Final write-up 13.01.16

Final write-up 13.01.16

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  1 | Pa g e A holistic inventory of mammalian species at Reserva Natural Laguna Blanca, Paraguay. By Simon Lowen BSc (Internship period: 06.02.15 - 06.05.15)
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  2 | Pa g e Section1:Introduction The surge incamera trap use as a surveymethodfor conservationandecologyishighlightedbya50% annual increase inthe numberof scientificpublicationsthathave adoptedthemasresearchtoolsbetween 1998 and 2008 (Rowcliffe &Carbone,2008). Ironically,cameratrapapplicationsforanalysingspecies abundance originatedfromsporthuntingdevices(Rovero etal.,2013). Such demandforpassive animal observationhasincreasedthe commercialavailabilityof cameratechnologiestothe extentthatreliable, digital cameratraps are now generallyavailable forbothscientificandnon-scientificpurposes. Cameratrap applicationsrange throughoutfieldresearchincludingcollectingspeciesinventoriesincluding the detectionof elusive species(Sanderson&Trolle,2005; Tobler et al.,2008); investigatingactivitypatterns (Jacomoet al., 2003); and estimatinganimal density/relativeabundance predominantlythrough photographicrate calculations(Carbone etal.,2001). For example,Jacomo etal (2003) successfully combinedcameratrapswithcanidscat analysistobetterdefinethe niche breadthsandresultantecological overlapsof the ManedWolf,Crab-eatingFox andHoary Fox inEmas National Park,Brazil. The rise of thistechnologyhasopenedavenuesinecology,especiallyinthe detectionof elusive species (Kelly,2008).There are howeveraplethoraof methodologicalissues,primarilyregardingthe mannerin whichthe camerasare set inthe field(Trolle &Kéry.,2003).The surroundingenvironmentandthe height& angle at whichthe camerasare setcan bothimpactthe observationalefficiencyof variouslysizedmammals. Likewise,changesinclimate have beenknowntoinadvertentlytriggercamera-traps(Acrenaz etal.,2012). Conversely,cameratrapsdo bringa numberof benefitstofieldbiology.They have beennotablyvaluableas theyprovide aconstantlevel of effort,looselydependonthe surveyor’sexperience withanimal observationsandcanbe leftinthe fieldforlongperiodswithoutrequiringbreaksbetweensurveys(Silveira et al., 2003). Similarly,the passive nature of thismethodallowsthe surveyortoobserve large areasovera relativelyshortperiodparticularlyinenvironmentswithproblematicterrainwhere otherfieldmethodshave commonlyfailed(O’Brien,Kinnaird&Wibisono.,2003). As withmanyyoungersurveytechnologies,the mannerinwhichcameratraps can be adoptedforspecies inventoriesisfarfromuniform.Some investigationssuggestplacingcamerasinconjunctionwithscentlures as to enhance animal detection(Giman etal.,2007). Othersquestionthe validityof suchmethodsand suggestthe stringentuse of anoverlyinggridsystemtoestablishcameralocationsforrepresentative data (Séquin etal.,2003). The literature advisescontinuedinvestigationintothe use of cameratrapsfor improvedguidelinesregardlessof the targetspecies’orecosystem(Hamel etal.,2013). In thisreport,a continuationof the initial cameratrapinventoryatReservaNatural LagunaBlanca(RNLB), Paraguayis evaluatedfocusingpurelyonthe detectionof mediumtolarge mammals.Datawas separately consideredfromsubsidiaryherpetologyandprimatologyprojects.Environmental andanthropogenic stressorsthatmay have createdany fluctuationinmammalianpopulationsare consideredinthe analysis. The on-site environmental station,ParaLa Tierra(PLT),was establishedin2010. Priorto this,there hadbeen no scientificinvestigationintothe ecologyof the reserve leavingagreatneedto establishanon-going inventoryof the environment.Anunderlyingaimof thisprojectistoconsiderpreviouscomplicationswith camera trappingsurveysandtoestablishasetof standardisedandreliable methodsallowingforcontinued analysis.
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  3 | Pa g e Section2:Sampling design 2a. Study area The study wasimplementedatParaLa Tierra Ecological Station(PLT) locatedwithinReservaNatural Laguna Blanca (RNLB) (S23°48’45.4”, W56°17’41.7”). The reserve ispositionedroughly25km eastof Santa RosaDel Aguarayin SanPedroDepartment,Paraguay.There are three majorhabitatswithinthe reserve:The Cerrado,the AtlanticForestandan area of transitional forestlocatedontheirfrontier.The majorityof the northernsectionof the reserve isoccupiedbythe southernextentof the Cerradohabitatwhichcontinues north intoBrazil;thisenvironmentischaracterisedbyscruband grasslandgrowingondry,nutrient-deficient earth(Hebblethwaite,2014).To the southof the reserve liesapatchof degradedbutrehabilitatingAtlantic Forestwhichhaspreviouslybeenencroacheduponbysoyfielddevelopmentonthe eastandwesternsides. A patch of semi-deciduous,semi-humidgallery‘TransitionalForest’(Smith etal.,2014) separatesthe Cerradofrom Paraguay’sonlyspring-fednatural lake namedLagunaBlanca.
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  4 | Pa g e The holisticapproachadoptedforthis studyalloweddatatobe gatheredfroma varietyof flexible sampling designsasdescribedbelow.Eachdesigniswrittenasastandardisedguideline due tothe variable nature of camera trap surveyingandthe affectsthatmanyanthropogenicandnatural confoundingfactorshave on surveyingefficiency.Theyare alsowrittentoallow forfuture continuationsof the inventory. The studyran throughoutthe periodof 07/02/15 to the 06/05/15. 2b. Primary design Representative samplingsitesweredevelopedbydividing the reserve into33 0.5km2 grid-squares.The coordinatesfor the central pointof each square were calculatedandnoted as the camera sites(figure i). Thissamplingdesignwas inspiredbythe initial inventoryasperformedbyDeclan Crace duringthe period05/04/12 to 14/06/12. A total of 6 ‘Bushnell TrophyCam’cameratraps were available atthe disposal of thisproject.The initialaimwasto obtain5 nightsof data (cameranights) ateach surveysite. Thiswas consideredflexible due tothe requirementsof side projectsandgeneral maintenance issueswiththe cameras. The cameras were installed underthe recommendationsof Rovero& Marshall (2009) and Tobler et al (2008). The formersuggestedthatcameraplacementshouldbe opportunisticusingwildlife trailsandsleeping/eatingsitesas camera trap locations.The latterconcludedthatinstallinga dual camera set-upinsteadof asingle camerawill increase capture probabilitybylessthan50%; efficiencyistherefore enhancedwhenadoptingasingle cameramethod. In lightof the literature,single cameraswere installedat‘areas of interest’ locatedwithin50mof the specificcoordinatesastokeepthe datarepresentative.Human manipulationsuchasbaitingwas avoidedwhilstestablishingthese ’areasof interest’ astoavoidissueswith ecological validity.Once apositionwaslocated,the cameraswere mainlysecuredtotreeshoweverother such reliable structurescouldbe useddependingonavailability.The cameraswere secured20-30cm off the groundas advisedbyTobleret al (2008). Figure i: Camera trap positioning as dictated by the 0.5km2 grid-square sampling design
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  5 | Pa g e 2c. Secondary design Betweenthe dates09/03/15 and 22/03/15, additional datawasgatheredfroma separate camera trap studyfocusingonobservationsof lizards,frogsandotherherpetofaunathatpopulate a seasonal pondlocatednearthe lake’sedge (S23°49’30.6” W56°17’24.97”). The cameraswere primarilysetusingbamboostructures (figure ii) alongpathwaysor inopenspaces.The ponds’ surroundingfoliagee.g.trees,shrubs&reedbeds were occasionallyusedasstructurestobindthe camerasto. The cameraswere setat dusk and removedatdawnin harmonywithanimal behaviour and to avoidtheftof equipment.The locationsof the camerasaround the pondwere notpre-emptedbut placedinareas of interestwhere previoussightings had occurredas dictatedbythe herpetologistwho managedthe study. 2d. Tertiary design The final portionof data wasextractedfroman ongoingprimatologystudybasedaroundman-madebaiting platforms.The platformsstoodapproximately2moff the groundand were capable of carryingthe weightof several monkeysplusthe desiredbait(oftencornorfruit.) Twobaitingtableswere locatedinthe southern and northernsectionsof the AtlanticForestrespectively,innatural openingswithenoughcanopyand reliable structurestosuitablypositionthe cameratraps.The heightandangle at whichthe traps were bound to the tree was assessedonsitedependingontheirorientationtothe baitingtablesasshowninfigure iii. Figure ii: An image showing a typical bamboo structure Figure iii: An image showing a camera set-up at one of the baiting stations in the Atlantic Forest Baiting station Camera-trap positioning
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  6 | Pa g e Section3:Results 3a. Primary results This‘secondaryinventory’(14.02.15– 21.04.15) observedatotal of 5 differentmediumtolarge mammalian species.Thissuggestsasignificantdecreaseinthe numberof speciesoccupyingthe reservecomparedto Declan’sprevious survey(initialinventory) whichobserved10speciesduringthe period 04.04.12 to 20.06.12. Despite the lackof diversity,abundance levelswere higherforsome speciesinthe second inventoryasdemonstratedinfigureiv. Species Initial inventory sightings Secondary inventory sightings Crab-eating Fox 12 20 Red Brocket Deer 5 4 Brazilian Cotton Tail 4 1 Six-Banded Armadillo 3 n/a Azara’s Agouti 2 4 Nine-banded Armadillo 2 6 Coati 2 n/a Lesser Grison 1 n/a Little Spotted Cat 1 n/a Southern Tamandua 1 n/a Both inventoriessuggestahighabundance of the Crab-eatingFox;thiswasmore significantforthe second inventoryhowevercomparedtoall otherspeciesthroughoutbothinventories,thiswasdefinitelythe most common. Figure iv: A table listing the species sightings for each inventory
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  7 | Pa g e The initial inventoryadoptedtwenty-sevencameratraplocations,6 of whichwere inthe Transitional Forest, fourteeninthe Cerrado and7 inthe AtlanticForest. Similarly,the secondaryinventoryhadatotal of thirty- three trap locationswithaspatial variationof 5 locationsinthe Transitional Forest,eighteeninthe Cerrado and 10 inthe AtlanticForest.The meantime acamera wasset at each locationwas4.9 (2 s.f.) camera nights. Fluctuationsbetweenthe three distincthabitatsare demonstratedinfigure R3.The mostsignificant variationisbetweenthe initialandsecondaryinventoriesof the Cerradohabitat.Despitethe vastlygreater surveyeffort,only1species(Crab-eatingFox)wasobservedinthe secondaryinventorycomparedto8 speciesin the initial inventory;thisconstituted80% of the total numberof speciessightedthroughoutthat inventory.The trendshowninfigure R3wouldsuggestanegative correlationbetweenthe numberof camera trap locationsandthe numberof speciesobserved withinthathabitat. 0 2 4 6 8 10 12 4 16 28 36 48 60 72 84 96 108 120 132 144 156 168 180 188 200 212 224 236 248 260 272 Numberofspecies Survey effort (camera nights) Initial inventory Secondary inventory 0 1 2 3 4 5 6 7 8 9 0 5 10 15 20 Numberofspecies Number of camera trap locations Initial Cerrado inventory Initial Atlantic Forest inventory Initial Transitional Forest inventory Secondary Transitional Forest inventory Secondary Atlantic Forest inventory Secondary Cerrado inventory Figure v: Comparative species abundance lines for two inventories at Reserva Natural Laguna Blanca, Paraguay. Figure vi: A chartto show thenumberof camera trap locations for each habitat compared to the number of observed species
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  8 | Pa g e 3b. Secondary results There wasonly1 recordedindividual thatfell underthe categoryof ‘mediumtolarge mammal’throughout the durationof thissubsidiaryproject.2birds-eye viewedimagesshowedthe backandfront feetof a potential adultSouthernRaccoon.Thisispossiblythe firstpictorial evidenceof the specieswithinthe reserve;furtherscientificverificationisongoingtoconfirmthe sighting. 3c. Tertiary results The ongoingprimatologyprojectatLagunaBlanca has deducedthe currentCapuchinMonkeypopulation withinthe reserve tobe fifteenin‘groupO’and8 in‘groupF’. The surveyperiodconsideredforthisresearch ran from 07.02.15 to 06.05.15. Duringthistime there were 3 recordedvisitstothe feedingtables.One by‘F group’at the tablesinthe northernsectionof the AtlanticForestinwhichthere were 4sightedindividuals. The other 2 recordedvisitsoccurredat the feedingtablesinthe southernsectionof the AtlanticForest; Acrossthese 2 visitsthere were 9sightedindividualsall of whichwere membersof ‘Ogroup’.Itshouldbe notedthat the camera trapswere not active throughoutthe entire surveyperiod;the datawas extrapolated froma total of sixteencameranights.We candeduce that50% of ‘groupO’ and 60% of ‘groupF’ were observedduringthe periodof thisstudy.
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  9 | Pa g e Section4:Discussion 4a. Variation as a result of changing habitats ReservaNatural LagunaBlanca boastsa varietyof natural habitatsand sub-habitats. Figurevi suggesta significantvariationinuseable observationsbetweenthe Cerrado,AtlanticForestandTransitional Forest.Thisvariationislikelythe resultof species nichesandrespectivepopulationswithinthe givenhabitat e.g. the RedBrocket Deerwere onlyrecordedinthe degradedsectionof the AtlanticForestwhere vegetationissparse;thisispossiblythe resultof responsetohighground-waterlevelsinthe southof the AtlanticForesthoweversuchungulatesare comfortable eatingsuchawoodybrowse diet(Bodmer,1990). Similarly,the Azara’sAgouti were only recordedinthe Transitional Forestandthe AtlanticForest due to theirfrugivorousdiet;evolutionaryadaptationshave giventhisspeciesexceptionallysharpteethallowing themto eatfallennutsandseedswhichare commoninthese dense environments(Ribeiro&Vieira,2014). As the resultssuggestfromfigure vi,anegative correlationwasfoundbetweenthe numberof camera trap locationsandthe numberof recordedspeciesforeachgivenhabitat.Perhapsalackof data frommany of the locationscouldbe avoidedbyfollowinga ‘clustered’designwhere thereare high concentrationsof cameratrap locationsasopposedtothe representativedesignadheredto inthis inventory.These ‘clustered’designsshouldbe positionedineachof the sub-habitatsastoincrease the recordingefficiencyof specieswithvariedecological characteristics(Kelly&Holub,2008) e.g.the Atlantic Forestshouldbe dividedintothe FloodedForest,NorthAtlantic,SouthAtlanticanddegradedterritories. As discussedinsection2b, ‘areasof interest’ werelocatedonsiteandinmostcaseswere unnatural influencerssuchasroads or footpaths. Itwaspre-emptedthatthese pathswere active habitatcorridorsfor mammalsespeciallyindensefloral environments(Bennett,1990). It wasconverselynotedthat the majority of recordings were limited tonighttime possiblythe resultof alearnedresponsebymammalianspeciesto recognise andavoidodoursof sympatricpredators(Swihart,1991). In lightof this,itis advisedthat‘areasof interest’are more carefullypre-emptedandpositionedinharmonywiththe respectivenatural attractorsof each sub-habitat.
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  10 | Pa g e 4b. Targeting specific species Whencomparingthe primaryand secondaryresultstothe tertiaryresultsitisclearthat the efficiencyof lureswhenattractingmammalscannotbe understated.Thisisdemonstratedbyhigh photographicdetectionsuccessof the reserve’sAzara’sCapuchinpopulationsoverarelativelyshort samplingperiod(16cameranights).The corn usedinthisprimatologyprojectnotonlyattractedthe monkeysbutduringeachvisit,there were sightingsof atleasttwo PlushCrestedJays. Conversely,the results fromthe primaryandsecondarydesignsare comparativelyrandomindetectingmammalse.g.the initial and secondaryinventory fromthe primarydesign sharedthe same numberof cameranightshoweverthe numberof speciesrecordedissignificantlygreaterinthe initial inventory (figure v).We candeduce that luresprovide sufficientincentive inanatural environmenttoincrease capture rate andimprove survey reliability. Roveroet al., (2010) useda varietyof scentedluresatan Acaciaplantationincentral Sarawakto monitorterrestrial mammal populations.Some luresfaileddue toheavyrainfallhoweveroilyluressuchas FishOilse.g.Magna Glandproducesa strong odourwhichremainedpungenttothe researcherforone monthevenduringthe rainyseason. The benefitsof natural luressuchas Wheat,oats,millet,canaryseed, and hemp are effective inincreasingcapture ratesof elusive species(Blair, 1941) howeverintroducing anthropogenicelements couldcreate unrepresentative datawhenresearchingpopulationdynamicsas intelligentspecies mayreturntothe pointof attraction as some capture-recapture modelswouldsuggest (Karanthet al., 2004) . In thisproject,lureswould have beenbeneficial asthe aim wasto furthera basic inventoryasopposedto understandingthe complexitiesof variousmammalianspecies. From an ethical standpoint,the importance of choosingappropriate lureshasbeenhighlightedby Schlexer(2008).His research considersthe serioushealththreatsassociatedwithraw fishbait,commonly usedto surveycarnivores.Salmonpoisoningdisease (SPD) andElokominfluke fever(EFF) were bothshown to be acute infectiousdiseasestransferredthroughthe ingestionof salmonandothercommonfish,‘SPDcan kill upto 90% of infectedanimals,while EFFusuallymanifestsinamilderform’ (Aiello,1998). To avoidthis ethical issue,researchingthe dietof eachtargeted mammal speciescouldoffereducated insightintosafe and effective lures.
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  11 | Pa g e 4c. Survey intensification Figuresv& vi wouldbothsuggestnopositive relationshipbetween increasedsurveyeffortand mammal capture rate. The data fromfigure vi wouldactuallysuggesta significantnegativecorrelation betweenthe numberof onsite cameralocationsand the numberof speciesobservede.g.only4of the 18 camera sitesinthe secondaryCerradoinventoryrecordedcollectabledata.Consideringeachsite was allocated≥5 camera nights,the resultswouldsuggestaninefficiencyof surveyeffort. As a relativelymodern ecological tool there are still questionsregardingthe ideal surveydesignto adoptfor camera trap studies.Soria-Diazetal., (2010) discussedthe variationof abundance anddensityof Puma concolorinzonesof highand lowconcentration of cameratraps inSierraNanchititlaNatural Reserve, Central Mexicoi.e.arepresentative orintensivesurveydesign.The representative designwasmore expansive(anaverage distance betweencameras of 4.6 km with56-618 trap days) comparedto the intensivedesign (anaverage distance betweencamerasof1.6kmwith 491-618 trap days).The representative designrecordedanaverage of 1.21±0.41 (individuals/100km²) whereasthe representative designrecordedanaverage of 5.49±0.54 (individuals/100km²). In the case of Soria-Diazetal., (2010) we can deduce thata greaterconcentrationof camera traps/unitareadid increase the likelihoodof registeringthe targetspecies.Itwasnotedhowever that considerationforthe niche widthandtherefore recapture probabilityisessential toconfirmimproved surveydesignthroughintensification(Maffei etal., 2004). In thisstudy,recapture ratesare lessof a concern as the aim was to establish abasicinventory whilstconfiguringthe niche widthof everypossiblemammalian speciesonthe reserve wouldbe ineffective.A more intensive design(asopposedtothe 0.5km2 grid system) wouldtherefore offershorterdistancesbetweencamerasites(allowingforsimplerretrieval of devices& checkingfortechnical issues) whilstpossibly improvingcapture ratesaspreviouslydiscussed.
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  12 | Pa g e 4d. Future research As previouslydiscussed,atheme of this projectwasto finda setof standardisedmethodstoallow forthe continuousrecordingof mammalianspeciesonthe reserve. The core inventoriesof the ‘primary methodology’were comparabledue totheirsimilarsamplingdesigns. The resultsof the primaryand secondaryinventories suggestsignificantvariationinspeciesabundance whencomparedagainstsurvey effortandhabitat (figuresv& vi).The 0.5km2 grid-square samplingdesignissoundwhenassessingageneral inventoryof the reserve.Anyfutureinventoriesusingthisstyle will onlyvalidate the previouslydiscussed results.Standardisingandcarefullyrecitingthismethodologywill alsocreate opportunitiesforvolunteers whovisitthe reserve tocontribute toan ongoingscientificprojectwithoutnecessarilypossessing previous experience inEcological principles. A separate camera-trapprojectcouldbe undertakento improve the capture rate of mammalsthroughoutthe various habitatsas depictedinfigure vii.The ‘location indicators’ suggestsites forcameratraps that are to be accompaniedby non-lethal lures.Surroundingcamerasshouldpointinwardsto the central camera. The central camera shouldbe pointing in the optimumdirectionasdictatedbythe geographyof the respective site. Toestablishvalidresults,7-10camera night surveysshouldbe implementedin eachof the varioussub- habitatsthat the Cerrado,AtlanticForestandTransitional forestpossesse.g.the Cerradoischaracterisedbycampotype vegetation,galleryforestandwell-drainedsavannahtype habitats.Thisdesignwill givebettersurveyarea intensification whilststill consideringthe shiftingmosaicthatthese habitats possess(Soria-Diazet al., 2010). A less generalised survey area design will also improvethe efficiency of the surveyor’s time in placingcameras in thefield therefore increasingthenumber of recordablecamera nights; an essential element of reliabledata (Rovero et al., 2010). Future developments to the herpetology camera trap study should involvechanges to the bamboo structures that held the cameras in place.It was often the casethat the structurehad collapsed duringthe night possibly dueto passing animals.Many of the pictures were also unclear becausethe cameras were set too closeto the ground. Amending these issues with higher standing,more physically reliable structures could improvethe capture rate of small herpetofauna; however, a significantchangein survey design would have to occur to improve the capture rate of mammals,specifically larger mammals thatwould not fitthrough small ‘gate’structures as shown in figure ii. With regards to the primatology camera trap project, any considerationsfor future research should be sourced from the on-site primatologist.Dueto the longstandingnatureof this study, the Azara’s Capuchin population of ‘Group O’ and ‘Group F’ are reliably understood.Currently the project focuses on feeding behaviours as opposed to population changes. Perhaps a future projectmight involveusingcamera traps to observe the less studied Howler Monkey population thatare occasionally heard in the Flooded Forest sub-habitatatthe southern extent of the Atlantic Forest. Unfortunately, such research might be difficultdueto the elusivenature of the species and the inaccessible environment that they populate. Figure vii: A possible sampling design in light of research performed by Soria-Diaz et al., (2010) & Rovero et al., (2010) Location Indicators
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  13 | Pa g e Conclusion Thisprojectgivesevidence of the needfor careful considerationwhenestablishingan appropriate surveydesignforan inventory. The holisticapproachgave insightintoavarietyof techniquesforrecording mediumtolarge mammals usingcameratraps. Regardingthe primarydesign,the initial inventoryobtained agreateryieldof usable images thanthe secondaryinventory.The nearidentical designs usedforbothinventories wouldsuggestthatthisvariationis the resultof natural factors as opposedtoany anthropogenicissues.Fromatechnical perspective,the initial inventorypromotedthe use of pathsandroadsas attractors formammalswhereasthe secondaryinventory suggestedthatnatural attractors suchas the educatedimplementationof diet-specificlures wouldoffera well-roundedviewof the mammalianpopulationsof eachhabitat. Furtherresearchintoavarietyof attractors isrequiredtobetterunderstand where the camerasshouldbe pointedtowards.The grid-square designisunarguablyrepresentativeasitcoversthe entire reserve inanequal manner;itshouldhoweverbe notedthat 9 of the 33 camera trap locationsfromthe secondaryinventoryyieldednousable images,a numberthatmightbe improvedbyadoptingamore concentratedsurvey design. The secondarysurveydesigndemonstratedanalternative meansof hoistingthe cameratraps.The vertical imageswere oftenuncleardue tothe reflectionfrom the groundandthe relativelysmall bamboo framesmeantthat a lotof the largermammals wouldnotbe able to fitthroughthe structures. It shouldbe notedthat thiscouldbe considered the mostsuccessfulof the three designsdue tothe possible sightingof a previouslyunseenspeciesonthe reserve;aSouthernRacoon. The tertiarydesign demonstratedhow beneficial lurescanbe inenticingmammals.Inthiscase,the use of feedingtablesrestrictedthe benefitsof the lure tomobile speciessuchasCapuchinMonkeys.A more accessible structure andamore universal lure e.g.fruits&nuts mightdemonstrate the true potentialof luresinestablishingathoroughinventory. Overall,continuedsurveyingof the reserve isessentialincompletingafull speciesinventory.Itis clearthat there are still unrecordedspeciesthatinhabitthe reserveasdemonstratedbysection3b.A continuationof the primarydesignwouldbe advisedhoweverconsiderationforwhere the camerasare placedisimportantto improve surveyefficiency.Itisalsoimportantthatdata iscollectedfromoutside of the summermonthsto allowforseasonal variation.Incorporatingall of the suggestionsfromeachof the surveydesignsdiscussedwithinthisprojectshouldgivesufficientguidance toestablishasetof reliable methodsallowingforcontinuedanalysisof mammalian populationsonthe reserve.
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  14 | Pa g e Bibliography Ancrenaz, M., Hearn, A. J., Ross, J., Sollmann, R., & Wilting, A. (2012). Handbook for wildlife monitoring using camera-traps. Sabah, Malaysai. Carbone, C., Christie, S., Conforti, K., Coulson, T., Franklin, N., Ginsberg, J. R., & Wan Shahruddin, W. N. (2001). The use of photographic rates to estimate densities of tigers and other cryptic mammals. Animal Conservation, 4(01), 75-79. De Almeida Jácomo, A. T., Silveira, L., & Diniz-Filho, J. A. F. (2004). Niche separation between the maned wolf (Chrysocyon brachyurus), the crab-eating fox (Dusicyon thous) and the hoary fox (Dusicyon vetulus) in central Brazil. Journal of Zoology, 262(01), 99-106. Giman, B., Stuebing, R., Megum, N., Mcshea, W. J., & Stewart, C. M. (2007). A camera trapping inventory for mammals in a mixed use planted forest in Sarawak. Raffles Bulletin of Zoology, 55(1), 209-215. Hamel, S., Killengreen, S. T., Henden, J. A., Eide, N. E., Roed‐Eriksen, L., Ims, R. A., & Yoccoz, N. G. (2013). Towards good practice guidance in using camera‐traps in ecology: influence of sampling design on validity of ecological inferences. Methods in Ecology and Evolution, 4(2), 105-113. Kelly, M. J. (2008). Design, evaluate, refine: camera trap studies for elusive species. Animal Conservation, 11(3), 182-184. O'Brien, T. G., Kinnaird, M. F., & Wibisono, H. T. (2003). Crouching tigers, hidden prey: Sumatran tiger and prey populations in a tropical forest landscape.Animal Conservation, 6(02), 131-139. Rowcliffe, J. M., & Carbone, C. (2008). Surveys using camera traps: are we looking to a brighter future?. Animal Conservation, 11(3), 185-186. Rovero, F., Zimmermann, F., Berzi, D., & Meek, P. (2013). " Which camera trap type and how many do I need?" A review of camera features and study designs for a range of wildlife research applications. Hystrix, the Italian Journal of Mammalogy, 24(2), 148-156. Sanderson, J. G., & Trolle, M. (2005). Monitoring Elusive Mammals-Unattended cameras reveal secrets of some of the world's wildest places. American Scientist, 93(2), 148-155. Séquin, E. S., Jaeger, M. M., Brussard, P. F., & Barrett, R. H. (2003). Wariness of coyotes to camera traps relative to social status and territory boundaries. Canadian Journal of Zoology, 81(12), 2015-2025. Silveira, L., Jacomo, A. T., & Diniz-Filho, J. A. F. (2003). Camera trap, line transect census and track surveys: a comparative evaluation. Biological Conservation, 114(3), 351-355. Tobler, M. W., Carrillo‐Percastegui, S. E., Leite Pitman, R., Mares, R., & Powell, G. (2008). An evaluation of camera traps for inventorying large‐and medium‐sized terrestrial rainforest mammals. Animal Conservation, 11(3), 169-178. Trolle, M., & Kéry, M. (2003). Estimation of ocelot density in the Pantanal using capture-recapture analysis of camera-trapping data. Journal of mammalogy, 84(2), 607-614. Tobler, M. W., Carrillo‐Percastegui, S. E., Leite Pitman, R., Mares, R., & Powell, G. (2008). An evaluation of camera traps for inventorying large‐and medium‐sized terrestrial rainforest mammals. Animal Conservation, 11(3), 169-178. Rovero, F., & Marshall, A. R. (2009). Camera trapping photographic rate as an index of density in forest ungulates. Journal of Applied Ecology, 46(5), 1011-1017. Tobler, M. W., Carrillo‐Percastegui, S. E., Leite Pitman, R., Mares, R., & Powell, G. (2008). Further notes on the analysis of mammal inventory data collected with camera traps. Animal Conservation, 11(3), 187-189.
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  15 | Pa g e Hebblethwaite, M. (2014). Paraguay. Bradt travel guides. Smith, P., Cacciali, P., Scott, N., del Castillo, H., Pheasey, H., & Atkinson, K. (2014). First record of the globally-threatened Cerrado endemic snake Philodryas livida (Amaral, 1923) (Serpentes, Dipsadidae) from Paraguay, and the importance of the Reserva Natural Laguna Blanca to its conservation. Cuadernos de Herpetología, 28(2). Soria-Díaz,L., Monroy-Vilchis,O.,Rodríguez-Soto,C.,Zarco-González,M.M., & Urios, V.(2010). Variationof abundance anddensityof Puma concolorinzonesof highandlow concentrationof camera trapsin Central Mexico. AnimalBiology,60(4), 361-371. Rovero, F., Tobler, M., & Sanderson, J. (2010). Camera-trapping for inventorying terrestrial vertebrates. Manual on field recording techniques and protocols for All Taxa Biodiversity Inventories and Monitoring. The Belgian National Focal Point to the Global Taxonomy Initiative, 100-128. Schlexer,F.V.(2008). Attractinganimalstodetectiondevices. Noninvasivesurvey methodsforcarnivores, 263-292. Blair,W. F. (1941). Techniquesforthe studyof mammal populations. Journalof Mammalogy,22(2),148-157. Aiello,S.E.(1998).The Merck VeterinaryManual,8thEdition,Merck& Co.,Inc, USA. Maffei ,L. & Noss, A. (2008) How small istoo small?Cameratrap surveyareasanddensityestimatesfor ocelotsinthe BolivianChaco.Biotropica,40(1), 71-75. Karanth, K. U., Chundawat, R. S., Nichols, J. D., & Kumar, N. (2004). Estimation of tiger densities in the tropical dry forests of Panna, Central India, using photographic capture–recapture sampling. Animal Conservation, 7(03), 285-290. Bodmer, R. E. (1990). Responses of ungulates to seasonal inundations in the Amazon floodplain. Journal of tropical Ecology, 6(02), 191-201. Ribeiro, J. F., & Vieira, E. M. (2014). Interactions between a seed‐eating neotropical rodent, the Azara's agouti (Dasyprocta azarae), and the Brazilian ‘pine’Araucaria angustifolia. Austral Ecology, 39(3), 279-287. Kelly, M. J., & Holub, E. L. (2008). Camera trapping of carnivores: trap success among camera types and across species, and habitat selection by species, on Salt Pond Mountain, Giles County, Virginia. Northeastern Naturalist, 15(2), 249-262. Bennett, A. F. (1990). Habitat corridors and the conservation of small mammals in a fragmented forest environment. Landscape Ecology, 4(2-3), 109-122. Swihart, R. K., Pignatello, J. J., & Mattina, M. J. I. (1991). Aversive responses of white-tailed deer, Odocoileus virginianus, to predator urines. Journal of Chemical Ecology, 17(4), 767-777.