5

Ctenotus robustus Carlia munda

b e ) )

Heteronotiabinoei Menetia greyii c )

Morethiataeniopleura f )

Accepted This article isprotected rights by All copyright. reserved. Article g )

a )

Accepted This article isprotected rights by All copyright. reserved. Article Figure6

Gehyradubia

b )

Cryptoblepharus australis c )