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Research priorities for the northern quoll (Dasyurus hallucatus) in the Pilbara region of Western Australia Viki A. Cramer A , Judy Dunlop A , Rob Davis B , Ryan Ellis C , Belinda Barnett D , Annette Cook A , Keith Morris A and Stephen van Leeuwen A,E A Science and Conservation Division, Department of Parks and Wildlife, Locked Bag 104, Bentley Delivery Centre, WA 6983, Australia. B School of Natural Sciences, Edith Cowan University, 270 Joondalup Drive, Joondalup, WA 6027, Australia. C Department of Terrestrial Zoology, Western Australian Museum, 49 Kew Street, Welshpool, WA 6106, Australia. D Environment Department, BHP Billiton Iron Ore, 125 St Georges Terrace, Perth, WA 6000, Australia. E Corresponding author. Email: [email protected] Abstract. The Pilbara population of the northern quoll (Dasyurus hallucatus) has been seldom studied, and the impacts of threats such as altered fire regimes, total grazing pressure, predation and mining and infrastructure development are not well understood. While the Pilbara was once thought likely to provide refuge for northern quolls from the poisonous cane toad (Rhinella marina), recent modelling suggests that cane toads will invade the region. The environmental approvals process for mining development in the Pilbara has generated considerable offset funds that are to be directed towards research on the northern quoll. In an effort to identify future research priorities for this species in the Pilbara through a collaborative process, the Western Australian Department of Parks and Wildlife hosted a workshop attended by scientists, environmental consultants, mining proponents and state and federal regulators. Participants at the workshop identified five key areas for future research effort: (1) develop appropriate and standardised survey and monitoring methods; (2) define areas of critical habitat and better understand how disturbance affects habitat quality; (3) improve our understanding of population dynamics; (4) better understand the key threats to the northern quoll and the interactions between these threats in the Pilbara; and (5) determine whether the northern quoll will colonise restored areas or artificial habitat. We provide the expected timelines and current allocation of resources to these research priorities over the next 10 years. We reflect on the lessons learnt from the workshop process and consider ways to improve the outcomes of such collaborative exercises. Additional keywords: cane toads, den sites, habitat quality, monitoring, population structure, predators. Received 27 March 2015, accepted 14 November 2015, published online 5 February 2016 Introduction The northern quoll (Dasyurus hallucatus) is the largest predatory dasyurid marsupial remaining in northern Australia. It was formerly distributed from the Pilbara and Kimberley across the Top End to southern Queensland, but has now contracted to several disjunct populations (Braithwaite and Griffiths 1994). Several interacting ecological factors, primarily predation by cats (Felis catus) and foxes (Vulpes vulpes) and changed fire regimes, are currently causing rapid and severe declines in the small- and medium-sized mammal fauna of northern Australia (Woinarski et al. 2014, 2015). In addition, an alarming decrease or complete collapse in some of the once locally abundant populations of the northern quoll in Queensland and the Northern Territory has occurred as a direct result of the invasion of the cane toad (Rhinella marina) (Burnett 1997; Oakwood and Foster 2008; Woinarski et al. 2010). While the Pilbara region and its offshore islands were formerly thought likely to remain free of cane toads, and therefore provide a ‘stronghold’ for the northern quoll, it is now predicted that cane toads will invade the Pilbara, and may even be able to reach its offshore islands (Kearney et al. 2008; Tingley et al. 2013). Owing to these factors, the northern quoll is listed as Endangered under both the Commonwealth’s Environment Protection and Biodiversity Conservation Act 1999 (EPBC Act 1999) and the Western Australian Wildlife Conservation Act 1950. While the ecology of the northern quoll has been studied in the Northern Territory (Begg 1981; Braithwaite and Griffiths 1994; Oakwood 1997, 2000, 2002), and to a lesser extent in the Kimberley (Schmitt et al. 1989; How et al. 2009; Cook 2010), few studies have been undertaken on the northern quoll in the Pilbara. A 10-year monitoring program that includes the collection of demographic and habitat data is being undertaken by the Western Australia Department of Parks and Wildlife (hereafter Parks and Wildlife), and commenced in 2014. The extent of occurrence of the northern quoll in the Pilbara and Journal compilation Ó Australian Mammal Society 2016 www.publish.csiro.au/journals/am CSIRO PUBLISHING Australian Mammalogy Review http://dx.doi.org/10.1071/AM15005

Research priorities for the northern quoll (Dasyurus ......The Pilbara population of the northern quoll (Dasyurus hallucatus) has been seldom studied, and the impacts of threats such

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Page 1: Research priorities for the northern quoll (Dasyurus ......The Pilbara population of the northern quoll (Dasyurus hallucatus) has been seldom studied, and the impacts of threats such

Research priorities for the northern quoll (Dasyurus hallucatus)in the Pilbara region of Western Australia

Viki A CramerA Judy DunlopA Rob DavisB Ryan EllisC Belinda BarnettD Annette CookAKeith MorrisA and Stephen van LeeuwenAE

AScience and ConservationDivision Department of Parks andWildlife Locked Bag 104 Bentley Delivery CentreWA 6983 Australia

BSchool of Natural Sciences Edith Cowan University 270 Joondalup Drive Joondalup WA 6027 AustraliaCDepartment of Terrestrial Zoology Western Australian Museum 49 Kew Street Welshpool WA 6106 AustraliaDEnvironment Department BHP Billiton Iron Ore 125 St Georges Terrace Perth WA 6000 AustraliaECorresponding author Email stephenvanleeuwendpawwagovau

Abstract The Pilbara population of the northern quoll (Dasyurus hallucatus) has been seldom studied and the impactsof threats such as altered fire regimes total grazing pressure predation and mining and infrastructure developmentare not well understood While the Pilbara was once thought likely to provide refuge for northern quolls from thepoisonous cane toad (Rhinella marina) recent modelling suggests that cane toads will invade the region The environmentalapprovals process for mining development in the Pilbara has generated considerable offset funds that are to be directedtowards research on the northern quoll In an effort to identify future research priorities for this species in the Pilbarathrough a collaborative process the Western Australian Department of Parks and Wildlife hosted a workshop attended byscientists environmental consultants mining proponents and state and federal regulators Participants at the workshopidentified five key areas for future research effort (1) develop appropriate and standardised survey and monitoringmethods (2) define areas of critical habitat and better understand how disturbance affects habitat quality (3) improve ourunderstanding of population dynamics (4) better understand the key threats to the northern quoll and the interactionsbetween these threats in the Pilbara and (5) determine whether the northern quoll will colonise restored areas or artificialhabitat We provide the expected timelines and current allocation of resources to these research priorities over the next10 years We reflect on the lessons learnt from the workshop process and consider ways to improve the outcomes ofsuch collaborative exercises

Additional keywords cane toads den sites habitat quality monitoring population structure predators

Received 27 March 2015 accepted 14 November 2015 published online 5 February 2016

Introduction

The northern quoll (Dasyurus hallucatus) is the largest predatorydasyurid marsupial remaining in northern Australia It wasformerly distributed from the Pilbara and Kimberley acrossthe Top End to southern Queensland but has now contracted toseveral disjunct populations (Braithwaite and Griffiths 1994)Several interacting ecological factors primarily predation bycats (Felis catus) and foxes (Vulpes vulpes) and changed fireregimes are currently causing rapid and severe declines inthe small- and medium-sized mammal fauna of northernAustralia (Woinarski et al 2014 2015) In addition an alarmingdecrease or complete collapse in some of the once locallyabundant populations of the northern quoll in Queensland andthe Northern Territory has occurred as a direct result of theinvasion of the cane toad (Rhinella marina) (Burnett 1997Oakwood and Foster 2008 Woinarski et al 2010) While thePilbara region and its offshore islands were formerly thoughtlikely to remain free of cane toads and therefore provide a

lsquostrongholdrsquo for the northern quoll it is now predicted thatcane toads will invade the Pilbara and may even be able toreach its offshore islands (Kearney et al 2008 Tingley et al2013) Owing to these factors the northern quoll is listed asEndangered under both the Commonwealthrsquos EnvironmentProtection and Biodiversity Conservation Act 1999 (EPBC Act1999) and the Western Australian Wildlife Conservation Act1950

While the ecology of the northern quoll has been studied inthe Northern Territory (Begg 1981 Braithwaite and Griffiths1994 Oakwood 1997 2000 2002) and to a lesser extent in theKimberley (Schmitt et al 1989 How et al 2009 Cook 2010)few studies have been undertaken on the northern quoll inthe Pilbara A 10-year monitoring program that includes thecollection of demographic and habitat data is being undertakenby the Western Australia Department of Parks and Wildlife(hereafter Parks and Wildlife) and commenced in 2014 Theextent of occurrence of the northern quoll in the Pilbara and

Journal compilation Australian Mammal Society 2016 wwwpublishcsiroaujournalsam

CSIRO PUBLISHING

Australian Mammalogy Reviewhttpdxdoiorg101071AM15005

Kimberley is estimated to be ~12 000 000 ha but the area ofoccupancy and the number of mature individuals within thetwo regions are not known (Parks and Wildlife unpubl data)Approximately 2500 records for the northern quoll (many ofwhich are highly clustered) are held in the NatureMap database(httpnaturemapdpawwagovau) these include records fromthe Western Australian Museum and Parks and Wildlifeand fauna returns from surveys undertaken by private groups(mostly environmental consultants) These location recordsindicate that quolls are more prevalent in complex rocky areas inthe north central and west Pilbara and are less likely to occurthrough the south and east areas of the Hamersley Rangesand Fortescue Marsh (Fig 1) The Pilbara population isgenetically distinct from the Kimberley and Northern Territorypopulations (Woolley et al 2015) and is the least diversewith ~77 heterozygosity (103 alleles) compared with84 heterozygosity (111 alleles) in the Kimberley mainlandpopulation (How et al 2009 Spencer et al 2013)

Habitat change due to altered fire regimes weed invasionand grazing predation by cats foxes and dingoeswild dogsthe future invasion of cane toads and the interactive effectsbetween these as well as habitat fragmentation and loss bymining and infrastructure development have been identified

as key threats for the northern quoll in the region (Woinarskiet al 2014) While the potential invasion of the cane toad isundoubtedly the most significant future threat to the northernquoll in the Pilbara there is little knowledge of what has been therelative impact or the interactive effects of each of the otherkey threats on the northern quoll in the region or how these willaffect northern quoll populations into the future

The areas of highest habitat quality for the northern quollare considered to be rugged rocky areas often in closeassociation with permanent water (Begg 1981 Schmitt et al1989 Braithwaite and Griffiths 1994 Oakwood 1997 Pollock1999) These rocky landforms provide a diversity ofenvironments denning opportunities protection from weatherand diurnal predators and immediate refuge from fire(Braithwaite and Griffiths 1994 Oakwood 2000 Cook 2010)In the Pilbara the ridges and mesas of channel-iron depositsand banded iron formation ranges are often the primary focusof iron-ore extraction (Ramanaidou and Morris 2010) whileexposed granite outcrops are quarried for road and rail beds Thenorthern quoll is therefore a key consideration in most miningproject assessments under the EPBC Act 1999 in the Pilbara aswell as two strategic environmental assessments examining thecumulative impacts of mining in the region (BHP Billiton Iron

116degdeg 117deg 118deg 119deg 120deg 121deg

116deg115deg

ndash23deg

ndash22deg

ndash21deg

ndash20deg

ndash23deg

ndash22deg

ndash21deg

ndash20deg

117deg 118deg 119deg 120deg 121deg

Fig 1 Records (black dots) of the northern quoll in the Pilbara obtained from the Department of Parks and WildlifeNatureMap database (httpnaturemapdpawwagovau) The shaded areas indicate the IBRA subregions and the hatched areasthe conservation estate within the region Much of the recent survey effort has occurred in areas with mining activity or mining-related infrastructure eg the rail corridors running south of Port Hedland

B Australian Mammalogy V A Cramer et al

Ore httpwwwenvironmentgovaunode18597 HamersleyIron Pty Ltd (Rio Tinto) httpwwwenvironmentgovaunode18608)

Biodiversity offsets are increasingly used as a policy toolto permit development but ensure that any residual adverseimpacts of the development on the environment arecompensated (Bull et al 2013) In best practice offsets areapplied as part of a mitigation hierarchy where all reasonableefforts to avoid or minimise impacts upon the environment arefirst considered (Kiesecker et al 2010) Yet the limited data onthe distribution and status of the northern quoll in the Pilbarathe poor understanding of critical habitat in relation to futuremining and infrastructure development as well as uncertaintyaround the level of impact from other threats has meant thatuncertainty around how to create direct ecologically equivalentoffsets is high Under the Australian Governmentrsquos offsetpolicy (Department of Sustainability Environment WaterPopulation and Communities 2012) compensatory offsetssuch as directing a proportion of offset funds towards scientificresearch are considered as an acceptable component of anoffsets package in these situations This has been the case forthe northern quoll in the Pilbara with approximately AU$175Mof offset funds committed to research on the species over thenext decade (although not all of this amount is currently active)

The dramatic increase in fauna surveys associated withenvironmental impact assessments in the Pilbara has generateda valuable dataset on the northern quoll and other threatenedspecies (Carwardine et al 2014) Collation of these datacould greatly improve our knowledge of speciesrsquo ranges andcritical habitats yet most of these data are not within thepublic domain (Carwardine et al 2014) Further it has beenargued that much conservation research is having little directinfluence on the achievement of on-ground conservationoutcomes in part because the conceptual novelty valued bynational funding agencies may not align with the moreapplied needs of practitioners (Knight et al 2008 Lauranceet al 2012) A potential solution to overcoming the lsquoknowledgendashimplementationrsquo or lsquoknowingndashdoingrsquo gap is to encouragecollaboration between scientists policy makers and practitionerswhen research projects are developed (Knight et al 2008Sutherland et al 2011 Laurance et al 2012) In an effort toencourage data-sharing and to foster a sense of ownership overthe research agenda that will be funded by offset provisionsParks and Wildlife instigated a collaborative process betweenscientists practitioners and mining proponents to identifyand rank research priorities for the northern quoll in the PilbaraIn this paper we present the research priorities and the keyquestions associated with each priority as identified byworkshop participants This is the first step in developing aresearch program to increase our knowledge of the northernquoll in the Pilbara and to better understand the impacts ofcurrent and future threats so that a viable population ismaintained into the future using evidence-based management

Methods

Parks and Wildlife hosted a workshop attended by researchscientists employed in the university or government sector (13people) research scientists and environmental consultants

employed in the private sector (30 people) mining industryrepresentatives (22 people) and representatives of WesternAustralian and Australian government departments responsiblefor environmental regulation and approvals (9 people) all ofwhom have knowledge of the northern quoll mining andassociated activities andor the Pilbara region The workshopwas facilitated by an independent professional facilitator Theprocess to define research priorities was as follows (1) After aseries of presentations by experts the group as a whole discussedthe key threats to the northern quoll in the Pilbara This includedboth current and future threats (eg climate change cane toadinvasion) and both well- and lesser-understood threats (eg theimpact of disease) (2) Participants were then allocated to one ofeight breakout groups so that each group contained membersfrom the range of affiliations present Each group then identifieda series of research priorities that they considered would bestaddress the current gaps in knowledge that are the main barriersto the effective management and conservation of the northernquoll in the Pilbara both in terms of better understanding theirdistribution and ecology and in mitigating the key threats Theresearch priorities from each group were then presented to allparticipants and collated by the facilitator (3) Each group thenranked the sequence in which the research priorities should beaddressed based on which research needs they considered weremost pressing (4) The rankings were then compared and collatedacross groups to define the final ranking of research priorities Inmost cases there was agreement across the groups as to thesequence in which the research priorities should be addressed(5) Finally new breakout groups were formed based onexpertise and interest to discuss and identify the key researchquestions that needed to be addressed under each researchpriority

Research priorities

Research priorities are presented below in the order in whichthey should be addressed based on the ranking of the workshopparticipants Key research questions aligned with each researchpriority are presented in Table 1 We also provide a brief reviewof the current evidence that will further guide the development ofa research program to answer these questions

Priority 1 Develop appropriate and standardised surveyand monitoring methods

Developing a standard survey and monitoring protocol for thenorthern quoll in the Pilbara based on detection and occupancyprobabilities (Wintle et al 2012) was identified as the highestresearch priority by the group A review of over 200 surveyreports most of which were related to mining activities foundthat the sampling effort and methods varied widely betweensurveys and were often not adequate to detect northern quolls(A Cook unpubl data) Inadequate survey effort andor usingan inappropriate survey method introduce uncertainty as towhether the failure to detect the northern quoll within an areareflects a true absence of the species (Wintle et al 2012) Thedata gathered from surveys that vary widely in terms of effortand method may have limited comparability and incurs a lostopportunity to gather regional metadata for the northern quollin an area where any survey effort represents a significant

Research priorities for the northern quoll Australian Mammalogy C

Tab

le1

Researchpriorities

fortheno

rthern

quollan

dkeyqu

estion

sassociated

witheach

priorityan

das

compa

redwiththeinform

ationrequ

ired

undersimila

rthem

esan

dthepriority

givento

thisin

Woina

rski

etal(201

4)

Rank

Theme

Researchpriority

Key

questio

nsInform

ationrequired

andpriority

givenundersimilarthem

ein

Woinarski

etal(2014)

1Surveyand

mon

itoring

Develop

appropriate

andstandardised

survey

and

mon

itoring

metho

ds

Whatistheefficacy

ofthemajor

survey

metho

dscurrently

used

forno

rthern

quollsin

the

Pilb

ara

How

dovariou

scombinatio

nsoftrap

typesarraysdurationoftrappingbaitsetc

compareWhich

camera-trap

protocol

ismosteffectivefor(a)identifying

individu

alno

rthern

quollsin

thePilb

araand(b)estim

atingpo

pulatio

nsizes

Which

methodismost

cost-effectiv

eandefficientin

variou

sPilb

araland

scapes

Whatisthedetectionprobability

ofno

rthern

quollsin

each

habitattypeBased

onprior

surveywhatitthepriorprobability

ofoccupancyin

thesehabitattypes

Based

onthis

know

ledge

how

manynon-detections

ofquollsarerequired

before

wecanstatewith

prespecified

confi

dencethat

northern

quollsareabsent

from

asite

Surveyto

betterdefine

fine-scale

distributio

nandnumberof

individu

alsin

subpop

ulations

mediumndashhigh

Establishor

enhancemon

itoring

program

bydesigningintegrated

mon

itoring

prog

ramsmedium

2Habitat

requirem

ents

Define

areasof

critical

habitatforno

rthern

quollin

thePilb

ara

How

donorthern

quollsusehabitatinthePilb

ara

How

arequollpopulations

distributedin

relatio

nto

habitatq

uality(egh

ighqu

ality

low

quality

orsink

habitats)

How

ishabitat

selectioninfluenced

bypredationrisk

andsocial

factors

Whatinformationisrequ

ired

tofurtherrefine

anSDM

forno

rthern

quollsin

thePilb

ara

How

canthisinform

ationbe

mosteffectivelyused

toim

provemonito

ring

effortand

outcom

es

How

domod

elsvary

across

thePilb

aramainlandandoffsho

reislandsaccountin

gfor

clim

atic

gradientsandtopography

Whatarefuture

predictio

nsfornorthern

quollpopulatio

ndistributio

nsu

nder

modelsof

clim

atechange

andcane

toad

invasion

Assesshabitatrequirem

entsna

Und

erstandho

wdisturbanceaffects

habitatqu

ality

and

conn

ectiv

ity

How

does

fire

andgrazingaffecthabitatq

ualityandhabitatu

seby

northern

quollsin

the

Pilb

ara

How

importantisfire

asabarrierto

dispersal

How

importantishabitatconnectivity

form

etapopulations

ofnorthernquollinthePilb

ara

How

does

habitattypeaffecttheconnectiv

ityof

metapopulationsHow

does

disturbance

both

discrete

(eginfrastructure

such

asrailandroads)

anddiffuse(egchangesin

vegetatio

ncompositio

nandstructure)alterhabitatconnectiv

ity

Assessrelativ

eim

pact

ofthreats

quantifyrelativ

eim

pact

offire

regimesmdashhigh

Assesseffectivenessof

threat

mitigatio

nop

tionsland

scape-scale

optio

nsforretentionof

longer-

unburntpatchesmdash

high

3

Pop

ulation

dynamics

Betterunderstand

the

popu

latio

ndy

namicsof

the

northern

quollinthe

Pilb

ara

How

manysubpop

ulations

aretherein

thePilb

ara

How

dotheseinteractAre

subpopulations

demographically

stablein

spaceandtim

eHow

does

habitatand

clim

ate

affect

popu

latio

ndy

namics

How

does

disturbanceaffect

demog

raphic

stability

Assessdietlifehistory

na

Investigatepo

pulatio

nstructureandthe

interactionbetween

popu

latio

ns

How

domeasuresofgenetic

distance

relatetospatialecology

asmeasuredby

otherm

eans

iedoes

inferred

genetic

connectiv

ityrelate

toobserved

dispersal

Can

genetic

inform

ationbe

used

todeterm

inehabitatbarriers

Resolve

taxonomic

uncertainties

assess

genetic

relatednessof

spatially

disparatepo

pulatio

nsto

identifywhether

thereare

genetically

distinct

subpop

ulationsmdashlowndashmedium

D Australian Mammalogy V A Cramer et al

4Key

threats

Introduced

predators

Doespredationsignificantly

affecttheno

rthern

quollinthePilb

ara

How

does

thisvary

across

habitattypes

Doescompetitionforresourceswith

foxesor

catsim

pact

onsubpopulationsWhich

lifestages

aremostvulnerable

Given

thattheno

rthern

quollisalso

amesop

redatordo

esthepresence

ofding

oeseither

directly(through

predation)

orindirectly(through

theregulatio

nof

foxesandcats)impact

onsubpop

ulations

inthePilb

ara

Assessrelativ

eim

pact

ofthreats

predationby

feralcatsanddogsmdash

high

interactions

betweenferal

catsdo

gsdingoes

andfoxesmdash

mediumndashhigh

Assesseffectivenessof

threat

mitigatio

nop

tionsassess

which

feasible

managem

entoptio

nsfor

feralcatsanddo

gscanlead

topo

pulatio

nrecoverymdashmediumndash

high

Develop

new

orenhanceexistin

gmanagem

entmechanism

sfor

broad-scale

targetedferalcat

controlmdash

medium

Canetoads

Will

theclosureof

artifi

cialwater

bodies

preventor

simplyslow

therateofinv

asionof

cane

toadsinto

thePilb

ara

Whatisthedegree

ofoverlapbetweenSDMfundamentalniche

modelsforn

orthernquolls

andcane

toadsin

thePilb

ara

Can

potentialrefug

eareasforno

rthern

quollb

eidentifi

ed

IsCTAlik

elytobe

effectivewhentaste-aversion

baits

aredeliv

ered

underfi

eldcond

itions

Atwhatscaleisbaitdeliv

eryfeasible

Assesseffectivenessof

threat

mitigatio

nop

tionsassess

feasibility

ofbroad-scaletaste-

aversion

lsquotrainingrsquo

ofqu

ollsmdash

mediumassess

local-scale

exclusionof

toadsor

preventio

nof

colonisatio

n(m

ainlandsites)mdash

medium

Infrastructure

developm

entand

interactions

with

humans

Are

road-related

deaths

ofno

rthern

quollasign

ificant

causeof

mortalityin

thePilb

ara

Dorailw

aylin

esinfrastructurecorridorsor

wideroadspresenta

barrierto

northern

quoll

mov

ement

Whatmethods

canbe

used

tomitigate

theim

pactsof

infrastructure

developm

entand

associated

machinery

operation

Not

addressed

Interactions

between

threats

Dofrequent

andintensefireslead

toreducedstructural

complexity

loss

ofvegetatio

ncoverandsubsequentincreasedpredationof

northern

quollsparticularlyof

males

during

andor

afterbreeding

Whatarethecumulativeim

pactsof

mortalitycaused

bycane

toad

ingestionandpredation

bycatsdo

gsdingoes

andfoxes

See

inform

ationon

introd

uced

predatorsandcane

toadsabove

5Recolon

isationof

restored

orartifi

cial

habitat

Determinetheability

oftheno

rthern

quoll

torecolonise

disturbedareasor

colonise

artifi

cial

habitat

Whatarethekeyelem

entsof

northern

quollhabitatthatneedtobe

restoredcreated

sothat

habitatcomplexity

andprod

uctiv

ityismaxim

ised

Whatlevelo

flandscapeconn

ectiv

ityisrequired

arou

ndrestoredartificialhabitatso

that

genetic

connectiv

itybetweensubpopulations

canbe

maintained

How

long

should

restored

orartifi

cial

habitatbe

directly

managedparticularly

for

predators

How

cantheretentionof

largebouldersof

wasterock

beincorporated

into

rehabilitation

plans

Not

addressed

Research priorities for the northern quoll Australian Mammalogy E

investment of time and resources The limited amount anduncertain quality of occurrence data has been a major restrictionon the utility of a species distribution model (SDM) and apopulation viability analysis (PVA) for the northern quoll in thePilbara (Cadenhead et al 2015 see below) While survey andmonitoring per se is not a research activity that directly informsmanagement or threat mitigation carefully implementedmonitoring programs are essential to identify many elements ofa speciesrsquo ecology to detect changes in population and toquantify the ecological effectiveness and cost-effectivenessof management interventions (Lindenmayer et al 2013Lindenmayer 2015)

Parks and Wildlife recently released revised protocols forthe long-term monitoring of known populations of thenorthern quoll (Dunlop et al 2014b) and the effectiveness ofthese protocols in presenceabsence surveys should be testedThe cost-effectiveness of using camera traps for wildlifesurveys in remote locations such as the Pilbara makes their useappealing Hohnen et al (2013) described how the strategicplacement of cameras and baits combined with taking a greaternumber of photographs per trigger allows the capture ofimages that can be used to identify individual northern quollsfrom their spot patterns Remote cameras have also been shownto be a viable method for estimating population sizes of thenorthern quoll in areas with both dense (Austin 2014) andsmall populations (Webb et al 2015) Further testing of a rangeof parameters (eg see Nelson et al 2014) to develop a robuststandard protocol for the Pilbara is required along with acomparison of the cost-effectiveness and any ethicalreducedhandling benefits of using remote cameras versus live-trappingfor estimating population sizes in Pilbara landscapes

Priority 2 Improve our understanding of habitatrequirements

Define areas of critical habitat for the northern quollin the Pilbara

The Department of Agriculture and Food Western Australiahas mapped 102 land systems across the Pilbara bioregionbased on several biophysical components including landformgeology soils vegetation and drainage patterns (Van Vreeswyket al 2004) A review of records for the northern quoll in thePilbara showed that 65 of records were from the RobeCapricorn and Boolaloo land systems (Table 2) which compriselow plateaux mesas hills and ridges of varying geology(limonite granite and sandstone respectively) Most recentsurvey effort in the Pilbara is biased towards mineral-bearingformations or areas of mining-related infrastructure (eg railand road corridors) (Fig 1) Further investigation of theserecords is required to assess whether these land systems areindeed preferred habitat for northern quolls as opposed to anartefact of survey effort The comparatively fewer records thatcoincide with the conservation estate in the region (Fig 1) arealso likely an artefact of survey effort

The first step in defining areas of critical habitat for thenorthern quoll in the Pilbara is to improve our understanding ofhabitat use particularly in terms of defining habitat suitability atfiner scales than is currently available As part of the strategic

environmental assessments being undertaken for the Pilbarabioregion an SDM has been developed for the northern quoll(Cadenhead et al 2015) Uncertainty about the quality of theoccurrence data available the limited amount of data availableas well as the bias towards data collection in areas proposed fordevelopment constrained the level of certainty around theoutcomes of the model Improved model certainty requiresfurther collection of contemporary species-occurrence data andthe development of finer-resolution GIS layers particularly onethat captures finer-scale geological attributes such as mesas orrocky outcrops considered as highest-quality habitat for thenorthern quoll (Cadenhead et al 2015)

Better definition of the habitat attributes favoured by thenorthern quoll will allow further refinement of the GIS variablesincluded in the SDM or allow the development of predictivesite-based habitat models Information on how northern quollsuse habitat can be gained through direct observation of howthey interact with habitat features or from the association ofhabitat features with the presence of quolls as determined byGPS receivers (Gaillard et al 2010) This level of detail iscurrently not available from survey records returned as part ofenvironmental impact assessments but collection of habitatdata using the protocols outlined by Dunlop et al (2014b) infuture surveys would rapidly increase the quality of dataavailable

Understand how disturbance affects habitat qualityand connectivity

The alteration of fire regimes has been the most pervasivelandscape-scale change in the Pilbara Between 1993 and 2006over 72 of the region was burnt with more than 28 of thearea burnt two or more times (McKenzie et al 2009) Woinarskiet al (2004) found that northern quolls may be able to tolerate amoderately high frequency of fire as long as these fires werelow-intensity early-season burns and the impacts of fire onvegetation structure and composition were not exacerbatedby the grazing of introduced herbivores In the Pilbarahowever as with many other grazing regions of Australia earlypastoralists often overestimated the productivity and resilienceof native vegetation to sheep and cattle grazing which led tosevere degradation of vegetation particularly along riverfrontages (Hennig 2004) The soil degradation and loss ofvegetative cover caused by intense grazing pressure in thesouth Kimberley is thought likely to have contributed to thesubstantial decline or disappearance of medium-sized animalsincluding the northern quoll from the Bungle Bungle area(Woinarski et al 1992) Death by predation is most likely in openareas (Oakwood 2000) and therefore the effects of fire andgrazing on northern quoll populations cannot be consideredwithout also taking account of likely increases in mortality due topredation (see lsquoIntroduced predatorsrsquo below)

Understanding how the localised but intensive activityassociated with mining affects habitat quality and habitat useby the northern quoll in areas surrounding development islimited A recent study that examined the use of ballast quarriesand paired undisturbed sites by northern quolls found that theywere foraging denning and showing signs of breeding at thequarry sites where greater numbers of quolls were captured(Dunlop et al 2014a) This was likely due to the combination of

F Australian Mammalogy V A Cramer et al

habitat complexity (eg diversity of boulder sizes) and thepresence of permanent water in the quarry which may haveincreased prey availability at these sites

Movement of animals through the landscape for dispersaland repopulation is another important consideration for a speciesin which both sexes tend towards semelparity and is thereforepredisposed to local extinction (Oakwood 1997) Improvedunderstanding of how the northern quoll (particularly dispersingmales) uses areas of lower habitat quality and the identificationof such habitat in habitat-suitability models will allow futurelandscape-scale strategic assessments to better predict thecombined impacts of disturbance on the ability of northern quolls

to move through the landscape Analysis of regional populationgenetics will also improve our understanding of the level ofdispersal occurring across the landscape

Priority 3 Population dynamics

Better understand the population dynamics of thenorthern quoll in the Pilbara

Little information is available on the number ofsubpopulations of northern quoll in the Pilbara how thesesubpopulations interact how stable they are in space and timeand how they respond to disturbance As no demographic

Table 2 Records of the northern quoll in each land system type in the PilbaraRecords were current as of 23 May 2014

Land system Area (km2) No of records Land system description

Augustus 61 1 Rugged ranges hills ridges and plateaux supporting mulga shrublands and hard spinifex grasslandsBoolaloo 1502 245 Granite hills domes and tor fields and sandy plains with shrubby spinifex grasslandsBoolgeeda 7748 29 Stony lower slopes and plains below hill systems supporting hard and soft spinifex grasslands andmulga

shrublandsCalcrete 1444 2 Low calcrete platforms and plains supporting shrubby hard spinifex grasslandsCallawa 1003 9 Highly dissected low hills mesas and gravelly plains of sandstone and conglomerate supporting soft and

hard spinifex grasslandsCapricorn 5296 257 Hills and ridges of sandstone and dolomite shrubby hard and soft spinifex grasslandsCoongimah 3244 12 Plateau surfaces low hills with steep slopes and undulating uplands supporting hard spinifex grasslandsDune 138 1 Dune fields supporting soft spinifex grasslandsGranitic 4020 1 Rugged granitic hills supporting shrubby hard and soft spinifex grasslandsHorseflat 1261 4 Gilgaied clay plains supporting tussock grasslands and minor grassy snakewood shrublandsLittoral 1577 3 Bare coastal mudflats with mangroves on seaward fringes samphire flats sandy islands coastal dunes

and beachesMacroy 13 095 94 Stony plains and occasional tor fields based on granite supporting hard and soft spinifex grasslandsMcKay 4202 21 Hills ridges plateau remnants and breakaways of metasedimentary rocks supporting hard spinifex

grasslandsMallina 2557 8 Sandy surfaced alluvial plains supporting soft spinifex (and occasionally hard spinifex) grasslandsMarandoo 459 2 Basalt hills and restricted stony plains supporting grassy mulga shrublandsMosquito 1840 3 Stony plains and prominent ridges of schist and other metamorphic rocks supporting hard spinifex

grasslandsNewman 14 580 81 Rugged jaspilite plateaux ridges and mountains supporting hard spinifex grasslandsOnslow 424 1 Sandplains dunes and clay plains supporting soft spinifex grasslands and minor tussock grasslandsParaburdoo 565 3 Basalt-derived stony gilgai plains and stony plains supporting snakewood and mulga shrublands with

spinifex and tussock grasslandsParadise 1479 8 Alluvial plains supporting soft spinifex grasslands and tussock grasslandsRiver 4088 37 Active flood plains major rivers supporting grassy eucalypt woodlands tussock grasslands and soft

spinifex grasslandsRobe 865 369 Low limonite mesas and buttes supporting soft spinifex (and occasionally hard spinifex) grasslandsRocklea 22 993 40 Basalt hills plateaux lower slopes and minor stony plains supporting hard spinifex (and occasionally

soft spinifex) grasslandsRuth 346 36 Hills and ridges of volcanic and other rocks supporting hard spinifex (occasionally soft spinifex)

grasslandsStuart 1794 3 Gently undulating stony plains supporting hard and soft spinifex grassland and snakewood shrublandsTaylor 129 3 Stony plains and isolated low hills of sedimentary rocks supporting hard and soft spinifex grasslandsTalga 2124 9 Hills and ridges of greenstone and chert and stony plains supporting hard and soft spinifex grasslandsTuree 581 2 Stony alluvial plains with gilgaied and non-gilgaied surfaces supporting tussock grasslands and grassy

shrublandsUaroo 7681 37 Broad sandy plains supporting shrubby hard and soft spinifex grasslandsUrandy 1311 6 Stony plains alluvial plains and drainage lines supporting shrubby soft spinifex grasslandsWhite Springs 266 4 Stony gilgai plains supporting tussock grasslands and hard spinifex grasslandsWona 1815 10 Basalt upland gilgai plains supporting tussock grasslands and minor hard spinifex grasslandsYamerina 1207 1 Floodplains and deltaic deposits supporting tussock grasslands grassy woodlands and minor halophytic

low shrublands

Research priorities for the northern quoll Australian Mammalogy G

studies have been undertaken in the region Cadenhead et al(2015) used demographic information from the NorthernTerritory and Kimberley populations in their PVA Uncertaintyabout data quality and the limited information available ondemographic parameters for the Pilbara constrained the level ofcertainty around the outcomes of the PVA The model doeshowever provide a framework into which new data can beincorporated as they become available For example preliminarydata from the Pilbara monitoring program suggest that theproportion of male die-off in the Pilbara population is lowerthan in the Kimberley and Northern Territory populations(Dunlop unpubl data) Cadenhead et al (2015) stress the needfor more demographic and occurrence data to be collectedbefore further development decisions are made (see lsquoDevelopappropriate and standardised survey and monitoring methodsrsquoand lsquoImprove our understanding of habitat requirementsrsquo above)

Investigate population structure and the interactionbetween populations

Analysis of the genetic diversity and population structure ofthe Pilbara population of the northern quoll is ongoingOnly small differences in genetic diversity as measured byheterozygosity were found between samples obtained at 16locations within the Pilbara with the exception of the isolatedpopulation on Dolphin Island (Spencer et al 2013) This resultindicates a high level of movement across the landscape byindividuals during mating and dispersal phases Spatial structureanalysis of samples collected from around Pannawonicasuggested a lsquoneighbour sizersquo of ~15 km (Spencer et al 2013)Current data show no evidence of genetic bottleneckssuggesting that if there has been a population decline in thePilbara that this has not affected genetic diversity A usefulapproach would be to combine genetic information with datafrom radio-tracking or GPS collars to better understand theuse of lsquolandscapersquo space and how this influences populationdynamics

Priority 4 Key threats and the interaction of these threats

Introduced predators

Introduced cats and foxes are considered to be one of themain causes of the decline and extinction of the critical-weight-range mammals (35 g to 55 kg) in Australia (Short and Smith1994 McKenzie et al 2007 Woinarski et al 2015) but theirimpacts on the northern quoll are less clear A recent review offeral cat diets across Australia found that northern quolls werepresent in the diet of feral cats (Doherty et al 2015a) andWoinarski et al (2014) consider the threat posed by cats to thenorthern quoll to be severe although the impacts of cats may bereduced in rugged refuge areas The impacts of cats on othermuch larger species of quoll appear to be inconsistent withsome evidence of coexistence between cats and quolls beforethe arrival of foxes (Glen and Dickman 2008 and referencescontained within) Due to the limited overlap in their distributionthroughout Australia the threat posed by foxes to northernquolls is less understood than for cats and Woinarski et al(2014) consider the threat posed by foxes to be moderate on anational level Foxes are common in coastal near-coastal and

riparian areas of the Pilbara and extend as far north as Broome(King and Smith 1985 Kendrick and Stanley 2002) They arerenowned for their aggression towards smaller carnivores(Palomares and Caro 1999) and neither the larger chuditch(Dasyurus geoffroii) nor spotted-tail quoll (Dasyurus maculatus)are known to coexist with large numbers of foxes (A Glen perscomm) The interactions between quolls cats and foxesare potentially more complex than for other medium-sizedmarsupials due to niche overlap in diet and competition for densites with the larger spotted-tailed quoll and chuditch bothexperiencing exploitation and interference competition fromfoxes (Morris et al 2003 Glen et al 2011) Competitionbetween feral cats and the northern quoll may potentially beintense during periods of low rainfall in the Pilbara when preyavailability is low Understanding whether foxes and cats aresignificant predators of or competitors with the northern quollin the Pilbara and how this affects their population viabilitywill clarify the level of risk posed by these predators Thesausage-style baits used for feral cats (eg Eradicatreg andCuriosityreg) are potentially attractive to the northern quoll withthe risk of mortality or reduced fitness Trials to assess themagnitude of the risks from Eradicatreg baits are currentlyunderway in the Pilbara

The role of dingoes and wild dogs in the mortality of andinteractions between mesopredators (both native and introduced)requires further investigation Dingoes are known to kill northernquolls butmay not eat these kills (Oakwood 1997) A recent studyin the Northern Territory found that camp dogs and dingoes werethe major source of mortality for northern quolls at the study site(Webb et al 2015) and roaming domestic dogs were identified asa cause of mortality for northern quolls in Queensland (Pollock1999) Human resource subsidies (eg stock carrion in rangelandareas) can artificially support predator populations thus indirectlyincreasing predation pressure on prey (Newsome et al 2015)A dynamic debate currently exists around the role of the dingo inregulating populations of feral cats and foxes and the implicationsof this for the conservation of the critical-weight-range mammalsin Australia (see Johnson et al 2007 Letnic et al 2009 Allenet al 2011 2013 Kennedy et al 2012) Understanding thepopulation density at which the role of the dingo as a regulator ofintroduced predators outweighs its own impact as a direct predatorof the northern quoll will inform whether dingowild dog controlwould benefit the northern quoll

Cane toads

The introduced cane toad now occupies more than half thehistoric range of the northern quoll and has caused an immediatecollapse or extirpation of northern quoll populations in theareas it has invaded (Burnett 1997 Oakwood and Foster2008 Woinarski et al 2008 2010) Earlier modelling work thatconsidered only climate parameters as well as more recentwork that incorporates future climate change suggest thatcane toads will not invade the Pilbara (Sutherst et al 1996Pavey and Bastin 2014) Other recent work that considers thefundamental niche of the cane toad and incorporates the locationof artificial watering points predicts that they will colonisethe Pilbara from the Kimberley via the coastal edge of theGreat Sandy Desert (Kearney et al 2008 Florance et al 2011

H Australian Mammalogy V A Cramer et al

Tingley et al 2013) This has created some level of uncertaintyabout the likely impact of cane toads on northern quolls in thePilbara Strategies for cane toad management were not rankedhighly by participants in the threat-management frameworkdeveloped by Carwardine et al (2014) which the authorsexplained as possibly due to uncertainty about how far canetoads will spread into the region

The model developed by Tingley et al (2013) suggested thatthe closure of 100 artificial water bodies at one of three potentiallocations along the coastal corridor between the Kimberley andthe Pilbara would prevent the invasion of cane toads into thePilbara While their model did account for uncharacteristicallywet years every three decades the dispersal of toads betweenwater bodies was determined by the number of rainy daysoccurring during the year (ie toads could only disperse betweenwater bodies for three days following rainfall) rather thanrainfall amount the associated likelihood of flooding and theavailability of free water across the landscape The cyclonic andthunderstorm nature of much of the rainfall in the north-west ofWestern Australia means that the region is prone to significantand sometimes extended flooding In April 1934 the countryaround Anna Plains (250 km south of Broome) was flooded forhundreds of miles when 27 inches (686mm) of rain fell infour days (Anon 1934) In March 2004 over 350mm of rain wasrecorded within a 48-h period at Telfer on the southern edge ofthe Great Sandy Desert due to Tropical Cyclone Fay Thiscaused the flooding of the De Grey and Oakover river systemsand flooded 140 km of the Telfer Access Road with floodwaterstaking several months to subside (wwwbomgovau) Extendingthe model of Tingley et al (2013) to account for such events willhelp clarify whether closing artificial water bodies is likely toprevent the invasion of cane toads into the Pilbara or may onlyassist in delaying their inevitable invasion

Comparison of SDMs (eg Cadenhead et al 2015) fornorthern quoll with fundamental niche models for cane toads(eg see Kearney et al 2008) within the Pilbara may identifypotential refuge areas for the northern quoll and these areasshould be prioritised for conservation actions Because theinvasion point of cane toads into the Pilbara is likely to be narrow(beginning around the mouth of the De Grey River Tingleyet al 2013) a proactive approach to training northern quollsto avoid cane toads may prove beneficial Conditioned taste-aversion (CTA) has been used successfully to train captive-reared northern quolls to avoid cane toads by offering themsmall dead toads infused with thiabendazole before theirrelease into the wild (OrsquoDonnell et al 2010)Monitoring of thesequolls after their release showed that some of their offspringhad learnt to avoid eating cane toads either through ingestionof small non-lethal toads or via social learning from theirmothers (Webb et al 2015) Fields trials of toad-aversion baits(either sausages or meatballs made from toad flesh andcontaining thiabendazole) are currently underway in the centralKimberley where cane toads were expected to invade in late2015 (Webb et al 2015) Further testing of CTA approachesunder a variety of field conditions is required The developmentof stable long-lasting baits suitable for aerial deployment isconsidered a pressing need as well as a captive colony ofnorthern quolls upon which to test the efficacy of toad-aversionbaits for inducing aversion to live toads (Webb et al 2015)

Infrastructure development and interactionswith humans

Infrastructure development may increase the likelihood ofnorthern quoll deaths due to motor vehicle strikes or beingcrushed in machinery Significant chuditch mortality wasobserved along mine access roads in the south-west of WesternAustralia (McGregor et al 2014) and the widening and sealingof an access road dramatically increased the number of roadkills and led to the extirpation of the resident population ofeastern quolls (D viverrinus) in Tasmania (Jones 2000)Infrastructure corridors such as high-tension power-lineeasements and ore-haulage railways are increasingly beinginstalled throughout the range of the northern quoll Althoughthese do not create the same mortality risk as roads they bisectand fragment habitat create large open spaces and potentiallyincrease predation risk

A recent small-scale radio-tracking study (see Dunlop et al2014a) around several quarries in the Pilbara found that northernquolls crossed roads but not railway lines The short-term natureof this work and the limited number of quolls able to be trackedmeans that the influence of roads railway lines and infrastructurecorridors on the movement and mortality of northern quollsrequires further assessment Using radio-tracking techniques todetermine crossing rates and road kill events may be difficultover large areas so population studies comparing groups ofquolls near major roads with those not affected by infrastructure(eg McCall et al 2010) may provide a more feasible meansof assessing the impacts of roads If vehicle strikes are foundto significantly contribute to the mortality of a local populationof the northern quoll then mitigation methods should betrialled on sections of road identified as road-kill hotspotsCulverts and underpasses have been shown to be used by avariety of small and medium-sized mammals and frogs (Taylorand Goldingay 2003) while installing rumble strips on roadshas led to decreased rates of road-kill of nocturnal animals(Lester 2015)

Interactions between threats

At present wildfire the subsequent loss of vegetation coverand increased predation of northern quolls is likely to be themost significant of the interactions between threats The directeffects of wildfire have been found to be less significant inthe subsequent decline of small to medium-sized mammalsparticularly for denning species such as the northern quoll thanthe indirect consequences of reduced resource availability andthe increased risk of predation due to lack of cover (Oakwood2000 Letnic et al 2005 McKenzie et al 2007 Legge et al2008 Doherty et al 2015b) Male northern quolls are perhapsmost vulnerable to such losses of cover during the breedingseason when their range can extend to over 100 ha (Oakwood2002) or where males survive after breeding their foragingrange is extended to gain the nutrition required to recover fromtheir poor condition (Cook 2010) Loss of cover due to late dry-season fires at East Alligator River in the Northern Territorymeant little cover was available for young quolls when theybegan foraging with dog and dingo predation a major source ofmortality (Webb et al 2015) The cumulative impacts of canetoad poisoning fire and predation are highly likely to lead to the

Research priorities for the northern quoll Australian Mammalogy I

extirpation of small and disjunct groups of northern quoll in thePilbara Population viability modelling of the East AlligatorRiver population suggested that high mortality from predationcombined with late dry-season burning was preventing therecovery of northern quoll populations after cane toad invasion

(Webb et al 2015) Improved knowledge of the severity of thesethreats to northern quolls in the Pilbara and their incorporationinto the SDM and PVA framework developed by Cadenheadet al (2015) will allow for better identification of priority areasfor integrated cane toad fire and predator management

Table 3 Expected timelines for the undertaking of research under each priority over the next 10 years and the funds currently allocated fromthe Pilbara Northern Quoll Offset Trust under each research priority

Rank Theme Research priority Current or proposed researchactivity

Time over next10 years

Current funding allocation(approximate)

1 Survey andmonitoring

Develop appropriate andstandardised survey andmonitoring methods

Develop monitoring protocol(traps and cameras) andinvestigate new methods

Ongoing survey and monitoringin conjunction with ResearchThemes 2 and 3 to better mapnorthern quoll distribution andto determine long-termpopulation trends

Years 1ndash2Years 3ndash10

$200 000 per annum over10 years for ResearchThemes 1 2 and 3(inclusive of salaries)

2 Habitatrequirements

Define areas of criticalhabitat for northernquoll in the Pilbara

Identify and describe criticalhabitat and undertake predictivemodelling of unsurveyed areasto inform further survey effort

Years 3ndash4 $50 000 for predictivemodelling (EdithCowan University)

Understand howdisturbance affectshabitat quality andconnectivity

Ongoing collection of data ondisturbance and habitat qualityin conjunction with ResearchThemes 1 and 4

Incorporation of new data intopredictive models

Years 4ndash10 See above

3 Populationdynamics

Better understand thepopulation dynamics ofthe northern quoll in thePilbara

Collect data on demographicparameters breeding dispersaland home-range use toincorporate into future PVA

Continued collectionof data inconjunction withpriorities 1 and 2during years 3ndash10

See above

Investigate populationstructure and theinteraction betweenpopulations

Analysis of existing tissuesamples and newly collectedsamples

Years 1ndash10 $120 000 for initial roundof genetic analysis(Years 3ndash5 MurdochUniversity)

4 Key threats Introduced predators Spatio-temporal interactionsbetween northern quolls andintroduced predators

Other projects as they arise

Years 2ndash4 (currentPhD projectUniversity ofQueensland)

$12 000 contribution tofield expenses

Cane toads Predictive modelling of canetoad distribution in the Pilbara

Years 3ndash4 Part of predictivemodelling by EdithCowan University

Infrastructuredevelopment andinteractions withhumans

Investigation of impacts oflinear infrastructure on northernquoll movement

Year 4 (currentHonours projectEdith CowanUniversity)

$15 000

Interactions betweenthreats

No on-ground projects currentlyplanned

Potential to be assessed throughPVA as data are collected underother research priorities

No funding currentlyallocated

5 Recolonisation ofrestored orartificial habitat

Determine the ability ofthe northern quoll torecolonise disturbedareas or coloniseartificial habitat

As projects arise Years 5ndash8 incollaboration withminesite restorationprograms

No funding currentlyallocated

J Australian Mammalogy V A Cramer et al

Priority 5 Recolonisation of restored andor artificial habitat

While anecdotal evidence suggests that northern quolls will denin waste rock dumps whether artificial or restored habitatprovides for long-term population viability is not known andlong-term assessment of created habitat is required Weacknowledge that a large gap exists between the desire to userestored or artificial habitat as an ecologically equivalent offsetand our limited ability to actually restore or re-create habitat thatis equivalent in structure composition and function particularlyon highly disturbed sites (Maron et al 2012) Yet there is a futureopportunity in the Pilbara to conduct lsquonaturalrsquo experiments onthe recolonisation of quolls after mining ceases including howto best design waste rock dumps so that habitat complexityand productivity is maximised by for example experimentingwith the size and positioning of boulders and their spatialarrangement in relation to surrounding landscape features Theuse of such artificial habitat should be tested in the first stagethrough direct monitoring of animals that have colonised orhave been translocated to the site Such experimentation isunderway at Atlas Ironrsquos Mt Dove operation (Atlas Iron 2012)Population modelling can then be used to predict futurepopulation sizes rates of growth and spread and the probabilityof long-term persistence in the context of various managementactions (eg predator control Armstrong and Reynolds 2012)

Resource allocation and timelines

While the allocation of resources and funding and the periodover which each of the research priorities was expected to beaddressed was not explicitly discussed during the workshopthe ranking of the priorities was taken to indicate the sequencein which the priorities should be addressed At the time of theworkshop some of the research questions identified werealready being investigated within various projects and a portionof the Pilbara Northern Quoll Offset Trust allocated to these InTable 3 we indicate the expected timelines over which someof the research programs are expected to be undertaken WhileAU$175M has been allocated to northern quoll research in thePilbara it is important to note that much of this amount is notcurrently available and will not become available until severalapproved mine developments have become operational InTable 3 we also indicate the proportion of offset funds expectedto be allocated to each research priority Research undertaken inother regions (eg research into CTA in the Kimberley regionahead of the cane toad invasion front) cannot be funded from thePilbara Northern Quoll Offset Trust but will help address someof the research questions identified here

Lessons learnt from the workshop process

The workshop approach used here was less formal than recentlarge-scale exercises used to elicit research priorities for emergingissues in science and policy at national or global levels (reviewedin Sutherland et al 2011) The focus in our (much smaller)workshop was on inclusivity rather than identifying participantsbased on purposive sampling of expertise or organisationalaffiliation We believe that the inclusive nature of the workshopwas a strength but we also recognise that this perhaps createdsome level of bias when ranking the research prioritiesAcknowledging both the experience and knowledge requirements

of those who work regularly in the Pilbara is an importantcomponent of improving the design of research programs that aimto improve conservation management in the region Yet theresearch priorities and their rankings are likely strongly reflectiveof the current information needs of most participants and maynot align with what conservation scientists believe to be themost critical areas of research for northern quoll conservation intothe future Baseline survey and monitoring are often the firstrequirements of a management or research plan for the northernquoll when offset conditions are prescribed and this was reflectedin the primary ranking given to the development of standardsurvey and monitoring methods We acknowledge the concernthat this could simply lead to offset funding being used to lsquomonitorthe extinctionrsquo of the northern quoll particularly after canetoads invade the Pilbara unless these data are used to triggermanagement actions (Oakwood and Foster 2008 Lindenmayeret al 2013) Clearly the currently limited information on habitatuse and demographic parameters for the northern quoll in thePilbara restricts the utility of more sophisticated researchapproaches such as developing robust SDM and PVA While thesevere threat that cane toads pose to the northern quoll wasrecognised this threat is not immediate in the region and mayexplain the less prominent ranking given to research to mitigatethe impacts of cane toads

The northern quoll has become emblematic of the dramaticand rapid species declines occurring in northern Australia andthe level of interest in this species was reflected in the highnumber of participants attending the workshop (one of six runby Parks and Wildlife) Given the number of attendees betterutilisation of the time available would have been gained byproviding participants with a brief before the workshopand asking them to contribute potential research questionsanonymously for interactive review by other participants Inthe exercises reviewed by Sutherland et al (2011) questionswere then discarded if they were not considered to meet criteriathat were previously agreed upon Some areas of researchthat gained some attention on the day may have been discardedif given greater consideration or interrogation before theworkshop For example there was some discussion duringthe workshop of investigating the use of habitat surrogates tomonitor the northern quoll yet research has shown that there islittle validity in using surrogates to monitor the abundance of asingle species (Pierson et al 2015)

Conclusions

Offset funds generated under the environmental approvalsprocess provide an opportunity to mobilise considerablefinancial resources for conservation (Kiesecker et al 2010) Yetthere remain significant conceptual and practical limitations tobe resolved to ensure that direct offsets provide measurableconservation outcomes that are ecologically equivalent to thelosses sustained by proceeding with the development (Maronet al 2012 Bull et al 2013) The collaborative identificationand ranking of research priorities undertaken here is the firststep in the process of developing and evaluating potentialconservation actions for the northern quoll in the Pilbara andserves as a mechanism to ensure that the research priorities ofstate and federal agencies reflects the knowledge needs of

Research priorities for the northern quoll Australian Mammalogy K

practitioners and proponents currently operating in the regionThe provision of offset funds over the next decade providesa unique opportunity to undertake research on a species ofconservation significance at a regional scale and over a longperiod through an integrated and strategic program of researchrather than the collection of ad hoc data While a morestructured approach to defining research questions before theworkshop may have led to greater critical reflection upon anddiscussion of some of the research priorities on the day theworkshop provided an opportunity for an open and diverse-ranging discussion in a collaborative environment amongstthe broader community of scientists environmental consultantsand mining proponents involved in the management andconservation of the northern quoll in the Pilbara

Acknowledgements

We gratefully acknowledge the financial support of Roy Hill who sponsoredthe workshop and partially funded VACrsquos salary We thank the Departmentof the Environmentrsquos Pilbara Taskforce especially Veronica Ritchie fortheir support of interested or affected parties The success of the workshopwas in great part due to the facilitation skills of Craig Salt of SustainableConsulting We thank all of the workshop participants for their open andcollaborative approach to the process Mike Bamford Eddie van EttenDavid Peacock and Peter Lyon provided useful comments on an earlier draftWe are grateful to John Woinarski for his constructive advice thatsubstantially improved the structure and clarity of the manuscript and ananonymous reviewer for careful and critical reading of the previousversion Lesley Gibson Russell Palmer and Al Glen generously discussedvarious aspects of the paper with VAC The map was prepared by CristinaRamalho

References

Allen B L Engeman R M and Allen L R (2011) Wild dogma anexamination of recent lsquoevidencersquo for dingo regulation of invasivemesopredator release in Australia Current Zoology 57 568ndash583

Allen B L Fleming P J Allen L R Engeman R M Ballard G andLeung L K-P (2013) As clear as mud a critical review of evidencefor the ecological roles of Australian dingoes Biological Conservation159 158ndash174 doi101016jbiocon201212004

Anon (1934) Floods in Western Australia Town isolated for week PerthMonday The Sydney Morning Herald 9

Armstrong D P and Reynolds M H (2012) Modelling reintroducedpopulations the state of the art and future directions In lsquoReintroductionBiology Integrating Science and Managementrsquo (Eds J G Ewen DP Armstrong K A Parker and P J Seddon) pp 165ndash222 (Wiley-Blackwell UK)

Atlas Iron (2012) Scope of works ndash Mount Dove artificial quoll habitatUnpublished report Atlas Iron Perth Available at httpwwwatlasironcomauIRMCompanyShowPageaspxCategoryId=190ampCPID=5424ampEID=94365410 [accessed 19 August 2015]

Austin C (2014) Can remote cameras accurately estimate populations ofthe endangered northern quoll BSc(Honours) Thesis University ofTechnology Sydney

Begg R J (1981) The small mammals of Little Nourlangie RockNT III Ecology ofDasyurus hallucatus the northern quoll (MarsupialiaDasyuridae) Wildlife Research 8 73ndash85 doi101071WR9810073

Braithwaite R W and Griffiths A D (1994) Demographic variationand range contraction in the northern quoll Dasyurus hallucatus(Marsupialia Dasyuridae)Wildlife Research 21 203ndash217 doi101071WR9940203

Bull J W Suttle K B Gordon A Singh N J and Milner-Gulland E J(2013) Biodiversity offsets in theory and practice Oryx 47 369ndash380doi101017S003060531200172X

Burnett S (1997) Colonizing cane toads cause population declines innative predators reliable anecdotal information and managementimplications Pacific Conservation Biology 3 65ndash72

Cadenhead N C R White A L and Wintle B A (2015) Identifyingconservation priorities and assessing impacts of proposed futuredevelopment in the Pilbara Bioregion in Western Australia Report bythe National Environment Research Program Decisions Hub to theDepartment of the Environment The University of MelbourneMelbourne

Carwardine J Nicol S Van Leeuwen S Walters B Firn J Reeson AMartin T G and Chades I (2014) Priority threat management forPilbara species of conservation significance CSIRO EcosystemSciences Brisbane

Cook A (2010) Habitat use and home-range of the northern quollDasyurushallucatus effects of fire MSc Thesis University ofWestern AustraliaPerth

Department of Sustainability Environment Water Population andCommunities (2012) Environment and Biodiversity Conservation Act1999 Environmental Offsets Policy Department of SustainabilityEnvironment Water Population and Communities Canberra

Doherty T S Davis R A Etten E J Algar D Collier N DickmanC R Edwards G Masters P Palmer R and Robinson S (2015a)A continental-scale analysis of feral cat diet in Australia Journal ofBiogeography 42 964ndash975 doi101111jbi12469

Doherty T S Dickman C R Nimmo D G and Ritchie E G (2015b)Multiple threats or multiplying the threats Interactions betweeninvasive predators and other ecological disturbances BiologicalConservation 190 60ndash68 doi101016jbiocon201505013

Dunlop J Cook A and Lees J (2014a) BHP Billiton Iron Ore Pilbaranorthern quoll monitoring project Final report May 2010ndashJune 2013Unpublished report prepared for BHP Billiton Iron Ore Department ofParks and Wildlife Perth

Dunlop J Cook A and Morris K (2014b) Pilbara northern quollproject surveying and monitoring Dasyurus hallucatus in the PilbaraWestern Australia Department of Parks and Wildlife Perth Availableat librarydpawwagovau

Florance D Webb J K Dempster T Kearney M R Worthing A andLetnic M (2011) Excluding access to invasion hubs can contain thespread of an invasive vertebrate Proceedings of the Royal Society ofLondon B Biological Sciences 278 2900ndash2908 doi101098rspb20110032

Gaillard J-M Hebblewhite M Loison A Fuller M Powell R BasilleM and Van Moorter B (2010) Habitatndashperformance relationshipsfinding the right metric at a given spatial scale PhilosophicalTransactions of the Royal Society of London Series B BiologicalSciences 365 2255ndash2265 doi101098rstb20100085

Glen A S and Dickman C R (2008) Niche overlap between marsupialand eutherian carnivores does competition threaten the endangeredspotted-tailed quoll Journal of Applied Ecology 45 700ndash707 doi101111j1365-2664200701449x

Glen A S Pennay M Dickman C R Wintle B A and Firestone K B(2011) Diets of sympatric native and introduced carnivores in theBarrington Tops eastern Australia Austral Ecology 36 290ndash296doi101111j1442-9993201002149x

Hennig P (2004) A brief land use history In lsquoAn Inventory and ConditionSurvey of the Pilbara Region Western Australiarsquo (Eds A M E VanVreeswyk A L Payne K A Leighton and P Hennig) TechnicalBulletin No 92 pp 13ndash18 (Western Australian Department ofAgriculture Perth)

Hohnen R Ashby J Tuft K and McGregor H (2013) Individualidentification of northern quolls (Dasyurus hallucatus) using remotecameras Australian Mammalogy 35 131ndash135 doi101071AM12015

L Australian Mammalogy V A Cramer et al

How R A Spencer P B S and Schmitt L H (2009) Island populationshave high conservation value for northern Australiarsquos top marsupialpredator ahead of a threatening process Journal of Zoology 278206ndash217 doi101111j1469-7998200900569x

Johnson C N Isaac J L and Fisher D O (2007) Rarity of a top predatortriggers continent-wide collapse of mammal prey dingoes andmarsupials in Australia Proceedings of the Royal Society of LondonSeries B Biological Sciences 274 341ndash346 doi101098rspb20063711

Jones M E (2000) Road upgrade road mortality and remedial measuresimpacts on a population of eastern quolls and Tasmanian devilsWildlifeResearch 27 289ndash296 doi101071WR98069

Kearney M Phillips B L Tracy C R Christian K A Betts G andPorter W P (2008) Modelling species distributions without usingspecies distributions the cane toad in Australia under current andfuture climates Ecography 31 423ndash434 doi101111j0906-7590200805457x

Kendrick P and Stanley F (2002) Pilbara 4 (PIL4 ndash Roebourne synopsis)In lsquoA Biodiversity Audit of Western Australiarsquos 53 BiogeographicalSubregions in 2002rsquo (Eds J E May and N L McKenzie) pp 581ndash593(Department of Conservation and Land Management Perth)

Kennedy M Phillips B L Legge S Murphy S A and Faulkner R A(2012) Do dingoes suppress the activity of feral cats in northernAustralia Austral Ecology 37 134ndash139 doi101111j1442-9993201102256x

Kiesecker J M Copeland H Pocewicz A and McKenney B (2010)Development by design blending landscape-level planning with themitigation hierarchy Frontiers in Ecology and the Environment 8261ndash266 doi101890090005

King D R and Smith L A (1985) The distribution of the European red fox(Vulpes vulpes) in Western Australia Records of the Western AustralianMuseum 12 197ndash205

Knight A T Cowling R M Rouget M Balmford A Lombard A Tand Campbell B M (2008) Knowing but not doing selecting priorityconservation areas and the researchndashimplementation gap ConservationBiology 22 610ndash617 doi101111j1523-1739200800914x

Laurance W F Koster H Grooten M Anderson A B Zuidema P AZwick S Zagt R J Lynam A J Linkie M and Anten N P (2012)Making conservation research more relevant for conservationpractitioners Biological Conservation 153 164ndash168 doi101016jbiocon201205012

Legge S Murphy S Heathcote J Flaxman E Augusteyn J andCrossman M (2008) The short-term effects of an extensive and high-intensity fire on vertebrates in the tropical savannas of the centralKimberley northern Australia Wildlife Research 35 33ndash43 doi101071WR07016

Lester D (2015) Effective wildlife roadkill mitigation Journal of Trafficand Transportation Engineering 3 42ndash51 doi10172652328-2142201501005

Letnic M Tamayo B and Dickman C R (2005) The responsesof mammals to La Nintildea (El Nintildeo Southern Oscillation)-associatedrainfall predation and wildfire in central Australia Journal ofMammalogy 86 689ndash703 doi1016441545-1542(2005)086[0689TROMTL]20CO2

Letnic M Koch F Gordon C Crowther M S and Dickman C R(2009) Keystone effects of an alien top-predator stem extinctions ofnative mammals Proceedings of the Royal Society of London Series BBiological Sciences 276 3249ndash3256 doi101098rspb20090574

Lindenmayer D B (2015) Continental-level biodiversity collapseProceedings of the National Academy of Sciences of the United States ofAmerica 112 4514ndash4515 doi101073pnas1502766112

Lindenmayer D B Piggott M P and Wintle B A (2013) Counting thebooks while the library burns why conservation monitoring programs

need a plan for action Frontiers in Ecology and the Environment 11549ndash555 doi101890120220

Maron M Hobbs R J Moilanen A Matthews J W Christie KGardner T A Keith D A Lindenmayer D B and McAlpine C A(2012) Faustian bargains Restoration realities in the context ofbiodiversity offset policies Biological Conservation 155 141ndash148doi101016jbiocon201206003

McCall S C McCarthy M A van der Ree R Harper M J Cesarini Sand Soanes K (2010) Evidence that a highway reduces apparentsurvival rates of squirrel gliders Ecology and Society 15 27

McGregor R A Stokes V L and Craig M D (2014) Does forestrestoration in fragmented landscapes provide habitat for a wide-ranging carnivore Animal Conservation 17 467ndash475 doi101111acv12112

McKenzie N L Burbidge A A Baynes A Brereton R N DickmanC R Gordon G Gibson L A Menkhorst P W Robinson A CWilliams M R and Woinarski J C Z (2007) Analysis of factorsimplicated in the recent decline of Australiarsquos mammal fauna Journal ofBiogeography 34 597ndash611 doi101111j1365-2699200601639x

McKenzie N L van Leeuwen S and Pinder AM (2009) Introduction tothe Pilbara Biodiversity Survey 2002ndash2007 Records of the WesternAustralian Museum 78(Suppl) 3ndash89 doi1018195issn0313-122x78(1)2009003-089

Morris K Johnson B Orell P Gaikhorst G Wane A and MoroD (2003) Recovery of the threatened chuditch (Dasyurus geoffroii)a case study In lsquoPredators with Pouches The Biology of CarnivorousMarsupialsrsquo (Eds M Jones C Dickman and M Archer) pp 435ndash451(CSIRO Publishing Melbourne)

Nelson J L Scroggie M P and Belcher C A (2014) Developing acamera trap survey protocol to detect a rare marsupial carnivore thespotted-tailed quoll (Dasyurus maculatus) In lsquoCamera TrappingWildlife Management and Researchrsquo (Eds P D Meek and PJ Fleming) pp 271ndash279 (CSIRO Publishing Melbourne)

Newsome T M Dellinger J A Pavey C R Ripple W J Shores C RWirsing A J and Dickman C R (2015) The ecological effects ofproviding resource subsidies to predators Global Ecology andBiogeography 24 1ndash11 doi101111geb12236

OrsquoDonnell S Webb J K and Shine R (2010) Conditioned tasteaversion enhances the survival of an endangered predator imperilled bya toxic invader Journal of Applied Ecology 47 558ndash565 doi101111j1365-2664201001802x

Oakwood M (1997) The ecology of the northern quoll Dasyurushallucatus PhD Thesis Australian National University Canberra

Oakwood M (2000) Reproduction and demography of the northern quollDasyurus hallucatus in the lowland savanna of northern AustraliaAustralian Journal of Zoology 48 519ndash539 doi101071ZO00028

Oakwood M (2002) Spatial and social organization of a carnivorousmarsupial Dasyurus hallucatus (Marsupialia Dasyuridae) Journal ofZoology 257 237ndash248 doi101017S0952836902000833

Oakwood M and Foster P (2008) Monitoring extinction of the northernquoll Australian Academy of Science Newsletter 71 6

Palomares F and Caro T M (1999) Interspecific killing amongmammalian carnivores American Naturalist 153 492ndash508 doi101086303189

Pavey C R and Bastin G (2014) lsquoAustralian Rangelands and ClimateChange ndash Invasive Animalsrsquo (Ninti One Limited amp CSIRO AliceSprings)

Pierson J C Barton P S Lane PW and Lindenmayer D B (2015) Canhabitat surrogates predict the response of target species to landscapechange Biological Conservation 184 1ndash10 doi101016jbiocon201412017

Pollock A B (1999) Notes on status distribution and diet of northernquoll Dasyurus hallucatus in the MackayndashBowen area mideastern

Research priorities for the northern quoll Australian Mammalogy M

Queensland Australian Zoologist 31 388ndash395 doi107882AZ1999040

Ramanaidou E R and Morris R C (2010) A synopsis of the channeliron deposits of the Hamersley Province Western AustraliaTransactions of the Institutions of Mining and Metallurgy Section BApplied Earth Science 119 56ndash59 doi101179037174510X12853354810624

Schmitt L H Bradley A J Kemper C M Kitchener D J HumphreysW F and How R A (1989) Ecology and physiology of the northernquoll Dasyurus hallucatus (Marsupialia Dasyuridae) at MitchellPlateau Kimberley Western Australia Journal of Zoology 217539ndash558 doi101111j1469-79981989tb02510x

Short J and Smith A (1994) Mammal decline and recovery in AustraliaJournal of Mammalogy 75 288ndash297 doi1023071382547

Spencer P How R Hillyer M Cook A Morris K Stevenson C andUmbrello L (2013) Genetic analysis of northern quolls from the PilbaraRegion ofWestern Australia Year one ndash final report Unpublished reportDepartment of Sustainability Environment Water Population andCommunities Canberra and Department of Parks and Wildlife Perth

Sutherland W J Fleishman E Mascia M B Pretty J and Rudd M A(2011) Methods for collaboratively identifying research priorities andemerging issues in science and policyMethods in Ecology and Evolution2 238ndash247 doi101111j2041-210X201000083x

Sutherst R W Floyd R B and Maywald G F (1996) The potentialgeographical distribution of the cane toad Bufo marinus L in AustraliaConservation Biology 10 294ndash299 doi101046j1523-1739199610010294x

Taylor B D and Goldingay R L (2003) Cutting the carnage wildlifeusage of road culverts in north-eastern New South Wales WildlifeResearch 30 529ndash537 doi101071WR01062

Tingley R Phillips B L Letnic M Brown G P Shine R and BairdS J (2013) Identifying optimal barriers to halt the invasion of canetoads Rhinella marina in arid Australia Journal of Applied Ecology 50129ndash137 doi1011111365-266412021

Van Vreeswyk A M E Payne A L Leighton K A and HennigP (2004) An Inventory and Condition Survey of the Pilbara RegionWestern Australia Technical Bulletin No 92 Department of AgricultureWestern Australia Perth

Webb J Legge S Tuft K Cremona T and Austin C (2015) Can wemitigate cane toad impacts on northern quolls Final report National

Environmental Research Program Northern Australia Hub CharlesDarwin University Darwin

Wintle B A Walshe T V Parris K M and McCarthy M A (2012)Designing occupancy surveys and interpreting non-detection whenobservations are imperfect Diversity amp Distributions 18 417ndash424doi101111j1472-4642201100874x

Woinarski J Menkhorst K Gambold N and Braithwaite R W (1992)Mammals of the Bungle Bungle area In lsquoA Survey of the Wildlife andVegetation of Purnululu (Bungle Bungle) National Park and AdjacentArearsquo (Ed J C Z Woinarski) Research Bulletin No 6 pp 53ndash67(CSIRO Division of Wildlife and Ecology Darwin)

Woinarski J C Z Risler J and Kean L (2004) Response of vegetationand vertebrate fauna to 23 years of fire exclusion in a tropical Eucalyptusopen forest Northern Territory Australia Austral Ecology 29 156ndash176doi101111j1442-9993200401333x

Woinarski J C Z Oakwood M Winter J Burnett S Milne D FosterP Myles H and Holmes B (2008) Surviving the toads patterns ofpersistence of the northern quoll Dasyurus hallucatus in QueenslandReport submitted to the Natural Heritage Trust Strategic ReserveProgram as a component of project 2005162 Monitoring andManagement of Cane Toad Impact in theNorthern Territory Available athttplrmntgovauplants-and-animalsinformation-and-publicationspublications2008 [accessed 14 November 2013]

Woinarski J C Z Armstrong M Brennan K Fisher A Griffiths A DHill B Milne D J Palmer C Ward S Watson M Winderlich Sand Young S (2010) Monitoring indicates rapid and severe decline ofnative small mammals in Kakadu National Park northern AustraliaWildlife Research 37 116ndash126 doi101071WR09125

Woinarski J Burbidge A and Harrison P (2014) lsquoAction Plan forAustralian Mammals 2012rsquo (CSIRO Publishing Melbourne)

Woinarski J C Burbidge A A and Harrison P L (2015) Ongoingunraveling of a continental fauna decline and extinction of Australianmammals since European settlement Proceedings of the NationalAcademy of Sciences of the United States of America 112 4531ndash4540doi101073pnas1417301112

Woolley P A Krajewski C and Westerman M (2015) Phylogeneticrelationships within Dasyurus (Dasyuromorphia Dasyuridae) quollsystematics based on molecular evidence and male characteristicsJournal of Mammalogy 96 37ndash46 doi101093jmammalgyu028

N Australian Mammalogy V A Cramer et al

wwwpublishcsiroaujournalsam

Page 2: Research priorities for the northern quoll (Dasyurus ......The Pilbara population of the northern quoll (Dasyurus hallucatus) has been seldom studied, and the impacts of threats such

Kimberley is estimated to be ~12 000 000 ha but the area ofoccupancy and the number of mature individuals within thetwo regions are not known (Parks and Wildlife unpubl data)Approximately 2500 records for the northern quoll (many ofwhich are highly clustered) are held in the NatureMap database(httpnaturemapdpawwagovau) these include records fromthe Western Australian Museum and Parks and Wildlifeand fauna returns from surveys undertaken by private groups(mostly environmental consultants) These location recordsindicate that quolls are more prevalent in complex rocky areas inthe north central and west Pilbara and are less likely to occurthrough the south and east areas of the Hamersley Rangesand Fortescue Marsh (Fig 1) The Pilbara population isgenetically distinct from the Kimberley and Northern Territorypopulations (Woolley et al 2015) and is the least diversewith ~77 heterozygosity (103 alleles) compared with84 heterozygosity (111 alleles) in the Kimberley mainlandpopulation (How et al 2009 Spencer et al 2013)

Habitat change due to altered fire regimes weed invasionand grazing predation by cats foxes and dingoeswild dogsthe future invasion of cane toads and the interactive effectsbetween these as well as habitat fragmentation and loss bymining and infrastructure development have been identified

as key threats for the northern quoll in the region (Woinarskiet al 2014) While the potential invasion of the cane toad isundoubtedly the most significant future threat to the northernquoll in the Pilbara there is little knowledge of what has been therelative impact or the interactive effects of each of the otherkey threats on the northern quoll in the region or how these willaffect northern quoll populations into the future

The areas of highest habitat quality for the northern quollare considered to be rugged rocky areas often in closeassociation with permanent water (Begg 1981 Schmitt et al1989 Braithwaite and Griffiths 1994 Oakwood 1997 Pollock1999) These rocky landforms provide a diversity ofenvironments denning opportunities protection from weatherand diurnal predators and immediate refuge from fire(Braithwaite and Griffiths 1994 Oakwood 2000 Cook 2010)In the Pilbara the ridges and mesas of channel-iron depositsand banded iron formation ranges are often the primary focusof iron-ore extraction (Ramanaidou and Morris 2010) whileexposed granite outcrops are quarried for road and rail beds Thenorthern quoll is therefore a key consideration in most miningproject assessments under the EPBC Act 1999 in the Pilbara aswell as two strategic environmental assessments examining thecumulative impacts of mining in the region (BHP Billiton Iron

116degdeg 117deg 118deg 119deg 120deg 121deg

116deg115deg

ndash23deg

ndash22deg

ndash21deg

ndash20deg

ndash23deg

ndash22deg

ndash21deg

ndash20deg

117deg 118deg 119deg 120deg 121deg

Fig 1 Records (black dots) of the northern quoll in the Pilbara obtained from the Department of Parks and WildlifeNatureMap database (httpnaturemapdpawwagovau) The shaded areas indicate the IBRA subregions and the hatched areasthe conservation estate within the region Much of the recent survey effort has occurred in areas with mining activity or mining-related infrastructure eg the rail corridors running south of Port Hedland

B Australian Mammalogy V A Cramer et al

Ore httpwwwenvironmentgovaunode18597 HamersleyIron Pty Ltd (Rio Tinto) httpwwwenvironmentgovaunode18608)

Biodiversity offsets are increasingly used as a policy toolto permit development but ensure that any residual adverseimpacts of the development on the environment arecompensated (Bull et al 2013) In best practice offsets areapplied as part of a mitigation hierarchy where all reasonableefforts to avoid or minimise impacts upon the environment arefirst considered (Kiesecker et al 2010) Yet the limited data onthe distribution and status of the northern quoll in the Pilbarathe poor understanding of critical habitat in relation to futuremining and infrastructure development as well as uncertaintyaround the level of impact from other threats has meant thatuncertainty around how to create direct ecologically equivalentoffsets is high Under the Australian Governmentrsquos offsetpolicy (Department of Sustainability Environment WaterPopulation and Communities 2012) compensatory offsetssuch as directing a proportion of offset funds towards scientificresearch are considered as an acceptable component of anoffsets package in these situations This has been the case forthe northern quoll in the Pilbara with approximately AU$175Mof offset funds committed to research on the species over thenext decade (although not all of this amount is currently active)

The dramatic increase in fauna surveys associated withenvironmental impact assessments in the Pilbara has generateda valuable dataset on the northern quoll and other threatenedspecies (Carwardine et al 2014) Collation of these datacould greatly improve our knowledge of speciesrsquo ranges andcritical habitats yet most of these data are not within thepublic domain (Carwardine et al 2014) Further it has beenargued that much conservation research is having little directinfluence on the achievement of on-ground conservationoutcomes in part because the conceptual novelty valued bynational funding agencies may not align with the moreapplied needs of practitioners (Knight et al 2008 Lauranceet al 2012) A potential solution to overcoming the lsquoknowledgendashimplementationrsquo or lsquoknowingndashdoingrsquo gap is to encouragecollaboration between scientists policy makers and practitionerswhen research projects are developed (Knight et al 2008Sutherland et al 2011 Laurance et al 2012) In an effort toencourage data-sharing and to foster a sense of ownership overthe research agenda that will be funded by offset provisionsParks and Wildlife instigated a collaborative process betweenscientists practitioners and mining proponents to identifyand rank research priorities for the northern quoll in the PilbaraIn this paper we present the research priorities and the keyquestions associated with each priority as identified byworkshop participants This is the first step in developing aresearch program to increase our knowledge of the northernquoll in the Pilbara and to better understand the impacts ofcurrent and future threats so that a viable population ismaintained into the future using evidence-based management

Methods

Parks and Wildlife hosted a workshop attended by researchscientists employed in the university or government sector (13people) research scientists and environmental consultants

employed in the private sector (30 people) mining industryrepresentatives (22 people) and representatives of WesternAustralian and Australian government departments responsiblefor environmental regulation and approvals (9 people) all ofwhom have knowledge of the northern quoll mining andassociated activities andor the Pilbara region The workshopwas facilitated by an independent professional facilitator Theprocess to define research priorities was as follows (1) After aseries of presentations by experts the group as a whole discussedthe key threats to the northern quoll in the Pilbara This includedboth current and future threats (eg climate change cane toadinvasion) and both well- and lesser-understood threats (eg theimpact of disease) (2) Participants were then allocated to one ofeight breakout groups so that each group contained membersfrom the range of affiliations present Each group then identifieda series of research priorities that they considered would bestaddress the current gaps in knowledge that are the main barriersto the effective management and conservation of the northernquoll in the Pilbara both in terms of better understanding theirdistribution and ecology and in mitigating the key threats Theresearch priorities from each group were then presented to allparticipants and collated by the facilitator (3) Each group thenranked the sequence in which the research priorities should beaddressed based on which research needs they considered weremost pressing (4) The rankings were then compared and collatedacross groups to define the final ranking of research priorities Inmost cases there was agreement across the groups as to thesequence in which the research priorities should be addressed(5) Finally new breakout groups were formed based onexpertise and interest to discuss and identify the key researchquestions that needed to be addressed under each researchpriority

Research priorities

Research priorities are presented below in the order in whichthey should be addressed based on the ranking of the workshopparticipants Key research questions aligned with each researchpriority are presented in Table 1 We also provide a brief reviewof the current evidence that will further guide the development ofa research program to answer these questions

Priority 1 Develop appropriate and standardised surveyand monitoring methods

Developing a standard survey and monitoring protocol for thenorthern quoll in the Pilbara based on detection and occupancyprobabilities (Wintle et al 2012) was identified as the highestresearch priority by the group A review of over 200 surveyreports most of which were related to mining activities foundthat the sampling effort and methods varied widely betweensurveys and were often not adequate to detect northern quolls(A Cook unpubl data) Inadequate survey effort andor usingan inappropriate survey method introduce uncertainty as towhether the failure to detect the northern quoll within an areareflects a true absence of the species (Wintle et al 2012) Thedata gathered from surveys that vary widely in terms of effortand method may have limited comparability and incurs a lostopportunity to gather regional metadata for the northern quollin an area where any survey effort represents a significant

Research priorities for the northern quoll Australian Mammalogy C

Tab

le1

Researchpriorities

fortheno

rthern

quollan

dkeyqu

estion

sassociated

witheach

priorityan

das

compa

redwiththeinform

ationrequ

ired

undersimila

rthem

esan

dthepriority

givento

thisin

Woina

rski

etal(201

4)

Rank

Theme

Researchpriority

Key

questio

nsInform

ationrequired

andpriority

givenundersimilarthem

ein

Woinarski

etal(2014)

1Surveyand

mon

itoring

Develop

appropriate

andstandardised

survey

and

mon

itoring

metho

ds

Whatistheefficacy

ofthemajor

survey

metho

dscurrently

used

forno

rthern

quollsin

the

Pilb

ara

How

dovariou

scombinatio

nsoftrap

typesarraysdurationoftrappingbaitsetc

compareWhich

camera-trap

protocol

ismosteffectivefor(a)identifying

individu

alno

rthern

quollsin

thePilb

araand(b)estim

atingpo

pulatio

nsizes

Which

methodismost

cost-effectiv

eandefficientin

variou

sPilb

araland

scapes

Whatisthedetectionprobability

ofno

rthern

quollsin

each

habitattypeBased

onprior

surveywhatitthepriorprobability

ofoccupancyin

thesehabitattypes

Based

onthis

know

ledge

how

manynon-detections

ofquollsarerequired

before

wecanstatewith

prespecified

confi

dencethat

northern

quollsareabsent

from

asite

Surveyto

betterdefine

fine-scale

distributio

nandnumberof

individu

alsin

subpop

ulations

mediumndashhigh

Establishor

enhancemon

itoring

program

bydesigningintegrated

mon

itoring

prog

ramsmedium

2Habitat

requirem

ents

Define

areasof

critical

habitatforno

rthern

quollin

thePilb

ara

How

donorthern

quollsusehabitatinthePilb

ara

How

arequollpopulations

distributedin

relatio

nto

habitatq

uality(egh

ighqu

ality

low

quality

orsink

habitats)

How

ishabitat

selectioninfluenced

bypredationrisk

andsocial

factors

Whatinformationisrequ

ired

tofurtherrefine

anSDM

forno

rthern

quollsin

thePilb

ara

How

canthisinform

ationbe

mosteffectivelyused

toim

provemonito

ring

effortand

outcom

es

How

domod

elsvary

across

thePilb

aramainlandandoffsho

reislandsaccountin

gfor

clim

atic

gradientsandtopography

Whatarefuture

predictio

nsfornorthern

quollpopulatio

ndistributio

nsu

nder

modelsof

clim

atechange

andcane

toad

invasion

Assesshabitatrequirem

entsna

Und

erstandho

wdisturbanceaffects

habitatqu

ality

and

conn

ectiv

ity

How

does

fire

andgrazingaffecthabitatq

ualityandhabitatu

seby

northern

quollsin

the

Pilb

ara

How

importantisfire

asabarrierto

dispersal

How

importantishabitatconnectivity

form

etapopulations

ofnorthernquollinthePilb

ara

How

does

habitattypeaffecttheconnectiv

ityof

metapopulationsHow

does

disturbance

both

discrete

(eginfrastructure

such

asrailandroads)

anddiffuse(egchangesin

vegetatio

ncompositio

nandstructure)alterhabitatconnectiv

ity

Assessrelativ

eim

pact

ofthreats

quantifyrelativ

eim

pact

offire

regimesmdashhigh

Assesseffectivenessof

threat

mitigatio

nop

tionsland

scape-scale

optio

nsforretentionof

longer-

unburntpatchesmdash

high

3

Pop

ulation

dynamics

Betterunderstand

the

popu

latio

ndy

namicsof

the

northern

quollinthe

Pilb

ara

How

manysubpop

ulations

aretherein

thePilb

ara

How

dotheseinteractAre

subpopulations

demographically

stablein

spaceandtim

eHow

does

habitatand

clim

ate

affect

popu

latio

ndy

namics

How

does

disturbanceaffect

demog

raphic

stability

Assessdietlifehistory

na

Investigatepo

pulatio

nstructureandthe

interactionbetween

popu

latio

ns

How

domeasuresofgenetic

distance

relatetospatialecology

asmeasuredby

otherm

eans

iedoes

inferred

genetic

connectiv

ityrelate

toobserved

dispersal

Can

genetic

inform

ationbe

used

todeterm

inehabitatbarriers

Resolve

taxonomic

uncertainties

assess

genetic

relatednessof

spatially

disparatepo

pulatio

nsto

identifywhether

thereare

genetically

distinct

subpop

ulationsmdashlowndashmedium

D Australian Mammalogy V A Cramer et al

4Key

threats

Introduced

predators

Doespredationsignificantly

affecttheno

rthern

quollinthePilb

ara

How

does

thisvary

across

habitattypes

Doescompetitionforresourceswith

foxesor

catsim

pact

onsubpopulationsWhich

lifestages

aremostvulnerable

Given

thattheno

rthern

quollisalso

amesop

redatordo

esthepresence

ofding

oeseither

directly(through

predation)

orindirectly(through

theregulatio

nof

foxesandcats)impact

onsubpop

ulations

inthePilb

ara

Assessrelativ

eim

pact

ofthreats

predationby

feralcatsanddogsmdash

high

interactions

betweenferal

catsdo

gsdingoes

andfoxesmdash

mediumndashhigh

Assesseffectivenessof

threat

mitigatio

nop

tionsassess

which

feasible

managem

entoptio

nsfor

feralcatsanddo

gscanlead

topo

pulatio

nrecoverymdashmediumndash

high

Develop

new

orenhanceexistin

gmanagem

entmechanism

sfor

broad-scale

targetedferalcat

controlmdash

medium

Canetoads

Will

theclosureof

artifi

cialwater

bodies

preventor

simplyslow

therateofinv

asionof

cane

toadsinto

thePilb

ara

Whatisthedegree

ofoverlapbetweenSDMfundamentalniche

modelsforn

orthernquolls

andcane

toadsin

thePilb

ara

Can

potentialrefug

eareasforno

rthern

quollb

eidentifi

ed

IsCTAlik

elytobe

effectivewhentaste-aversion

baits

aredeliv

ered

underfi

eldcond

itions

Atwhatscaleisbaitdeliv

eryfeasible

Assesseffectivenessof

threat

mitigatio

nop

tionsassess

feasibility

ofbroad-scaletaste-

aversion

lsquotrainingrsquo

ofqu

ollsmdash

mediumassess

local-scale

exclusionof

toadsor

preventio

nof

colonisatio

n(m

ainlandsites)mdash

medium

Infrastructure

developm

entand

interactions

with

humans

Are

road-related

deaths

ofno

rthern

quollasign

ificant

causeof

mortalityin

thePilb

ara

Dorailw

aylin

esinfrastructurecorridorsor

wideroadspresenta

barrierto

northern

quoll

mov

ement

Whatmethods

canbe

used

tomitigate

theim

pactsof

infrastructure

developm

entand

associated

machinery

operation

Not

addressed

Interactions

between

threats

Dofrequent

andintensefireslead

toreducedstructural

complexity

loss

ofvegetatio

ncoverandsubsequentincreasedpredationof

northern

quollsparticularlyof

males

during

andor

afterbreeding

Whatarethecumulativeim

pactsof

mortalitycaused

bycane

toad

ingestionandpredation

bycatsdo

gsdingoes

andfoxes

See

inform

ationon

introd

uced

predatorsandcane

toadsabove

5Recolon

isationof

restored

orartifi

cial

habitat

Determinetheability

oftheno

rthern

quoll

torecolonise

disturbedareasor

colonise

artifi

cial

habitat

Whatarethekeyelem

entsof

northern

quollhabitatthatneedtobe

restoredcreated

sothat

habitatcomplexity

andprod

uctiv

ityismaxim

ised

Whatlevelo

flandscapeconn

ectiv

ityisrequired

arou

ndrestoredartificialhabitatso

that

genetic

connectiv

itybetweensubpopulations

canbe

maintained

How

long

should

restored

orartifi

cial

habitatbe

directly

managedparticularly

for

predators

How

cantheretentionof

largebouldersof

wasterock

beincorporated

into

rehabilitation

plans

Not

addressed

Research priorities for the northern quoll Australian Mammalogy E

investment of time and resources The limited amount anduncertain quality of occurrence data has been a major restrictionon the utility of a species distribution model (SDM) and apopulation viability analysis (PVA) for the northern quoll in thePilbara (Cadenhead et al 2015 see below) While survey andmonitoring per se is not a research activity that directly informsmanagement or threat mitigation carefully implementedmonitoring programs are essential to identify many elements ofa speciesrsquo ecology to detect changes in population and toquantify the ecological effectiveness and cost-effectivenessof management interventions (Lindenmayer et al 2013Lindenmayer 2015)

Parks and Wildlife recently released revised protocols forthe long-term monitoring of known populations of thenorthern quoll (Dunlop et al 2014b) and the effectiveness ofthese protocols in presenceabsence surveys should be testedThe cost-effectiveness of using camera traps for wildlifesurveys in remote locations such as the Pilbara makes their useappealing Hohnen et al (2013) described how the strategicplacement of cameras and baits combined with taking a greaternumber of photographs per trigger allows the capture ofimages that can be used to identify individual northern quollsfrom their spot patterns Remote cameras have also been shownto be a viable method for estimating population sizes of thenorthern quoll in areas with both dense (Austin 2014) andsmall populations (Webb et al 2015) Further testing of a rangeof parameters (eg see Nelson et al 2014) to develop a robuststandard protocol for the Pilbara is required along with acomparison of the cost-effectiveness and any ethicalreducedhandling benefits of using remote cameras versus live-trappingfor estimating population sizes in Pilbara landscapes

Priority 2 Improve our understanding of habitatrequirements

Define areas of critical habitat for the northern quollin the Pilbara

The Department of Agriculture and Food Western Australiahas mapped 102 land systems across the Pilbara bioregionbased on several biophysical components including landformgeology soils vegetation and drainage patterns (Van Vreeswyket al 2004) A review of records for the northern quoll in thePilbara showed that 65 of records were from the RobeCapricorn and Boolaloo land systems (Table 2) which compriselow plateaux mesas hills and ridges of varying geology(limonite granite and sandstone respectively) Most recentsurvey effort in the Pilbara is biased towards mineral-bearingformations or areas of mining-related infrastructure (eg railand road corridors) (Fig 1) Further investigation of theserecords is required to assess whether these land systems areindeed preferred habitat for northern quolls as opposed to anartefact of survey effort The comparatively fewer records thatcoincide with the conservation estate in the region (Fig 1) arealso likely an artefact of survey effort

The first step in defining areas of critical habitat for thenorthern quoll in the Pilbara is to improve our understanding ofhabitat use particularly in terms of defining habitat suitability atfiner scales than is currently available As part of the strategic

environmental assessments being undertaken for the Pilbarabioregion an SDM has been developed for the northern quoll(Cadenhead et al 2015) Uncertainty about the quality of theoccurrence data available the limited amount of data availableas well as the bias towards data collection in areas proposed fordevelopment constrained the level of certainty around theoutcomes of the model Improved model certainty requiresfurther collection of contemporary species-occurrence data andthe development of finer-resolution GIS layers particularly onethat captures finer-scale geological attributes such as mesas orrocky outcrops considered as highest-quality habitat for thenorthern quoll (Cadenhead et al 2015)

Better definition of the habitat attributes favoured by thenorthern quoll will allow further refinement of the GIS variablesincluded in the SDM or allow the development of predictivesite-based habitat models Information on how northern quollsuse habitat can be gained through direct observation of howthey interact with habitat features or from the association ofhabitat features with the presence of quolls as determined byGPS receivers (Gaillard et al 2010) This level of detail iscurrently not available from survey records returned as part ofenvironmental impact assessments but collection of habitatdata using the protocols outlined by Dunlop et al (2014b) infuture surveys would rapidly increase the quality of dataavailable

Understand how disturbance affects habitat qualityand connectivity

The alteration of fire regimes has been the most pervasivelandscape-scale change in the Pilbara Between 1993 and 2006over 72 of the region was burnt with more than 28 of thearea burnt two or more times (McKenzie et al 2009) Woinarskiet al (2004) found that northern quolls may be able to tolerate amoderately high frequency of fire as long as these fires werelow-intensity early-season burns and the impacts of fire onvegetation structure and composition were not exacerbatedby the grazing of introduced herbivores In the Pilbarahowever as with many other grazing regions of Australia earlypastoralists often overestimated the productivity and resilienceof native vegetation to sheep and cattle grazing which led tosevere degradation of vegetation particularly along riverfrontages (Hennig 2004) The soil degradation and loss ofvegetative cover caused by intense grazing pressure in thesouth Kimberley is thought likely to have contributed to thesubstantial decline or disappearance of medium-sized animalsincluding the northern quoll from the Bungle Bungle area(Woinarski et al 1992) Death by predation is most likely in openareas (Oakwood 2000) and therefore the effects of fire andgrazing on northern quoll populations cannot be consideredwithout also taking account of likely increases in mortality due topredation (see lsquoIntroduced predatorsrsquo below)

Understanding how the localised but intensive activityassociated with mining affects habitat quality and habitat useby the northern quoll in areas surrounding development islimited A recent study that examined the use of ballast quarriesand paired undisturbed sites by northern quolls found that theywere foraging denning and showing signs of breeding at thequarry sites where greater numbers of quolls were captured(Dunlop et al 2014a) This was likely due to the combination of

F Australian Mammalogy V A Cramer et al

habitat complexity (eg diversity of boulder sizes) and thepresence of permanent water in the quarry which may haveincreased prey availability at these sites

Movement of animals through the landscape for dispersaland repopulation is another important consideration for a speciesin which both sexes tend towards semelparity and is thereforepredisposed to local extinction (Oakwood 1997) Improvedunderstanding of how the northern quoll (particularly dispersingmales) uses areas of lower habitat quality and the identificationof such habitat in habitat-suitability models will allow futurelandscape-scale strategic assessments to better predict thecombined impacts of disturbance on the ability of northern quolls

to move through the landscape Analysis of regional populationgenetics will also improve our understanding of the level ofdispersal occurring across the landscape

Priority 3 Population dynamics

Better understand the population dynamics of thenorthern quoll in the Pilbara

Little information is available on the number ofsubpopulations of northern quoll in the Pilbara how thesesubpopulations interact how stable they are in space and timeand how they respond to disturbance As no demographic

Table 2 Records of the northern quoll in each land system type in the PilbaraRecords were current as of 23 May 2014

Land system Area (km2) No of records Land system description

Augustus 61 1 Rugged ranges hills ridges and plateaux supporting mulga shrublands and hard spinifex grasslandsBoolaloo 1502 245 Granite hills domes and tor fields and sandy plains with shrubby spinifex grasslandsBoolgeeda 7748 29 Stony lower slopes and plains below hill systems supporting hard and soft spinifex grasslands andmulga

shrublandsCalcrete 1444 2 Low calcrete platforms and plains supporting shrubby hard spinifex grasslandsCallawa 1003 9 Highly dissected low hills mesas and gravelly plains of sandstone and conglomerate supporting soft and

hard spinifex grasslandsCapricorn 5296 257 Hills and ridges of sandstone and dolomite shrubby hard and soft spinifex grasslandsCoongimah 3244 12 Plateau surfaces low hills with steep slopes and undulating uplands supporting hard spinifex grasslandsDune 138 1 Dune fields supporting soft spinifex grasslandsGranitic 4020 1 Rugged granitic hills supporting shrubby hard and soft spinifex grasslandsHorseflat 1261 4 Gilgaied clay plains supporting tussock grasslands and minor grassy snakewood shrublandsLittoral 1577 3 Bare coastal mudflats with mangroves on seaward fringes samphire flats sandy islands coastal dunes

and beachesMacroy 13 095 94 Stony plains and occasional tor fields based on granite supporting hard and soft spinifex grasslandsMcKay 4202 21 Hills ridges plateau remnants and breakaways of metasedimentary rocks supporting hard spinifex

grasslandsMallina 2557 8 Sandy surfaced alluvial plains supporting soft spinifex (and occasionally hard spinifex) grasslandsMarandoo 459 2 Basalt hills and restricted stony plains supporting grassy mulga shrublandsMosquito 1840 3 Stony plains and prominent ridges of schist and other metamorphic rocks supporting hard spinifex

grasslandsNewman 14 580 81 Rugged jaspilite plateaux ridges and mountains supporting hard spinifex grasslandsOnslow 424 1 Sandplains dunes and clay plains supporting soft spinifex grasslands and minor tussock grasslandsParaburdoo 565 3 Basalt-derived stony gilgai plains and stony plains supporting snakewood and mulga shrublands with

spinifex and tussock grasslandsParadise 1479 8 Alluvial plains supporting soft spinifex grasslands and tussock grasslandsRiver 4088 37 Active flood plains major rivers supporting grassy eucalypt woodlands tussock grasslands and soft

spinifex grasslandsRobe 865 369 Low limonite mesas and buttes supporting soft spinifex (and occasionally hard spinifex) grasslandsRocklea 22 993 40 Basalt hills plateaux lower slopes and minor stony plains supporting hard spinifex (and occasionally

soft spinifex) grasslandsRuth 346 36 Hills and ridges of volcanic and other rocks supporting hard spinifex (occasionally soft spinifex)

grasslandsStuart 1794 3 Gently undulating stony plains supporting hard and soft spinifex grassland and snakewood shrublandsTaylor 129 3 Stony plains and isolated low hills of sedimentary rocks supporting hard and soft spinifex grasslandsTalga 2124 9 Hills and ridges of greenstone and chert and stony plains supporting hard and soft spinifex grasslandsTuree 581 2 Stony alluvial plains with gilgaied and non-gilgaied surfaces supporting tussock grasslands and grassy

shrublandsUaroo 7681 37 Broad sandy plains supporting shrubby hard and soft spinifex grasslandsUrandy 1311 6 Stony plains alluvial plains and drainage lines supporting shrubby soft spinifex grasslandsWhite Springs 266 4 Stony gilgai plains supporting tussock grasslands and hard spinifex grasslandsWona 1815 10 Basalt upland gilgai plains supporting tussock grasslands and minor hard spinifex grasslandsYamerina 1207 1 Floodplains and deltaic deposits supporting tussock grasslands grassy woodlands and minor halophytic

low shrublands

Research priorities for the northern quoll Australian Mammalogy G

studies have been undertaken in the region Cadenhead et al(2015) used demographic information from the NorthernTerritory and Kimberley populations in their PVA Uncertaintyabout data quality and the limited information available ondemographic parameters for the Pilbara constrained the level ofcertainty around the outcomes of the PVA The model doeshowever provide a framework into which new data can beincorporated as they become available For example preliminarydata from the Pilbara monitoring program suggest that theproportion of male die-off in the Pilbara population is lowerthan in the Kimberley and Northern Territory populations(Dunlop unpubl data) Cadenhead et al (2015) stress the needfor more demographic and occurrence data to be collectedbefore further development decisions are made (see lsquoDevelopappropriate and standardised survey and monitoring methodsrsquoand lsquoImprove our understanding of habitat requirementsrsquo above)

Investigate population structure and the interactionbetween populations

Analysis of the genetic diversity and population structure ofthe Pilbara population of the northern quoll is ongoingOnly small differences in genetic diversity as measured byheterozygosity were found between samples obtained at 16locations within the Pilbara with the exception of the isolatedpopulation on Dolphin Island (Spencer et al 2013) This resultindicates a high level of movement across the landscape byindividuals during mating and dispersal phases Spatial structureanalysis of samples collected from around Pannawonicasuggested a lsquoneighbour sizersquo of ~15 km (Spencer et al 2013)Current data show no evidence of genetic bottleneckssuggesting that if there has been a population decline in thePilbara that this has not affected genetic diversity A usefulapproach would be to combine genetic information with datafrom radio-tracking or GPS collars to better understand theuse of lsquolandscapersquo space and how this influences populationdynamics

Priority 4 Key threats and the interaction of these threats

Introduced predators

Introduced cats and foxes are considered to be one of themain causes of the decline and extinction of the critical-weight-range mammals (35 g to 55 kg) in Australia (Short and Smith1994 McKenzie et al 2007 Woinarski et al 2015) but theirimpacts on the northern quoll are less clear A recent review offeral cat diets across Australia found that northern quolls werepresent in the diet of feral cats (Doherty et al 2015a) andWoinarski et al (2014) consider the threat posed by cats to thenorthern quoll to be severe although the impacts of cats may bereduced in rugged refuge areas The impacts of cats on othermuch larger species of quoll appear to be inconsistent withsome evidence of coexistence between cats and quolls beforethe arrival of foxes (Glen and Dickman 2008 and referencescontained within) Due to the limited overlap in their distributionthroughout Australia the threat posed by foxes to northernquolls is less understood than for cats and Woinarski et al(2014) consider the threat posed by foxes to be moderate on anational level Foxes are common in coastal near-coastal and

riparian areas of the Pilbara and extend as far north as Broome(King and Smith 1985 Kendrick and Stanley 2002) They arerenowned for their aggression towards smaller carnivores(Palomares and Caro 1999) and neither the larger chuditch(Dasyurus geoffroii) nor spotted-tail quoll (Dasyurus maculatus)are known to coexist with large numbers of foxes (A Glen perscomm) The interactions between quolls cats and foxesare potentially more complex than for other medium-sizedmarsupials due to niche overlap in diet and competition for densites with the larger spotted-tailed quoll and chuditch bothexperiencing exploitation and interference competition fromfoxes (Morris et al 2003 Glen et al 2011) Competitionbetween feral cats and the northern quoll may potentially beintense during periods of low rainfall in the Pilbara when preyavailability is low Understanding whether foxes and cats aresignificant predators of or competitors with the northern quollin the Pilbara and how this affects their population viabilitywill clarify the level of risk posed by these predators Thesausage-style baits used for feral cats (eg Eradicatreg andCuriosityreg) are potentially attractive to the northern quoll withthe risk of mortality or reduced fitness Trials to assess themagnitude of the risks from Eradicatreg baits are currentlyunderway in the Pilbara

The role of dingoes and wild dogs in the mortality of andinteractions between mesopredators (both native and introduced)requires further investigation Dingoes are known to kill northernquolls butmay not eat these kills (Oakwood 1997) A recent studyin the Northern Territory found that camp dogs and dingoes werethe major source of mortality for northern quolls at the study site(Webb et al 2015) and roaming domestic dogs were identified asa cause of mortality for northern quolls in Queensland (Pollock1999) Human resource subsidies (eg stock carrion in rangelandareas) can artificially support predator populations thus indirectlyincreasing predation pressure on prey (Newsome et al 2015)A dynamic debate currently exists around the role of the dingo inregulating populations of feral cats and foxes and the implicationsof this for the conservation of the critical-weight-range mammalsin Australia (see Johnson et al 2007 Letnic et al 2009 Allenet al 2011 2013 Kennedy et al 2012) Understanding thepopulation density at which the role of the dingo as a regulator ofintroduced predators outweighs its own impact as a direct predatorof the northern quoll will inform whether dingowild dog controlwould benefit the northern quoll

Cane toads

The introduced cane toad now occupies more than half thehistoric range of the northern quoll and has caused an immediatecollapse or extirpation of northern quoll populations in theareas it has invaded (Burnett 1997 Oakwood and Foster2008 Woinarski et al 2008 2010) Earlier modelling work thatconsidered only climate parameters as well as more recentwork that incorporates future climate change suggest thatcane toads will not invade the Pilbara (Sutherst et al 1996Pavey and Bastin 2014) Other recent work that considers thefundamental niche of the cane toad and incorporates the locationof artificial watering points predicts that they will colonisethe Pilbara from the Kimberley via the coastal edge of theGreat Sandy Desert (Kearney et al 2008 Florance et al 2011

H Australian Mammalogy V A Cramer et al

Tingley et al 2013) This has created some level of uncertaintyabout the likely impact of cane toads on northern quolls in thePilbara Strategies for cane toad management were not rankedhighly by participants in the threat-management frameworkdeveloped by Carwardine et al (2014) which the authorsexplained as possibly due to uncertainty about how far canetoads will spread into the region

The model developed by Tingley et al (2013) suggested thatthe closure of 100 artificial water bodies at one of three potentiallocations along the coastal corridor between the Kimberley andthe Pilbara would prevent the invasion of cane toads into thePilbara While their model did account for uncharacteristicallywet years every three decades the dispersal of toads betweenwater bodies was determined by the number of rainy daysoccurring during the year (ie toads could only disperse betweenwater bodies for three days following rainfall) rather thanrainfall amount the associated likelihood of flooding and theavailability of free water across the landscape The cyclonic andthunderstorm nature of much of the rainfall in the north-west ofWestern Australia means that the region is prone to significantand sometimes extended flooding In April 1934 the countryaround Anna Plains (250 km south of Broome) was flooded forhundreds of miles when 27 inches (686mm) of rain fell infour days (Anon 1934) In March 2004 over 350mm of rain wasrecorded within a 48-h period at Telfer on the southern edge ofthe Great Sandy Desert due to Tropical Cyclone Fay Thiscaused the flooding of the De Grey and Oakover river systemsand flooded 140 km of the Telfer Access Road with floodwaterstaking several months to subside (wwwbomgovau) Extendingthe model of Tingley et al (2013) to account for such events willhelp clarify whether closing artificial water bodies is likely toprevent the invasion of cane toads into the Pilbara or may onlyassist in delaying their inevitable invasion

Comparison of SDMs (eg Cadenhead et al 2015) fornorthern quoll with fundamental niche models for cane toads(eg see Kearney et al 2008) within the Pilbara may identifypotential refuge areas for the northern quoll and these areasshould be prioritised for conservation actions Because theinvasion point of cane toads into the Pilbara is likely to be narrow(beginning around the mouth of the De Grey River Tingleyet al 2013) a proactive approach to training northern quollsto avoid cane toads may prove beneficial Conditioned taste-aversion (CTA) has been used successfully to train captive-reared northern quolls to avoid cane toads by offering themsmall dead toads infused with thiabendazole before theirrelease into the wild (OrsquoDonnell et al 2010)Monitoring of thesequolls after their release showed that some of their offspringhad learnt to avoid eating cane toads either through ingestionof small non-lethal toads or via social learning from theirmothers (Webb et al 2015) Fields trials of toad-aversion baits(either sausages or meatballs made from toad flesh andcontaining thiabendazole) are currently underway in the centralKimberley where cane toads were expected to invade in late2015 (Webb et al 2015) Further testing of CTA approachesunder a variety of field conditions is required The developmentof stable long-lasting baits suitable for aerial deployment isconsidered a pressing need as well as a captive colony ofnorthern quolls upon which to test the efficacy of toad-aversionbaits for inducing aversion to live toads (Webb et al 2015)

Infrastructure development and interactionswith humans

Infrastructure development may increase the likelihood ofnorthern quoll deaths due to motor vehicle strikes or beingcrushed in machinery Significant chuditch mortality wasobserved along mine access roads in the south-west of WesternAustralia (McGregor et al 2014) and the widening and sealingof an access road dramatically increased the number of roadkills and led to the extirpation of the resident population ofeastern quolls (D viverrinus) in Tasmania (Jones 2000)Infrastructure corridors such as high-tension power-lineeasements and ore-haulage railways are increasingly beinginstalled throughout the range of the northern quoll Althoughthese do not create the same mortality risk as roads they bisectand fragment habitat create large open spaces and potentiallyincrease predation risk

A recent small-scale radio-tracking study (see Dunlop et al2014a) around several quarries in the Pilbara found that northernquolls crossed roads but not railway lines The short-term natureof this work and the limited number of quolls able to be trackedmeans that the influence of roads railway lines and infrastructurecorridors on the movement and mortality of northern quollsrequires further assessment Using radio-tracking techniques todetermine crossing rates and road kill events may be difficultover large areas so population studies comparing groups ofquolls near major roads with those not affected by infrastructure(eg McCall et al 2010) may provide a more feasible meansof assessing the impacts of roads If vehicle strikes are foundto significantly contribute to the mortality of a local populationof the northern quoll then mitigation methods should betrialled on sections of road identified as road-kill hotspotsCulverts and underpasses have been shown to be used by avariety of small and medium-sized mammals and frogs (Taylorand Goldingay 2003) while installing rumble strips on roadshas led to decreased rates of road-kill of nocturnal animals(Lester 2015)

Interactions between threats

At present wildfire the subsequent loss of vegetation coverand increased predation of northern quolls is likely to be themost significant of the interactions between threats The directeffects of wildfire have been found to be less significant inthe subsequent decline of small to medium-sized mammalsparticularly for denning species such as the northern quoll thanthe indirect consequences of reduced resource availability andthe increased risk of predation due to lack of cover (Oakwood2000 Letnic et al 2005 McKenzie et al 2007 Legge et al2008 Doherty et al 2015b) Male northern quolls are perhapsmost vulnerable to such losses of cover during the breedingseason when their range can extend to over 100 ha (Oakwood2002) or where males survive after breeding their foragingrange is extended to gain the nutrition required to recover fromtheir poor condition (Cook 2010) Loss of cover due to late dry-season fires at East Alligator River in the Northern Territorymeant little cover was available for young quolls when theybegan foraging with dog and dingo predation a major source ofmortality (Webb et al 2015) The cumulative impacts of canetoad poisoning fire and predation are highly likely to lead to the

Research priorities for the northern quoll Australian Mammalogy I

extirpation of small and disjunct groups of northern quoll in thePilbara Population viability modelling of the East AlligatorRiver population suggested that high mortality from predationcombined with late dry-season burning was preventing therecovery of northern quoll populations after cane toad invasion

(Webb et al 2015) Improved knowledge of the severity of thesethreats to northern quolls in the Pilbara and their incorporationinto the SDM and PVA framework developed by Cadenheadet al (2015) will allow for better identification of priority areasfor integrated cane toad fire and predator management

Table 3 Expected timelines for the undertaking of research under each priority over the next 10 years and the funds currently allocated fromthe Pilbara Northern Quoll Offset Trust under each research priority

Rank Theme Research priority Current or proposed researchactivity

Time over next10 years

Current funding allocation(approximate)

1 Survey andmonitoring

Develop appropriate andstandardised survey andmonitoring methods

Develop monitoring protocol(traps and cameras) andinvestigate new methods

Ongoing survey and monitoringin conjunction with ResearchThemes 2 and 3 to better mapnorthern quoll distribution andto determine long-termpopulation trends

Years 1ndash2Years 3ndash10

$200 000 per annum over10 years for ResearchThemes 1 2 and 3(inclusive of salaries)

2 Habitatrequirements

Define areas of criticalhabitat for northernquoll in the Pilbara

Identify and describe criticalhabitat and undertake predictivemodelling of unsurveyed areasto inform further survey effort

Years 3ndash4 $50 000 for predictivemodelling (EdithCowan University)

Understand howdisturbance affectshabitat quality andconnectivity

Ongoing collection of data ondisturbance and habitat qualityin conjunction with ResearchThemes 1 and 4

Incorporation of new data intopredictive models

Years 4ndash10 See above

3 Populationdynamics

Better understand thepopulation dynamics ofthe northern quoll in thePilbara

Collect data on demographicparameters breeding dispersaland home-range use toincorporate into future PVA

Continued collectionof data inconjunction withpriorities 1 and 2during years 3ndash10

See above

Investigate populationstructure and theinteraction betweenpopulations

Analysis of existing tissuesamples and newly collectedsamples

Years 1ndash10 $120 000 for initial roundof genetic analysis(Years 3ndash5 MurdochUniversity)

4 Key threats Introduced predators Spatio-temporal interactionsbetween northern quolls andintroduced predators

Other projects as they arise

Years 2ndash4 (currentPhD projectUniversity ofQueensland)

$12 000 contribution tofield expenses

Cane toads Predictive modelling of canetoad distribution in the Pilbara

Years 3ndash4 Part of predictivemodelling by EdithCowan University

Infrastructuredevelopment andinteractions withhumans

Investigation of impacts oflinear infrastructure on northernquoll movement

Year 4 (currentHonours projectEdith CowanUniversity)

$15 000

Interactions betweenthreats

No on-ground projects currentlyplanned

Potential to be assessed throughPVA as data are collected underother research priorities

No funding currentlyallocated

5 Recolonisation ofrestored orartificial habitat

Determine the ability ofthe northern quoll torecolonise disturbedareas or coloniseartificial habitat

As projects arise Years 5ndash8 incollaboration withminesite restorationprograms

No funding currentlyallocated

J Australian Mammalogy V A Cramer et al

Priority 5 Recolonisation of restored andor artificial habitat

While anecdotal evidence suggests that northern quolls will denin waste rock dumps whether artificial or restored habitatprovides for long-term population viability is not known andlong-term assessment of created habitat is required Weacknowledge that a large gap exists between the desire to userestored or artificial habitat as an ecologically equivalent offsetand our limited ability to actually restore or re-create habitat thatis equivalent in structure composition and function particularlyon highly disturbed sites (Maron et al 2012) Yet there is a futureopportunity in the Pilbara to conduct lsquonaturalrsquo experiments onthe recolonisation of quolls after mining ceases including howto best design waste rock dumps so that habitat complexityand productivity is maximised by for example experimentingwith the size and positioning of boulders and their spatialarrangement in relation to surrounding landscape features Theuse of such artificial habitat should be tested in the first stagethrough direct monitoring of animals that have colonised orhave been translocated to the site Such experimentation isunderway at Atlas Ironrsquos Mt Dove operation (Atlas Iron 2012)Population modelling can then be used to predict futurepopulation sizes rates of growth and spread and the probabilityof long-term persistence in the context of various managementactions (eg predator control Armstrong and Reynolds 2012)

Resource allocation and timelines

While the allocation of resources and funding and the periodover which each of the research priorities was expected to beaddressed was not explicitly discussed during the workshopthe ranking of the priorities was taken to indicate the sequencein which the priorities should be addressed At the time of theworkshop some of the research questions identified werealready being investigated within various projects and a portionof the Pilbara Northern Quoll Offset Trust allocated to these InTable 3 we indicate the expected timelines over which someof the research programs are expected to be undertaken WhileAU$175M has been allocated to northern quoll research in thePilbara it is important to note that much of this amount is notcurrently available and will not become available until severalapproved mine developments have become operational InTable 3 we also indicate the proportion of offset funds expectedto be allocated to each research priority Research undertaken inother regions (eg research into CTA in the Kimberley regionahead of the cane toad invasion front) cannot be funded from thePilbara Northern Quoll Offset Trust but will help address someof the research questions identified here

Lessons learnt from the workshop process

The workshop approach used here was less formal than recentlarge-scale exercises used to elicit research priorities for emergingissues in science and policy at national or global levels (reviewedin Sutherland et al 2011) The focus in our (much smaller)workshop was on inclusivity rather than identifying participantsbased on purposive sampling of expertise or organisationalaffiliation We believe that the inclusive nature of the workshopwas a strength but we also recognise that this perhaps createdsome level of bias when ranking the research prioritiesAcknowledging both the experience and knowledge requirements

of those who work regularly in the Pilbara is an importantcomponent of improving the design of research programs that aimto improve conservation management in the region Yet theresearch priorities and their rankings are likely strongly reflectiveof the current information needs of most participants and maynot align with what conservation scientists believe to be themost critical areas of research for northern quoll conservation intothe future Baseline survey and monitoring are often the firstrequirements of a management or research plan for the northernquoll when offset conditions are prescribed and this was reflectedin the primary ranking given to the development of standardsurvey and monitoring methods We acknowledge the concernthat this could simply lead to offset funding being used to lsquomonitorthe extinctionrsquo of the northern quoll particularly after canetoads invade the Pilbara unless these data are used to triggermanagement actions (Oakwood and Foster 2008 Lindenmayeret al 2013) Clearly the currently limited information on habitatuse and demographic parameters for the northern quoll in thePilbara restricts the utility of more sophisticated researchapproaches such as developing robust SDM and PVA While thesevere threat that cane toads pose to the northern quoll wasrecognised this threat is not immediate in the region and mayexplain the less prominent ranking given to research to mitigatethe impacts of cane toads

The northern quoll has become emblematic of the dramaticand rapid species declines occurring in northern Australia andthe level of interest in this species was reflected in the highnumber of participants attending the workshop (one of six runby Parks and Wildlife) Given the number of attendees betterutilisation of the time available would have been gained byproviding participants with a brief before the workshopand asking them to contribute potential research questionsanonymously for interactive review by other participants Inthe exercises reviewed by Sutherland et al (2011) questionswere then discarded if they were not considered to meet criteriathat were previously agreed upon Some areas of researchthat gained some attention on the day may have been discardedif given greater consideration or interrogation before theworkshop For example there was some discussion duringthe workshop of investigating the use of habitat surrogates tomonitor the northern quoll yet research has shown that there islittle validity in using surrogates to monitor the abundance of asingle species (Pierson et al 2015)

Conclusions

Offset funds generated under the environmental approvalsprocess provide an opportunity to mobilise considerablefinancial resources for conservation (Kiesecker et al 2010) Yetthere remain significant conceptual and practical limitations tobe resolved to ensure that direct offsets provide measurableconservation outcomes that are ecologically equivalent to thelosses sustained by proceeding with the development (Maronet al 2012 Bull et al 2013) The collaborative identificationand ranking of research priorities undertaken here is the firststep in the process of developing and evaluating potentialconservation actions for the northern quoll in the Pilbara andserves as a mechanism to ensure that the research priorities ofstate and federal agencies reflects the knowledge needs of

Research priorities for the northern quoll Australian Mammalogy K

practitioners and proponents currently operating in the regionThe provision of offset funds over the next decade providesa unique opportunity to undertake research on a species ofconservation significance at a regional scale and over a longperiod through an integrated and strategic program of researchrather than the collection of ad hoc data While a morestructured approach to defining research questions before theworkshop may have led to greater critical reflection upon anddiscussion of some of the research priorities on the day theworkshop provided an opportunity for an open and diverse-ranging discussion in a collaborative environment amongstthe broader community of scientists environmental consultantsand mining proponents involved in the management andconservation of the northern quoll in the Pilbara

Acknowledgements

We gratefully acknowledge the financial support of Roy Hill who sponsoredthe workshop and partially funded VACrsquos salary We thank the Departmentof the Environmentrsquos Pilbara Taskforce especially Veronica Ritchie fortheir support of interested or affected parties The success of the workshopwas in great part due to the facilitation skills of Craig Salt of SustainableConsulting We thank all of the workshop participants for their open andcollaborative approach to the process Mike Bamford Eddie van EttenDavid Peacock and Peter Lyon provided useful comments on an earlier draftWe are grateful to John Woinarski for his constructive advice thatsubstantially improved the structure and clarity of the manuscript and ananonymous reviewer for careful and critical reading of the previousversion Lesley Gibson Russell Palmer and Al Glen generously discussedvarious aspects of the paper with VAC The map was prepared by CristinaRamalho

References

Allen B L Engeman R M and Allen L R (2011) Wild dogma anexamination of recent lsquoevidencersquo for dingo regulation of invasivemesopredator release in Australia Current Zoology 57 568ndash583

Allen B L Fleming P J Allen L R Engeman R M Ballard G andLeung L K-P (2013) As clear as mud a critical review of evidencefor the ecological roles of Australian dingoes Biological Conservation159 158ndash174 doi101016jbiocon201212004

Anon (1934) Floods in Western Australia Town isolated for week PerthMonday The Sydney Morning Herald 9

Armstrong D P and Reynolds M H (2012) Modelling reintroducedpopulations the state of the art and future directions In lsquoReintroductionBiology Integrating Science and Managementrsquo (Eds J G Ewen DP Armstrong K A Parker and P J Seddon) pp 165ndash222 (Wiley-Blackwell UK)

Atlas Iron (2012) Scope of works ndash Mount Dove artificial quoll habitatUnpublished report Atlas Iron Perth Available at httpwwwatlasironcomauIRMCompanyShowPageaspxCategoryId=190ampCPID=5424ampEID=94365410 [accessed 19 August 2015]

Austin C (2014) Can remote cameras accurately estimate populations ofthe endangered northern quoll BSc(Honours) Thesis University ofTechnology Sydney

Begg R J (1981) The small mammals of Little Nourlangie RockNT III Ecology ofDasyurus hallucatus the northern quoll (MarsupialiaDasyuridae) Wildlife Research 8 73ndash85 doi101071WR9810073

Braithwaite R W and Griffiths A D (1994) Demographic variationand range contraction in the northern quoll Dasyurus hallucatus(Marsupialia Dasyuridae)Wildlife Research 21 203ndash217 doi101071WR9940203

Bull J W Suttle K B Gordon A Singh N J and Milner-Gulland E J(2013) Biodiversity offsets in theory and practice Oryx 47 369ndash380doi101017S003060531200172X

Burnett S (1997) Colonizing cane toads cause population declines innative predators reliable anecdotal information and managementimplications Pacific Conservation Biology 3 65ndash72

Cadenhead N C R White A L and Wintle B A (2015) Identifyingconservation priorities and assessing impacts of proposed futuredevelopment in the Pilbara Bioregion in Western Australia Report bythe National Environment Research Program Decisions Hub to theDepartment of the Environment The University of MelbourneMelbourne

Carwardine J Nicol S Van Leeuwen S Walters B Firn J Reeson AMartin T G and Chades I (2014) Priority threat management forPilbara species of conservation significance CSIRO EcosystemSciences Brisbane

Cook A (2010) Habitat use and home-range of the northern quollDasyurushallucatus effects of fire MSc Thesis University ofWestern AustraliaPerth

Department of Sustainability Environment Water Population andCommunities (2012) Environment and Biodiversity Conservation Act1999 Environmental Offsets Policy Department of SustainabilityEnvironment Water Population and Communities Canberra

Doherty T S Davis R A Etten E J Algar D Collier N DickmanC R Edwards G Masters P Palmer R and Robinson S (2015a)A continental-scale analysis of feral cat diet in Australia Journal ofBiogeography 42 964ndash975 doi101111jbi12469

Doherty T S Dickman C R Nimmo D G and Ritchie E G (2015b)Multiple threats or multiplying the threats Interactions betweeninvasive predators and other ecological disturbances BiologicalConservation 190 60ndash68 doi101016jbiocon201505013

Dunlop J Cook A and Lees J (2014a) BHP Billiton Iron Ore Pilbaranorthern quoll monitoring project Final report May 2010ndashJune 2013Unpublished report prepared for BHP Billiton Iron Ore Department ofParks and Wildlife Perth

Dunlop J Cook A and Morris K (2014b) Pilbara northern quollproject surveying and monitoring Dasyurus hallucatus in the PilbaraWestern Australia Department of Parks and Wildlife Perth Availableat librarydpawwagovau

Florance D Webb J K Dempster T Kearney M R Worthing A andLetnic M (2011) Excluding access to invasion hubs can contain thespread of an invasive vertebrate Proceedings of the Royal Society ofLondon B Biological Sciences 278 2900ndash2908 doi101098rspb20110032

Gaillard J-M Hebblewhite M Loison A Fuller M Powell R BasilleM and Van Moorter B (2010) Habitatndashperformance relationshipsfinding the right metric at a given spatial scale PhilosophicalTransactions of the Royal Society of London Series B BiologicalSciences 365 2255ndash2265 doi101098rstb20100085

Glen A S and Dickman C R (2008) Niche overlap between marsupialand eutherian carnivores does competition threaten the endangeredspotted-tailed quoll Journal of Applied Ecology 45 700ndash707 doi101111j1365-2664200701449x

Glen A S Pennay M Dickman C R Wintle B A and Firestone K B(2011) Diets of sympatric native and introduced carnivores in theBarrington Tops eastern Australia Austral Ecology 36 290ndash296doi101111j1442-9993201002149x

Hennig P (2004) A brief land use history In lsquoAn Inventory and ConditionSurvey of the Pilbara Region Western Australiarsquo (Eds A M E VanVreeswyk A L Payne K A Leighton and P Hennig) TechnicalBulletin No 92 pp 13ndash18 (Western Australian Department ofAgriculture Perth)

Hohnen R Ashby J Tuft K and McGregor H (2013) Individualidentification of northern quolls (Dasyurus hallucatus) using remotecameras Australian Mammalogy 35 131ndash135 doi101071AM12015

L Australian Mammalogy V A Cramer et al

How R A Spencer P B S and Schmitt L H (2009) Island populationshave high conservation value for northern Australiarsquos top marsupialpredator ahead of a threatening process Journal of Zoology 278206ndash217 doi101111j1469-7998200900569x

Johnson C N Isaac J L and Fisher D O (2007) Rarity of a top predatortriggers continent-wide collapse of mammal prey dingoes andmarsupials in Australia Proceedings of the Royal Society of LondonSeries B Biological Sciences 274 341ndash346 doi101098rspb20063711

Jones M E (2000) Road upgrade road mortality and remedial measuresimpacts on a population of eastern quolls and Tasmanian devilsWildlifeResearch 27 289ndash296 doi101071WR98069

Kearney M Phillips B L Tracy C R Christian K A Betts G andPorter W P (2008) Modelling species distributions without usingspecies distributions the cane toad in Australia under current andfuture climates Ecography 31 423ndash434 doi101111j0906-7590200805457x

Kendrick P and Stanley F (2002) Pilbara 4 (PIL4 ndash Roebourne synopsis)In lsquoA Biodiversity Audit of Western Australiarsquos 53 BiogeographicalSubregions in 2002rsquo (Eds J E May and N L McKenzie) pp 581ndash593(Department of Conservation and Land Management Perth)

Kennedy M Phillips B L Legge S Murphy S A and Faulkner R A(2012) Do dingoes suppress the activity of feral cats in northernAustralia Austral Ecology 37 134ndash139 doi101111j1442-9993201102256x

Kiesecker J M Copeland H Pocewicz A and McKenney B (2010)Development by design blending landscape-level planning with themitigation hierarchy Frontiers in Ecology and the Environment 8261ndash266 doi101890090005

King D R and Smith L A (1985) The distribution of the European red fox(Vulpes vulpes) in Western Australia Records of the Western AustralianMuseum 12 197ndash205

Knight A T Cowling R M Rouget M Balmford A Lombard A Tand Campbell B M (2008) Knowing but not doing selecting priorityconservation areas and the researchndashimplementation gap ConservationBiology 22 610ndash617 doi101111j1523-1739200800914x

Laurance W F Koster H Grooten M Anderson A B Zuidema P AZwick S Zagt R J Lynam A J Linkie M and Anten N P (2012)Making conservation research more relevant for conservationpractitioners Biological Conservation 153 164ndash168 doi101016jbiocon201205012

Legge S Murphy S Heathcote J Flaxman E Augusteyn J andCrossman M (2008) The short-term effects of an extensive and high-intensity fire on vertebrates in the tropical savannas of the centralKimberley northern Australia Wildlife Research 35 33ndash43 doi101071WR07016

Lester D (2015) Effective wildlife roadkill mitigation Journal of Trafficand Transportation Engineering 3 42ndash51 doi10172652328-2142201501005

Letnic M Tamayo B and Dickman C R (2005) The responsesof mammals to La Nintildea (El Nintildeo Southern Oscillation)-associatedrainfall predation and wildfire in central Australia Journal ofMammalogy 86 689ndash703 doi1016441545-1542(2005)086[0689TROMTL]20CO2

Letnic M Koch F Gordon C Crowther M S and Dickman C R(2009) Keystone effects of an alien top-predator stem extinctions ofnative mammals Proceedings of the Royal Society of London Series BBiological Sciences 276 3249ndash3256 doi101098rspb20090574

Lindenmayer D B (2015) Continental-level biodiversity collapseProceedings of the National Academy of Sciences of the United States ofAmerica 112 4514ndash4515 doi101073pnas1502766112

Lindenmayer D B Piggott M P and Wintle B A (2013) Counting thebooks while the library burns why conservation monitoring programs

need a plan for action Frontiers in Ecology and the Environment 11549ndash555 doi101890120220

Maron M Hobbs R J Moilanen A Matthews J W Christie KGardner T A Keith D A Lindenmayer D B and McAlpine C A(2012) Faustian bargains Restoration realities in the context ofbiodiversity offset policies Biological Conservation 155 141ndash148doi101016jbiocon201206003

McCall S C McCarthy M A van der Ree R Harper M J Cesarini Sand Soanes K (2010) Evidence that a highway reduces apparentsurvival rates of squirrel gliders Ecology and Society 15 27

McGregor R A Stokes V L and Craig M D (2014) Does forestrestoration in fragmented landscapes provide habitat for a wide-ranging carnivore Animal Conservation 17 467ndash475 doi101111acv12112

McKenzie N L Burbidge A A Baynes A Brereton R N DickmanC R Gordon G Gibson L A Menkhorst P W Robinson A CWilliams M R and Woinarski J C Z (2007) Analysis of factorsimplicated in the recent decline of Australiarsquos mammal fauna Journal ofBiogeography 34 597ndash611 doi101111j1365-2699200601639x

McKenzie N L van Leeuwen S and Pinder AM (2009) Introduction tothe Pilbara Biodiversity Survey 2002ndash2007 Records of the WesternAustralian Museum 78(Suppl) 3ndash89 doi1018195issn0313-122x78(1)2009003-089

Morris K Johnson B Orell P Gaikhorst G Wane A and MoroD (2003) Recovery of the threatened chuditch (Dasyurus geoffroii)a case study In lsquoPredators with Pouches The Biology of CarnivorousMarsupialsrsquo (Eds M Jones C Dickman and M Archer) pp 435ndash451(CSIRO Publishing Melbourne)

Nelson J L Scroggie M P and Belcher C A (2014) Developing acamera trap survey protocol to detect a rare marsupial carnivore thespotted-tailed quoll (Dasyurus maculatus) In lsquoCamera TrappingWildlife Management and Researchrsquo (Eds P D Meek and PJ Fleming) pp 271ndash279 (CSIRO Publishing Melbourne)

Newsome T M Dellinger J A Pavey C R Ripple W J Shores C RWirsing A J and Dickman C R (2015) The ecological effects ofproviding resource subsidies to predators Global Ecology andBiogeography 24 1ndash11 doi101111geb12236

OrsquoDonnell S Webb J K and Shine R (2010) Conditioned tasteaversion enhances the survival of an endangered predator imperilled bya toxic invader Journal of Applied Ecology 47 558ndash565 doi101111j1365-2664201001802x

Oakwood M (1997) The ecology of the northern quoll Dasyurushallucatus PhD Thesis Australian National University Canberra

Oakwood M (2000) Reproduction and demography of the northern quollDasyurus hallucatus in the lowland savanna of northern AustraliaAustralian Journal of Zoology 48 519ndash539 doi101071ZO00028

Oakwood M (2002) Spatial and social organization of a carnivorousmarsupial Dasyurus hallucatus (Marsupialia Dasyuridae) Journal ofZoology 257 237ndash248 doi101017S0952836902000833

Oakwood M and Foster P (2008) Monitoring extinction of the northernquoll Australian Academy of Science Newsletter 71 6

Palomares F and Caro T M (1999) Interspecific killing amongmammalian carnivores American Naturalist 153 492ndash508 doi101086303189

Pavey C R and Bastin G (2014) lsquoAustralian Rangelands and ClimateChange ndash Invasive Animalsrsquo (Ninti One Limited amp CSIRO AliceSprings)

Pierson J C Barton P S Lane PW and Lindenmayer D B (2015) Canhabitat surrogates predict the response of target species to landscapechange Biological Conservation 184 1ndash10 doi101016jbiocon201412017

Pollock A B (1999) Notes on status distribution and diet of northernquoll Dasyurus hallucatus in the MackayndashBowen area mideastern

Research priorities for the northern quoll Australian Mammalogy M

Queensland Australian Zoologist 31 388ndash395 doi107882AZ1999040

Ramanaidou E R and Morris R C (2010) A synopsis of the channeliron deposits of the Hamersley Province Western AustraliaTransactions of the Institutions of Mining and Metallurgy Section BApplied Earth Science 119 56ndash59 doi101179037174510X12853354810624

Schmitt L H Bradley A J Kemper C M Kitchener D J HumphreysW F and How R A (1989) Ecology and physiology of the northernquoll Dasyurus hallucatus (Marsupialia Dasyuridae) at MitchellPlateau Kimberley Western Australia Journal of Zoology 217539ndash558 doi101111j1469-79981989tb02510x

Short J and Smith A (1994) Mammal decline and recovery in AustraliaJournal of Mammalogy 75 288ndash297 doi1023071382547

Spencer P How R Hillyer M Cook A Morris K Stevenson C andUmbrello L (2013) Genetic analysis of northern quolls from the PilbaraRegion ofWestern Australia Year one ndash final report Unpublished reportDepartment of Sustainability Environment Water Population andCommunities Canberra and Department of Parks and Wildlife Perth

Sutherland W J Fleishman E Mascia M B Pretty J and Rudd M A(2011) Methods for collaboratively identifying research priorities andemerging issues in science and policyMethods in Ecology and Evolution2 238ndash247 doi101111j2041-210X201000083x

Sutherst R W Floyd R B and Maywald G F (1996) The potentialgeographical distribution of the cane toad Bufo marinus L in AustraliaConservation Biology 10 294ndash299 doi101046j1523-1739199610010294x

Taylor B D and Goldingay R L (2003) Cutting the carnage wildlifeusage of road culverts in north-eastern New South Wales WildlifeResearch 30 529ndash537 doi101071WR01062

Tingley R Phillips B L Letnic M Brown G P Shine R and BairdS J (2013) Identifying optimal barriers to halt the invasion of canetoads Rhinella marina in arid Australia Journal of Applied Ecology 50129ndash137 doi1011111365-266412021

Van Vreeswyk A M E Payne A L Leighton K A and HennigP (2004) An Inventory and Condition Survey of the Pilbara RegionWestern Australia Technical Bulletin No 92 Department of AgricultureWestern Australia Perth

Webb J Legge S Tuft K Cremona T and Austin C (2015) Can wemitigate cane toad impacts on northern quolls Final report National

Environmental Research Program Northern Australia Hub CharlesDarwin University Darwin

Wintle B A Walshe T V Parris K M and McCarthy M A (2012)Designing occupancy surveys and interpreting non-detection whenobservations are imperfect Diversity amp Distributions 18 417ndash424doi101111j1472-4642201100874x

Woinarski J Menkhorst K Gambold N and Braithwaite R W (1992)Mammals of the Bungle Bungle area In lsquoA Survey of the Wildlife andVegetation of Purnululu (Bungle Bungle) National Park and AdjacentArearsquo (Ed J C Z Woinarski) Research Bulletin No 6 pp 53ndash67(CSIRO Division of Wildlife and Ecology Darwin)

Woinarski J C Z Risler J and Kean L (2004) Response of vegetationand vertebrate fauna to 23 years of fire exclusion in a tropical Eucalyptusopen forest Northern Territory Australia Austral Ecology 29 156ndash176doi101111j1442-9993200401333x

Woinarski J C Z Oakwood M Winter J Burnett S Milne D FosterP Myles H and Holmes B (2008) Surviving the toads patterns ofpersistence of the northern quoll Dasyurus hallucatus in QueenslandReport submitted to the Natural Heritage Trust Strategic ReserveProgram as a component of project 2005162 Monitoring andManagement of Cane Toad Impact in theNorthern Territory Available athttplrmntgovauplants-and-animalsinformation-and-publicationspublications2008 [accessed 14 November 2013]

Woinarski J C Z Armstrong M Brennan K Fisher A Griffiths A DHill B Milne D J Palmer C Ward S Watson M Winderlich Sand Young S (2010) Monitoring indicates rapid and severe decline ofnative small mammals in Kakadu National Park northern AustraliaWildlife Research 37 116ndash126 doi101071WR09125

Woinarski J Burbidge A and Harrison P (2014) lsquoAction Plan forAustralian Mammals 2012rsquo (CSIRO Publishing Melbourne)

Woinarski J C Burbidge A A and Harrison P L (2015) Ongoingunraveling of a continental fauna decline and extinction of Australianmammals since European settlement Proceedings of the NationalAcademy of Sciences of the United States of America 112 4531ndash4540doi101073pnas1417301112

Woolley P A Krajewski C and Westerman M (2015) Phylogeneticrelationships within Dasyurus (Dasyuromorphia Dasyuridae) quollsystematics based on molecular evidence and male characteristicsJournal of Mammalogy 96 37ndash46 doi101093jmammalgyu028

N Australian Mammalogy V A Cramer et al

wwwpublishcsiroaujournalsam

Page 3: Research priorities for the northern quoll (Dasyurus ......The Pilbara population of the northern quoll (Dasyurus hallucatus) has been seldom studied, and the impacts of threats such

Ore httpwwwenvironmentgovaunode18597 HamersleyIron Pty Ltd (Rio Tinto) httpwwwenvironmentgovaunode18608)

Biodiversity offsets are increasingly used as a policy toolto permit development but ensure that any residual adverseimpacts of the development on the environment arecompensated (Bull et al 2013) In best practice offsets areapplied as part of a mitigation hierarchy where all reasonableefforts to avoid or minimise impacts upon the environment arefirst considered (Kiesecker et al 2010) Yet the limited data onthe distribution and status of the northern quoll in the Pilbarathe poor understanding of critical habitat in relation to futuremining and infrastructure development as well as uncertaintyaround the level of impact from other threats has meant thatuncertainty around how to create direct ecologically equivalentoffsets is high Under the Australian Governmentrsquos offsetpolicy (Department of Sustainability Environment WaterPopulation and Communities 2012) compensatory offsetssuch as directing a proportion of offset funds towards scientificresearch are considered as an acceptable component of anoffsets package in these situations This has been the case forthe northern quoll in the Pilbara with approximately AU$175Mof offset funds committed to research on the species over thenext decade (although not all of this amount is currently active)

The dramatic increase in fauna surveys associated withenvironmental impact assessments in the Pilbara has generateda valuable dataset on the northern quoll and other threatenedspecies (Carwardine et al 2014) Collation of these datacould greatly improve our knowledge of speciesrsquo ranges andcritical habitats yet most of these data are not within thepublic domain (Carwardine et al 2014) Further it has beenargued that much conservation research is having little directinfluence on the achievement of on-ground conservationoutcomes in part because the conceptual novelty valued bynational funding agencies may not align with the moreapplied needs of practitioners (Knight et al 2008 Lauranceet al 2012) A potential solution to overcoming the lsquoknowledgendashimplementationrsquo or lsquoknowingndashdoingrsquo gap is to encouragecollaboration between scientists policy makers and practitionerswhen research projects are developed (Knight et al 2008Sutherland et al 2011 Laurance et al 2012) In an effort toencourage data-sharing and to foster a sense of ownership overthe research agenda that will be funded by offset provisionsParks and Wildlife instigated a collaborative process betweenscientists practitioners and mining proponents to identifyand rank research priorities for the northern quoll in the PilbaraIn this paper we present the research priorities and the keyquestions associated with each priority as identified byworkshop participants This is the first step in developing aresearch program to increase our knowledge of the northernquoll in the Pilbara and to better understand the impacts ofcurrent and future threats so that a viable population ismaintained into the future using evidence-based management

Methods

Parks and Wildlife hosted a workshop attended by researchscientists employed in the university or government sector (13people) research scientists and environmental consultants

employed in the private sector (30 people) mining industryrepresentatives (22 people) and representatives of WesternAustralian and Australian government departments responsiblefor environmental regulation and approvals (9 people) all ofwhom have knowledge of the northern quoll mining andassociated activities andor the Pilbara region The workshopwas facilitated by an independent professional facilitator Theprocess to define research priorities was as follows (1) After aseries of presentations by experts the group as a whole discussedthe key threats to the northern quoll in the Pilbara This includedboth current and future threats (eg climate change cane toadinvasion) and both well- and lesser-understood threats (eg theimpact of disease) (2) Participants were then allocated to one ofeight breakout groups so that each group contained membersfrom the range of affiliations present Each group then identifieda series of research priorities that they considered would bestaddress the current gaps in knowledge that are the main barriersto the effective management and conservation of the northernquoll in the Pilbara both in terms of better understanding theirdistribution and ecology and in mitigating the key threats Theresearch priorities from each group were then presented to allparticipants and collated by the facilitator (3) Each group thenranked the sequence in which the research priorities should beaddressed based on which research needs they considered weremost pressing (4) The rankings were then compared and collatedacross groups to define the final ranking of research priorities Inmost cases there was agreement across the groups as to thesequence in which the research priorities should be addressed(5) Finally new breakout groups were formed based onexpertise and interest to discuss and identify the key researchquestions that needed to be addressed under each researchpriority

Research priorities

Research priorities are presented below in the order in whichthey should be addressed based on the ranking of the workshopparticipants Key research questions aligned with each researchpriority are presented in Table 1 We also provide a brief reviewof the current evidence that will further guide the development ofa research program to answer these questions

Priority 1 Develop appropriate and standardised surveyand monitoring methods

Developing a standard survey and monitoring protocol for thenorthern quoll in the Pilbara based on detection and occupancyprobabilities (Wintle et al 2012) was identified as the highestresearch priority by the group A review of over 200 surveyreports most of which were related to mining activities foundthat the sampling effort and methods varied widely betweensurveys and were often not adequate to detect northern quolls(A Cook unpubl data) Inadequate survey effort andor usingan inappropriate survey method introduce uncertainty as towhether the failure to detect the northern quoll within an areareflects a true absence of the species (Wintle et al 2012) Thedata gathered from surveys that vary widely in terms of effortand method may have limited comparability and incurs a lostopportunity to gather regional metadata for the northern quollin an area where any survey effort represents a significant

Research priorities for the northern quoll Australian Mammalogy C

Tab

le1

Researchpriorities

fortheno

rthern

quollan

dkeyqu

estion

sassociated

witheach

priorityan

das

compa

redwiththeinform

ationrequ

ired

undersimila

rthem

esan

dthepriority

givento

thisin

Woina

rski

etal(201

4)

Rank

Theme

Researchpriority

Key

questio

nsInform

ationrequired

andpriority

givenundersimilarthem

ein

Woinarski

etal(2014)

1Surveyand

mon

itoring

Develop

appropriate

andstandardised

survey

and

mon

itoring

metho

ds

Whatistheefficacy

ofthemajor

survey

metho

dscurrently

used

forno

rthern

quollsin

the

Pilb

ara

How

dovariou

scombinatio

nsoftrap

typesarraysdurationoftrappingbaitsetc

compareWhich

camera-trap

protocol

ismosteffectivefor(a)identifying

individu

alno

rthern

quollsin

thePilb

araand(b)estim

atingpo

pulatio

nsizes

Which

methodismost

cost-effectiv

eandefficientin

variou

sPilb

araland

scapes

Whatisthedetectionprobability

ofno

rthern

quollsin

each

habitattypeBased

onprior

surveywhatitthepriorprobability

ofoccupancyin

thesehabitattypes

Based

onthis

know

ledge

how

manynon-detections

ofquollsarerequired

before

wecanstatewith

prespecified

confi

dencethat

northern

quollsareabsent

from

asite

Surveyto

betterdefine

fine-scale

distributio

nandnumberof

individu

alsin

subpop

ulations

mediumndashhigh

Establishor

enhancemon

itoring

program

bydesigningintegrated

mon

itoring

prog

ramsmedium

2Habitat

requirem

ents

Define

areasof

critical

habitatforno

rthern

quollin

thePilb

ara

How

donorthern

quollsusehabitatinthePilb

ara

How

arequollpopulations

distributedin

relatio

nto

habitatq

uality(egh

ighqu

ality

low

quality

orsink

habitats)

How

ishabitat

selectioninfluenced

bypredationrisk

andsocial

factors

Whatinformationisrequ

ired

tofurtherrefine

anSDM

forno

rthern

quollsin

thePilb

ara

How

canthisinform

ationbe

mosteffectivelyused

toim

provemonito

ring

effortand

outcom

es

How

domod

elsvary

across

thePilb

aramainlandandoffsho

reislandsaccountin

gfor

clim

atic

gradientsandtopography

Whatarefuture

predictio

nsfornorthern

quollpopulatio

ndistributio

nsu

nder

modelsof

clim

atechange

andcane

toad

invasion

Assesshabitatrequirem

entsna

Und

erstandho

wdisturbanceaffects

habitatqu

ality

and

conn

ectiv

ity

How

does

fire

andgrazingaffecthabitatq

ualityandhabitatu

seby

northern

quollsin

the

Pilb

ara

How

importantisfire

asabarrierto

dispersal

How

importantishabitatconnectivity

form

etapopulations

ofnorthernquollinthePilb

ara

How

does

habitattypeaffecttheconnectiv

ityof

metapopulationsHow

does

disturbance

both

discrete

(eginfrastructure

such

asrailandroads)

anddiffuse(egchangesin

vegetatio

ncompositio

nandstructure)alterhabitatconnectiv

ity

Assessrelativ

eim

pact

ofthreats

quantifyrelativ

eim

pact

offire

regimesmdashhigh

Assesseffectivenessof

threat

mitigatio

nop

tionsland

scape-scale

optio

nsforretentionof

longer-

unburntpatchesmdash

high

3

Pop

ulation

dynamics

Betterunderstand

the

popu

latio

ndy

namicsof

the

northern

quollinthe

Pilb

ara

How

manysubpop

ulations

aretherein

thePilb

ara

How

dotheseinteractAre

subpopulations

demographically

stablein

spaceandtim

eHow

does

habitatand

clim

ate

affect

popu

latio

ndy

namics

How

does

disturbanceaffect

demog

raphic

stability

Assessdietlifehistory

na

Investigatepo

pulatio

nstructureandthe

interactionbetween

popu

latio

ns

How

domeasuresofgenetic

distance

relatetospatialecology

asmeasuredby

otherm

eans

iedoes

inferred

genetic

connectiv

ityrelate

toobserved

dispersal

Can

genetic

inform

ationbe

used

todeterm

inehabitatbarriers

Resolve

taxonomic

uncertainties

assess

genetic

relatednessof

spatially

disparatepo

pulatio

nsto

identifywhether

thereare

genetically

distinct

subpop

ulationsmdashlowndashmedium

D Australian Mammalogy V A Cramer et al

4Key

threats

Introduced

predators

Doespredationsignificantly

affecttheno

rthern

quollinthePilb

ara

How

does

thisvary

across

habitattypes

Doescompetitionforresourceswith

foxesor

catsim

pact

onsubpopulationsWhich

lifestages

aremostvulnerable

Given

thattheno

rthern

quollisalso

amesop

redatordo

esthepresence

ofding

oeseither

directly(through

predation)

orindirectly(through

theregulatio

nof

foxesandcats)impact

onsubpop

ulations

inthePilb

ara

Assessrelativ

eim

pact

ofthreats

predationby

feralcatsanddogsmdash

high

interactions

betweenferal

catsdo

gsdingoes

andfoxesmdash

mediumndashhigh

Assesseffectivenessof

threat

mitigatio

nop

tionsassess

which

feasible

managem

entoptio

nsfor

feralcatsanddo

gscanlead

topo

pulatio

nrecoverymdashmediumndash

high

Develop

new

orenhanceexistin

gmanagem

entmechanism

sfor

broad-scale

targetedferalcat

controlmdash

medium

Canetoads

Will

theclosureof

artifi

cialwater

bodies

preventor

simplyslow

therateofinv

asionof

cane

toadsinto

thePilb

ara

Whatisthedegree

ofoverlapbetweenSDMfundamentalniche

modelsforn

orthernquolls

andcane

toadsin

thePilb

ara

Can

potentialrefug

eareasforno

rthern

quollb

eidentifi

ed

IsCTAlik

elytobe

effectivewhentaste-aversion

baits

aredeliv

ered

underfi

eldcond

itions

Atwhatscaleisbaitdeliv

eryfeasible

Assesseffectivenessof

threat

mitigatio

nop

tionsassess

feasibility

ofbroad-scaletaste-

aversion

lsquotrainingrsquo

ofqu

ollsmdash

mediumassess

local-scale

exclusionof

toadsor

preventio

nof

colonisatio

n(m

ainlandsites)mdash

medium

Infrastructure

developm

entand

interactions

with

humans

Are

road-related

deaths

ofno

rthern

quollasign

ificant

causeof

mortalityin

thePilb

ara

Dorailw

aylin

esinfrastructurecorridorsor

wideroadspresenta

barrierto

northern

quoll

mov

ement

Whatmethods

canbe

used

tomitigate

theim

pactsof

infrastructure

developm

entand

associated

machinery

operation

Not

addressed

Interactions

between

threats

Dofrequent

andintensefireslead

toreducedstructural

complexity

loss

ofvegetatio

ncoverandsubsequentincreasedpredationof

northern

quollsparticularlyof

males

during

andor

afterbreeding

Whatarethecumulativeim

pactsof

mortalitycaused

bycane

toad

ingestionandpredation

bycatsdo

gsdingoes

andfoxes

See

inform

ationon

introd

uced

predatorsandcane

toadsabove

5Recolon

isationof

restored

orartifi

cial

habitat

Determinetheability

oftheno

rthern

quoll

torecolonise

disturbedareasor

colonise

artifi

cial

habitat

Whatarethekeyelem

entsof

northern

quollhabitatthatneedtobe

restoredcreated

sothat

habitatcomplexity

andprod

uctiv

ityismaxim

ised

Whatlevelo

flandscapeconn

ectiv

ityisrequired

arou

ndrestoredartificialhabitatso

that

genetic

connectiv

itybetweensubpopulations

canbe

maintained

How

long

should

restored

orartifi

cial

habitatbe

directly

managedparticularly

for

predators

How

cantheretentionof

largebouldersof

wasterock

beincorporated

into

rehabilitation

plans

Not

addressed

Research priorities for the northern quoll Australian Mammalogy E

investment of time and resources The limited amount anduncertain quality of occurrence data has been a major restrictionon the utility of a species distribution model (SDM) and apopulation viability analysis (PVA) for the northern quoll in thePilbara (Cadenhead et al 2015 see below) While survey andmonitoring per se is not a research activity that directly informsmanagement or threat mitigation carefully implementedmonitoring programs are essential to identify many elements ofa speciesrsquo ecology to detect changes in population and toquantify the ecological effectiveness and cost-effectivenessof management interventions (Lindenmayer et al 2013Lindenmayer 2015)

Parks and Wildlife recently released revised protocols forthe long-term monitoring of known populations of thenorthern quoll (Dunlop et al 2014b) and the effectiveness ofthese protocols in presenceabsence surveys should be testedThe cost-effectiveness of using camera traps for wildlifesurveys in remote locations such as the Pilbara makes their useappealing Hohnen et al (2013) described how the strategicplacement of cameras and baits combined with taking a greaternumber of photographs per trigger allows the capture ofimages that can be used to identify individual northern quollsfrom their spot patterns Remote cameras have also been shownto be a viable method for estimating population sizes of thenorthern quoll in areas with both dense (Austin 2014) andsmall populations (Webb et al 2015) Further testing of a rangeof parameters (eg see Nelson et al 2014) to develop a robuststandard protocol for the Pilbara is required along with acomparison of the cost-effectiveness and any ethicalreducedhandling benefits of using remote cameras versus live-trappingfor estimating population sizes in Pilbara landscapes

Priority 2 Improve our understanding of habitatrequirements

Define areas of critical habitat for the northern quollin the Pilbara

The Department of Agriculture and Food Western Australiahas mapped 102 land systems across the Pilbara bioregionbased on several biophysical components including landformgeology soils vegetation and drainage patterns (Van Vreeswyket al 2004) A review of records for the northern quoll in thePilbara showed that 65 of records were from the RobeCapricorn and Boolaloo land systems (Table 2) which compriselow plateaux mesas hills and ridges of varying geology(limonite granite and sandstone respectively) Most recentsurvey effort in the Pilbara is biased towards mineral-bearingformations or areas of mining-related infrastructure (eg railand road corridors) (Fig 1) Further investigation of theserecords is required to assess whether these land systems areindeed preferred habitat for northern quolls as opposed to anartefact of survey effort The comparatively fewer records thatcoincide with the conservation estate in the region (Fig 1) arealso likely an artefact of survey effort

The first step in defining areas of critical habitat for thenorthern quoll in the Pilbara is to improve our understanding ofhabitat use particularly in terms of defining habitat suitability atfiner scales than is currently available As part of the strategic

environmental assessments being undertaken for the Pilbarabioregion an SDM has been developed for the northern quoll(Cadenhead et al 2015) Uncertainty about the quality of theoccurrence data available the limited amount of data availableas well as the bias towards data collection in areas proposed fordevelopment constrained the level of certainty around theoutcomes of the model Improved model certainty requiresfurther collection of contemporary species-occurrence data andthe development of finer-resolution GIS layers particularly onethat captures finer-scale geological attributes such as mesas orrocky outcrops considered as highest-quality habitat for thenorthern quoll (Cadenhead et al 2015)

Better definition of the habitat attributes favoured by thenorthern quoll will allow further refinement of the GIS variablesincluded in the SDM or allow the development of predictivesite-based habitat models Information on how northern quollsuse habitat can be gained through direct observation of howthey interact with habitat features or from the association ofhabitat features with the presence of quolls as determined byGPS receivers (Gaillard et al 2010) This level of detail iscurrently not available from survey records returned as part ofenvironmental impact assessments but collection of habitatdata using the protocols outlined by Dunlop et al (2014b) infuture surveys would rapidly increase the quality of dataavailable

Understand how disturbance affects habitat qualityand connectivity

The alteration of fire regimes has been the most pervasivelandscape-scale change in the Pilbara Between 1993 and 2006over 72 of the region was burnt with more than 28 of thearea burnt two or more times (McKenzie et al 2009) Woinarskiet al (2004) found that northern quolls may be able to tolerate amoderately high frequency of fire as long as these fires werelow-intensity early-season burns and the impacts of fire onvegetation structure and composition were not exacerbatedby the grazing of introduced herbivores In the Pilbarahowever as with many other grazing regions of Australia earlypastoralists often overestimated the productivity and resilienceof native vegetation to sheep and cattle grazing which led tosevere degradation of vegetation particularly along riverfrontages (Hennig 2004) The soil degradation and loss ofvegetative cover caused by intense grazing pressure in thesouth Kimberley is thought likely to have contributed to thesubstantial decline or disappearance of medium-sized animalsincluding the northern quoll from the Bungle Bungle area(Woinarski et al 1992) Death by predation is most likely in openareas (Oakwood 2000) and therefore the effects of fire andgrazing on northern quoll populations cannot be consideredwithout also taking account of likely increases in mortality due topredation (see lsquoIntroduced predatorsrsquo below)

Understanding how the localised but intensive activityassociated with mining affects habitat quality and habitat useby the northern quoll in areas surrounding development islimited A recent study that examined the use of ballast quarriesand paired undisturbed sites by northern quolls found that theywere foraging denning and showing signs of breeding at thequarry sites where greater numbers of quolls were captured(Dunlop et al 2014a) This was likely due to the combination of

F Australian Mammalogy V A Cramer et al

habitat complexity (eg diversity of boulder sizes) and thepresence of permanent water in the quarry which may haveincreased prey availability at these sites

Movement of animals through the landscape for dispersaland repopulation is another important consideration for a speciesin which both sexes tend towards semelparity and is thereforepredisposed to local extinction (Oakwood 1997) Improvedunderstanding of how the northern quoll (particularly dispersingmales) uses areas of lower habitat quality and the identificationof such habitat in habitat-suitability models will allow futurelandscape-scale strategic assessments to better predict thecombined impacts of disturbance on the ability of northern quolls

to move through the landscape Analysis of regional populationgenetics will also improve our understanding of the level ofdispersal occurring across the landscape

Priority 3 Population dynamics

Better understand the population dynamics of thenorthern quoll in the Pilbara

Little information is available on the number ofsubpopulations of northern quoll in the Pilbara how thesesubpopulations interact how stable they are in space and timeand how they respond to disturbance As no demographic

Table 2 Records of the northern quoll in each land system type in the PilbaraRecords were current as of 23 May 2014

Land system Area (km2) No of records Land system description

Augustus 61 1 Rugged ranges hills ridges and plateaux supporting mulga shrublands and hard spinifex grasslandsBoolaloo 1502 245 Granite hills domes and tor fields and sandy plains with shrubby spinifex grasslandsBoolgeeda 7748 29 Stony lower slopes and plains below hill systems supporting hard and soft spinifex grasslands andmulga

shrublandsCalcrete 1444 2 Low calcrete platforms and plains supporting shrubby hard spinifex grasslandsCallawa 1003 9 Highly dissected low hills mesas and gravelly plains of sandstone and conglomerate supporting soft and

hard spinifex grasslandsCapricorn 5296 257 Hills and ridges of sandstone and dolomite shrubby hard and soft spinifex grasslandsCoongimah 3244 12 Plateau surfaces low hills with steep slopes and undulating uplands supporting hard spinifex grasslandsDune 138 1 Dune fields supporting soft spinifex grasslandsGranitic 4020 1 Rugged granitic hills supporting shrubby hard and soft spinifex grasslandsHorseflat 1261 4 Gilgaied clay plains supporting tussock grasslands and minor grassy snakewood shrublandsLittoral 1577 3 Bare coastal mudflats with mangroves on seaward fringes samphire flats sandy islands coastal dunes

and beachesMacroy 13 095 94 Stony plains and occasional tor fields based on granite supporting hard and soft spinifex grasslandsMcKay 4202 21 Hills ridges plateau remnants and breakaways of metasedimentary rocks supporting hard spinifex

grasslandsMallina 2557 8 Sandy surfaced alluvial plains supporting soft spinifex (and occasionally hard spinifex) grasslandsMarandoo 459 2 Basalt hills and restricted stony plains supporting grassy mulga shrublandsMosquito 1840 3 Stony plains and prominent ridges of schist and other metamorphic rocks supporting hard spinifex

grasslandsNewman 14 580 81 Rugged jaspilite plateaux ridges and mountains supporting hard spinifex grasslandsOnslow 424 1 Sandplains dunes and clay plains supporting soft spinifex grasslands and minor tussock grasslandsParaburdoo 565 3 Basalt-derived stony gilgai plains and stony plains supporting snakewood and mulga shrublands with

spinifex and tussock grasslandsParadise 1479 8 Alluvial plains supporting soft spinifex grasslands and tussock grasslandsRiver 4088 37 Active flood plains major rivers supporting grassy eucalypt woodlands tussock grasslands and soft

spinifex grasslandsRobe 865 369 Low limonite mesas and buttes supporting soft spinifex (and occasionally hard spinifex) grasslandsRocklea 22 993 40 Basalt hills plateaux lower slopes and minor stony plains supporting hard spinifex (and occasionally

soft spinifex) grasslandsRuth 346 36 Hills and ridges of volcanic and other rocks supporting hard spinifex (occasionally soft spinifex)

grasslandsStuart 1794 3 Gently undulating stony plains supporting hard and soft spinifex grassland and snakewood shrublandsTaylor 129 3 Stony plains and isolated low hills of sedimentary rocks supporting hard and soft spinifex grasslandsTalga 2124 9 Hills and ridges of greenstone and chert and stony plains supporting hard and soft spinifex grasslandsTuree 581 2 Stony alluvial plains with gilgaied and non-gilgaied surfaces supporting tussock grasslands and grassy

shrublandsUaroo 7681 37 Broad sandy plains supporting shrubby hard and soft spinifex grasslandsUrandy 1311 6 Stony plains alluvial plains and drainage lines supporting shrubby soft spinifex grasslandsWhite Springs 266 4 Stony gilgai plains supporting tussock grasslands and hard spinifex grasslandsWona 1815 10 Basalt upland gilgai plains supporting tussock grasslands and minor hard spinifex grasslandsYamerina 1207 1 Floodplains and deltaic deposits supporting tussock grasslands grassy woodlands and minor halophytic

low shrublands

Research priorities for the northern quoll Australian Mammalogy G

studies have been undertaken in the region Cadenhead et al(2015) used demographic information from the NorthernTerritory and Kimberley populations in their PVA Uncertaintyabout data quality and the limited information available ondemographic parameters for the Pilbara constrained the level ofcertainty around the outcomes of the PVA The model doeshowever provide a framework into which new data can beincorporated as they become available For example preliminarydata from the Pilbara monitoring program suggest that theproportion of male die-off in the Pilbara population is lowerthan in the Kimberley and Northern Territory populations(Dunlop unpubl data) Cadenhead et al (2015) stress the needfor more demographic and occurrence data to be collectedbefore further development decisions are made (see lsquoDevelopappropriate and standardised survey and monitoring methodsrsquoand lsquoImprove our understanding of habitat requirementsrsquo above)

Investigate population structure and the interactionbetween populations

Analysis of the genetic diversity and population structure ofthe Pilbara population of the northern quoll is ongoingOnly small differences in genetic diversity as measured byheterozygosity were found between samples obtained at 16locations within the Pilbara with the exception of the isolatedpopulation on Dolphin Island (Spencer et al 2013) This resultindicates a high level of movement across the landscape byindividuals during mating and dispersal phases Spatial structureanalysis of samples collected from around Pannawonicasuggested a lsquoneighbour sizersquo of ~15 km (Spencer et al 2013)Current data show no evidence of genetic bottleneckssuggesting that if there has been a population decline in thePilbara that this has not affected genetic diversity A usefulapproach would be to combine genetic information with datafrom radio-tracking or GPS collars to better understand theuse of lsquolandscapersquo space and how this influences populationdynamics

Priority 4 Key threats and the interaction of these threats

Introduced predators

Introduced cats and foxes are considered to be one of themain causes of the decline and extinction of the critical-weight-range mammals (35 g to 55 kg) in Australia (Short and Smith1994 McKenzie et al 2007 Woinarski et al 2015) but theirimpacts on the northern quoll are less clear A recent review offeral cat diets across Australia found that northern quolls werepresent in the diet of feral cats (Doherty et al 2015a) andWoinarski et al (2014) consider the threat posed by cats to thenorthern quoll to be severe although the impacts of cats may bereduced in rugged refuge areas The impacts of cats on othermuch larger species of quoll appear to be inconsistent withsome evidence of coexistence between cats and quolls beforethe arrival of foxes (Glen and Dickman 2008 and referencescontained within) Due to the limited overlap in their distributionthroughout Australia the threat posed by foxes to northernquolls is less understood than for cats and Woinarski et al(2014) consider the threat posed by foxes to be moderate on anational level Foxes are common in coastal near-coastal and

riparian areas of the Pilbara and extend as far north as Broome(King and Smith 1985 Kendrick and Stanley 2002) They arerenowned for their aggression towards smaller carnivores(Palomares and Caro 1999) and neither the larger chuditch(Dasyurus geoffroii) nor spotted-tail quoll (Dasyurus maculatus)are known to coexist with large numbers of foxes (A Glen perscomm) The interactions between quolls cats and foxesare potentially more complex than for other medium-sizedmarsupials due to niche overlap in diet and competition for densites with the larger spotted-tailed quoll and chuditch bothexperiencing exploitation and interference competition fromfoxes (Morris et al 2003 Glen et al 2011) Competitionbetween feral cats and the northern quoll may potentially beintense during periods of low rainfall in the Pilbara when preyavailability is low Understanding whether foxes and cats aresignificant predators of or competitors with the northern quollin the Pilbara and how this affects their population viabilitywill clarify the level of risk posed by these predators Thesausage-style baits used for feral cats (eg Eradicatreg andCuriosityreg) are potentially attractive to the northern quoll withthe risk of mortality or reduced fitness Trials to assess themagnitude of the risks from Eradicatreg baits are currentlyunderway in the Pilbara

The role of dingoes and wild dogs in the mortality of andinteractions between mesopredators (both native and introduced)requires further investigation Dingoes are known to kill northernquolls butmay not eat these kills (Oakwood 1997) A recent studyin the Northern Territory found that camp dogs and dingoes werethe major source of mortality for northern quolls at the study site(Webb et al 2015) and roaming domestic dogs were identified asa cause of mortality for northern quolls in Queensland (Pollock1999) Human resource subsidies (eg stock carrion in rangelandareas) can artificially support predator populations thus indirectlyincreasing predation pressure on prey (Newsome et al 2015)A dynamic debate currently exists around the role of the dingo inregulating populations of feral cats and foxes and the implicationsof this for the conservation of the critical-weight-range mammalsin Australia (see Johnson et al 2007 Letnic et al 2009 Allenet al 2011 2013 Kennedy et al 2012) Understanding thepopulation density at which the role of the dingo as a regulator ofintroduced predators outweighs its own impact as a direct predatorof the northern quoll will inform whether dingowild dog controlwould benefit the northern quoll

Cane toads

The introduced cane toad now occupies more than half thehistoric range of the northern quoll and has caused an immediatecollapse or extirpation of northern quoll populations in theareas it has invaded (Burnett 1997 Oakwood and Foster2008 Woinarski et al 2008 2010) Earlier modelling work thatconsidered only climate parameters as well as more recentwork that incorporates future climate change suggest thatcane toads will not invade the Pilbara (Sutherst et al 1996Pavey and Bastin 2014) Other recent work that considers thefundamental niche of the cane toad and incorporates the locationof artificial watering points predicts that they will colonisethe Pilbara from the Kimberley via the coastal edge of theGreat Sandy Desert (Kearney et al 2008 Florance et al 2011

H Australian Mammalogy V A Cramer et al

Tingley et al 2013) This has created some level of uncertaintyabout the likely impact of cane toads on northern quolls in thePilbara Strategies for cane toad management were not rankedhighly by participants in the threat-management frameworkdeveloped by Carwardine et al (2014) which the authorsexplained as possibly due to uncertainty about how far canetoads will spread into the region

The model developed by Tingley et al (2013) suggested thatthe closure of 100 artificial water bodies at one of three potentiallocations along the coastal corridor between the Kimberley andthe Pilbara would prevent the invasion of cane toads into thePilbara While their model did account for uncharacteristicallywet years every three decades the dispersal of toads betweenwater bodies was determined by the number of rainy daysoccurring during the year (ie toads could only disperse betweenwater bodies for three days following rainfall) rather thanrainfall amount the associated likelihood of flooding and theavailability of free water across the landscape The cyclonic andthunderstorm nature of much of the rainfall in the north-west ofWestern Australia means that the region is prone to significantand sometimes extended flooding In April 1934 the countryaround Anna Plains (250 km south of Broome) was flooded forhundreds of miles when 27 inches (686mm) of rain fell infour days (Anon 1934) In March 2004 over 350mm of rain wasrecorded within a 48-h period at Telfer on the southern edge ofthe Great Sandy Desert due to Tropical Cyclone Fay Thiscaused the flooding of the De Grey and Oakover river systemsand flooded 140 km of the Telfer Access Road with floodwaterstaking several months to subside (wwwbomgovau) Extendingthe model of Tingley et al (2013) to account for such events willhelp clarify whether closing artificial water bodies is likely toprevent the invasion of cane toads into the Pilbara or may onlyassist in delaying their inevitable invasion

Comparison of SDMs (eg Cadenhead et al 2015) fornorthern quoll with fundamental niche models for cane toads(eg see Kearney et al 2008) within the Pilbara may identifypotential refuge areas for the northern quoll and these areasshould be prioritised for conservation actions Because theinvasion point of cane toads into the Pilbara is likely to be narrow(beginning around the mouth of the De Grey River Tingleyet al 2013) a proactive approach to training northern quollsto avoid cane toads may prove beneficial Conditioned taste-aversion (CTA) has been used successfully to train captive-reared northern quolls to avoid cane toads by offering themsmall dead toads infused with thiabendazole before theirrelease into the wild (OrsquoDonnell et al 2010)Monitoring of thesequolls after their release showed that some of their offspringhad learnt to avoid eating cane toads either through ingestionof small non-lethal toads or via social learning from theirmothers (Webb et al 2015) Fields trials of toad-aversion baits(either sausages or meatballs made from toad flesh andcontaining thiabendazole) are currently underway in the centralKimberley where cane toads were expected to invade in late2015 (Webb et al 2015) Further testing of CTA approachesunder a variety of field conditions is required The developmentof stable long-lasting baits suitable for aerial deployment isconsidered a pressing need as well as a captive colony ofnorthern quolls upon which to test the efficacy of toad-aversionbaits for inducing aversion to live toads (Webb et al 2015)

Infrastructure development and interactionswith humans

Infrastructure development may increase the likelihood ofnorthern quoll deaths due to motor vehicle strikes or beingcrushed in machinery Significant chuditch mortality wasobserved along mine access roads in the south-west of WesternAustralia (McGregor et al 2014) and the widening and sealingof an access road dramatically increased the number of roadkills and led to the extirpation of the resident population ofeastern quolls (D viverrinus) in Tasmania (Jones 2000)Infrastructure corridors such as high-tension power-lineeasements and ore-haulage railways are increasingly beinginstalled throughout the range of the northern quoll Althoughthese do not create the same mortality risk as roads they bisectand fragment habitat create large open spaces and potentiallyincrease predation risk

A recent small-scale radio-tracking study (see Dunlop et al2014a) around several quarries in the Pilbara found that northernquolls crossed roads but not railway lines The short-term natureof this work and the limited number of quolls able to be trackedmeans that the influence of roads railway lines and infrastructurecorridors on the movement and mortality of northern quollsrequires further assessment Using radio-tracking techniques todetermine crossing rates and road kill events may be difficultover large areas so population studies comparing groups ofquolls near major roads with those not affected by infrastructure(eg McCall et al 2010) may provide a more feasible meansof assessing the impacts of roads If vehicle strikes are foundto significantly contribute to the mortality of a local populationof the northern quoll then mitigation methods should betrialled on sections of road identified as road-kill hotspotsCulverts and underpasses have been shown to be used by avariety of small and medium-sized mammals and frogs (Taylorand Goldingay 2003) while installing rumble strips on roadshas led to decreased rates of road-kill of nocturnal animals(Lester 2015)

Interactions between threats

At present wildfire the subsequent loss of vegetation coverand increased predation of northern quolls is likely to be themost significant of the interactions between threats The directeffects of wildfire have been found to be less significant inthe subsequent decline of small to medium-sized mammalsparticularly for denning species such as the northern quoll thanthe indirect consequences of reduced resource availability andthe increased risk of predation due to lack of cover (Oakwood2000 Letnic et al 2005 McKenzie et al 2007 Legge et al2008 Doherty et al 2015b) Male northern quolls are perhapsmost vulnerable to such losses of cover during the breedingseason when their range can extend to over 100 ha (Oakwood2002) or where males survive after breeding their foragingrange is extended to gain the nutrition required to recover fromtheir poor condition (Cook 2010) Loss of cover due to late dry-season fires at East Alligator River in the Northern Territorymeant little cover was available for young quolls when theybegan foraging with dog and dingo predation a major source ofmortality (Webb et al 2015) The cumulative impacts of canetoad poisoning fire and predation are highly likely to lead to the

Research priorities for the northern quoll Australian Mammalogy I

extirpation of small and disjunct groups of northern quoll in thePilbara Population viability modelling of the East AlligatorRiver population suggested that high mortality from predationcombined with late dry-season burning was preventing therecovery of northern quoll populations after cane toad invasion

(Webb et al 2015) Improved knowledge of the severity of thesethreats to northern quolls in the Pilbara and their incorporationinto the SDM and PVA framework developed by Cadenheadet al (2015) will allow for better identification of priority areasfor integrated cane toad fire and predator management

Table 3 Expected timelines for the undertaking of research under each priority over the next 10 years and the funds currently allocated fromthe Pilbara Northern Quoll Offset Trust under each research priority

Rank Theme Research priority Current or proposed researchactivity

Time over next10 years

Current funding allocation(approximate)

1 Survey andmonitoring

Develop appropriate andstandardised survey andmonitoring methods

Develop monitoring protocol(traps and cameras) andinvestigate new methods

Ongoing survey and monitoringin conjunction with ResearchThemes 2 and 3 to better mapnorthern quoll distribution andto determine long-termpopulation trends

Years 1ndash2Years 3ndash10

$200 000 per annum over10 years for ResearchThemes 1 2 and 3(inclusive of salaries)

2 Habitatrequirements

Define areas of criticalhabitat for northernquoll in the Pilbara

Identify and describe criticalhabitat and undertake predictivemodelling of unsurveyed areasto inform further survey effort

Years 3ndash4 $50 000 for predictivemodelling (EdithCowan University)

Understand howdisturbance affectshabitat quality andconnectivity

Ongoing collection of data ondisturbance and habitat qualityin conjunction with ResearchThemes 1 and 4

Incorporation of new data intopredictive models

Years 4ndash10 See above

3 Populationdynamics

Better understand thepopulation dynamics ofthe northern quoll in thePilbara

Collect data on demographicparameters breeding dispersaland home-range use toincorporate into future PVA

Continued collectionof data inconjunction withpriorities 1 and 2during years 3ndash10

See above

Investigate populationstructure and theinteraction betweenpopulations

Analysis of existing tissuesamples and newly collectedsamples

Years 1ndash10 $120 000 for initial roundof genetic analysis(Years 3ndash5 MurdochUniversity)

4 Key threats Introduced predators Spatio-temporal interactionsbetween northern quolls andintroduced predators

Other projects as they arise

Years 2ndash4 (currentPhD projectUniversity ofQueensland)

$12 000 contribution tofield expenses

Cane toads Predictive modelling of canetoad distribution in the Pilbara

Years 3ndash4 Part of predictivemodelling by EdithCowan University

Infrastructuredevelopment andinteractions withhumans

Investigation of impacts oflinear infrastructure on northernquoll movement

Year 4 (currentHonours projectEdith CowanUniversity)

$15 000

Interactions betweenthreats

No on-ground projects currentlyplanned

Potential to be assessed throughPVA as data are collected underother research priorities

No funding currentlyallocated

5 Recolonisation ofrestored orartificial habitat

Determine the ability ofthe northern quoll torecolonise disturbedareas or coloniseartificial habitat

As projects arise Years 5ndash8 incollaboration withminesite restorationprograms

No funding currentlyallocated

J Australian Mammalogy V A Cramer et al

Priority 5 Recolonisation of restored andor artificial habitat

While anecdotal evidence suggests that northern quolls will denin waste rock dumps whether artificial or restored habitatprovides for long-term population viability is not known andlong-term assessment of created habitat is required Weacknowledge that a large gap exists between the desire to userestored or artificial habitat as an ecologically equivalent offsetand our limited ability to actually restore or re-create habitat thatis equivalent in structure composition and function particularlyon highly disturbed sites (Maron et al 2012) Yet there is a futureopportunity in the Pilbara to conduct lsquonaturalrsquo experiments onthe recolonisation of quolls after mining ceases including howto best design waste rock dumps so that habitat complexityand productivity is maximised by for example experimentingwith the size and positioning of boulders and their spatialarrangement in relation to surrounding landscape features Theuse of such artificial habitat should be tested in the first stagethrough direct monitoring of animals that have colonised orhave been translocated to the site Such experimentation isunderway at Atlas Ironrsquos Mt Dove operation (Atlas Iron 2012)Population modelling can then be used to predict futurepopulation sizes rates of growth and spread and the probabilityof long-term persistence in the context of various managementactions (eg predator control Armstrong and Reynolds 2012)

Resource allocation and timelines

While the allocation of resources and funding and the periodover which each of the research priorities was expected to beaddressed was not explicitly discussed during the workshopthe ranking of the priorities was taken to indicate the sequencein which the priorities should be addressed At the time of theworkshop some of the research questions identified werealready being investigated within various projects and a portionof the Pilbara Northern Quoll Offset Trust allocated to these InTable 3 we indicate the expected timelines over which someof the research programs are expected to be undertaken WhileAU$175M has been allocated to northern quoll research in thePilbara it is important to note that much of this amount is notcurrently available and will not become available until severalapproved mine developments have become operational InTable 3 we also indicate the proportion of offset funds expectedto be allocated to each research priority Research undertaken inother regions (eg research into CTA in the Kimberley regionahead of the cane toad invasion front) cannot be funded from thePilbara Northern Quoll Offset Trust but will help address someof the research questions identified here

Lessons learnt from the workshop process

The workshop approach used here was less formal than recentlarge-scale exercises used to elicit research priorities for emergingissues in science and policy at national or global levels (reviewedin Sutherland et al 2011) The focus in our (much smaller)workshop was on inclusivity rather than identifying participantsbased on purposive sampling of expertise or organisationalaffiliation We believe that the inclusive nature of the workshopwas a strength but we also recognise that this perhaps createdsome level of bias when ranking the research prioritiesAcknowledging both the experience and knowledge requirements

of those who work regularly in the Pilbara is an importantcomponent of improving the design of research programs that aimto improve conservation management in the region Yet theresearch priorities and their rankings are likely strongly reflectiveof the current information needs of most participants and maynot align with what conservation scientists believe to be themost critical areas of research for northern quoll conservation intothe future Baseline survey and monitoring are often the firstrequirements of a management or research plan for the northernquoll when offset conditions are prescribed and this was reflectedin the primary ranking given to the development of standardsurvey and monitoring methods We acknowledge the concernthat this could simply lead to offset funding being used to lsquomonitorthe extinctionrsquo of the northern quoll particularly after canetoads invade the Pilbara unless these data are used to triggermanagement actions (Oakwood and Foster 2008 Lindenmayeret al 2013) Clearly the currently limited information on habitatuse and demographic parameters for the northern quoll in thePilbara restricts the utility of more sophisticated researchapproaches such as developing robust SDM and PVA While thesevere threat that cane toads pose to the northern quoll wasrecognised this threat is not immediate in the region and mayexplain the less prominent ranking given to research to mitigatethe impacts of cane toads

The northern quoll has become emblematic of the dramaticand rapid species declines occurring in northern Australia andthe level of interest in this species was reflected in the highnumber of participants attending the workshop (one of six runby Parks and Wildlife) Given the number of attendees betterutilisation of the time available would have been gained byproviding participants with a brief before the workshopand asking them to contribute potential research questionsanonymously for interactive review by other participants Inthe exercises reviewed by Sutherland et al (2011) questionswere then discarded if they were not considered to meet criteriathat were previously agreed upon Some areas of researchthat gained some attention on the day may have been discardedif given greater consideration or interrogation before theworkshop For example there was some discussion duringthe workshop of investigating the use of habitat surrogates tomonitor the northern quoll yet research has shown that there islittle validity in using surrogates to monitor the abundance of asingle species (Pierson et al 2015)

Conclusions

Offset funds generated under the environmental approvalsprocess provide an opportunity to mobilise considerablefinancial resources for conservation (Kiesecker et al 2010) Yetthere remain significant conceptual and practical limitations tobe resolved to ensure that direct offsets provide measurableconservation outcomes that are ecologically equivalent to thelosses sustained by proceeding with the development (Maronet al 2012 Bull et al 2013) The collaborative identificationand ranking of research priorities undertaken here is the firststep in the process of developing and evaluating potentialconservation actions for the northern quoll in the Pilbara andserves as a mechanism to ensure that the research priorities ofstate and federal agencies reflects the knowledge needs of

Research priorities for the northern quoll Australian Mammalogy K

practitioners and proponents currently operating in the regionThe provision of offset funds over the next decade providesa unique opportunity to undertake research on a species ofconservation significance at a regional scale and over a longperiod through an integrated and strategic program of researchrather than the collection of ad hoc data While a morestructured approach to defining research questions before theworkshop may have led to greater critical reflection upon anddiscussion of some of the research priorities on the day theworkshop provided an opportunity for an open and diverse-ranging discussion in a collaborative environment amongstthe broader community of scientists environmental consultantsand mining proponents involved in the management andconservation of the northern quoll in the Pilbara

Acknowledgements

We gratefully acknowledge the financial support of Roy Hill who sponsoredthe workshop and partially funded VACrsquos salary We thank the Departmentof the Environmentrsquos Pilbara Taskforce especially Veronica Ritchie fortheir support of interested or affected parties The success of the workshopwas in great part due to the facilitation skills of Craig Salt of SustainableConsulting We thank all of the workshop participants for their open andcollaborative approach to the process Mike Bamford Eddie van EttenDavid Peacock and Peter Lyon provided useful comments on an earlier draftWe are grateful to John Woinarski for his constructive advice thatsubstantially improved the structure and clarity of the manuscript and ananonymous reviewer for careful and critical reading of the previousversion Lesley Gibson Russell Palmer and Al Glen generously discussedvarious aspects of the paper with VAC The map was prepared by CristinaRamalho

References

Allen B L Engeman R M and Allen L R (2011) Wild dogma anexamination of recent lsquoevidencersquo for dingo regulation of invasivemesopredator release in Australia Current Zoology 57 568ndash583

Allen B L Fleming P J Allen L R Engeman R M Ballard G andLeung L K-P (2013) As clear as mud a critical review of evidencefor the ecological roles of Australian dingoes Biological Conservation159 158ndash174 doi101016jbiocon201212004

Anon (1934) Floods in Western Australia Town isolated for week PerthMonday The Sydney Morning Herald 9

Armstrong D P and Reynolds M H (2012) Modelling reintroducedpopulations the state of the art and future directions In lsquoReintroductionBiology Integrating Science and Managementrsquo (Eds J G Ewen DP Armstrong K A Parker and P J Seddon) pp 165ndash222 (Wiley-Blackwell UK)

Atlas Iron (2012) Scope of works ndash Mount Dove artificial quoll habitatUnpublished report Atlas Iron Perth Available at httpwwwatlasironcomauIRMCompanyShowPageaspxCategoryId=190ampCPID=5424ampEID=94365410 [accessed 19 August 2015]

Austin C (2014) Can remote cameras accurately estimate populations ofthe endangered northern quoll BSc(Honours) Thesis University ofTechnology Sydney

Begg R J (1981) The small mammals of Little Nourlangie RockNT III Ecology ofDasyurus hallucatus the northern quoll (MarsupialiaDasyuridae) Wildlife Research 8 73ndash85 doi101071WR9810073

Braithwaite R W and Griffiths A D (1994) Demographic variationand range contraction in the northern quoll Dasyurus hallucatus(Marsupialia Dasyuridae)Wildlife Research 21 203ndash217 doi101071WR9940203

Bull J W Suttle K B Gordon A Singh N J and Milner-Gulland E J(2013) Biodiversity offsets in theory and practice Oryx 47 369ndash380doi101017S003060531200172X

Burnett S (1997) Colonizing cane toads cause population declines innative predators reliable anecdotal information and managementimplications Pacific Conservation Biology 3 65ndash72

Cadenhead N C R White A L and Wintle B A (2015) Identifyingconservation priorities and assessing impacts of proposed futuredevelopment in the Pilbara Bioregion in Western Australia Report bythe National Environment Research Program Decisions Hub to theDepartment of the Environment The University of MelbourneMelbourne

Carwardine J Nicol S Van Leeuwen S Walters B Firn J Reeson AMartin T G and Chades I (2014) Priority threat management forPilbara species of conservation significance CSIRO EcosystemSciences Brisbane

Cook A (2010) Habitat use and home-range of the northern quollDasyurushallucatus effects of fire MSc Thesis University ofWestern AustraliaPerth

Department of Sustainability Environment Water Population andCommunities (2012) Environment and Biodiversity Conservation Act1999 Environmental Offsets Policy Department of SustainabilityEnvironment Water Population and Communities Canberra

Doherty T S Davis R A Etten E J Algar D Collier N DickmanC R Edwards G Masters P Palmer R and Robinson S (2015a)A continental-scale analysis of feral cat diet in Australia Journal ofBiogeography 42 964ndash975 doi101111jbi12469

Doherty T S Dickman C R Nimmo D G and Ritchie E G (2015b)Multiple threats or multiplying the threats Interactions betweeninvasive predators and other ecological disturbances BiologicalConservation 190 60ndash68 doi101016jbiocon201505013

Dunlop J Cook A and Lees J (2014a) BHP Billiton Iron Ore Pilbaranorthern quoll monitoring project Final report May 2010ndashJune 2013Unpublished report prepared for BHP Billiton Iron Ore Department ofParks and Wildlife Perth

Dunlop J Cook A and Morris K (2014b) Pilbara northern quollproject surveying and monitoring Dasyurus hallucatus in the PilbaraWestern Australia Department of Parks and Wildlife Perth Availableat librarydpawwagovau

Florance D Webb J K Dempster T Kearney M R Worthing A andLetnic M (2011) Excluding access to invasion hubs can contain thespread of an invasive vertebrate Proceedings of the Royal Society ofLondon B Biological Sciences 278 2900ndash2908 doi101098rspb20110032

Gaillard J-M Hebblewhite M Loison A Fuller M Powell R BasilleM and Van Moorter B (2010) Habitatndashperformance relationshipsfinding the right metric at a given spatial scale PhilosophicalTransactions of the Royal Society of London Series B BiologicalSciences 365 2255ndash2265 doi101098rstb20100085

Glen A S and Dickman C R (2008) Niche overlap between marsupialand eutherian carnivores does competition threaten the endangeredspotted-tailed quoll Journal of Applied Ecology 45 700ndash707 doi101111j1365-2664200701449x

Glen A S Pennay M Dickman C R Wintle B A and Firestone K B(2011) Diets of sympatric native and introduced carnivores in theBarrington Tops eastern Australia Austral Ecology 36 290ndash296doi101111j1442-9993201002149x

Hennig P (2004) A brief land use history In lsquoAn Inventory and ConditionSurvey of the Pilbara Region Western Australiarsquo (Eds A M E VanVreeswyk A L Payne K A Leighton and P Hennig) TechnicalBulletin No 92 pp 13ndash18 (Western Australian Department ofAgriculture Perth)

Hohnen R Ashby J Tuft K and McGregor H (2013) Individualidentification of northern quolls (Dasyurus hallucatus) using remotecameras Australian Mammalogy 35 131ndash135 doi101071AM12015

L Australian Mammalogy V A Cramer et al

How R A Spencer P B S and Schmitt L H (2009) Island populationshave high conservation value for northern Australiarsquos top marsupialpredator ahead of a threatening process Journal of Zoology 278206ndash217 doi101111j1469-7998200900569x

Johnson C N Isaac J L and Fisher D O (2007) Rarity of a top predatortriggers continent-wide collapse of mammal prey dingoes andmarsupials in Australia Proceedings of the Royal Society of LondonSeries B Biological Sciences 274 341ndash346 doi101098rspb20063711

Jones M E (2000) Road upgrade road mortality and remedial measuresimpacts on a population of eastern quolls and Tasmanian devilsWildlifeResearch 27 289ndash296 doi101071WR98069

Kearney M Phillips B L Tracy C R Christian K A Betts G andPorter W P (2008) Modelling species distributions without usingspecies distributions the cane toad in Australia under current andfuture climates Ecography 31 423ndash434 doi101111j0906-7590200805457x

Kendrick P and Stanley F (2002) Pilbara 4 (PIL4 ndash Roebourne synopsis)In lsquoA Biodiversity Audit of Western Australiarsquos 53 BiogeographicalSubregions in 2002rsquo (Eds J E May and N L McKenzie) pp 581ndash593(Department of Conservation and Land Management Perth)

Kennedy M Phillips B L Legge S Murphy S A and Faulkner R A(2012) Do dingoes suppress the activity of feral cats in northernAustralia Austral Ecology 37 134ndash139 doi101111j1442-9993201102256x

Kiesecker J M Copeland H Pocewicz A and McKenney B (2010)Development by design blending landscape-level planning with themitigation hierarchy Frontiers in Ecology and the Environment 8261ndash266 doi101890090005

King D R and Smith L A (1985) The distribution of the European red fox(Vulpes vulpes) in Western Australia Records of the Western AustralianMuseum 12 197ndash205

Knight A T Cowling R M Rouget M Balmford A Lombard A Tand Campbell B M (2008) Knowing but not doing selecting priorityconservation areas and the researchndashimplementation gap ConservationBiology 22 610ndash617 doi101111j1523-1739200800914x

Laurance W F Koster H Grooten M Anderson A B Zuidema P AZwick S Zagt R J Lynam A J Linkie M and Anten N P (2012)Making conservation research more relevant for conservationpractitioners Biological Conservation 153 164ndash168 doi101016jbiocon201205012

Legge S Murphy S Heathcote J Flaxman E Augusteyn J andCrossman M (2008) The short-term effects of an extensive and high-intensity fire on vertebrates in the tropical savannas of the centralKimberley northern Australia Wildlife Research 35 33ndash43 doi101071WR07016

Lester D (2015) Effective wildlife roadkill mitigation Journal of Trafficand Transportation Engineering 3 42ndash51 doi10172652328-2142201501005

Letnic M Tamayo B and Dickman C R (2005) The responsesof mammals to La Nintildea (El Nintildeo Southern Oscillation)-associatedrainfall predation and wildfire in central Australia Journal ofMammalogy 86 689ndash703 doi1016441545-1542(2005)086[0689TROMTL]20CO2

Letnic M Koch F Gordon C Crowther M S and Dickman C R(2009) Keystone effects of an alien top-predator stem extinctions ofnative mammals Proceedings of the Royal Society of London Series BBiological Sciences 276 3249ndash3256 doi101098rspb20090574

Lindenmayer D B (2015) Continental-level biodiversity collapseProceedings of the National Academy of Sciences of the United States ofAmerica 112 4514ndash4515 doi101073pnas1502766112

Lindenmayer D B Piggott M P and Wintle B A (2013) Counting thebooks while the library burns why conservation monitoring programs

need a plan for action Frontiers in Ecology and the Environment 11549ndash555 doi101890120220

Maron M Hobbs R J Moilanen A Matthews J W Christie KGardner T A Keith D A Lindenmayer D B and McAlpine C A(2012) Faustian bargains Restoration realities in the context ofbiodiversity offset policies Biological Conservation 155 141ndash148doi101016jbiocon201206003

McCall S C McCarthy M A van der Ree R Harper M J Cesarini Sand Soanes K (2010) Evidence that a highway reduces apparentsurvival rates of squirrel gliders Ecology and Society 15 27

McGregor R A Stokes V L and Craig M D (2014) Does forestrestoration in fragmented landscapes provide habitat for a wide-ranging carnivore Animal Conservation 17 467ndash475 doi101111acv12112

McKenzie N L Burbidge A A Baynes A Brereton R N DickmanC R Gordon G Gibson L A Menkhorst P W Robinson A CWilliams M R and Woinarski J C Z (2007) Analysis of factorsimplicated in the recent decline of Australiarsquos mammal fauna Journal ofBiogeography 34 597ndash611 doi101111j1365-2699200601639x

McKenzie N L van Leeuwen S and Pinder AM (2009) Introduction tothe Pilbara Biodiversity Survey 2002ndash2007 Records of the WesternAustralian Museum 78(Suppl) 3ndash89 doi1018195issn0313-122x78(1)2009003-089

Morris K Johnson B Orell P Gaikhorst G Wane A and MoroD (2003) Recovery of the threatened chuditch (Dasyurus geoffroii)a case study In lsquoPredators with Pouches The Biology of CarnivorousMarsupialsrsquo (Eds M Jones C Dickman and M Archer) pp 435ndash451(CSIRO Publishing Melbourne)

Nelson J L Scroggie M P and Belcher C A (2014) Developing acamera trap survey protocol to detect a rare marsupial carnivore thespotted-tailed quoll (Dasyurus maculatus) In lsquoCamera TrappingWildlife Management and Researchrsquo (Eds P D Meek and PJ Fleming) pp 271ndash279 (CSIRO Publishing Melbourne)

Newsome T M Dellinger J A Pavey C R Ripple W J Shores C RWirsing A J and Dickman C R (2015) The ecological effects ofproviding resource subsidies to predators Global Ecology andBiogeography 24 1ndash11 doi101111geb12236

OrsquoDonnell S Webb J K and Shine R (2010) Conditioned tasteaversion enhances the survival of an endangered predator imperilled bya toxic invader Journal of Applied Ecology 47 558ndash565 doi101111j1365-2664201001802x

Oakwood M (1997) The ecology of the northern quoll Dasyurushallucatus PhD Thesis Australian National University Canberra

Oakwood M (2000) Reproduction and demography of the northern quollDasyurus hallucatus in the lowland savanna of northern AustraliaAustralian Journal of Zoology 48 519ndash539 doi101071ZO00028

Oakwood M (2002) Spatial and social organization of a carnivorousmarsupial Dasyurus hallucatus (Marsupialia Dasyuridae) Journal ofZoology 257 237ndash248 doi101017S0952836902000833

Oakwood M and Foster P (2008) Monitoring extinction of the northernquoll Australian Academy of Science Newsletter 71 6

Palomares F and Caro T M (1999) Interspecific killing amongmammalian carnivores American Naturalist 153 492ndash508 doi101086303189

Pavey C R and Bastin G (2014) lsquoAustralian Rangelands and ClimateChange ndash Invasive Animalsrsquo (Ninti One Limited amp CSIRO AliceSprings)

Pierson J C Barton P S Lane PW and Lindenmayer D B (2015) Canhabitat surrogates predict the response of target species to landscapechange Biological Conservation 184 1ndash10 doi101016jbiocon201412017

Pollock A B (1999) Notes on status distribution and diet of northernquoll Dasyurus hallucatus in the MackayndashBowen area mideastern

Research priorities for the northern quoll Australian Mammalogy M

Queensland Australian Zoologist 31 388ndash395 doi107882AZ1999040

Ramanaidou E R and Morris R C (2010) A synopsis of the channeliron deposits of the Hamersley Province Western AustraliaTransactions of the Institutions of Mining and Metallurgy Section BApplied Earth Science 119 56ndash59 doi101179037174510X12853354810624

Schmitt L H Bradley A J Kemper C M Kitchener D J HumphreysW F and How R A (1989) Ecology and physiology of the northernquoll Dasyurus hallucatus (Marsupialia Dasyuridae) at MitchellPlateau Kimberley Western Australia Journal of Zoology 217539ndash558 doi101111j1469-79981989tb02510x

Short J and Smith A (1994) Mammal decline and recovery in AustraliaJournal of Mammalogy 75 288ndash297 doi1023071382547

Spencer P How R Hillyer M Cook A Morris K Stevenson C andUmbrello L (2013) Genetic analysis of northern quolls from the PilbaraRegion ofWestern Australia Year one ndash final report Unpublished reportDepartment of Sustainability Environment Water Population andCommunities Canberra and Department of Parks and Wildlife Perth

Sutherland W J Fleishman E Mascia M B Pretty J and Rudd M A(2011) Methods for collaboratively identifying research priorities andemerging issues in science and policyMethods in Ecology and Evolution2 238ndash247 doi101111j2041-210X201000083x

Sutherst R W Floyd R B and Maywald G F (1996) The potentialgeographical distribution of the cane toad Bufo marinus L in AustraliaConservation Biology 10 294ndash299 doi101046j1523-1739199610010294x

Taylor B D and Goldingay R L (2003) Cutting the carnage wildlifeusage of road culverts in north-eastern New South Wales WildlifeResearch 30 529ndash537 doi101071WR01062

Tingley R Phillips B L Letnic M Brown G P Shine R and BairdS J (2013) Identifying optimal barriers to halt the invasion of canetoads Rhinella marina in arid Australia Journal of Applied Ecology 50129ndash137 doi1011111365-266412021

Van Vreeswyk A M E Payne A L Leighton K A and HennigP (2004) An Inventory and Condition Survey of the Pilbara RegionWestern Australia Technical Bulletin No 92 Department of AgricultureWestern Australia Perth

Webb J Legge S Tuft K Cremona T and Austin C (2015) Can wemitigate cane toad impacts on northern quolls Final report National

Environmental Research Program Northern Australia Hub CharlesDarwin University Darwin

Wintle B A Walshe T V Parris K M and McCarthy M A (2012)Designing occupancy surveys and interpreting non-detection whenobservations are imperfect Diversity amp Distributions 18 417ndash424doi101111j1472-4642201100874x

Woinarski J Menkhorst K Gambold N and Braithwaite R W (1992)Mammals of the Bungle Bungle area In lsquoA Survey of the Wildlife andVegetation of Purnululu (Bungle Bungle) National Park and AdjacentArearsquo (Ed J C Z Woinarski) Research Bulletin No 6 pp 53ndash67(CSIRO Division of Wildlife and Ecology Darwin)

Woinarski J C Z Risler J and Kean L (2004) Response of vegetationand vertebrate fauna to 23 years of fire exclusion in a tropical Eucalyptusopen forest Northern Territory Australia Austral Ecology 29 156ndash176doi101111j1442-9993200401333x

Woinarski J C Z Oakwood M Winter J Burnett S Milne D FosterP Myles H and Holmes B (2008) Surviving the toads patterns ofpersistence of the northern quoll Dasyurus hallucatus in QueenslandReport submitted to the Natural Heritage Trust Strategic ReserveProgram as a component of project 2005162 Monitoring andManagement of Cane Toad Impact in theNorthern Territory Available athttplrmntgovauplants-and-animalsinformation-and-publicationspublications2008 [accessed 14 November 2013]

Woinarski J C Z Armstrong M Brennan K Fisher A Griffiths A DHill B Milne D J Palmer C Ward S Watson M Winderlich Sand Young S (2010) Monitoring indicates rapid and severe decline ofnative small mammals in Kakadu National Park northern AustraliaWildlife Research 37 116ndash126 doi101071WR09125

Woinarski J Burbidge A and Harrison P (2014) lsquoAction Plan forAustralian Mammals 2012rsquo (CSIRO Publishing Melbourne)

Woinarski J C Burbidge A A and Harrison P L (2015) Ongoingunraveling of a continental fauna decline and extinction of Australianmammals since European settlement Proceedings of the NationalAcademy of Sciences of the United States of America 112 4531ndash4540doi101073pnas1417301112

Woolley P A Krajewski C and Westerman M (2015) Phylogeneticrelationships within Dasyurus (Dasyuromorphia Dasyuridae) quollsystematics based on molecular evidence and male characteristicsJournal of Mammalogy 96 37ndash46 doi101093jmammalgyu028

N Australian Mammalogy V A Cramer et al

wwwpublishcsiroaujournalsam

Page 4: Research priorities for the northern quoll (Dasyurus ......The Pilbara population of the northern quoll (Dasyurus hallucatus) has been seldom studied, and the impacts of threats such

Tab

le1

Researchpriorities

fortheno

rthern

quollan

dkeyqu

estion

sassociated

witheach

priorityan

das

compa

redwiththeinform

ationrequ

ired

undersimila

rthem

esan

dthepriority

givento

thisin

Woina

rski

etal(201

4)

Rank

Theme

Researchpriority

Key

questio

nsInform

ationrequired

andpriority

givenundersimilarthem

ein

Woinarski

etal(2014)

1Surveyand

mon

itoring

Develop

appropriate

andstandardised

survey

and

mon

itoring

metho

ds

Whatistheefficacy

ofthemajor

survey

metho

dscurrently

used

forno

rthern

quollsin

the

Pilb

ara

How

dovariou

scombinatio

nsoftrap

typesarraysdurationoftrappingbaitsetc

compareWhich

camera-trap

protocol

ismosteffectivefor(a)identifying

individu

alno

rthern

quollsin

thePilb

araand(b)estim

atingpo

pulatio

nsizes

Which

methodismost

cost-effectiv

eandefficientin

variou

sPilb

araland

scapes

Whatisthedetectionprobability

ofno

rthern

quollsin

each

habitattypeBased

onprior

surveywhatitthepriorprobability

ofoccupancyin

thesehabitattypes

Based

onthis

know

ledge

how

manynon-detections

ofquollsarerequired

before

wecanstatewith

prespecified

confi

dencethat

northern

quollsareabsent

from

asite

Surveyto

betterdefine

fine-scale

distributio

nandnumberof

individu

alsin

subpop

ulations

mediumndashhigh

Establishor

enhancemon

itoring

program

bydesigningintegrated

mon

itoring

prog

ramsmedium

2Habitat

requirem

ents

Define

areasof

critical

habitatforno

rthern

quollin

thePilb

ara

How

donorthern

quollsusehabitatinthePilb

ara

How

arequollpopulations

distributedin

relatio

nto

habitatq

uality(egh

ighqu

ality

low

quality

orsink

habitats)

How

ishabitat

selectioninfluenced

bypredationrisk

andsocial

factors

Whatinformationisrequ

ired

tofurtherrefine

anSDM

forno

rthern

quollsin

thePilb

ara

How

canthisinform

ationbe

mosteffectivelyused

toim

provemonito

ring

effortand

outcom

es

How

domod

elsvary

across

thePilb

aramainlandandoffsho

reislandsaccountin

gfor

clim

atic

gradientsandtopography

Whatarefuture

predictio

nsfornorthern

quollpopulatio

ndistributio

nsu

nder

modelsof

clim

atechange

andcane

toad

invasion

Assesshabitatrequirem

entsna

Und

erstandho

wdisturbanceaffects

habitatqu

ality

and

conn

ectiv

ity

How

does

fire

andgrazingaffecthabitatq

ualityandhabitatu

seby

northern

quollsin

the

Pilb

ara

How

importantisfire

asabarrierto

dispersal

How

importantishabitatconnectivity

form

etapopulations

ofnorthernquollinthePilb

ara

How

does

habitattypeaffecttheconnectiv

ityof

metapopulationsHow

does

disturbance

both

discrete

(eginfrastructure

such

asrailandroads)

anddiffuse(egchangesin

vegetatio

ncompositio

nandstructure)alterhabitatconnectiv

ity

Assessrelativ

eim

pact

ofthreats

quantifyrelativ

eim

pact

offire

regimesmdashhigh

Assesseffectivenessof

threat

mitigatio

nop

tionsland

scape-scale

optio

nsforretentionof

longer-

unburntpatchesmdash

high

3

Pop

ulation

dynamics

Betterunderstand

the

popu

latio

ndy

namicsof

the

northern

quollinthe

Pilb

ara

How

manysubpop

ulations

aretherein

thePilb

ara

How

dotheseinteractAre

subpopulations

demographically

stablein

spaceandtim

eHow

does

habitatand

clim

ate

affect

popu

latio

ndy

namics

How

does

disturbanceaffect

demog

raphic

stability

Assessdietlifehistory

na

Investigatepo

pulatio

nstructureandthe

interactionbetween

popu

latio

ns

How

domeasuresofgenetic

distance

relatetospatialecology

asmeasuredby

otherm

eans

iedoes

inferred

genetic

connectiv

ityrelate

toobserved

dispersal

Can

genetic

inform

ationbe

used

todeterm

inehabitatbarriers

Resolve

taxonomic

uncertainties

assess

genetic

relatednessof

spatially

disparatepo

pulatio

nsto

identifywhether

thereare

genetically

distinct

subpop

ulationsmdashlowndashmedium

D Australian Mammalogy V A Cramer et al

4Key

threats

Introduced

predators

Doespredationsignificantly

affecttheno

rthern

quollinthePilb

ara

How

does

thisvary

across

habitattypes

Doescompetitionforresourceswith

foxesor

catsim

pact

onsubpopulationsWhich

lifestages

aremostvulnerable

Given

thattheno

rthern

quollisalso

amesop

redatordo

esthepresence

ofding

oeseither

directly(through

predation)

orindirectly(through

theregulatio

nof

foxesandcats)impact

onsubpop

ulations

inthePilb

ara

Assessrelativ

eim

pact

ofthreats

predationby

feralcatsanddogsmdash

high

interactions

betweenferal

catsdo

gsdingoes

andfoxesmdash

mediumndashhigh

Assesseffectivenessof

threat

mitigatio

nop

tionsassess

which

feasible

managem

entoptio

nsfor

feralcatsanddo

gscanlead

topo

pulatio

nrecoverymdashmediumndash

high

Develop

new

orenhanceexistin

gmanagem

entmechanism

sfor

broad-scale

targetedferalcat

controlmdash

medium

Canetoads

Will

theclosureof

artifi

cialwater

bodies

preventor

simplyslow

therateofinv

asionof

cane

toadsinto

thePilb

ara

Whatisthedegree

ofoverlapbetweenSDMfundamentalniche

modelsforn

orthernquolls

andcane

toadsin

thePilb

ara

Can

potentialrefug

eareasforno

rthern

quollb

eidentifi

ed

IsCTAlik

elytobe

effectivewhentaste-aversion

baits

aredeliv

ered

underfi

eldcond

itions

Atwhatscaleisbaitdeliv

eryfeasible

Assesseffectivenessof

threat

mitigatio

nop

tionsassess

feasibility

ofbroad-scaletaste-

aversion

lsquotrainingrsquo

ofqu

ollsmdash

mediumassess

local-scale

exclusionof

toadsor

preventio

nof

colonisatio

n(m

ainlandsites)mdash

medium

Infrastructure

developm

entand

interactions

with

humans

Are

road-related

deaths

ofno

rthern

quollasign

ificant

causeof

mortalityin

thePilb

ara

Dorailw

aylin

esinfrastructurecorridorsor

wideroadspresenta

barrierto

northern

quoll

mov

ement

Whatmethods

canbe

used

tomitigate

theim

pactsof

infrastructure

developm

entand

associated

machinery

operation

Not

addressed

Interactions

between

threats

Dofrequent

andintensefireslead

toreducedstructural

complexity

loss

ofvegetatio

ncoverandsubsequentincreasedpredationof

northern

quollsparticularlyof

males

during

andor

afterbreeding

Whatarethecumulativeim

pactsof

mortalitycaused

bycane

toad

ingestionandpredation

bycatsdo

gsdingoes

andfoxes

See

inform

ationon

introd

uced

predatorsandcane

toadsabove

5Recolon

isationof

restored

orartifi

cial

habitat

Determinetheability

oftheno

rthern

quoll

torecolonise

disturbedareasor

colonise

artifi

cial

habitat

Whatarethekeyelem

entsof

northern

quollhabitatthatneedtobe

restoredcreated

sothat

habitatcomplexity

andprod

uctiv

ityismaxim

ised

Whatlevelo

flandscapeconn

ectiv

ityisrequired

arou

ndrestoredartificialhabitatso

that

genetic

connectiv

itybetweensubpopulations

canbe

maintained

How

long

should

restored

orartifi

cial

habitatbe

directly

managedparticularly

for

predators

How

cantheretentionof

largebouldersof

wasterock

beincorporated

into

rehabilitation

plans

Not

addressed

Research priorities for the northern quoll Australian Mammalogy E

investment of time and resources The limited amount anduncertain quality of occurrence data has been a major restrictionon the utility of a species distribution model (SDM) and apopulation viability analysis (PVA) for the northern quoll in thePilbara (Cadenhead et al 2015 see below) While survey andmonitoring per se is not a research activity that directly informsmanagement or threat mitigation carefully implementedmonitoring programs are essential to identify many elements ofa speciesrsquo ecology to detect changes in population and toquantify the ecological effectiveness and cost-effectivenessof management interventions (Lindenmayer et al 2013Lindenmayer 2015)

Parks and Wildlife recently released revised protocols forthe long-term monitoring of known populations of thenorthern quoll (Dunlop et al 2014b) and the effectiveness ofthese protocols in presenceabsence surveys should be testedThe cost-effectiveness of using camera traps for wildlifesurveys in remote locations such as the Pilbara makes their useappealing Hohnen et al (2013) described how the strategicplacement of cameras and baits combined with taking a greaternumber of photographs per trigger allows the capture ofimages that can be used to identify individual northern quollsfrom their spot patterns Remote cameras have also been shownto be a viable method for estimating population sizes of thenorthern quoll in areas with both dense (Austin 2014) andsmall populations (Webb et al 2015) Further testing of a rangeof parameters (eg see Nelson et al 2014) to develop a robuststandard protocol for the Pilbara is required along with acomparison of the cost-effectiveness and any ethicalreducedhandling benefits of using remote cameras versus live-trappingfor estimating population sizes in Pilbara landscapes

Priority 2 Improve our understanding of habitatrequirements

Define areas of critical habitat for the northern quollin the Pilbara

The Department of Agriculture and Food Western Australiahas mapped 102 land systems across the Pilbara bioregionbased on several biophysical components including landformgeology soils vegetation and drainage patterns (Van Vreeswyket al 2004) A review of records for the northern quoll in thePilbara showed that 65 of records were from the RobeCapricorn and Boolaloo land systems (Table 2) which compriselow plateaux mesas hills and ridges of varying geology(limonite granite and sandstone respectively) Most recentsurvey effort in the Pilbara is biased towards mineral-bearingformations or areas of mining-related infrastructure (eg railand road corridors) (Fig 1) Further investigation of theserecords is required to assess whether these land systems areindeed preferred habitat for northern quolls as opposed to anartefact of survey effort The comparatively fewer records thatcoincide with the conservation estate in the region (Fig 1) arealso likely an artefact of survey effort

The first step in defining areas of critical habitat for thenorthern quoll in the Pilbara is to improve our understanding ofhabitat use particularly in terms of defining habitat suitability atfiner scales than is currently available As part of the strategic

environmental assessments being undertaken for the Pilbarabioregion an SDM has been developed for the northern quoll(Cadenhead et al 2015) Uncertainty about the quality of theoccurrence data available the limited amount of data availableas well as the bias towards data collection in areas proposed fordevelopment constrained the level of certainty around theoutcomes of the model Improved model certainty requiresfurther collection of contemporary species-occurrence data andthe development of finer-resolution GIS layers particularly onethat captures finer-scale geological attributes such as mesas orrocky outcrops considered as highest-quality habitat for thenorthern quoll (Cadenhead et al 2015)

Better definition of the habitat attributes favoured by thenorthern quoll will allow further refinement of the GIS variablesincluded in the SDM or allow the development of predictivesite-based habitat models Information on how northern quollsuse habitat can be gained through direct observation of howthey interact with habitat features or from the association ofhabitat features with the presence of quolls as determined byGPS receivers (Gaillard et al 2010) This level of detail iscurrently not available from survey records returned as part ofenvironmental impact assessments but collection of habitatdata using the protocols outlined by Dunlop et al (2014b) infuture surveys would rapidly increase the quality of dataavailable

Understand how disturbance affects habitat qualityand connectivity

The alteration of fire regimes has been the most pervasivelandscape-scale change in the Pilbara Between 1993 and 2006over 72 of the region was burnt with more than 28 of thearea burnt two or more times (McKenzie et al 2009) Woinarskiet al (2004) found that northern quolls may be able to tolerate amoderately high frequency of fire as long as these fires werelow-intensity early-season burns and the impacts of fire onvegetation structure and composition were not exacerbatedby the grazing of introduced herbivores In the Pilbarahowever as with many other grazing regions of Australia earlypastoralists often overestimated the productivity and resilienceof native vegetation to sheep and cattle grazing which led tosevere degradation of vegetation particularly along riverfrontages (Hennig 2004) The soil degradation and loss ofvegetative cover caused by intense grazing pressure in thesouth Kimberley is thought likely to have contributed to thesubstantial decline or disappearance of medium-sized animalsincluding the northern quoll from the Bungle Bungle area(Woinarski et al 1992) Death by predation is most likely in openareas (Oakwood 2000) and therefore the effects of fire andgrazing on northern quoll populations cannot be consideredwithout also taking account of likely increases in mortality due topredation (see lsquoIntroduced predatorsrsquo below)

Understanding how the localised but intensive activityassociated with mining affects habitat quality and habitat useby the northern quoll in areas surrounding development islimited A recent study that examined the use of ballast quarriesand paired undisturbed sites by northern quolls found that theywere foraging denning and showing signs of breeding at thequarry sites where greater numbers of quolls were captured(Dunlop et al 2014a) This was likely due to the combination of

F Australian Mammalogy V A Cramer et al

habitat complexity (eg diversity of boulder sizes) and thepresence of permanent water in the quarry which may haveincreased prey availability at these sites

Movement of animals through the landscape for dispersaland repopulation is another important consideration for a speciesin which both sexes tend towards semelparity and is thereforepredisposed to local extinction (Oakwood 1997) Improvedunderstanding of how the northern quoll (particularly dispersingmales) uses areas of lower habitat quality and the identificationof such habitat in habitat-suitability models will allow futurelandscape-scale strategic assessments to better predict thecombined impacts of disturbance on the ability of northern quolls

to move through the landscape Analysis of regional populationgenetics will also improve our understanding of the level ofdispersal occurring across the landscape

Priority 3 Population dynamics

Better understand the population dynamics of thenorthern quoll in the Pilbara

Little information is available on the number ofsubpopulations of northern quoll in the Pilbara how thesesubpopulations interact how stable they are in space and timeand how they respond to disturbance As no demographic

Table 2 Records of the northern quoll in each land system type in the PilbaraRecords were current as of 23 May 2014

Land system Area (km2) No of records Land system description

Augustus 61 1 Rugged ranges hills ridges and plateaux supporting mulga shrublands and hard spinifex grasslandsBoolaloo 1502 245 Granite hills domes and tor fields and sandy plains with shrubby spinifex grasslandsBoolgeeda 7748 29 Stony lower slopes and plains below hill systems supporting hard and soft spinifex grasslands andmulga

shrublandsCalcrete 1444 2 Low calcrete platforms and plains supporting shrubby hard spinifex grasslandsCallawa 1003 9 Highly dissected low hills mesas and gravelly plains of sandstone and conglomerate supporting soft and

hard spinifex grasslandsCapricorn 5296 257 Hills and ridges of sandstone and dolomite shrubby hard and soft spinifex grasslandsCoongimah 3244 12 Plateau surfaces low hills with steep slopes and undulating uplands supporting hard spinifex grasslandsDune 138 1 Dune fields supporting soft spinifex grasslandsGranitic 4020 1 Rugged granitic hills supporting shrubby hard and soft spinifex grasslandsHorseflat 1261 4 Gilgaied clay plains supporting tussock grasslands and minor grassy snakewood shrublandsLittoral 1577 3 Bare coastal mudflats with mangroves on seaward fringes samphire flats sandy islands coastal dunes

and beachesMacroy 13 095 94 Stony plains and occasional tor fields based on granite supporting hard and soft spinifex grasslandsMcKay 4202 21 Hills ridges plateau remnants and breakaways of metasedimentary rocks supporting hard spinifex

grasslandsMallina 2557 8 Sandy surfaced alluvial plains supporting soft spinifex (and occasionally hard spinifex) grasslandsMarandoo 459 2 Basalt hills and restricted stony plains supporting grassy mulga shrublandsMosquito 1840 3 Stony plains and prominent ridges of schist and other metamorphic rocks supporting hard spinifex

grasslandsNewman 14 580 81 Rugged jaspilite plateaux ridges and mountains supporting hard spinifex grasslandsOnslow 424 1 Sandplains dunes and clay plains supporting soft spinifex grasslands and minor tussock grasslandsParaburdoo 565 3 Basalt-derived stony gilgai plains and stony plains supporting snakewood and mulga shrublands with

spinifex and tussock grasslandsParadise 1479 8 Alluvial plains supporting soft spinifex grasslands and tussock grasslandsRiver 4088 37 Active flood plains major rivers supporting grassy eucalypt woodlands tussock grasslands and soft

spinifex grasslandsRobe 865 369 Low limonite mesas and buttes supporting soft spinifex (and occasionally hard spinifex) grasslandsRocklea 22 993 40 Basalt hills plateaux lower slopes and minor stony plains supporting hard spinifex (and occasionally

soft spinifex) grasslandsRuth 346 36 Hills and ridges of volcanic and other rocks supporting hard spinifex (occasionally soft spinifex)

grasslandsStuart 1794 3 Gently undulating stony plains supporting hard and soft spinifex grassland and snakewood shrublandsTaylor 129 3 Stony plains and isolated low hills of sedimentary rocks supporting hard and soft spinifex grasslandsTalga 2124 9 Hills and ridges of greenstone and chert and stony plains supporting hard and soft spinifex grasslandsTuree 581 2 Stony alluvial plains with gilgaied and non-gilgaied surfaces supporting tussock grasslands and grassy

shrublandsUaroo 7681 37 Broad sandy plains supporting shrubby hard and soft spinifex grasslandsUrandy 1311 6 Stony plains alluvial plains and drainage lines supporting shrubby soft spinifex grasslandsWhite Springs 266 4 Stony gilgai plains supporting tussock grasslands and hard spinifex grasslandsWona 1815 10 Basalt upland gilgai plains supporting tussock grasslands and minor hard spinifex grasslandsYamerina 1207 1 Floodplains and deltaic deposits supporting tussock grasslands grassy woodlands and minor halophytic

low shrublands

Research priorities for the northern quoll Australian Mammalogy G

studies have been undertaken in the region Cadenhead et al(2015) used demographic information from the NorthernTerritory and Kimberley populations in their PVA Uncertaintyabout data quality and the limited information available ondemographic parameters for the Pilbara constrained the level ofcertainty around the outcomes of the PVA The model doeshowever provide a framework into which new data can beincorporated as they become available For example preliminarydata from the Pilbara monitoring program suggest that theproportion of male die-off in the Pilbara population is lowerthan in the Kimberley and Northern Territory populations(Dunlop unpubl data) Cadenhead et al (2015) stress the needfor more demographic and occurrence data to be collectedbefore further development decisions are made (see lsquoDevelopappropriate and standardised survey and monitoring methodsrsquoand lsquoImprove our understanding of habitat requirementsrsquo above)

Investigate population structure and the interactionbetween populations

Analysis of the genetic diversity and population structure ofthe Pilbara population of the northern quoll is ongoingOnly small differences in genetic diversity as measured byheterozygosity were found between samples obtained at 16locations within the Pilbara with the exception of the isolatedpopulation on Dolphin Island (Spencer et al 2013) This resultindicates a high level of movement across the landscape byindividuals during mating and dispersal phases Spatial structureanalysis of samples collected from around Pannawonicasuggested a lsquoneighbour sizersquo of ~15 km (Spencer et al 2013)Current data show no evidence of genetic bottleneckssuggesting that if there has been a population decline in thePilbara that this has not affected genetic diversity A usefulapproach would be to combine genetic information with datafrom radio-tracking or GPS collars to better understand theuse of lsquolandscapersquo space and how this influences populationdynamics

Priority 4 Key threats and the interaction of these threats

Introduced predators

Introduced cats and foxes are considered to be one of themain causes of the decline and extinction of the critical-weight-range mammals (35 g to 55 kg) in Australia (Short and Smith1994 McKenzie et al 2007 Woinarski et al 2015) but theirimpacts on the northern quoll are less clear A recent review offeral cat diets across Australia found that northern quolls werepresent in the diet of feral cats (Doherty et al 2015a) andWoinarski et al (2014) consider the threat posed by cats to thenorthern quoll to be severe although the impacts of cats may bereduced in rugged refuge areas The impacts of cats on othermuch larger species of quoll appear to be inconsistent withsome evidence of coexistence between cats and quolls beforethe arrival of foxes (Glen and Dickman 2008 and referencescontained within) Due to the limited overlap in their distributionthroughout Australia the threat posed by foxes to northernquolls is less understood than for cats and Woinarski et al(2014) consider the threat posed by foxes to be moderate on anational level Foxes are common in coastal near-coastal and

riparian areas of the Pilbara and extend as far north as Broome(King and Smith 1985 Kendrick and Stanley 2002) They arerenowned for their aggression towards smaller carnivores(Palomares and Caro 1999) and neither the larger chuditch(Dasyurus geoffroii) nor spotted-tail quoll (Dasyurus maculatus)are known to coexist with large numbers of foxes (A Glen perscomm) The interactions between quolls cats and foxesare potentially more complex than for other medium-sizedmarsupials due to niche overlap in diet and competition for densites with the larger spotted-tailed quoll and chuditch bothexperiencing exploitation and interference competition fromfoxes (Morris et al 2003 Glen et al 2011) Competitionbetween feral cats and the northern quoll may potentially beintense during periods of low rainfall in the Pilbara when preyavailability is low Understanding whether foxes and cats aresignificant predators of or competitors with the northern quollin the Pilbara and how this affects their population viabilitywill clarify the level of risk posed by these predators Thesausage-style baits used for feral cats (eg Eradicatreg andCuriosityreg) are potentially attractive to the northern quoll withthe risk of mortality or reduced fitness Trials to assess themagnitude of the risks from Eradicatreg baits are currentlyunderway in the Pilbara

The role of dingoes and wild dogs in the mortality of andinteractions between mesopredators (both native and introduced)requires further investigation Dingoes are known to kill northernquolls butmay not eat these kills (Oakwood 1997) A recent studyin the Northern Territory found that camp dogs and dingoes werethe major source of mortality for northern quolls at the study site(Webb et al 2015) and roaming domestic dogs were identified asa cause of mortality for northern quolls in Queensland (Pollock1999) Human resource subsidies (eg stock carrion in rangelandareas) can artificially support predator populations thus indirectlyincreasing predation pressure on prey (Newsome et al 2015)A dynamic debate currently exists around the role of the dingo inregulating populations of feral cats and foxes and the implicationsof this for the conservation of the critical-weight-range mammalsin Australia (see Johnson et al 2007 Letnic et al 2009 Allenet al 2011 2013 Kennedy et al 2012) Understanding thepopulation density at which the role of the dingo as a regulator ofintroduced predators outweighs its own impact as a direct predatorof the northern quoll will inform whether dingowild dog controlwould benefit the northern quoll

Cane toads

The introduced cane toad now occupies more than half thehistoric range of the northern quoll and has caused an immediatecollapse or extirpation of northern quoll populations in theareas it has invaded (Burnett 1997 Oakwood and Foster2008 Woinarski et al 2008 2010) Earlier modelling work thatconsidered only climate parameters as well as more recentwork that incorporates future climate change suggest thatcane toads will not invade the Pilbara (Sutherst et al 1996Pavey and Bastin 2014) Other recent work that considers thefundamental niche of the cane toad and incorporates the locationof artificial watering points predicts that they will colonisethe Pilbara from the Kimberley via the coastal edge of theGreat Sandy Desert (Kearney et al 2008 Florance et al 2011

H Australian Mammalogy V A Cramer et al

Tingley et al 2013) This has created some level of uncertaintyabout the likely impact of cane toads on northern quolls in thePilbara Strategies for cane toad management were not rankedhighly by participants in the threat-management frameworkdeveloped by Carwardine et al (2014) which the authorsexplained as possibly due to uncertainty about how far canetoads will spread into the region

The model developed by Tingley et al (2013) suggested thatthe closure of 100 artificial water bodies at one of three potentiallocations along the coastal corridor between the Kimberley andthe Pilbara would prevent the invasion of cane toads into thePilbara While their model did account for uncharacteristicallywet years every three decades the dispersal of toads betweenwater bodies was determined by the number of rainy daysoccurring during the year (ie toads could only disperse betweenwater bodies for three days following rainfall) rather thanrainfall amount the associated likelihood of flooding and theavailability of free water across the landscape The cyclonic andthunderstorm nature of much of the rainfall in the north-west ofWestern Australia means that the region is prone to significantand sometimes extended flooding In April 1934 the countryaround Anna Plains (250 km south of Broome) was flooded forhundreds of miles when 27 inches (686mm) of rain fell infour days (Anon 1934) In March 2004 over 350mm of rain wasrecorded within a 48-h period at Telfer on the southern edge ofthe Great Sandy Desert due to Tropical Cyclone Fay Thiscaused the flooding of the De Grey and Oakover river systemsand flooded 140 km of the Telfer Access Road with floodwaterstaking several months to subside (wwwbomgovau) Extendingthe model of Tingley et al (2013) to account for such events willhelp clarify whether closing artificial water bodies is likely toprevent the invasion of cane toads into the Pilbara or may onlyassist in delaying their inevitable invasion

Comparison of SDMs (eg Cadenhead et al 2015) fornorthern quoll with fundamental niche models for cane toads(eg see Kearney et al 2008) within the Pilbara may identifypotential refuge areas for the northern quoll and these areasshould be prioritised for conservation actions Because theinvasion point of cane toads into the Pilbara is likely to be narrow(beginning around the mouth of the De Grey River Tingleyet al 2013) a proactive approach to training northern quollsto avoid cane toads may prove beneficial Conditioned taste-aversion (CTA) has been used successfully to train captive-reared northern quolls to avoid cane toads by offering themsmall dead toads infused with thiabendazole before theirrelease into the wild (OrsquoDonnell et al 2010)Monitoring of thesequolls after their release showed that some of their offspringhad learnt to avoid eating cane toads either through ingestionof small non-lethal toads or via social learning from theirmothers (Webb et al 2015) Fields trials of toad-aversion baits(either sausages or meatballs made from toad flesh andcontaining thiabendazole) are currently underway in the centralKimberley where cane toads were expected to invade in late2015 (Webb et al 2015) Further testing of CTA approachesunder a variety of field conditions is required The developmentof stable long-lasting baits suitable for aerial deployment isconsidered a pressing need as well as a captive colony ofnorthern quolls upon which to test the efficacy of toad-aversionbaits for inducing aversion to live toads (Webb et al 2015)

Infrastructure development and interactionswith humans

Infrastructure development may increase the likelihood ofnorthern quoll deaths due to motor vehicle strikes or beingcrushed in machinery Significant chuditch mortality wasobserved along mine access roads in the south-west of WesternAustralia (McGregor et al 2014) and the widening and sealingof an access road dramatically increased the number of roadkills and led to the extirpation of the resident population ofeastern quolls (D viverrinus) in Tasmania (Jones 2000)Infrastructure corridors such as high-tension power-lineeasements and ore-haulage railways are increasingly beinginstalled throughout the range of the northern quoll Althoughthese do not create the same mortality risk as roads they bisectand fragment habitat create large open spaces and potentiallyincrease predation risk

A recent small-scale radio-tracking study (see Dunlop et al2014a) around several quarries in the Pilbara found that northernquolls crossed roads but not railway lines The short-term natureof this work and the limited number of quolls able to be trackedmeans that the influence of roads railway lines and infrastructurecorridors on the movement and mortality of northern quollsrequires further assessment Using radio-tracking techniques todetermine crossing rates and road kill events may be difficultover large areas so population studies comparing groups ofquolls near major roads with those not affected by infrastructure(eg McCall et al 2010) may provide a more feasible meansof assessing the impacts of roads If vehicle strikes are foundto significantly contribute to the mortality of a local populationof the northern quoll then mitigation methods should betrialled on sections of road identified as road-kill hotspotsCulverts and underpasses have been shown to be used by avariety of small and medium-sized mammals and frogs (Taylorand Goldingay 2003) while installing rumble strips on roadshas led to decreased rates of road-kill of nocturnal animals(Lester 2015)

Interactions between threats

At present wildfire the subsequent loss of vegetation coverand increased predation of northern quolls is likely to be themost significant of the interactions between threats The directeffects of wildfire have been found to be less significant inthe subsequent decline of small to medium-sized mammalsparticularly for denning species such as the northern quoll thanthe indirect consequences of reduced resource availability andthe increased risk of predation due to lack of cover (Oakwood2000 Letnic et al 2005 McKenzie et al 2007 Legge et al2008 Doherty et al 2015b) Male northern quolls are perhapsmost vulnerable to such losses of cover during the breedingseason when their range can extend to over 100 ha (Oakwood2002) or where males survive after breeding their foragingrange is extended to gain the nutrition required to recover fromtheir poor condition (Cook 2010) Loss of cover due to late dry-season fires at East Alligator River in the Northern Territorymeant little cover was available for young quolls when theybegan foraging with dog and dingo predation a major source ofmortality (Webb et al 2015) The cumulative impacts of canetoad poisoning fire and predation are highly likely to lead to the

Research priorities for the northern quoll Australian Mammalogy I

extirpation of small and disjunct groups of northern quoll in thePilbara Population viability modelling of the East AlligatorRiver population suggested that high mortality from predationcombined with late dry-season burning was preventing therecovery of northern quoll populations after cane toad invasion

(Webb et al 2015) Improved knowledge of the severity of thesethreats to northern quolls in the Pilbara and their incorporationinto the SDM and PVA framework developed by Cadenheadet al (2015) will allow for better identification of priority areasfor integrated cane toad fire and predator management

Table 3 Expected timelines for the undertaking of research under each priority over the next 10 years and the funds currently allocated fromthe Pilbara Northern Quoll Offset Trust under each research priority

Rank Theme Research priority Current or proposed researchactivity

Time over next10 years

Current funding allocation(approximate)

1 Survey andmonitoring

Develop appropriate andstandardised survey andmonitoring methods

Develop monitoring protocol(traps and cameras) andinvestigate new methods

Ongoing survey and monitoringin conjunction with ResearchThemes 2 and 3 to better mapnorthern quoll distribution andto determine long-termpopulation trends

Years 1ndash2Years 3ndash10

$200 000 per annum over10 years for ResearchThemes 1 2 and 3(inclusive of salaries)

2 Habitatrequirements

Define areas of criticalhabitat for northernquoll in the Pilbara

Identify and describe criticalhabitat and undertake predictivemodelling of unsurveyed areasto inform further survey effort

Years 3ndash4 $50 000 for predictivemodelling (EdithCowan University)

Understand howdisturbance affectshabitat quality andconnectivity

Ongoing collection of data ondisturbance and habitat qualityin conjunction with ResearchThemes 1 and 4

Incorporation of new data intopredictive models

Years 4ndash10 See above

3 Populationdynamics

Better understand thepopulation dynamics ofthe northern quoll in thePilbara

Collect data on demographicparameters breeding dispersaland home-range use toincorporate into future PVA

Continued collectionof data inconjunction withpriorities 1 and 2during years 3ndash10

See above

Investigate populationstructure and theinteraction betweenpopulations

Analysis of existing tissuesamples and newly collectedsamples

Years 1ndash10 $120 000 for initial roundof genetic analysis(Years 3ndash5 MurdochUniversity)

4 Key threats Introduced predators Spatio-temporal interactionsbetween northern quolls andintroduced predators

Other projects as they arise

Years 2ndash4 (currentPhD projectUniversity ofQueensland)

$12 000 contribution tofield expenses

Cane toads Predictive modelling of canetoad distribution in the Pilbara

Years 3ndash4 Part of predictivemodelling by EdithCowan University

Infrastructuredevelopment andinteractions withhumans

Investigation of impacts oflinear infrastructure on northernquoll movement

Year 4 (currentHonours projectEdith CowanUniversity)

$15 000

Interactions betweenthreats

No on-ground projects currentlyplanned

Potential to be assessed throughPVA as data are collected underother research priorities

No funding currentlyallocated

5 Recolonisation ofrestored orartificial habitat

Determine the ability ofthe northern quoll torecolonise disturbedareas or coloniseartificial habitat

As projects arise Years 5ndash8 incollaboration withminesite restorationprograms

No funding currentlyallocated

J Australian Mammalogy V A Cramer et al

Priority 5 Recolonisation of restored andor artificial habitat

While anecdotal evidence suggests that northern quolls will denin waste rock dumps whether artificial or restored habitatprovides for long-term population viability is not known andlong-term assessment of created habitat is required Weacknowledge that a large gap exists between the desire to userestored or artificial habitat as an ecologically equivalent offsetand our limited ability to actually restore or re-create habitat thatis equivalent in structure composition and function particularlyon highly disturbed sites (Maron et al 2012) Yet there is a futureopportunity in the Pilbara to conduct lsquonaturalrsquo experiments onthe recolonisation of quolls after mining ceases including howto best design waste rock dumps so that habitat complexityand productivity is maximised by for example experimentingwith the size and positioning of boulders and their spatialarrangement in relation to surrounding landscape features Theuse of such artificial habitat should be tested in the first stagethrough direct monitoring of animals that have colonised orhave been translocated to the site Such experimentation isunderway at Atlas Ironrsquos Mt Dove operation (Atlas Iron 2012)Population modelling can then be used to predict futurepopulation sizes rates of growth and spread and the probabilityof long-term persistence in the context of various managementactions (eg predator control Armstrong and Reynolds 2012)

Resource allocation and timelines

While the allocation of resources and funding and the periodover which each of the research priorities was expected to beaddressed was not explicitly discussed during the workshopthe ranking of the priorities was taken to indicate the sequencein which the priorities should be addressed At the time of theworkshop some of the research questions identified werealready being investigated within various projects and a portionof the Pilbara Northern Quoll Offset Trust allocated to these InTable 3 we indicate the expected timelines over which someof the research programs are expected to be undertaken WhileAU$175M has been allocated to northern quoll research in thePilbara it is important to note that much of this amount is notcurrently available and will not become available until severalapproved mine developments have become operational InTable 3 we also indicate the proportion of offset funds expectedto be allocated to each research priority Research undertaken inother regions (eg research into CTA in the Kimberley regionahead of the cane toad invasion front) cannot be funded from thePilbara Northern Quoll Offset Trust but will help address someof the research questions identified here

Lessons learnt from the workshop process

The workshop approach used here was less formal than recentlarge-scale exercises used to elicit research priorities for emergingissues in science and policy at national or global levels (reviewedin Sutherland et al 2011) The focus in our (much smaller)workshop was on inclusivity rather than identifying participantsbased on purposive sampling of expertise or organisationalaffiliation We believe that the inclusive nature of the workshopwas a strength but we also recognise that this perhaps createdsome level of bias when ranking the research prioritiesAcknowledging both the experience and knowledge requirements

of those who work regularly in the Pilbara is an importantcomponent of improving the design of research programs that aimto improve conservation management in the region Yet theresearch priorities and their rankings are likely strongly reflectiveof the current information needs of most participants and maynot align with what conservation scientists believe to be themost critical areas of research for northern quoll conservation intothe future Baseline survey and monitoring are often the firstrequirements of a management or research plan for the northernquoll when offset conditions are prescribed and this was reflectedin the primary ranking given to the development of standardsurvey and monitoring methods We acknowledge the concernthat this could simply lead to offset funding being used to lsquomonitorthe extinctionrsquo of the northern quoll particularly after canetoads invade the Pilbara unless these data are used to triggermanagement actions (Oakwood and Foster 2008 Lindenmayeret al 2013) Clearly the currently limited information on habitatuse and demographic parameters for the northern quoll in thePilbara restricts the utility of more sophisticated researchapproaches such as developing robust SDM and PVA While thesevere threat that cane toads pose to the northern quoll wasrecognised this threat is not immediate in the region and mayexplain the less prominent ranking given to research to mitigatethe impacts of cane toads

The northern quoll has become emblematic of the dramaticand rapid species declines occurring in northern Australia andthe level of interest in this species was reflected in the highnumber of participants attending the workshop (one of six runby Parks and Wildlife) Given the number of attendees betterutilisation of the time available would have been gained byproviding participants with a brief before the workshopand asking them to contribute potential research questionsanonymously for interactive review by other participants Inthe exercises reviewed by Sutherland et al (2011) questionswere then discarded if they were not considered to meet criteriathat were previously agreed upon Some areas of researchthat gained some attention on the day may have been discardedif given greater consideration or interrogation before theworkshop For example there was some discussion duringthe workshop of investigating the use of habitat surrogates tomonitor the northern quoll yet research has shown that there islittle validity in using surrogates to monitor the abundance of asingle species (Pierson et al 2015)

Conclusions

Offset funds generated under the environmental approvalsprocess provide an opportunity to mobilise considerablefinancial resources for conservation (Kiesecker et al 2010) Yetthere remain significant conceptual and practical limitations tobe resolved to ensure that direct offsets provide measurableconservation outcomes that are ecologically equivalent to thelosses sustained by proceeding with the development (Maronet al 2012 Bull et al 2013) The collaborative identificationand ranking of research priorities undertaken here is the firststep in the process of developing and evaluating potentialconservation actions for the northern quoll in the Pilbara andserves as a mechanism to ensure that the research priorities ofstate and federal agencies reflects the knowledge needs of

Research priorities for the northern quoll Australian Mammalogy K

practitioners and proponents currently operating in the regionThe provision of offset funds over the next decade providesa unique opportunity to undertake research on a species ofconservation significance at a regional scale and over a longperiod through an integrated and strategic program of researchrather than the collection of ad hoc data While a morestructured approach to defining research questions before theworkshop may have led to greater critical reflection upon anddiscussion of some of the research priorities on the day theworkshop provided an opportunity for an open and diverse-ranging discussion in a collaborative environment amongstthe broader community of scientists environmental consultantsand mining proponents involved in the management andconservation of the northern quoll in the Pilbara

Acknowledgements

We gratefully acknowledge the financial support of Roy Hill who sponsoredthe workshop and partially funded VACrsquos salary We thank the Departmentof the Environmentrsquos Pilbara Taskforce especially Veronica Ritchie fortheir support of interested or affected parties The success of the workshopwas in great part due to the facilitation skills of Craig Salt of SustainableConsulting We thank all of the workshop participants for their open andcollaborative approach to the process Mike Bamford Eddie van EttenDavid Peacock and Peter Lyon provided useful comments on an earlier draftWe are grateful to John Woinarski for his constructive advice thatsubstantially improved the structure and clarity of the manuscript and ananonymous reviewer for careful and critical reading of the previousversion Lesley Gibson Russell Palmer and Al Glen generously discussedvarious aspects of the paper with VAC The map was prepared by CristinaRamalho

References

Allen B L Engeman R M and Allen L R (2011) Wild dogma anexamination of recent lsquoevidencersquo for dingo regulation of invasivemesopredator release in Australia Current Zoology 57 568ndash583

Allen B L Fleming P J Allen L R Engeman R M Ballard G andLeung L K-P (2013) As clear as mud a critical review of evidencefor the ecological roles of Australian dingoes Biological Conservation159 158ndash174 doi101016jbiocon201212004

Anon (1934) Floods in Western Australia Town isolated for week PerthMonday The Sydney Morning Herald 9

Armstrong D P and Reynolds M H (2012) Modelling reintroducedpopulations the state of the art and future directions In lsquoReintroductionBiology Integrating Science and Managementrsquo (Eds J G Ewen DP Armstrong K A Parker and P J Seddon) pp 165ndash222 (Wiley-Blackwell UK)

Atlas Iron (2012) Scope of works ndash Mount Dove artificial quoll habitatUnpublished report Atlas Iron Perth Available at httpwwwatlasironcomauIRMCompanyShowPageaspxCategoryId=190ampCPID=5424ampEID=94365410 [accessed 19 August 2015]

Austin C (2014) Can remote cameras accurately estimate populations ofthe endangered northern quoll BSc(Honours) Thesis University ofTechnology Sydney

Begg R J (1981) The small mammals of Little Nourlangie RockNT III Ecology ofDasyurus hallucatus the northern quoll (MarsupialiaDasyuridae) Wildlife Research 8 73ndash85 doi101071WR9810073

Braithwaite R W and Griffiths A D (1994) Demographic variationand range contraction in the northern quoll Dasyurus hallucatus(Marsupialia Dasyuridae)Wildlife Research 21 203ndash217 doi101071WR9940203

Bull J W Suttle K B Gordon A Singh N J and Milner-Gulland E J(2013) Biodiversity offsets in theory and practice Oryx 47 369ndash380doi101017S003060531200172X

Burnett S (1997) Colonizing cane toads cause population declines innative predators reliable anecdotal information and managementimplications Pacific Conservation Biology 3 65ndash72

Cadenhead N C R White A L and Wintle B A (2015) Identifyingconservation priorities and assessing impacts of proposed futuredevelopment in the Pilbara Bioregion in Western Australia Report bythe National Environment Research Program Decisions Hub to theDepartment of the Environment The University of MelbourneMelbourne

Carwardine J Nicol S Van Leeuwen S Walters B Firn J Reeson AMartin T G and Chades I (2014) Priority threat management forPilbara species of conservation significance CSIRO EcosystemSciences Brisbane

Cook A (2010) Habitat use and home-range of the northern quollDasyurushallucatus effects of fire MSc Thesis University ofWestern AustraliaPerth

Department of Sustainability Environment Water Population andCommunities (2012) Environment and Biodiversity Conservation Act1999 Environmental Offsets Policy Department of SustainabilityEnvironment Water Population and Communities Canberra

Doherty T S Davis R A Etten E J Algar D Collier N DickmanC R Edwards G Masters P Palmer R and Robinson S (2015a)A continental-scale analysis of feral cat diet in Australia Journal ofBiogeography 42 964ndash975 doi101111jbi12469

Doherty T S Dickman C R Nimmo D G and Ritchie E G (2015b)Multiple threats or multiplying the threats Interactions betweeninvasive predators and other ecological disturbances BiologicalConservation 190 60ndash68 doi101016jbiocon201505013

Dunlop J Cook A and Lees J (2014a) BHP Billiton Iron Ore Pilbaranorthern quoll monitoring project Final report May 2010ndashJune 2013Unpublished report prepared for BHP Billiton Iron Ore Department ofParks and Wildlife Perth

Dunlop J Cook A and Morris K (2014b) Pilbara northern quollproject surveying and monitoring Dasyurus hallucatus in the PilbaraWestern Australia Department of Parks and Wildlife Perth Availableat librarydpawwagovau

Florance D Webb J K Dempster T Kearney M R Worthing A andLetnic M (2011) Excluding access to invasion hubs can contain thespread of an invasive vertebrate Proceedings of the Royal Society ofLondon B Biological Sciences 278 2900ndash2908 doi101098rspb20110032

Gaillard J-M Hebblewhite M Loison A Fuller M Powell R BasilleM and Van Moorter B (2010) Habitatndashperformance relationshipsfinding the right metric at a given spatial scale PhilosophicalTransactions of the Royal Society of London Series B BiologicalSciences 365 2255ndash2265 doi101098rstb20100085

Glen A S and Dickman C R (2008) Niche overlap between marsupialand eutherian carnivores does competition threaten the endangeredspotted-tailed quoll Journal of Applied Ecology 45 700ndash707 doi101111j1365-2664200701449x

Glen A S Pennay M Dickman C R Wintle B A and Firestone K B(2011) Diets of sympatric native and introduced carnivores in theBarrington Tops eastern Australia Austral Ecology 36 290ndash296doi101111j1442-9993201002149x

Hennig P (2004) A brief land use history In lsquoAn Inventory and ConditionSurvey of the Pilbara Region Western Australiarsquo (Eds A M E VanVreeswyk A L Payne K A Leighton and P Hennig) TechnicalBulletin No 92 pp 13ndash18 (Western Australian Department ofAgriculture Perth)

Hohnen R Ashby J Tuft K and McGregor H (2013) Individualidentification of northern quolls (Dasyurus hallucatus) using remotecameras Australian Mammalogy 35 131ndash135 doi101071AM12015

L Australian Mammalogy V A Cramer et al

How R A Spencer P B S and Schmitt L H (2009) Island populationshave high conservation value for northern Australiarsquos top marsupialpredator ahead of a threatening process Journal of Zoology 278206ndash217 doi101111j1469-7998200900569x

Johnson C N Isaac J L and Fisher D O (2007) Rarity of a top predatortriggers continent-wide collapse of mammal prey dingoes andmarsupials in Australia Proceedings of the Royal Society of LondonSeries B Biological Sciences 274 341ndash346 doi101098rspb20063711

Jones M E (2000) Road upgrade road mortality and remedial measuresimpacts on a population of eastern quolls and Tasmanian devilsWildlifeResearch 27 289ndash296 doi101071WR98069

Kearney M Phillips B L Tracy C R Christian K A Betts G andPorter W P (2008) Modelling species distributions without usingspecies distributions the cane toad in Australia under current andfuture climates Ecography 31 423ndash434 doi101111j0906-7590200805457x

Kendrick P and Stanley F (2002) Pilbara 4 (PIL4 ndash Roebourne synopsis)In lsquoA Biodiversity Audit of Western Australiarsquos 53 BiogeographicalSubregions in 2002rsquo (Eds J E May and N L McKenzie) pp 581ndash593(Department of Conservation and Land Management Perth)

Kennedy M Phillips B L Legge S Murphy S A and Faulkner R A(2012) Do dingoes suppress the activity of feral cats in northernAustralia Austral Ecology 37 134ndash139 doi101111j1442-9993201102256x

Kiesecker J M Copeland H Pocewicz A and McKenney B (2010)Development by design blending landscape-level planning with themitigation hierarchy Frontiers in Ecology and the Environment 8261ndash266 doi101890090005

King D R and Smith L A (1985) The distribution of the European red fox(Vulpes vulpes) in Western Australia Records of the Western AustralianMuseum 12 197ndash205

Knight A T Cowling R M Rouget M Balmford A Lombard A Tand Campbell B M (2008) Knowing but not doing selecting priorityconservation areas and the researchndashimplementation gap ConservationBiology 22 610ndash617 doi101111j1523-1739200800914x

Laurance W F Koster H Grooten M Anderson A B Zuidema P AZwick S Zagt R J Lynam A J Linkie M and Anten N P (2012)Making conservation research more relevant for conservationpractitioners Biological Conservation 153 164ndash168 doi101016jbiocon201205012

Legge S Murphy S Heathcote J Flaxman E Augusteyn J andCrossman M (2008) The short-term effects of an extensive and high-intensity fire on vertebrates in the tropical savannas of the centralKimberley northern Australia Wildlife Research 35 33ndash43 doi101071WR07016

Lester D (2015) Effective wildlife roadkill mitigation Journal of Trafficand Transportation Engineering 3 42ndash51 doi10172652328-2142201501005

Letnic M Tamayo B and Dickman C R (2005) The responsesof mammals to La Nintildea (El Nintildeo Southern Oscillation)-associatedrainfall predation and wildfire in central Australia Journal ofMammalogy 86 689ndash703 doi1016441545-1542(2005)086[0689TROMTL]20CO2

Letnic M Koch F Gordon C Crowther M S and Dickman C R(2009) Keystone effects of an alien top-predator stem extinctions ofnative mammals Proceedings of the Royal Society of London Series BBiological Sciences 276 3249ndash3256 doi101098rspb20090574

Lindenmayer D B (2015) Continental-level biodiversity collapseProceedings of the National Academy of Sciences of the United States ofAmerica 112 4514ndash4515 doi101073pnas1502766112

Lindenmayer D B Piggott M P and Wintle B A (2013) Counting thebooks while the library burns why conservation monitoring programs

need a plan for action Frontiers in Ecology and the Environment 11549ndash555 doi101890120220

Maron M Hobbs R J Moilanen A Matthews J W Christie KGardner T A Keith D A Lindenmayer D B and McAlpine C A(2012) Faustian bargains Restoration realities in the context ofbiodiversity offset policies Biological Conservation 155 141ndash148doi101016jbiocon201206003

McCall S C McCarthy M A van der Ree R Harper M J Cesarini Sand Soanes K (2010) Evidence that a highway reduces apparentsurvival rates of squirrel gliders Ecology and Society 15 27

McGregor R A Stokes V L and Craig M D (2014) Does forestrestoration in fragmented landscapes provide habitat for a wide-ranging carnivore Animal Conservation 17 467ndash475 doi101111acv12112

McKenzie N L Burbidge A A Baynes A Brereton R N DickmanC R Gordon G Gibson L A Menkhorst P W Robinson A CWilliams M R and Woinarski J C Z (2007) Analysis of factorsimplicated in the recent decline of Australiarsquos mammal fauna Journal ofBiogeography 34 597ndash611 doi101111j1365-2699200601639x

McKenzie N L van Leeuwen S and Pinder AM (2009) Introduction tothe Pilbara Biodiversity Survey 2002ndash2007 Records of the WesternAustralian Museum 78(Suppl) 3ndash89 doi1018195issn0313-122x78(1)2009003-089

Morris K Johnson B Orell P Gaikhorst G Wane A and MoroD (2003) Recovery of the threatened chuditch (Dasyurus geoffroii)a case study In lsquoPredators with Pouches The Biology of CarnivorousMarsupialsrsquo (Eds M Jones C Dickman and M Archer) pp 435ndash451(CSIRO Publishing Melbourne)

Nelson J L Scroggie M P and Belcher C A (2014) Developing acamera trap survey protocol to detect a rare marsupial carnivore thespotted-tailed quoll (Dasyurus maculatus) In lsquoCamera TrappingWildlife Management and Researchrsquo (Eds P D Meek and PJ Fleming) pp 271ndash279 (CSIRO Publishing Melbourne)

Newsome T M Dellinger J A Pavey C R Ripple W J Shores C RWirsing A J and Dickman C R (2015) The ecological effects ofproviding resource subsidies to predators Global Ecology andBiogeography 24 1ndash11 doi101111geb12236

OrsquoDonnell S Webb J K and Shine R (2010) Conditioned tasteaversion enhances the survival of an endangered predator imperilled bya toxic invader Journal of Applied Ecology 47 558ndash565 doi101111j1365-2664201001802x

Oakwood M (1997) The ecology of the northern quoll Dasyurushallucatus PhD Thesis Australian National University Canberra

Oakwood M (2000) Reproduction and demography of the northern quollDasyurus hallucatus in the lowland savanna of northern AustraliaAustralian Journal of Zoology 48 519ndash539 doi101071ZO00028

Oakwood M (2002) Spatial and social organization of a carnivorousmarsupial Dasyurus hallucatus (Marsupialia Dasyuridae) Journal ofZoology 257 237ndash248 doi101017S0952836902000833

Oakwood M and Foster P (2008) Monitoring extinction of the northernquoll Australian Academy of Science Newsletter 71 6

Palomares F and Caro T M (1999) Interspecific killing amongmammalian carnivores American Naturalist 153 492ndash508 doi101086303189

Pavey C R and Bastin G (2014) lsquoAustralian Rangelands and ClimateChange ndash Invasive Animalsrsquo (Ninti One Limited amp CSIRO AliceSprings)

Pierson J C Barton P S Lane PW and Lindenmayer D B (2015) Canhabitat surrogates predict the response of target species to landscapechange Biological Conservation 184 1ndash10 doi101016jbiocon201412017

Pollock A B (1999) Notes on status distribution and diet of northernquoll Dasyurus hallucatus in the MackayndashBowen area mideastern

Research priorities for the northern quoll Australian Mammalogy M

Queensland Australian Zoologist 31 388ndash395 doi107882AZ1999040

Ramanaidou E R and Morris R C (2010) A synopsis of the channeliron deposits of the Hamersley Province Western AustraliaTransactions of the Institutions of Mining and Metallurgy Section BApplied Earth Science 119 56ndash59 doi101179037174510X12853354810624

Schmitt L H Bradley A J Kemper C M Kitchener D J HumphreysW F and How R A (1989) Ecology and physiology of the northernquoll Dasyurus hallucatus (Marsupialia Dasyuridae) at MitchellPlateau Kimberley Western Australia Journal of Zoology 217539ndash558 doi101111j1469-79981989tb02510x

Short J and Smith A (1994) Mammal decline and recovery in AustraliaJournal of Mammalogy 75 288ndash297 doi1023071382547

Spencer P How R Hillyer M Cook A Morris K Stevenson C andUmbrello L (2013) Genetic analysis of northern quolls from the PilbaraRegion ofWestern Australia Year one ndash final report Unpublished reportDepartment of Sustainability Environment Water Population andCommunities Canberra and Department of Parks and Wildlife Perth

Sutherland W J Fleishman E Mascia M B Pretty J and Rudd M A(2011) Methods for collaboratively identifying research priorities andemerging issues in science and policyMethods in Ecology and Evolution2 238ndash247 doi101111j2041-210X201000083x

Sutherst R W Floyd R B and Maywald G F (1996) The potentialgeographical distribution of the cane toad Bufo marinus L in AustraliaConservation Biology 10 294ndash299 doi101046j1523-1739199610010294x

Taylor B D and Goldingay R L (2003) Cutting the carnage wildlifeusage of road culverts in north-eastern New South Wales WildlifeResearch 30 529ndash537 doi101071WR01062

Tingley R Phillips B L Letnic M Brown G P Shine R and BairdS J (2013) Identifying optimal barriers to halt the invasion of canetoads Rhinella marina in arid Australia Journal of Applied Ecology 50129ndash137 doi1011111365-266412021

Van Vreeswyk A M E Payne A L Leighton K A and HennigP (2004) An Inventory and Condition Survey of the Pilbara RegionWestern Australia Technical Bulletin No 92 Department of AgricultureWestern Australia Perth

Webb J Legge S Tuft K Cremona T and Austin C (2015) Can wemitigate cane toad impacts on northern quolls Final report National

Environmental Research Program Northern Australia Hub CharlesDarwin University Darwin

Wintle B A Walshe T V Parris K M and McCarthy M A (2012)Designing occupancy surveys and interpreting non-detection whenobservations are imperfect Diversity amp Distributions 18 417ndash424doi101111j1472-4642201100874x

Woinarski J Menkhorst K Gambold N and Braithwaite R W (1992)Mammals of the Bungle Bungle area In lsquoA Survey of the Wildlife andVegetation of Purnululu (Bungle Bungle) National Park and AdjacentArearsquo (Ed J C Z Woinarski) Research Bulletin No 6 pp 53ndash67(CSIRO Division of Wildlife and Ecology Darwin)

Woinarski J C Z Risler J and Kean L (2004) Response of vegetationand vertebrate fauna to 23 years of fire exclusion in a tropical Eucalyptusopen forest Northern Territory Australia Austral Ecology 29 156ndash176doi101111j1442-9993200401333x

Woinarski J C Z Oakwood M Winter J Burnett S Milne D FosterP Myles H and Holmes B (2008) Surviving the toads patterns ofpersistence of the northern quoll Dasyurus hallucatus in QueenslandReport submitted to the Natural Heritage Trust Strategic ReserveProgram as a component of project 2005162 Monitoring andManagement of Cane Toad Impact in theNorthern Territory Available athttplrmntgovauplants-and-animalsinformation-and-publicationspublications2008 [accessed 14 November 2013]

Woinarski J C Z Armstrong M Brennan K Fisher A Griffiths A DHill B Milne D J Palmer C Ward S Watson M Winderlich Sand Young S (2010) Monitoring indicates rapid and severe decline ofnative small mammals in Kakadu National Park northern AustraliaWildlife Research 37 116ndash126 doi101071WR09125

Woinarski J Burbidge A and Harrison P (2014) lsquoAction Plan forAustralian Mammals 2012rsquo (CSIRO Publishing Melbourne)

Woinarski J C Burbidge A A and Harrison P L (2015) Ongoingunraveling of a continental fauna decline and extinction of Australianmammals since European settlement Proceedings of the NationalAcademy of Sciences of the United States of America 112 4531ndash4540doi101073pnas1417301112

Woolley P A Krajewski C and Westerman M (2015) Phylogeneticrelationships within Dasyurus (Dasyuromorphia Dasyuridae) quollsystematics based on molecular evidence and male characteristicsJournal of Mammalogy 96 37ndash46 doi101093jmammalgyu028

N Australian Mammalogy V A Cramer et al

wwwpublishcsiroaujournalsam

Page 5: Research priorities for the northern quoll (Dasyurus ......The Pilbara population of the northern quoll (Dasyurus hallucatus) has been seldom studied, and the impacts of threats such

4Key

threats

Introduced

predators

Doespredationsignificantly

affecttheno

rthern

quollinthePilb

ara

How

does

thisvary

across

habitattypes

Doescompetitionforresourceswith

foxesor

catsim

pact

onsubpopulationsWhich

lifestages

aremostvulnerable

Given

thattheno

rthern

quollisalso

amesop

redatordo

esthepresence

ofding

oeseither

directly(through

predation)

orindirectly(through

theregulatio

nof

foxesandcats)impact

onsubpop

ulations

inthePilb

ara

Assessrelativ

eim

pact

ofthreats

predationby

feralcatsanddogsmdash

high

interactions

betweenferal

catsdo

gsdingoes

andfoxesmdash

mediumndashhigh

Assesseffectivenessof

threat

mitigatio

nop

tionsassess

which

feasible

managem

entoptio

nsfor

feralcatsanddo

gscanlead

topo

pulatio

nrecoverymdashmediumndash

high

Develop

new

orenhanceexistin

gmanagem

entmechanism

sfor

broad-scale

targetedferalcat

controlmdash

medium

Canetoads

Will

theclosureof

artifi

cialwater

bodies

preventor

simplyslow

therateofinv

asionof

cane

toadsinto

thePilb

ara

Whatisthedegree

ofoverlapbetweenSDMfundamentalniche

modelsforn

orthernquolls

andcane

toadsin

thePilb

ara

Can

potentialrefug

eareasforno

rthern

quollb

eidentifi

ed

IsCTAlik

elytobe

effectivewhentaste-aversion

baits

aredeliv

ered

underfi

eldcond

itions

Atwhatscaleisbaitdeliv

eryfeasible

Assesseffectivenessof

threat

mitigatio

nop

tionsassess

feasibility

ofbroad-scaletaste-

aversion

lsquotrainingrsquo

ofqu

ollsmdash

mediumassess

local-scale

exclusionof

toadsor

preventio

nof

colonisatio

n(m

ainlandsites)mdash

medium

Infrastructure

developm

entand

interactions

with

humans

Are

road-related

deaths

ofno

rthern

quollasign

ificant

causeof

mortalityin

thePilb

ara

Dorailw

aylin

esinfrastructurecorridorsor

wideroadspresenta

barrierto

northern

quoll

mov

ement

Whatmethods

canbe

used

tomitigate

theim

pactsof

infrastructure

developm

entand

associated

machinery

operation

Not

addressed

Interactions

between

threats

Dofrequent

andintensefireslead

toreducedstructural

complexity

loss

ofvegetatio

ncoverandsubsequentincreasedpredationof

northern

quollsparticularlyof

males

during

andor

afterbreeding

Whatarethecumulativeim

pactsof

mortalitycaused

bycane

toad

ingestionandpredation

bycatsdo

gsdingoes

andfoxes

See

inform

ationon

introd

uced

predatorsandcane

toadsabove

5Recolon

isationof

restored

orartifi

cial

habitat

Determinetheability

oftheno

rthern

quoll

torecolonise

disturbedareasor

colonise

artifi

cial

habitat

Whatarethekeyelem

entsof

northern

quollhabitatthatneedtobe

restoredcreated

sothat

habitatcomplexity

andprod

uctiv

ityismaxim

ised

Whatlevelo

flandscapeconn

ectiv

ityisrequired

arou

ndrestoredartificialhabitatso

that

genetic

connectiv

itybetweensubpopulations

canbe

maintained

How

long

should

restored

orartifi

cial

habitatbe

directly

managedparticularly

for

predators

How

cantheretentionof

largebouldersof

wasterock

beincorporated

into

rehabilitation

plans

Not

addressed

Research priorities for the northern quoll Australian Mammalogy E

investment of time and resources The limited amount anduncertain quality of occurrence data has been a major restrictionon the utility of a species distribution model (SDM) and apopulation viability analysis (PVA) for the northern quoll in thePilbara (Cadenhead et al 2015 see below) While survey andmonitoring per se is not a research activity that directly informsmanagement or threat mitigation carefully implementedmonitoring programs are essential to identify many elements ofa speciesrsquo ecology to detect changes in population and toquantify the ecological effectiveness and cost-effectivenessof management interventions (Lindenmayer et al 2013Lindenmayer 2015)

Parks and Wildlife recently released revised protocols forthe long-term monitoring of known populations of thenorthern quoll (Dunlop et al 2014b) and the effectiveness ofthese protocols in presenceabsence surveys should be testedThe cost-effectiveness of using camera traps for wildlifesurveys in remote locations such as the Pilbara makes their useappealing Hohnen et al (2013) described how the strategicplacement of cameras and baits combined with taking a greaternumber of photographs per trigger allows the capture ofimages that can be used to identify individual northern quollsfrom their spot patterns Remote cameras have also been shownto be a viable method for estimating population sizes of thenorthern quoll in areas with both dense (Austin 2014) andsmall populations (Webb et al 2015) Further testing of a rangeof parameters (eg see Nelson et al 2014) to develop a robuststandard protocol for the Pilbara is required along with acomparison of the cost-effectiveness and any ethicalreducedhandling benefits of using remote cameras versus live-trappingfor estimating population sizes in Pilbara landscapes

Priority 2 Improve our understanding of habitatrequirements

Define areas of critical habitat for the northern quollin the Pilbara

The Department of Agriculture and Food Western Australiahas mapped 102 land systems across the Pilbara bioregionbased on several biophysical components including landformgeology soils vegetation and drainage patterns (Van Vreeswyket al 2004) A review of records for the northern quoll in thePilbara showed that 65 of records were from the RobeCapricorn and Boolaloo land systems (Table 2) which compriselow plateaux mesas hills and ridges of varying geology(limonite granite and sandstone respectively) Most recentsurvey effort in the Pilbara is biased towards mineral-bearingformations or areas of mining-related infrastructure (eg railand road corridors) (Fig 1) Further investigation of theserecords is required to assess whether these land systems areindeed preferred habitat for northern quolls as opposed to anartefact of survey effort The comparatively fewer records thatcoincide with the conservation estate in the region (Fig 1) arealso likely an artefact of survey effort

The first step in defining areas of critical habitat for thenorthern quoll in the Pilbara is to improve our understanding ofhabitat use particularly in terms of defining habitat suitability atfiner scales than is currently available As part of the strategic

environmental assessments being undertaken for the Pilbarabioregion an SDM has been developed for the northern quoll(Cadenhead et al 2015) Uncertainty about the quality of theoccurrence data available the limited amount of data availableas well as the bias towards data collection in areas proposed fordevelopment constrained the level of certainty around theoutcomes of the model Improved model certainty requiresfurther collection of contemporary species-occurrence data andthe development of finer-resolution GIS layers particularly onethat captures finer-scale geological attributes such as mesas orrocky outcrops considered as highest-quality habitat for thenorthern quoll (Cadenhead et al 2015)

Better definition of the habitat attributes favoured by thenorthern quoll will allow further refinement of the GIS variablesincluded in the SDM or allow the development of predictivesite-based habitat models Information on how northern quollsuse habitat can be gained through direct observation of howthey interact with habitat features or from the association ofhabitat features with the presence of quolls as determined byGPS receivers (Gaillard et al 2010) This level of detail iscurrently not available from survey records returned as part ofenvironmental impact assessments but collection of habitatdata using the protocols outlined by Dunlop et al (2014b) infuture surveys would rapidly increase the quality of dataavailable

Understand how disturbance affects habitat qualityand connectivity

The alteration of fire regimes has been the most pervasivelandscape-scale change in the Pilbara Between 1993 and 2006over 72 of the region was burnt with more than 28 of thearea burnt two or more times (McKenzie et al 2009) Woinarskiet al (2004) found that northern quolls may be able to tolerate amoderately high frequency of fire as long as these fires werelow-intensity early-season burns and the impacts of fire onvegetation structure and composition were not exacerbatedby the grazing of introduced herbivores In the Pilbarahowever as with many other grazing regions of Australia earlypastoralists often overestimated the productivity and resilienceof native vegetation to sheep and cattle grazing which led tosevere degradation of vegetation particularly along riverfrontages (Hennig 2004) The soil degradation and loss ofvegetative cover caused by intense grazing pressure in thesouth Kimberley is thought likely to have contributed to thesubstantial decline or disappearance of medium-sized animalsincluding the northern quoll from the Bungle Bungle area(Woinarski et al 1992) Death by predation is most likely in openareas (Oakwood 2000) and therefore the effects of fire andgrazing on northern quoll populations cannot be consideredwithout also taking account of likely increases in mortality due topredation (see lsquoIntroduced predatorsrsquo below)

Understanding how the localised but intensive activityassociated with mining affects habitat quality and habitat useby the northern quoll in areas surrounding development islimited A recent study that examined the use of ballast quarriesand paired undisturbed sites by northern quolls found that theywere foraging denning and showing signs of breeding at thequarry sites where greater numbers of quolls were captured(Dunlop et al 2014a) This was likely due to the combination of

F Australian Mammalogy V A Cramer et al

habitat complexity (eg diversity of boulder sizes) and thepresence of permanent water in the quarry which may haveincreased prey availability at these sites

Movement of animals through the landscape for dispersaland repopulation is another important consideration for a speciesin which both sexes tend towards semelparity and is thereforepredisposed to local extinction (Oakwood 1997) Improvedunderstanding of how the northern quoll (particularly dispersingmales) uses areas of lower habitat quality and the identificationof such habitat in habitat-suitability models will allow futurelandscape-scale strategic assessments to better predict thecombined impacts of disturbance on the ability of northern quolls

to move through the landscape Analysis of regional populationgenetics will also improve our understanding of the level ofdispersal occurring across the landscape

Priority 3 Population dynamics

Better understand the population dynamics of thenorthern quoll in the Pilbara

Little information is available on the number ofsubpopulations of northern quoll in the Pilbara how thesesubpopulations interact how stable they are in space and timeand how they respond to disturbance As no demographic

Table 2 Records of the northern quoll in each land system type in the PilbaraRecords were current as of 23 May 2014

Land system Area (km2) No of records Land system description

Augustus 61 1 Rugged ranges hills ridges and plateaux supporting mulga shrublands and hard spinifex grasslandsBoolaloo 1502 245 Granite hills domes and tor fields and sandy plains with shrubby spinifex grasslandsBoolgeeda 7748 29 Stony lower slopes and plains below hill systems supporting hard and soft spinifex grasslands andmulga

shrublandsCalcrete 1444 2 Low calcrete platforms and plains supporting shrubby hard spinifex grasslandsCallawa 1003 9 Highly dissected low hills mesas and gravelly plains of sandstone and conglomerate supporting soft and

hard spinifex grasslandsCapricorn 5296 257 Hills and ridges of sandstone and dolomite shrubby hard and soft spinifex grasslandsCoongimah 3244 12 Plateau surfaces low hills with steep slopes and undulating uplands supporting hard spinifex grasslandsDune 138 1 Dune fields supporting soft spinifex grasslandsGranitic 4020 1 Rugged granitic hills supporting shrubby hard and soft spinifex grasslandsHorseflat 1261 4 Gilgaied clay plains supporting tussock grasslands and minor grassy snakewood shrublandsLittoral 1577 3 Bare coastal mudflats with mangroves on seaward fringes samphire flats sandy islands coastal dunes

and beachesMacroy 13 095 94 Stony plains and occasional tor fields based on granite supporting hard and soft spinifex grasslandsMcKay 4202 21 Hills ridges plateau remnants and breakaways of metasedimentary rocks supporting hard spinifex

grasslandsMallina 2557 8 Sandy surfaced alluvial plains supporting soft spinifex (and occasionally hard spinifex) grasslandsMarandoo 459 2 Basalt hills and restricted stony plains supporting grassy mulga shrublandsMosquito 1840 3 Stony plains and prominent ridges of schist and other metamorphic rocks supporting hard spinifex

grasslandsNewman 14 580 81 Rugged jaspilite plateaux ridges and mountains supporting hard spinifex grasslandsOnslow 424 1 Sandplains dunes and clay plains supporting soft spinifex grasslands and minor tussock grasslandsParaburdoo 565 3 Basalt-derived stony gilgai plains and stony plains supporting snakewood and mulga shrublands with

spinifex and tussock grasslandsParadise 1479 8 Alluvial plains supporting soft spinifex grasslands and tussock grasslandsRiver 4088 37 Active flood plains major rivers supporting grassy eucalypt woodlands tussock grasslands and soft

spinifex grasslandsRobe 865 369 Low limonite mesas and buttes supporting soft spinifex (and occasionally hard spinifex) grasslandsRocklea 22 993 40 Basalt hills plateaux lower slopes and minor stony plains supporting hard spinifex (and occasionally

soft spinifex) grasslandsRuth 346 36 Hills and ridges of volcanic and other rocks supporting hard spinifex (occasionally soft spinifex)

grasslandsStuart 1794 3 Gently undulating stony plains supporting hard and soft spinifex grassland and snakewood shrublandsTaylor 129 3 Stony plains and isolated low hills of sedimentary rocks supporting hard and soft spinifex grasslandsTalga 2124 9 Hills and ridges of greenstone and chert and stony plains supporting hard and soft spinifex grasslandsTuree 581 2 Stony alluvial plains with gilgaied and non-gilgaied surfaces supporting tussock grasslands and grassy

shrublandsUaroo 7681 37 Broad sandy plains supporting shrubby hard and soft spinifex grasslandsUrandy 1311 6 Stony plains alluvial plains and drainage lines supporting shrubby soft spinifex grasslandsWhite Springs 266 4 Stony gilgai plains supporting tussock grasslands and hard spinifex grasslandsWona 1815 10 Basalt upland gilgai plains supporting tussock grasslands and minor hard spinifex grasslandsYamerina 1207 1 Floodplains and deltaic deposits supporting tussock grasslands grassy woodlands and minor halophytic

low shrublands

Research priorities for the northern quoll Australian Mammalogy G

studies have been undertaken in the region Cadenhead et al(2015) used demographic information from the NorthernTerritory and Kimberley populations in their PVA Uncertaintyabout data quality and the limited information available ondemographic parameters for the Pilbara constrained the level ofcertainty around the outcomes of the PVA The model doeshowever provide a framework into which new data can beincorporated as they become available For example preliminarydata from the Pilbara monitoring program suggest that theproportion of male die-off in the Pilbara population is lowerthan in the Kimberley and Northern Territory populations(Dunlop unpubl data) Cadenhead et al (2015) stress the needfor more demographic and occurrence data to be collectedbefore further development decisions are made (see lsquoDevelopappropriate and standardised survey and monitoring methodsrsquoand lsquoImprove our understanding of habitat requirementsrsquo above)

Investigate population structure and the interactionbetween populations

Analysis of the genetic diversity and population structure ofthe Pilbara population of the northern quoll is ongoingOnly small differences in genetic diversity as measured byheterozygosity were found between samples obtained at 16locations within the Pilbara with the exception of the isolatedpopulation on Dolphin Island (Spencer et al 2013) This resultindicates a high level of movement across the landscape byindividuals during mating and dispersal phases Spatial structureanalysis of samples collected from around Pannawonicasuggested a lsquoneighbour sizersquo of ~15 km (Spencer et al 2013)Current data show no evidence of genetic bottleneckssuggesting that if there has been a population decline in thePilbara that this has not affected genetic diversity A usefulapproach would be to combine genetic information with datafrom radio-tracking or GPS collars to better understand theuse of lsquolandscapersquo space and how this influences populationdynamics

Priority 4 Key threats and the interaction of these threats

Introduced predators

Introduced cats and foxes are considered to be one of themain causes of the decline and extinction of the critical-weight-range mammals (35 g to 55 kg) in Australia (Short and Smith1994 McKenzie et al 2007 Woinarski et al 2015) but theirimpacts on the northern quoll are less clear A recent review offeral cat diets across Australia found that northern quolls werepresent in the diet of feral cats (Doherty et al 2015a) andWoinarski et al (2014) consider the threat posed by cats to thenorthern quoll to be severe although the impacts of cats may bereduced in rugged refuge areas The impacts of cats on othermuch larger species of quoll appear to be inconsistent withsome evidence of coexistence between cats and quolls beforethe arrival of foxes (Glen and Dickman 2008 and referencescontained within) Due to the limited overlap in their distributionthroughout Australia the threat posed by foxes to northernquolls is less understood than for cats and Woinarski et al(2014) consider the threat posed by foxes to be moderate on anational level Foxes are common in coastal near-coastal and

riparian areas of the Pilbara and extend as far north as Broome(King and Smith 1985 Kendrick and Stanley 2002) They arerenowned for their aggression towards smaller carnivores(Palomares and Caro 1999) and neither the larger chuditch(Dasyurus geoffroii) nor spotted-tail quoll (Dasyurus maculatus)are known to coexist with large numbers of foxes (A Glen perscomm) The interactions between quolls cats and foxesare potentially more complex than for other medium-sizedmarsupials due to niche overlap in diet and competition for densites with the larger spotted-tailed quoll and chuditch bothexperiencing exploitation and interference competition fromfoxes (Morris et al 2003 Glen et al 2011) Competitionbetween feral cats and the northern quoll may potentially beintense during periods of low rainfall in the Pilbara when preyavailability is low Understanding whether foxes and cats aresignificant predators of or competitors with the northern quollin the Pilbara and how this affects their population viabilitywill clarify the level of risk posed by these predators Thesausage-style baits used for feral cats (eg Eradicatreg andCuriosityreg) are potentially attractive to the northern quoll withthe risk of mortality or reduced fitness Trials to assess themagnitude of the risks from Eradicatreg baits are currentlyunderway in the Pilbara

The role of dingoes and wild dogs in the mortality of andinteractions between mesopredators (both native and introduced)requires further investigation Dingoes are known to kill northernquolls butmay not eat these kills (Oakwood 1997) A recent studyin the Northern Territory found that camp dogs and dingoes werethe major source of mortality for northern quolls at the study site(Webb et al 2015) and roaming domestic dogs were identified asa cause of mortality for northern quolls in Queensland (Pollock1999) Human resource subsidies (eg stock carrion in rangelandareas) can artificially support predator populations thus indirectlyincreasing predation pressure on prey (Newsome et al 2015)A dynamic debate currently exists around the role of the dingo inregulating populations of feral cats and foxes and the implicationsof this for the conservation of the critical-weight-range mammalsin Australia (see Johnson et al 2007 Letnic et al 2009 Allenet al 2011 2013 Kennedy et al 2012) Understanding thepopulation density at which the role of the dingo as a regulator ofintroduced predators outweighs its own impact as a direct predatorof the northern quoll will inform whether dingowild dog controlwould benefit the northern quoll

Cane toads

The introduced cane toad now occupies more than half thehistoric range of the northern quoll and has caused an immediatecollapse or extirpation of northern quoll populations in theareas it has invaded (Burnett 1997 Oakwood and Foster2008 Woinarski et al 2008 2010) Earlier modelling work thatconsidered only climate parameters as well as more recentwork that incorporates future climate change suggest thatcane toads will not invade the Pilbara (Sutherst et al 1996Pavey and Bastin 2014) Other recent work that considers thefundamental niche of the cane toad and incorporates the locationof artificial watering points predicts that they will colonisethe Pilbara from the Kimberley via the coastal edge of theGreat Sandy Desert (Kearney et al 2008 Florance et al 2011

H Australian Mammalogy V A Cramer et al

Tingley et al 2013) This has created some level of uncertaintyabout the likely impact of cane toads on northern quolls in thePilbara Strategies for cane toad management were not rankedhighly by participants in the threat-management frameworkdeveloped by Carwardine et al (2014) which the authorsexplained as possibly due to uncertainty about how far canetoads will spread into the region

The model developed by Tingley et al (2013) suggested thatthe closure of 100 artificial water bodies at one of three potentiallocations along the coastal corridor between the Kimberley andthe Pilbara would prevent the invasion of cane toads into thePilbara While their model did account for uncharacteristicallywet years every three decades the dispersal of toads betweenwater bodies was determined by the number of rainy daysoccurring during the year (ie toads could only disperse betweenwater bodies for three days following rainfall) rather thanrainfall amount the associated likelihood of flooding and theavailability of free water across the landscape The cyclonic andthunderstorm nature of much of the rainfall in the north-west ofWestern Australia means that the region is prone to significantand sometimes extended flooding In April 1934 the countryaround Anna Plains (250 km south of Broome) was flooded forhundreds of miles when 27 inches (686mm) of rain fell infour days (Anon 1934) In March 2004 over 350mm of rain wasrecorded within a 48-h period at Telfer on the southern edge ofthe Great Sandy Desert due to Tropical Cyclone Fay Thiscaused the flooding of the De Grey and Oakover river systemsand flooded 140 km of the Telfer Access Road with floodwaterstaking several months to subside (wwwbomgovau) Extendingthe model of Tingley et al (2013) to account for such events willhelp clarify whether closing artificial water bodies is likely toprevent the invasion of cane toads into the Pilbara or may onlyassist in delaying their inevitable invasion

Comparison of SDMs (eg Cadenhead et al 2015) fornorthern quoll with fundamental niche models for cane toads(eg see Kearney et al 2008) within the Pilbara may identifypotential refuge areas for the northern quoll and these areasshould be prioritised for conservation actions Because theinvasion point of cane toads into the Pilbara is likely to be narrow(beginning around the mouth of the De Grey River Tingleyet al 2013) a proactive approach to training northern quollsto avoid cane toads may prove beneficial Conditioned taste-aversion (CTA) has been used successfully to train captive-reared northern quolls to avoid cane toads by offering themsmall dead toads infused with thiabendazole before theirrelease into the wild (OrsquoDonnell et al 2010)Monitoring of thesequolls after their release showed that some of their offspringhad learnt to avoid eating cane toads either through ingestionof small non-lethal toads or via social learning from theirmothers (Webb et al 2015) Fields trials of toad-aversion baits(either sausages or meatballs made from toad flesh andcontaining thiabendazole) are currently underway in the centralKimberley where cane toads were expected to invade in late2015 (Webb et al 2015) Further testing of CTA approachesunder a variety of field conditions is required The developmentof stable long-lasting baits suitable for aerial deployment isconsidered a pressing need as well as a captive colony ofnorthern quolls upon which to test the efficacy of toad-aversionbaits for inducing aversion to live toads (Webb et al 2015)

Infrastructure development and interactionswith humans

Infrastructure development may increase the likelihood ofnorthern quoll deaths due to motor vehicle strikes or beingcrushed in machinery Significant chuditch mortality wasobserved along mine access roads in the south-west of WesternAustralia (McGregor et al 2014) and the widening and sealingof an access road dramatically increased the number of roadkills and led to the extirpation of the resident population ofeastern quolls (D viverrinus) in Tasmania (Jones 2000)Infrastructure corridors such as high-tension power-lineeasements and ore-haulage railways are increasingly beinginstalled throughout the range of the northern quoll Althoughthese do not create the same mortality risk as roads they bisectand fragment habitat create large open spaces and potentiallyincrease predation risk

A recent small-scale radio-tracking study (see Dunlop et al2014a) around several quarries in the Pilbara found that northernquolls crossed roads but not railway lines The short-term natureof this work and the limited number of quolls able to be trackedmeans that the influence of roads railway lines and infrastructurecorridors on the movement and mortality of northern quollsrequires further assessment Using radio-tracking techniques todetermine crossing rates and road kill events may be difficultover large areas so population studies comparing groups ofquolls near major roads with those not affected by infrastructure(eg McCall et al 2010) may provide a more feasible meansof assessing the impacts of roads If vehicle strikes are foundto significantly contribute to the mortality of a local populationof the northern quoll then mitigation methods should betrialled on sections of road identified as road-kill hotspotsCulverts and underpasses have been shown to be used by avariety of small and medium-sized mammals and frogs (Taylorand Goldingay 2003) while installing rumble strips on roadshas led to decreased rates of road-kill of nocturnal animals(Lester 2015)

Interactions between threats

At present wildfire the subsequent loss of vegetation coverand increased predation of northern quolls is likely to be themost significant of the interactions between threats The directeffects of wildfire have been found to be less significant inthe subsequent decline of small to medium-sized mammalsparticularly for denning species such as the northern quoll thanthe indirect consequences of reduced resource availability andthe increased risk of predation due to lack of cover (Oakwood2000 Letnic et al 2005 McKenzie et al 2007 Legge et al2008 Doherty et al 2015b) Male northern quolls are perhapsmost vulnerable to such losses of cover during the breedingseason when their range can extend to over 100 ha (Oakwood2002) or where males survive after breeding their foragingrange is extended to gain the nutrition required to recover fromtheir poor condition (Cook 2010) Loss of cover due to late dry-season fires at East Alligator River in the Northern Territorymeant little cover was available for young quolls when theybegan foraging with dog and dingo predation a major source ofmortality (Webb et al 2015) The cumulative impacts of canetoad poisoning fire and predation are highly likely to lead to the

Research priorities for the northern quoll Australian Mammalogy I

extirpation of small and disjunct groups of northern quoll in thePilbara Population viability modelling of the East AlligatorRiver population suggested that high mortality from predationcombined with late dry-season burning was preventing therecovery of northern quoll populations after cane toad invasion

(Webb et al 2015) Improved knowledge of the severity of thesethreats to northern quolls in the Pilbara and their incorporationinto the SDM and PVA framework developed by Cadenheadet al (2015) will allow for better identification of priority areasfor integrated cane toad fire and predator management

Table 3 Expected timelines for the undertaking of research under each priority over the next 10 years and the funds currently allocated fromthe Pilbara Northern Quoll Offset Trust under each research priority

Rank Theme Research priority Current or proposed researchactivity

Time over next10 years

Current funding allocation(approximate)

1 Survey andmonitoring

Develop appropriate andstandardised survey andmonitoring methods

Develop monitoring protocol(traps and cameras) andinvestigate new methods

Ongoing survey and monitoringin conjunction with ResearchThemes 2 and 3 to better mapnorthern quoll distribution andto determine long-termpopulation trends

Years 1ndash2Years 3ndash10

$200 000 per annum over10 years for ResearchThemes 1 2 and 3(inclusive of salaries)

2 Habitatrequirements

Define areas of criticalhabitat for northernquoll in the Pilbara

Identify and describe criticalhabitat and undertake predictivemodelling of unsurveyed areasto inform further survey effort

Years 3ndash4 $50 000 for predictivemodelling (EdithCowan University)

Understand howdisturbance affectshabitat quality andconnectivity

Ongoing collection of data ondisturbance and habitat qualityin conjunction with ResearchThemes 1 and 4

Incorporation of new data intopredictive models

Years 4ndash10 See above

3 Populationdynamics

Better understand thepopulation dynamics ofthe northern quoll in thePilbara

Collect data on demographicparameters breeding dispersaland home-range use toincorporate into future PVA

Continued collectionof data inconjunction withpriorities 1 and 2during years 3ndash10

See above

Investigate populationstructure and theinteraction betweenpopulations

Analysis of existing tissuesamples and newly collectedsamples

Years 1ndash10 $120 000 for initial roundof genetic analysis(Years 3ndash5 MurdochUniversity)

4 Key threats Introduced predators Spatio-temporal interactionsbetween northern quolls andintroduced predators

Other projects as they arise

Years 2ndash4 (currentPhD projectUniversity ofQueensland)

$12 000 contribution tofield expenses

Cane toads Predictive modelling of canetoad distribution in the Pilbara

Years 3ndash4 Part of predictivemodelling by EdithCowan University

Infrastructuredevelopment andinteractions withhumans

Investigation of impacts oflinear infrastructure on northernquoll movement

Year 4 (currentHonours projectEdith CowanUniversity)

$15 000

Interactions betweenthreats

No on-ground projects currentlyplanned

Potential to be assessed throughPVA as data are collected underother research priorities

No funding currentlyallocated

5 Recolonisation ofrestored orartificial habitat

Determine the ability ofthe northern quoll torecolonise disturbedareas or coloniseartificial habitat

As projects arise Years 5ndash8 incollaboration withminesite restorationprograms

No funding currentlyallocated

J Australian Mammalogy V A Cramer et al

Priority 5 Recolonisation of restored andor artificial habitat

While anecdotal evidence suggests that northern quolls will denin waste rock dumps whether artificial or restored habitatprovides for long-term population viability is not known andlong-term assessment of created habitat is required Weacknowledge that a large gap exists between the desire to userestored or artificial habitat as an ecologically equivalent offsetand our limited ability to actually restore or re-create habitat thatis equivalent in structure composition and function particularlyon highly disturbed sites (Maron et al 2012) Yet there is a futureopportunity in the Pilbara to conduct lsquonaturalrsquo experiments onthe recolonisation of quolls after mining ceases including howto best design waste rock dumps so that habitat complexityand productivity is maximised by for example experimentingwith the size and positioning of boulders and their spatialarrangement in relation to surrounding landscape features Theuse of such artificial habitat should be tested in the first stagethrough direct monitoring of animals that have colonised orhave been translocated to the site Such experimentation isunderway at Atlas Ironrsquos Mt Dove operation (Atlas Iron 2012)Population modelling can then be used to predict futurepopulation sizes rates of growth and spread and the probabilityof long-term persistence in the context of various managementactions (eg predator control Armstrong and Reynolds 2012)

Resource allocation and timelines

While the allocation of resources and funding and the periodover which each of the research priorities was expected to beaddressed was not explicitly discussed during the workshopthe ranking of the priorities was taken to indicate the sequencein which the priorities should be addressed At the time of theworkshop some of the research questions identified werealready being investigated within various projects and a portionof the Pilbara Northern Quoll Offset Trust allocated to these InTable 3 we indicate the expected timelines over which someof the research programs are expected to be undertaken WhileAU$175M has been allocated to northern quoll research in thePilbara it is important to note that much of this amount is notcurrently available and will not become available until severalapproved mine developments have become operational InTable 3 we also indicate the proportion of offset funds expectedto be allocated to each research priority Research undertaken inother regions (eg research into CTA in the Kimberley regionahead of the cane toad invasion front) cannot be funded from thePilbara Northern Quoll Offset Trust but will help address someof the research questions identified here

Lessons learnt from the workshop process

The workshop approach used here was less formal than recentlarge-scale exercises used to elicit research priorities for emergingissues in science and policy at national or global levels (reviewedin Sutherland et al 2011) The focus in our (much smaller)workshop was on inclusivity rather than identifying participantsbased on purposive sampling of expertise or organisationalaffiliation We believe that the inclusive nature of the workshopwas a strength but we also recognise that this perhaps createdsome level of bias when ranking the research prioritiesAcknowledging both the experience and knowledge requirements

of those who work regularly in the Pilbara is an importantcomponent of improving the design of research programs that aimto improve conservation management in the region Yet theresearch priorities and their rankings are likely strongly reflectiveof the current information needs of most participants and maynot align with what conservation scientists believe to be themost critical areas of research for northern quoll conservation intothe future Baseline survey and monitoring are often the firstrequirements of a management or research plan for the northernquoll when offset conditions are prescribed and this was reflectedin the primary ranking given to the development of standardsurvey and monitoring methods We acknowledge the concernthat this could simply lead to offset funding being used to lsquomonitorthe extinctionrsquo of the northern quoll particularly after canetoads invade the Pilbara unless these data are used to triggermanagement actions (Oakwood and Foster 2008 Lindenmayeret al 2013) Clearly the currently limited information on habitatuse and demographic parameters for the northern quoll in thePilbara restricts the utility of more sophisticated researchapproaches such as developing robust SDM and PVA While thesevere threat that cane toads pose to the northern quoll wasrecognised this threat is not immediate in the region and mayexplain the less prominent ranking given to research to mitigatethe impacts of cane toads

The northern quoll has become emblematic of the dramaticand rapid species declines occurring in northern Australia andthe level of interest in this species was reflected in the highnumber of participants attending the workshop (one of six runby Parks and Wildlife) Given the number of attendees betterutilisation of the time available would have been gained byproviding participants with a brief before the workshopand asking them to contribute potential research questionsanonymously for interactive review by other participants Inthe exercises reviewed by Sutherland et al (2011) questionswere then discarded if they were not considered to meet criteriathat were previously agreed upon Some areas of researchthat gained some attention on the day may have been discardedif given greater consideration or interrogation before theworkshop For example there was some discussion duringthe workshop of investigating the use of habitat surrogates tomonitor the northern quoll yet research has shown that there islittle validity in using surrogates to monitor the abundance of asingle species (Pierson et al 2015)

Conclusions

Offset funds generated under the environmental approvalsprocess provide an opportunity to mobilise considerablefinancial resources for conservation (Kiesecker et al 2010) Yetthere remain significant conceptual and practical limitations tobe resolved to ensure that direct offsets provide measurableconservation outcomes that are ecologically equivalent to thelosses sustained by proceeding with the development (Maronet al 2012 Bull et al 2013) The collaborative identificationand ranking of research priorities undertaken here is the firststep in the process of developing and evaluating potentialconservation actions for the northern quoll in the Pilbara andserves as a mechanism to ensure that the research priorities ofstate and federal agencies reflects the knowledge needs of

Research priorities for the northern quoll Australian Mammalogy K

practitioners and proponents currently operating in the regionThe provision of offset funds over the next decade providesa unique opportunity to undertake research on a species ofconservation significance at a regional scale and over a longperiod through an integrated and strategic program of researchrather than the collection of ad hoc data While a morestructured approach to defining research questions before theworkshop may have led to greater critical reflection upon anddiscussion of some of the research priorities on the day theworkshop provided an opportunity for an open and diverse-ranging discussion in a collaborative environment amongstthe broader community of scientists environmental consultantsand mining proponents involved in the management andconservation of the northern quoll in the Pilbara

Acknowledgements

We gratefully acknowledge the financial support of Roy Hill who sponsoredthe workshop and partially funded VACrsquos salary We thank the Departmentof the Environmentrsquos Pilbara Taskforce especially Veronica Ritchie fortheir support of interested or affected parties The success of the workshopwas in great part due to the facilitation skills of Craig Salt of SustainableConsulting We thank all of the workshop participants for their open andcollaborative approach to the process Mike Bamford Eddie van EttenDavid Peacock and Peter Lyon provided useful comments on an earlier draftWe are grateful to John Woinarski for his constructive advice thatsubstantially improved the structure and clarity of the manuscript and ananonymous reviewer for careful and critical reading of the previousversion Lesley Gibson Russell Palmer and Al Glen generously discussedvarious aspects of the paper with VAC The map was prepared by CristinaRamalho

References

Allen B L Engeman R M and Allen L R (2011) Wild dogma anexamination of recent lsquoevidencersquo for dingo regulation of invasivemesopredator release in Australia Current Zoology 57 568ndash583

Allen B L Fleming P J Allen L R Engeman R M Ballard G andLeung L K-P (2013) As clear as mud a critical review of evidencefor the ecological roles of Australian dingoes Biological Conservation159 158ndash174 doi101016jbiocon201212004

Anon (1934) Floods in Western Australia Town isolated for week PerthMonday The Sydney Morning Herald 9

Armstrong D P and Reynolds M H (2012) Modelling reintroducedpopulations the state of the art and future directions In lsquoReintroductionBiology Integrating Science and Managementrsquo (Eds J G Ewen DP Armstrong K A Parker and P J Seddon) pp 165ndash222 (Wiley-Blackwell UK)

Atlas Iron (2012) Scope of works ndash Mount Dove artificial quoll habitatUnpublished report Atlas Iron Perth Available at httpwwwatlasironcomauIRMCompanyShowPageaspxCategoryId=190ampCPID=5424ampEID=94365410 [accessed 19 August 2015]

Austin C (2014) Can remote cameras accurately estimate populations ofthe endangered northern quoll BSc(Honours) Thesis University ofTechnology Sydney

Begg R J (1981) The small mammals of Little Nourlangie RockNT III Ecology ofDasyurus hallucatus the northern quoll (MarsupialiaDasyuridae) Wildlife Research 8 73ndash85 doi101071WR9810073

Braithwaite R W and Griffiths A D (1994) Demographic variationand range contraction in the northern quoll Dasyurus hallucatus(Marsupialia Dasyuridae)Wildlife Research 21 203ndash217 doi101071WR9940203

Bull J W Suttle K B Gordon A Singh N J and Milner-Gulland E J(2013) Biodiversity offsets in theory and practice Oryx 47 369ndash380doi101017S003060531200172X

Burnett S (1997) Colonizing cane toads cause population declines innative predators reliable anecdotal information and managementimplications Pacific Conservation Biology 3 65ndash72

Cadenhead N C R White A L and Wintle B A (2015) Identifyingconservation priorities and assessing impacts of proposed futuredevelopment in the Pilbara Bioregion in Western Australia Report bythe National Environment Research Program Decisions Hub to theDepartment of the Environment The University of MelbourneMelbourne

Carwardine J Nicol S Van Leeuwen S Walters B Firn J Reeson AMartin T G and Chades I (2014) Priority threat management forPilbara species of conservation significance CSIRO EcosystemSciences Brisbane

Cook A (2010) Habitat use and home-range of the northern quollDasyurushallucatus effects of fire MSc Thesis University ofWestern AustraliaPerth

Department of Sustainability Environment Water Population andCommunities (2012) Environment and Biodiversity Conservation Act1999 Environmental Offsets Policy Department of SustainabilityEnvironment Water Population and Communities Canberra

Doherty T S Davis R A Etten E J Algar D Collier N DickmanC R Edwards G Masters P Palmer R and Robinson S (2015a)A continental-scale analysis of feral cat diet in Australia Journal ofBiogeography 42 964ndash975 doi101111jbi12469

Doherty T S Dickman C R Nimmo D G and Ritchie E G (2015b)Multiple threats or multiplying the threats Interactions betweeninvasive predators and other ecological disturbances BiologicalConservation 190 60ndash68 doi101016jbiocon201505013

Dunlop J Cook A and Lees J (2014a) BHP Billiton Iron Ore Pilbaranorthern quoll monitoring project Final report May 2010ndashJune 2013Unpublished report prepared for BHP Billiton Iron Ore Department ofParks and Wildlife Perth

Dunlop J Cook A and Morris K (2014b) Pilbara northern quollproject surveying and monitoring Dasyurus hallucatus in the PilbaraWestern Australia Department of Parks and Wildlife Perth Availableat librarydpawwagovau

Florance D Webb J K Dempster T Kearney M R Worthing A andLetnic M (2011) Excluding access to invasion hubs can contain thespread of an invasive vertebrate Proceedings of the Royal Society ofLondon B Biological Sciences 278 2900ndash2908 doi101098rspb20110032

Gaillard J-M Hebblewhite M Loison A Fuller M Powell R BasilleM and Van Moorter B (2010) Habitatndashperformance relationshipsfinding the right metric at a given spatial scale PhilosophicalTransactions of the Royal Society of London Series B BiologicalSciences 365 2255ndash2265 doi101098rstb20100085

Glen A S and Dickman C R (2008) Niche overlap between marsupialand eutherian carnivores does competition threaten the endangeredspotted-tailed quoll Journal of Applied Ecology 45 700ndash707 doi101111j1365-2664200701449x

Glen A S Pennay M Dickman C R Wintle B A and Firestone K B(2011) Diets of sympatric native and introduced carnivores in theBarrington Tops eastern Australia Austral Ecology 36 290ndash296doi101111j1442-9993201002149x

Hennig P (2004) A brief land use history In lsquoAn Inventory and ConditionSurvey of the Pilbara Region Western Australiarsquo (Eds A M E VanVreeswyk A L Payne K A Leighton and P Hennig) TechnicalBulletin No 92 pp 13ndash18 (Western Australian Department ofAgriculture Perth)

Hohnen R Ashby J Tuft K and McGregor H (2013) Individualidentification of northern quolls (Dasyurus hallucatus) using remotecameras Australian Mammalogy 35 131ndash135 doi101071AM12015

L Australian Mammalogy V A Cramer et al

How R A Spencer P B S and Schmitt L H (2009) Island populationshave high conservation value for northern Australiarsquos top marsupialpredator ahead of a threatening process Journal of Zoology 278206ndash217 doi101111j1469-7998200900569x

Johnson C N Isaac J L and Fisher D O (2007) Rarity of a top predatortriggers continent-wide collapse of mammal prey dingoes andmarsupials in Australia Proceedings of the Royal Society of LondonSeries B Biological Sciences 274 341ndash346 doi101098rspb20063711

Jones M E (2000) Road upgrade road mortality and remedial measuresimpacts on a population of eastern quolls and Tasmanian devilsWildlifeResearch 27 289ndash296 doi101071WR98069

Kearney M Phillips B L Tracy C R Christian K A Betts G andPorter W P (2008) Modelling species distributions without usingspecies distributions the cane toad in Australia under current andfuture climates Ecography 31 423ndash434 doi101111j0906-7590200805457x

Kendrick P and Stanley F (2002) Pilbara 4 (PIL4 ndash Roebourne synopsis)In lsquoA Biodiversity Audit of Western Australiarsquos 53 BiogeographicalSubregions in 2002rsquo (Eds J E May and N L McKenzie) pp 581ndash593(Department of Conservation and Land Management Perth)

Kennedy M Phillips B L Legge S Murphy S A and Faulkner R A(2012) Do dingoes suppress the activity of feral cats in northernAustralia Austral Ecology 37 134ndash139 doi101111j1442-9993201102256x

Kiesecker J M Copeland H Pocewicz A and McKenney B (2010)Development by design blending landscape-level planning with themitigation hierarchy Frontiers in Ecology and the Environment 8261ndash266 doi101890090005

King D R and Smith L A (1985) The distribution of the European red fox(Vulpes vulpes) in Western Australia Records of the Western AustralianMuseum 12 197ndash205

Knight A T Cowling R M Rouget M Balmford A Lombard A Tand Campbell B M (2008) Knowing but not doing selecting priorityconservation areas and the researchndashimplementation gap ConservationBiology 22 610ndash617 doi101111j1523-1739200800914x

Laurance W F Koster H Grooten M Anderson A B Zuidema P AZwick S Zagt R J Lynam A J Linkie M and Anten N P (2012)Making conservation research more relevant for conservationpractitioners Biological Conservation 153 164ndash168 doi101016jbiocon201205012

Legge S Murphy S Heathcote J Flaxman E Augusteyn J andCrossman M (2008) The short-term effects of an extensive and high-intensity fire on vertebrates in the tropical savannas of the centralKimberley northern Australia Wildlife Research 35 33ndash43 doi101071WR07016

Lester D (2015) Effective wildlife roadkill mitigation Journal of Trafficand Transportation Engineering 3 42ndash51 doi10172652328-2142201501005

Letnic M Tamayo B and Dickman C R (2005) The responsesof mammals to La Nintildea (El Nintildeo Southern Oscillation)-associatedrainfall predation and wildfire in central Australia Journal ofMammalogy 86 689ndash703 doi1016441545-1542(2005)086[0689TROMTL]20CO2

Letnic M Koch F Gordon C Crowther M S and Dickman C R(2009) Keystone effects of an alien top-predator stem extinctions ofnative mammals Proceedings of the Royal Society of London Series BBiological Sciences 276 3249ndash3256 doi101098rspb20090574

Lindenmayer D B (2015) Continental-level biodiversity collapseProceedings of the National Academy of Sciences of the United States ofAmerica 112 4514ndash4515 doi101073pnas1502766112

Lindenmayer D B Piggott M P and Wintle B A (2013) Counting thebooks while the library burns why conservation monitoring programs

need a plan for action Frontiers in Ecology and the Environment 11549ndash555 doi101890120220

Maron M Hobbs R J Moilanen A Matthews J W Christie KGardner T A Keith D A Lindenmayer D B and McAlpine C A(2012) Faustian bargains Restoration realities in the context ofbiodiversity offset policies Biological Conservation 155 141ndash148doi101016jbiocon201206003

McCall S C McCarthy M A van der Ree R Harper M J Cesarini Sand Soanes K (2010) Evidence that a highway reduces apparentsurvival rates of squirrel gliders Ecology and Society 15 27

McGregor R A Stokes V L and Craig M D (2014) Does forestrestoration in fragmented landscapes provide habitat for a wide-ranging carnivore Animal Conservation 17 467ndash475 doi101111acv12112

McKenzie N L Burbidge A A Baynes A Brereton R N DickmanC R Gordon G Gibson L A Menkhorst P W Robinson A CWilliams M R and Woinarski J C Z (2007) Analysis of factorsimplicated in the recent decline of Australiarsquos mammal fauna Journal ofBiogeography 34 597ndash611 doi101111j1365-2699200601639x

McKenzie N L van Leeuwen S and Pinder AM (2009) Introduction tothe Pilbara Biodiversity Survey 2002ndash2007 Records of the WesternAustralian Museum 78(Suppl) 3ndash89 doi1018195issn0313-122x78(1)2009003-089

Morris K Johnson B Orell P Gaikhorst G Wane A and MoroD (2003) Recovery of the threatened chuditch (Dasyurus geoffroii)a case study In lsquoPredators with Pouches The Biology of CarnivorousMarsupialsrsquo (Eds M Jones C Dickman and M Archer) pp 435ndash451(CSIRO Publishing Melbourne)

Nelson J L Scroggie M P and Belcher C A (2014) Developing acamera trap survey protocol to detect a rare marsupial carnivore thespotted-tailed quoll (Dasyurus maculatus) In lsquoCamera TrappingWildlife Management and Researchrsquo (Eds P D Meek and PJ Fleming) pp 271ndash279 (CSIRO Publishing Melbourne)

Newsome T M Dellinger J A Pavey C R Ripple W J Shores C RWirsing A J and Dickman C R (2015) The ecological effects ofproviding resource subsidies to predators Global Ecology andBiogeography 24 1ndash11 doi101111geb12236

OrsquoDonnell S Webb J K and Shine R (2010) Conditioned tasteaversion enhances the survival of an endangered predator imperilled bya toxic invader Journal of Applied Ecology 47 558ndash565 doi101111j1365-2664201001802x

Oakwood M (1997) The ecology of the northern quoll Dasyurushallucatus PhD Thesis Australian National University Canberra

Oakwood M (2000) Reproduction and demography of the northern quollDasyurus hallucatus in the lowland savanna of northern AustraliaAustralian Journal of Zoology 48 519ndash539 doi101071ZO00028

Oakwood M (2002) Spatial and social organization of a carnivorousmarsupial Dasyurus hallucatus (Marsupialia Dasyuridae) Journal ofZoology 257 237ndash248 doi101017S0952836902000833

Oakwood M and Foster P (2008) Monitoring extinction of the northernquoll Australian Academy of Science Newsletter 71 6

Palomares F and Caro T M (1999) Interspecific killing amongmammalian carnivores American Naturalist 153 492ndash508 doi101086303189

Pavey C R and Bastin G (2014) lsquoAustralian Rangelands and ClimateChange ndash Invasive Animalsrsquo (Ninti One Limited amp CSIRO AliceSprings)

Pierson J C Barton P S Lane PW and Lindenmayer D B (2015) Canhabitat surrogates predict the response of target species to landscapechange Biological Conservation 184 1ndash10 doi101016jbiocon201412017

Pollock A B (1999) Notes on status distribution and diet of northernquoll Dasyurus hallucatus in the MackayndashBowen area mideastern

Research priorities for the northern quoll Australian Mammalogy M

Queensland Australian Zoologist 31 388ndash395 doi107882AZ1999040

Ramanaidou E R and Morris R C (2010) A synopsis of the channeliron deposits of the Hamersley Province Western AustraliaTransactions of the Institutions of Mining and Metallurgy Section BApplied Earth Science 119 56ndash59 doi101179037174510X12853354810624

Schmitt L H Bradley A J Kemper C M Kitchener D J HumphreysW F and How R A (1989) Ecology and physiology of the northernquoll Dasyurus hallucatus (Marsupialia Dasyuridae) at MitchellPlateau Kimberley Western Australia Journal of Zoology 217539ndash558 doi101111j1469-79981989tb02510x

Short J and Smith A (1994) Mammal decline and recovery in AustraliaJournal of Mammalogy 75 288ndash297 doi1023071382547

Spencer P How R Hillyer M Cook A Morris K Stevenson C andUmbrello L (2013) Genetic analysis of northern quolls from the PilbaraRegion ofWestern Australia Year one ndash final report Unpublished reportDepartment of Sustainability Environment Water Population andCommunities Canberra and Department of Parks and Wildlife Perth

Sutherland W J Fleishman E Mascia M B Pretty J and Rudd M A(2011) Methods for collaboratively identifying research priorities andemerging issues in science and policyMethods in Ecology and Evolution2 238ndash247 doi101111j2041-210X201000083x

Sutherst R W Floyd R B and Maywald G F (1996) The potentialgeographical distribution of the cane toad Bufo marinus L in AustraliaConservation Biology 10 294ndash299 doi101046j1523-1739199610010294x

Taylor B D and Goldingay R L (2003) Cutting the carnage wildlifeusage of road culverts in north-eastern New South Wales WildlifeResearch 30 529ndash537 doi101071WR01062

Tingley R Phillips B L Letnic M Brown G P Shine R and BairdS J (2013) Identifying optimal barriers to halt the invasion of canetoads Rhinella marina in arid Australia Journal of Applied Ecology 50129ndash137 doi1011111365-266412021

Van Vreeswyk A M E Payne A L Leighton K A and HennigP (2004) An Inventory and Condition Survey of the Pilbara RegionWestern Australia Technical Bulletin No 92 Department of AgricultureWestern Australia Perth

Webb J Legge S Tuft K Cremona T and Austin C (2015) Can wemitigate cane toad impacts on northern quolls Final report National

Environmental Research Program Northern Australia Hub CharlesDarwin University Darwin

Wintle B A Walshe T V Parris K M and McCarthy M A (2012)Designing occupancy surveys and interpreting non-detection whenobservations are imperfect Diversity amp Distributions 18 417ndash424doi101111j1472-4642201100874x

Woinarski J Menkhorst K Gambold N and Braithwaite R W (1992)Mammals of the Bungle Bungle area In lsquoA Survey of the Wildlife andVegetation of Purnululu (Bungle Bungle) National Park and AdjacentArearsquo (Ed J C Z Woinarski) Research Bulletin No 6 pp 53ndash67(CSIRO Division of Wildlife and Ecology Darwin)

Woinarski J C Z Risler J and Kean L (2004) Response of vegetationand vertebrate fauna to 23 years of fire exclusion in a tropical Eucalyptusopen forest Northern Territory Australia Austral Ecology 29 156ndash176doi101111j1442-9993200401333x

Woinarski J C Z Oakwood M Winter J Burnett S Milne D FosterP Myles H and Holmes B (2008) Surviving the toads patterns ofpersistence of the northern quoll Dasyurus hallucatus in QueenslandReport submitted to the Natural Heritage Trust Strategic ReserveProgram as a component of project 2005162 Monitoring andManagement of Cane Toad Impact in theNorthern Territory Available athttplrmntgovauplants-and-animalsinformation-and-publicationspublications2008 [accessed 14 November 2013]

Woinarski J C Z Armstrong M Brennan K Fisher A Griffiths A DHill B Milne D J Palmer C Ward S Watson M Winderlich Sand Young S (2010) Monitoring indicates rapid and severe decline ofnative small mammals in Kakadu National Park northern AustraliaWildlife Research 37 116ndash126 doi101071WR09125

Woinarski J Burbidge A and Harrison P (2014) lsquoAction Plan forAustralian Mammals 2012rsquo (CSIRO Publishing Melbourne)

Woinarski J C Burbidge A A and Harrison P L (2015) Ongoingunraveling of a continental fauna decline and extinction of Australianmammals since European settlement Proceedings of the NationalAcademy of Sciences of the United States of America 112 4531ndash4540doi101073pnas1417301112

Woolley P A Krajewski C and Westerman M (2015) Phylogeneticrelationships within Dasyurus (Dasyuromorphia Dasyuridae) quollsystematics based on molecular evidence and male characteristicsJournal of Mammalogy 96 37ndash46 doi101093jmammalgyu028

N Australian Mammalogy V A Cramer et al

wwwpublishcsiroaujournalsam

Page 6: Research priorities for the northern quoll (Dasyurus ......The Pilbara population of the northern quoll (Dasyurus hallucatus) has been seldom studied, and the impacts of threats such

investment of time and resources The limited amount anduncertain quality of occurrence data has been a major restrictionon the utility of a species distribution model (SDM) and apopulation viability analysis (PVA) for the northern quoll in thePilbara (Cadenhead et al 2015 see below) While survey andmonitoring per se is not a research activity that directly informsmanagement or threat mitigation carefully implementedmonitoring programs are essential to identify many elements ofa speciesrsquo ecology to detect changes in population and toquantify the ecological effectiveness and cost-effectivenessof management interventions (Lindenmayer et al 2013Lindenmayer 2015)

Parks and Wildlife recently released revised protocols forthe long-term monitoring of known populations of thenorthern quoll (Dunlop et al 2014b) and the effectiveness ofthese protocols in presenceabsence surveys should be testedThe cost-effectiveness of using camera traps for wildlifesurveys in remote locations such as the Pilbara makes their useappealing Hohnen et al (2013) described how the strategicplacement of cameras and baits combined with taking a greaternumber of photographs per trigger allows the capture ofimages that can be used to identify individual northern quollsfrom their spot patterns Remote cameras have also been shownto be a viable method for estimating population sizes of thenorthern quoll in areas with both dense (Austin 2014) andsmall populations (Webb et al 2015) Further testing of a rangeof parameters (eg see Nelson et al 2014) to develop a robuststandard protocol for the Pilbara is required along with acomparison of the cost-effectiveness and any ethicalreducedhandling benefits of using remote cameras versus live-trappingfor estimating population sizes in Pilbara landscapes

Priority 2 Improve our understanding of habitatrequirements

Define areas of critical habitat for the northern quollin the Pilbara

The Department of Agriculture and Food Western Australiahas mapped 102 land systems across the Pilbara bioregionbased on several biophysical components including landformgeology soils vegetation and drainage patterns (Van Vreeswyket al 2004) A review of records for the northern quoll in thePilbara showed that 65 of records were from the RobeCapricorn and Boolaloo land systems (Table 2) which compriselow plateaux mesas hills and ridges of varying geology(limonite granite and sandstone respectively) Most recentsurvey effort in the Pilbara is biased towards mineral-bearingformations or areas of mining-related infrastructure (eg railand road corridors) (Fig 1) Further investigation of theserecords is required to assess whether these land systems areindeed preferred habitat for northern quolls as opposed to anartefact of survey effort The comparatively fewer records thatcoincide with the conservation estate in the region (Fig 1) arealso likely an artefact of survey effort

The first step in defining areas of critical habitat for thenorthern quoll in the Pilbara is to improve our understanding ofhabitat use particularly in terms of defining habitat suitability atfiner scales than is currently available As part of the strategic

environmental assessments being undertaken for the Pilbarabioregion an SDM has been developed for the northern quoll(Cadenhead et al 2015) Uncertainty about the quality of theoccurrence data available the limited amount of data availableas well as the bias towards data collection in areas proposed fordevelopment constrained the level of certainty around theoutcomes of the model Improved model certainty requiresfurther collection of contemporary species-occurrence data andthe development of finer-resolution GIS layers particularly onethat captures finer-scale geological attributes such as mesas orrocky outcrops considered as highest-quality habitat for thenorthern quoll (Cadenhead et al 2015)

Better definition of the habitat attributes favoured by thenorthern quoll will allow further refinement of the GIS variablesincluded in the SDM or allow the development of predictivesite-based habitat models Information on how northern quollsuse habitat can be gained through direct observation of howthey interact with habitat features or from the association ofhabitat features with the presence of quolls as determined byGPS receivers (Gaillard et al 2010) This level of detail iscurrently not available from survey records returned as part ofenvironmental impact assessments but collection of habitatdata using the protocols outlined by Dunlop et al (2014b) infuture surveys would rapidly increase the quality of dataavailable

Understand how disturbance affects habitat qualityand connectivity

The alteration of fire regimes has been the most pervasivelandscape-scale change in the Pilbara Between 1993 and 2006over 72 of the region was burnt with more than 28 of thearea burnt two or more times (McKenzie et al 2009) Woinarskiet al (2004) found that northern quolls may be able to tolerate amoderately high frequency of fire as long as these fires werelow-intensity early-season burns and the impacts of fire onvegetation structure and composition were not exacerbatedby the grazing of introduced herbivores In the Pilbarahowever as with many other grazing regions of Australia earlypastoralists often overestimated the productivity and resilienceof native vegetation to sheep and cattle grazing which led tosevere degradation of vegetation particularly along riverfrontages (Hennig 2004) The soil degradation and loss ofvegetative cover caused by intense grazing pressure in thesouth Kimberley is thought likely to have contributed to thesubstantial decline or disappearance of medium-sized animalsincluding the northern quoll from the Bungle Bungle area(Woinarski et al 1992) Death by predation is most likely in openareas (Oakwood 2000) and therefore the effects of fire andgrazing on northern quoll populations cannot be consideredwithout also taking account of likely increases in mortality due topredation (see lsquoIntroduced predatorsrsquo below)

Understanding how the localised but intensive activityassociated with mining affects habitat quality and habitat useby the northern quoll in areas surrounding development islimited A recent study that examined the use of ballast quarriesand paired undisturbed sites by northern quolls found that theywere foraging denning and showing signs of breeding at thequarry sites where greater numbers of quolls were captured(Dunlop et al 2014a) This was likely due to the combination of

F Australian Mammalogy V A Cramer et al

habitat complexity (eg diversity of boulder sizes) and thepresence of permanent water in the quarry which may haveincreased prey availability at these sites

Movement of animals through the landscape for dispersaland repopulation is another important consideration for a speciesin which both sexes tend towards semelparity and is thereforepredisposed to local extinction (Oakwood 1997) Improvedunderstanding of how the northern quoll (particularly dispersingmales) uses areas of lower habitat quality and the identificationof such habitat in habitat-suitability models will allow futurelandscape-scale strategic assessments to better predict thecombined impacts of disturbance on the ability of northern quolls

to move through the landscape Analysis of regional populationgenetics will also improve our understanding of the level ofdispersal occurring across the landscape

Priority 3 Population dynamics

Better understand the population dynamics of thenorthern quoll in the Pilbara

Little information is available on the number ofsubpopulations of northern quoll in the Pilbara how thesesubpopulations interact how stable they are in space and timeand how they respond to disturbance As no demographic

Table 2 Records of the northern quoll in each land system type in the PilbaraRecords were current as of 23 May 2014

Land system Area (km2) No of records Land system description

Augustus 61 1 Rugged ranges hills ridges and plateaux supporting mulga shrublands and hard spinifex grasslandsBoolaloo 1502 245 Granite hills domes and tor fields and sandy plains with shrubby spinifex grasslandsBoolgeeda 7748 29 Stony lower slopes and plains below hill systems supporting hard and soft spinifex grasslands andmulga

shrublandsCalcrete 1444 2 Low calcrete platforms and plains supporting shrubby hard spinifex grasslandsCallawa 1003 9 Highly dissected low hills mesas and gravelly plains of sandstone and conglomerate supporting soft and

hard spinifex grasslandsCapricorn 5296 257 Hills and ridges of sandstone and dolomite shrubby hard and soft spinifex grasslandsCoongimah 3244 12 Plateau surfaces low hills with steep slopes and undulating uplands supporting hard spinifex grasslandsDune 138 1 Dune fields supporting soft spinifex grasslandsGranitic 4020 1 Rugged granitic hills supporting shrubby hard and soft spinifex grasslandsHorseflat 1261 4 Gilgaied clay plains supporting tussock grasslands and minor grassy snakewood shrublandsLittoral 1577 3 Bare coastal mudflats with mangroves on seaward fringes samphire flats sandy islands coastal dunes

and beachesMacroy 13 095 94 Stony plains and occasional tor fields based on granite supporting hard and soft spinifex grasslandsMcKay 4202 21 Hills ridges plateau remnants and breakaways of metasedimentary rocks supporting hard spinifex

grasslandsMallina 2557 8 Sandy surfaced alluvial plains supporting soft spinifex (and occasionally hard spinifex) grasslandsMarandoo 459 2 Basalt hills and restricted stony plains supporting grassy mulga shrublandsMosquito 1840 3 Stony plains and prominent ridges of schist and other metamorphic rocks supporting hard spinifex

grasslandsNewman 14 580 81 Rugged jaspilite plateaux ridges and mountains supporting hard spinifex grasslandsOnslow 424 1 Sandplains dunes and clay plains supporting soft spinifex grasslands and minor tussock grasslandsParaburdoo 565 3 Basalt-derived stony gilgai plains and stony plains supporting snakewood and mulga shrublands with

spinifex and tussock grasslandsParadise 1479 8 Alluvial plains supporting soft spinifex grasslands and tussock grasslandsRiver 4088 37 Active flood plains major rivers supporting grassy eucalypt woodlands tussock grasslands and soft

spinifex grasslandsRobe 865 369 Low limonite mesas and buttes supporting soft spinifex (and occasionally hard spinifex) grasslandsRocklea 22 993 40 Basalt hills plateaux lower slopes and minor stony plains supporting hard spinifex (and occasionally

soft spinifex) grasslandsRuth 346 36 Hills and ridges of volcanic and other rocks supporting hard spinifex (occasionally soft spinifex)

grasslandsStuart 1794 3 Gently undulating stony plains supporting hard and soft spinifex grassland and snakewood shrublandsTaylor 129 3 Stony plains and isolated low hills of sedimentary rocks supporting hard and soft spinifex grasslandsTalga 2124 9 Hills and ridges of greenstone and chert and stony plains supporting hard and soft spinifex grasslandsTuree 581 2 Stony alluvial plains with gilgaied and non-gilgaied surfaces supporting tussock grasslands and grassy

shrublandsUaroo 7681 37 Broad sandy plains supporting shrubby hard and soft spinifex grasslandsUrandy 1311 6 Stony plains alluvial plains and drainage lines supporting shrubby soft spinifex grasslandsWhite Springs 266 4 Stony gilgai plains supporting tussock grasslands and hard spinifex grasslandsWona 1815 10 Basalt upland gilgai plains supporting tussock grasslands and minor hard spinifex grasslandsYamerina 1207 1 Floodplains and deltaic deposits supporting tussock grasslands grassy woodlands and minor halophytic

low shrublands

Research priorities for the northern quoll Australian Mammalogy G

studies have been undertaken in the region Cadenhead et al(2015) used demographic information from the NorthernTerritory and Kimberley populations in their PVA Uncertaintyabout data quality and the limited information available ondemographic parameters for the Pilbara constrained the level ofcertainty around the outcomes of the PVA The model doeshowever provide a framework into which new data can beincorporated as they become available For example preliminarydata from the Pilbara monitoring program suggest that theproportion of male die-off in the Pilbara population is lowerthan in the Kimberley and Northern Territory populations(Dunlop unpubl data) Cadenhead et al (2015) stress the needfor more demographic and occurrence data to be collectedbefore further development decisions are made (see lsquoDevelopappropriate and standardised survey and monitoring methodsrsquoand lsquoImprove our understanding of habitat requirementsrsquo above)

Investigate population structure and the interactionbetween populations

Analysis of the genetic diversity and population structure ofthe Pilbara population of the northern quoll is ongoingOnly small differences in genetic diversity as measured byheterozygosity were found between samples obtained at 16locations within the Pilbara with the exception of the isolatedpopulation on Dolphin Island (Spencer et al 2013) This resultindicates a high level of movement across the landscape byindividuals during mating and dispersal phases Spatial structureanalysis of samples collected from around Pannawonicasuggested a lsquoneighbour sizersquo of ~15 km (Spencer et al 2013)Current data show no evidence of genetic bottleneckssuggesting that if there has been a population decline in thePilbara that this has not affected genetic diversity A usefulapproach would be to combine genetic information with datafrom radio-tracking or GPS collars to better understand theuse of lsquolandscapersquo space and how this influences populationdynamics

Priority 4 Key threats and the interaction of these threats

Introduced predators

Introduced cats and foxes are considered to be one of themain causes of the decline and extinction of the critical-weight-range mammals (35 g to 55 kg) in Australia (Short and Smith1994 McKenzie et al 2007 Woinarski et al 2015) but theirimpacts on the northern quoll are less clear A recent review offeral cat diets across Australia found that northern quolls werepresent in the diet of feral cats (Doherty et al 2015a) andWoinarski et al (2014) consider the threat posed by cats to thenorthern quoll to be severe although the impacts of cats may bereduced in rugged refuge areas The impacts of cats on othermuch larger species of quoll appear to be inconsistent withsome evidence of coexistence between cats and quolls beforethe arrival of foxes (Glen and Dickman 2008 and referencescontained within) Due to the limited overlap in their distributionthroughout Australia the threat posed by foxes to northernquolls is less understood than for cats and Woinarski et al(2014) consider the threat posed by foxes to be moderate on anational level Foxes are common in coastal near-coastal and

riparian areas of the Pilbara and extend as far north as Broome(King and Smith 1985 Kendrick and Stanley 2002) They arerenowned for their aggression towards smaller carnivores(Palomares and Caro 1999) and neither the larger chuditch(Dasyurus geoffroii) nor spotted-tail quoll (Dasyurus maculatus)are known to coexist with large numbers of foxes (A Glen perscomm) The interactions between quolls cats and foxesare potentially more complex than for other medium-sizedmarsupials due to niche overlap in diet and competition for densites with the larger spotted-tailed quoll and chuditch bothexperiencing exploitation and interference competition fromfoxes (Morris et al 2003 Glen et al 2011) Competitionbetween feral cats and the northern quoll may potentially beintense during periods of low rainfall in the Pilbara when preyavailability is low Understanding whether foxes and cats aresignificant predators of or competitors with the northern quollin the Pilbara and how this affects their population viabilitywill clarify the level of risk posed by these predators Thesausage-style baits used for feral cats (eg Eradicatreg andCuriosityreg) are potentially attractive to the northern quoll withthe risk of mortality or reduced fitness Trials to assess themagnitude of the risks from Eradicatreg baits are currentlyunderway in the Pilbara

The role of dingoes and wild dogs in the mortality of andinteractions between mesopredators (both native and introduced)requires further investigation Dingoes are known to kill northernquolls butmay not eat these kills (Oakwood 1997) A recent studyin the Northern Territory found that camp dogs and dingoes werethe major source of mortality for northern quolls at the study site(Webb et al 2015) and roaming domestic dogs were identified asa cause of mortality for northern quolls in Queensland (Pollock1999) Human resource subsidies (eg stock carrion in rangelandareas) can artificially support predator populations thus indirectlyincreasing predation pressure on prey (Newsome et al 2015)A dynamic debate currently exists around the role of the dingo inregulating populations of feral cats and foxes and the implicationsof this for the conservation of the critical-weight-range mammalsin Australia (see Johnson et al 2007 Letnic et al 2009 Allenet al 2011 2013 Kennedy et al 2012) Understanding thepopulation density at which the role of the dingo as a regulator ofintroduced predators outweighs its own impact as a direct predatorof the northern quoll will inform whether dingowild dog controlwould benefit the northern quoll

Cane toads

The introduced cane toad now occupies more than half thehistoric range of the northern quoll and has caused an immediatecollapse or extirpation of northern quoll populations in theareas it has invaded (Burnett 1997 Oakwood and Foster2008 Woinarski et al 2008 2010) Earlier modelling work thatconsidered only climate parameters as well as more recentwork that incorporates future climate change suggest thatcane toads will not invade the Pilbara (Sutherst et al 1996Pavey and Bastin 2014) Other recent work that considers thefundamental niche of the cane toad and incorporates the locationof artificial watering points predicts that they will colonisethe Pilbara from the Kimberley via the coastal edge of theGreat Sandy Desert (Kearney et al 2008 Florance et al 2011

H Australian Mammalogy V A Cramer et al

Tingley et al 2013) This has created some level of uncertaintyabout the likely impact of cane toads on northern quolls in thePilbara Strategies for cane toad management were not rankedhighly by participants in the threat-management frameworkdeveloped by Carwardine et al (2014) which the authorsexplained as possibly due to uncertainty about how far canetoads will spread into the region

The model developed by Tingley et al (2013) suggested thatthe closure of 100 artificial water bodies at one of three potentiallocations along the coastal corridor between the Kimberley andthe Pilbara would prevent the invasion of cane toads into thePilbara While their model did account for uncharacteristicallywet years every three decades the dispersal of toads betweenwater bodies was determined by the number of rainy daysoccurring during the year (ie toads could only disperse betweenwater bodies for three days following rainfall) rather thanrainfall amount the associated likelihood of flooding and theavailability of free water across the landscape The cyclonic andthunderstorm nature of much of the rainfall in the north-west ofWestern Australia means that the region is prone to significantand sometimes extended flooding In April 1934 the countryaround Anna Plains (250 km south of Broome) was flooded forhundreds of miles when 27 inches (686mm) of rain fell infour days (Anon 1934) In March 2004 over 350mm of rain wasrecorded within a 48-h period at Telfer on the southern edge ofthe Great Sandy Desert due to Tropical Cyclone Fay Thiscaused the flooding of the De Grey and Oakover river systemsand flooded 140 km of the Telfer Access Road with floodwaterstaking several months to subside (wwwbomgovau) Extendingthe model of Tingley et al (2013) to account for such events willhelp clarify whether closing artificial water bodies is likely toprevent the invasion of cane toads into the Pilbara or may onlyassist in delaying their inevitable invasion

Comparison of SDMs (eg Cadenhead et al 2015) fornorthern quoll with fundamental niche models for cane toads(eg see Kearney et al 2008) within the Pilbara may identifypotential refuge areas for the northern quoll and these areasshould be prioritised for conservation actions Because theinvasion point of cane toads into the Pilbara is likely to be narrow(beginning around the mouth of the De Grey River Tingleyet al 2013) a proactive approach to training northern quollsto avoid cane toads may prove beneficial Conditioned taste-aversion (CTA) has been used successfully to train captive-reared northern quolls to avoid cane toads by offering themsmall dead toads infused with thiabendazole before theirrelease into the wild (OrsquoDonnell et al 2010)Monitoring of thesequolls after their release showed that some of their offspringhad learnt to avoid eating cane toads either through ingestionof small non-lethal toads or via social learning from theirmothers (Webb et al 2015) Fields trials of toad-aversion baits(either sausages or meatballs made from toad flesh andcontaining thiabendazole) are currently underway in the centralKimberley where cane toads were expected to invade in late2015 (Webb et al 2015) Further testing of CTA approachesunder a variety of field conditions is required The developmentof stable long-lasting baits suitable for aerial deployment isconsidered a pressing need as well as a captive colony ofnorthern quolls upon which to test the efficacy of toad-aversionbaits for inducing aversion to live toads (Webb et al 2015)

Infrastructure development and interactionswith humans

Infrastructure development may increase the likelihood ofnorthern quoll deaths due to motor vehicle strikes or beingcrushed in machinery Significant chuditch mortality wasobserved along mine access roads in the south-west of WesternAustralia (McGregor et al 2014) and the widening and sealingof an access road dramatically increased the number of roadkills and led to the extirpation of the resident population ofeastern quolls (D viverrinus) in Tasmania (Jones 2000)Infrastructure corridors such as high-tension power-lineeasements and ore-haulage railways are increasingly beinginstalled throughout the range of the northern quoll Althoughthese do not create the same mortality risk as roads they bisectand fragment habitat create large open spaces and potentiallyincrease predation risk

A recent small-scale radio-tracking study (see Dunlop et al2014a) around several quarries in the Pilbara found that northernquolls crossed roads but not railway lines The short-term natureof this work and the limited number of quolls able to be trackedmeans that the influence of roads railway lines and infrastructurecorridors on the movement and mortality of northern quollsrequires further assessment Using radio-tracking techniques todetermine crossing rates and road kill events may be difficultover large areas so population studies comparing groups ofquolls near major roads with those not affected by infrastructure(eg McCall et al 2010) may provide a more feasible meansof assessing the impacts of roads If vehicle strikes are foundto significantly contribute to the mortality of a local populationof the northern quoll then mitigation methods should betrialled on sections of road identified as road-kill hotspotsCulverts and underpasses have been shown to be used by avariety of small and medium-sized mammals and frogs (Taylorand Goldingay 2003) while installing rumble strips on roadshas led to decreased rates of road-kill of nocturnal animals(Lester 2015)

Interactions between threats

At present wildfire the subsequent loss of vegetation coverand increased predation of northern quolls is likely to be themost significant of the interactions between threats The directeffects of wildfire have been found to be less significant inthe subsequent decline of small to medium-sized mammalsparticularly for denning species such as the northern quoll thanthe indirect consequences of reduced resource availability andthe increased risk of predation due to lack of cover (Oakwood2000 Letnic et al 2005 McKenzie et al 2007 Legge et al2008 Doherty et al 2015b) Male northern quolls are perhapsmost vulnerable to such losses of cover during the breedingseason when their range can extend to over 100 ha (Oakwood2002) or where males survive after breeding their foragingrange is extended to gain the nutrition required to recover fromtheir poor condition (Cook 2010) Loss of cover due to late dry-season fires at East Alligator River in the Northern Territorymeant little cover was available for young quolls when theybegan foraging with dog and dingo predation a major source ofmortality (Webb et al 2015) The cumulative impacts of canetoad poisoning fire and predation are highly likely to lead to the

Research priorities for the northern quoll Australian Mammalogy I

extirpation of small and disjunct groups of northern quoll in thePilbara Population viability modelling of the East AlligatorRiver population suggested that high mortality from predationcombined with late dry-season burning was preventing therecovery of northern quoll populations after cane toad invasion

(Webb et al 2015) Improved knowledge of the severity of thesethreats to northern quolls in the Pilbara and their incorporationinto the SDM and PVA framework developed by Cadenheadet al (2015) will allow for better identification of priority areasfor integrated cane toad fire and predator management

Table 3 Expected timelines for the undertaking of research under each priority over the next 10 years and the funds currently allocated fromthe Pilbara Northern Quoll Offset Trust under each research priority

Rank Theme Research priority Current or proposed researchactivity

Time over next10 years

Current funding allocation(approximate)

1 Survey andmonitoring

Develop appropriate andstandardised survey andmonitoring methods

Develop monitoring protocol(traps and cameras) andinvestigate new methods

Ongoing survey and monitoringin conjunction with ResearchThemes 2 and 3 to better mapnorthern quoll distribution andto determine long-termpopulation trends

Years 1ndash2Years 3ndash10

$200 000 per annum over10 years for ResearchThemes 1 2 and 3(inclusive of salaries)

2 Habitatrequirements

Define areas of criticalhabitat for northernquoll in the Pilbara

Identify and describe criticalhabitat and undertake predictivemodelling of unsurveyed areasto inform further survey effort

Years 3ndash4 $50 000 for predictivemodelling (EdithCowan University)

Understand howdisturbance affectshabitat quality andconnectivity

Ongoing collection of data ondisturbance and habitat qualityin conjunction with ResearchThemes 1 and 4

Incorporation of new data intopredictive models

Years 4ndash10 See above

3 Populationdynamics

Better understand thepopulation dynamics ofthe northern quoll in thePilbara

Collect data on demographicparameters breeding dispersaland home-range use toincorporate into future PVA

Continued collectionof data inconjunction withpriorities 1 and 2during years 3ndash10

See above

Investigate populationstructure and theinteraction betweenpopulations

Analysis of existing tissuesamples and newly collectedsamples

Years 1ndash10 $120 000 for initial roundof genetic analysis(Years 3ndash5 MurdochUniversity)

4 Key threats Introduced predators Spatio-temporal interactionsbetween northern quolls andintroduced predators

Other projects as they arise

Years 2ndash4 (currentPhD projectUniversity ofQueensland)

$12 000 contribution tofield expenses

Cane toads Predictive modelling of canetoad distribution in the Pilbara

Years 3ndash4 Part of predictivemodelling by EdithCowan University

Infrastructuredevelopment andinteractions withhumans

Investigation of impacts oflinear infrastructure on northernquoll movement

Year 4 (currentHonours projectEdith CowanUniversity)

$15 000

Interactions betweenthreats

No on-ground projects currentlyplanned

Potential to be assessed throughPVA as data are collected underother research priorities

No funding currentlyallocated

5 Recolonisation ofrestored orartificial habitat

Determine the ability ofthe northern quoll torecolonise disturbedareas or coloniseartificial habitat

As projects arise Years 5ndash8 incollaboration withminesite restorationprograms

No funding currentlyallocated

J Australian Mammalogy V A Cramer et al

Priority 5 Recolonisation of restored andor artificial habitat

While anecdotal evidence suggests that northern quolls will denin waste rock dumps whether artificial or restored habitatprovides for long-term population viability is not known andlong-term assessment of created habitat is required Weacknowledge that a large gap exists between the desire to userestored or artificial habitat as an ecologically equivalent offsetand our limited ability to actually restore or re-create habitat thatis equivalent in structure composition and function particularlyon highly disturbed sites (Maron et al 2012) Yet there is a futureopportunity in the Pilbara to conduct lsquonaturalrsquo experiments onthe recolonisation of quolls after mining ceases including howto best design waste rock dumps so that habitat complexityand productivity is maximised by for example experimentingwith the size and positioning of boulders and their spatialarrangement in relation to surrounding landscape features Theuse of such artificial habitat should be tested in the first stagethrough direct monitoring of animals that have colonised orhave been translocated to the site Such experimentation isunderway at Atlas Ironrsquos Mt Dove operation (Atlas Iron 2012)Population modelling can then be used to predict futurepopulation sizes rates of growth and spread and the probabilityof long-term persistence in the context of various managementactions (eg predator control Armstrong and Reynolds 2012)

Resource allocation and timelines

While the allocation of resources and funding and the periodover which each of the research priorities was expected to beaddressed was not explicitly discussed during the workshopthe ranking of the priorities was taken to indicate the sequencein which the priorities should be addressed At the time of theworkshop some of the research questions identified werealready being investigated within various projects and a portionof the Pilbara Northern Quoll Offset Trust allocated to these InTable 3 we indicate the expected timelines over which someof the research programs are expected to be undertaken WhileAU$175M has been allocated to northern quoll research in thePilbara it is important to note that much of this amount is notcurrently available and will not become available until severalapproved mine developments have become operational InTable 3 we also indicate the proportion of offset funds expectedto be allocated to each research priority Research undertaken inother regions (eg research into CTA in the Kimberley regionahead of the cane toad invasion front) cannot be funded from thePilbara Northern Quoll Offset Trust but will help address someof the research questions identified here

Lessons learnt from the workshop process

The workshop approach used here was less formal than recentlarge-scale exercises used to elicit research priorities for emergingissues in science and policy at national or global levels (reviewedin Sutherland et al 2011) The focus in our (much smaller)workshop was on inclusivity rather than identifying participantsbased on purposive sampling of expertise or organisationalaffiliation We believe that the inclusive nature of the workshopwas a strength but we also recognise that this perhaps createdsome level of bias when ranking the research prioritiesAcknowledging both the experience and knowledge requirements

of those who work regularly in the Pilbara is an importantcomponent of improving the design of research programs that aimto improve conservation management in the region Yet theresearch priorities and their rankings are likely strongly reflectiveof the current information needs of most participants and maynot align with what conservation scientists believe to be themost critical areas of research for northern quoll conservation intothe future Baseline survey and monitoring are often the firstrequirements of a management or research plan for the northernquoll when offset conditions are prescribed and this was reflectedin the primary ranking given to the development of standardsurvey and monitoring methods We acknowledge the concernthat this could simply lead to offset funding being used to lsquomonitorthe extinctionrsquo of the northern quoll particularly after canetoads invade the Pilbara unless these data are used to triggermanagement actions (Oakwood and Foster 2008 Lindenmayeret al 2013) Clearly the currently limited information on habitatuse and demographic parameters for the northern quoll in thePilbara restricts the utility of more sophisticated researchapproaches such as developing robust SDM and PVA While thesevere threat that cane toads pose to the northern quoll wasrecognised this threat is not immediate in the region and mayexplain the less prominent ranking given to research to mitigatethe impacts of cane toads

The northern quoll has become emblematic of the dramaticand rapid species declines occurring in northern Australia andthe level of interest in this species was reflected in the highnumber of participants attending the workshop (one of six runby Parks and Wildlife) Given the number of attendees betterutilisation of the time available would have been gained byproviding participants with a brief before the workshopand asking them to contribute potential research questionsanonymously for interactive review by other participants Inthe exercises reviewed by Sutherland et al (2011) questionswere then discarded if they were not considered to meet criteriathat were previously agreed upon Some areas of researchthat gained some attention on the day may have been discardedif given greater consideration or interrogation before theworkshop For example there was some discussion duringthe workshop of investigating the use of habitat surrogates tomonitor the northern quoll yet research has shown that there islittle validity in using surrogates to monitor the abundance of asingle species (Pierson et al 2015)

Conclusions

Offset funds generated under the environmental approvalsprocess provide an opportunity to mobilise considerablefinancial resources for conservation (Kiesecker et al 2010) Yetthere remain significant conceptual and practical limitations tobe resolved to ensure that direct offsets provide measurableconservation outcomes that are ecologically equivalent to thelosses sustained by proceeding with the development (Maronet al 2012 Bull et al 2013) The collaborative identificationand ranking of research priorities undertaken here is the firststep in the process of developing and evaluating potentialconservation actions for the northern quoll in the Pilbara andserves as a mechanism to ensure that the research priorities ofstate and federal agencies reflects the knowledge needs of

Research priorities for the northern quoll Australian Mammalogy K

practitioners and proponents currently operating in the regionThe provision of offset funds over the next decade providesa unique opportunity to undertake research on a species ofconservation significance at a regional scale and over a longperiod through an integrated and strategic program of researchrather than the collection of ad hoc data While a morestructured approach to defining research questions before theworkshop may have led to greater critical reflection upon anddiscussion of some of the research priorities on the day theworkshop provided an opportunity for an open and diverse-ranging discussion in a collaborative environment amongstthe broader community of scientists environmental consultantsand mining proponents involved in the management andconservation of the northern quoll in the Pilbara

Acknowledgements

We gratefully acknowledge the financial support of Roy Hill who sponsoredthe workshop and partially funded VACrsquos salary We thank the Departmentof the Environmentrsquos Pilbara Taskforce especially Veronica Ritchie fortheir support of interested or affected parties The success of the workshopwas in great part due to the facilitation skills of Craig Salt of SustainableConsulting We thank all of the workshop participants for their open andcollaborative approach to the process Mike Bamford Eddie van EttenDavid Peacock and Peter Lyon provided useful comments on an earlier draftWe are grateful to John Woinarski for his constructive advice thatsubstantially improved the structure and clarity of the manuscript and ananonymous reviewer for careful and critical reading of the previousversion Lesley Gibson Russell Palmer and Al Glen generously discussedvarious aspects of the paper with VAC The map was prepared by CristinaRamalho

References

Allen B L Engeman R M and Allen L R (2011) Wild dogma anexamination of recent lsquoevidencersquo for dingo regulation of invasivemesopredator release in Australia Current Zoology 57 568ndash583

Allen B L Fleming P J Allen L R Engeman R M Ballard G andLeung L K-P (2013) As clear as mud a critical review of evidencefor the ecological roles of Australian dingoes Biological Conservation159 158ndash174 doi101016jbiocon201212004

Anon (1934) Floods in Western Australia Town isolated for week PerthMonday The Sydney Morning Herald 9

Armstrong D P and Reynolds M H (2012) Modelling reintroducedpopulations the state of the art and future directions In lsquoReintroductionBiology Integrating Science and Managementrsquo (Eds J G Ewen DP Armstrong K A Parker and P J Seddon) pp 165ndash222 (Wiley-Blackwell UK)

Atlas Iron (2012) Scope of works ndash Mount Dove artificial quoll habitatUnpublished report Atlas Iron Perth Available at httpwwwatlasironcomauIRMCompanyShowPageaspxCategoryId=190ampCPID=5424ampEID=94365410 [accessed 19 August 2015]

Austin C (2014) Can remote cameras accurately estimate populations ofthe endangered northern quoll BSc(Honours) Thesis University ofTechnology Sydney

Begg R J (1981) The small mammals of Little Nourlangie RockNT III Ecology ofDasyurus hallucatus the northern quoll (MarsupialiaDasyuridae) Wildlife Research 8 73ndash85 doi101071WR9810073

Braithwaite R W and Griffiths A D (1994) Demographic variationand range contraction in the northern quoll Dasyurus hallucatus(Marsupialia Dasyuridae)Wildlife Research 21 203ndash217 doi101071WR9940203

Bull J W Suttle K B Gordon A Singh N J and Milner-Gulland E J(2013) Biodiversity offsets in theory and practice Oryx 47 369ndash380doi101017S003060531200172X

Burnett S (1997) Colonizing cane toads cause population declines innative predators reliable anecdotal information and managementimplications Pacific Conservation Biology 3 65ndash72

Cadenhead N C R White A L and Wintle B A (2015) Identifyingconservation priorities and assessing impacts of proposed futuredevelopment in the Pilbara Bioregion in Western Australia Report bythe National Environment Research Program Decisions Hub to theDepartment of the Environment The University of MelbourneMelbourne

Carwardine J Nicol S Van Leeuwen S Walters B Firn J Reeson AMartin T G and Chades I (2014) Priority threat management forPilbara species of conservation significance CSIRO EcosystemSciences Brisbane

Cook A (2010) Habitat use and home-range of the northern quollDasyurushallucatus effects of fire MSc Thesis University ofWestern AustraliaPerth

Department of Sustainability Environment Water Population andCommunities (2012) Environment and Biodiversity Conservation Act1999 Environmental Offsets Policy Department of SustainabilityEnvironment Water Population and Communities Canberra

Doherty T S Davis R A Etten E J Algar D Collier N DickmanC R Edwards G Masters P Palmer R and Robinson S (2015a)A continental-scale analysis of feral cat diet in Australia Journal ofBiogeography 42 964ndash975 doi101111jbi12469

Doherty T S Dickman C R Nimmo D G and Ritchie E G (2015b)Multiple threats or multiplying the threats Interactions betweeninvasive predators and other ecological disturbances BiologicalConservation 190 60ndash68 doi101016jbiocon201505013

Dunlop J Cook A and Lees J (2014a) BHP Billiton Iron Ore Pilbaranorthern quoll monitoring project Final report May 2010ndashJune 2013Unpublished report prepared for BHP Billiton Iron Ore Department ofParks and Wildlife Perth

Dunlop J Cook A and Morris K (2014b) Pilbara northern quollproject surveying and monitoring Dasyurus hallucatus in the PilbaraWestern Australia Department of Parks and Wildlife Perth Availableat librarydpawwagovau

Florance D Webb J K Dempster T Kearney M R Worthing A andLetnic M (2011) Excluding access to invasion hubs can contain thespread of an invasive vertebrate Proceedings of the Royal Society ofLondon B Biological Sciences 278 2900ndash2908 doi101098rspb20110032

Gaillard J-M Hebblewhite M Loison A Fuller M Powell R BasilleM and Van Moorter B (2010) Habitatndashperformance relationshipsfinding the right metric at a given spatial scale PhilosophicalTransactions of the Royal Society of London Series B BiologicalSciences 365 2255ndash2265 doi101098rstb20100085

Glen A S and Dickman C R (2008) Niche overlap between marsupialand eutherian carnivores does competition threaten the endangeredspotted-tailed quoll Journal of Applied Ecology 45 700ndash707 doi101111j1365-2664200701449x

Glen A S Pennay M Dickman C R Wintle B A and Firestone K B(2011) Diets of sympatric native and introduced carnivores in theBarrington Tops eastern Australia Austral Ecology 36 290ndash296doi101111j1442-9993201002149x

Hennig P (2004) A brief land use history In lsquoAn Inventory and ConditionSurvey of the Pilbara Region Western Australiarsquo (Eds A M E VanVreeswyk A L Payne K A Leighton and P Hennig) TechnicalBulletin No 92 pp 13ndash18 (Western Australian Department ofAgriculture Perth)

Hohnen R Ashby J Tuft K and McGregor H (2013) Individualidentification of northern quolls (Dasyurus hallucatus) using remotecameras Australian Mammalogy 35 131ndash135 doi101071AM12015

L Australian Mammalogy V A Cramer et al

How R A Spencer P B S and Schmitt L H (2009) Island populationshave high conservation value for northern Australiarsquos top marsupialpredator ahead of a threatening process Journal of Zoology 278206ndash217 doi101111j1469-7998200900569x

Johnson C N Isaac J L and Fisher D O (2007) Rarity of a top predatortriggers continent-wide collapse of mammal prey dingoes andmarsupials in Australia Proceedings of the Royal Society of LondonSeries B Biological Sciences 274 341ndash346 doi101098rspb20063711

Jones M E (2000) Road upgrade road mortality and remedial measuresimpacts on a population of eastern quolls and Tasmanian devilsWildlifeResearch 27 289ndash296 doi101071WR98069

Kearney M Phillips B L Tracy C R Christian K A Betts G andPorter W P (2008) Modelling species distributions without usingspecies distributions the cane toad in Australia under current andfuture climates Ecography 31 423ndash434 doi101111j0906-7590200805457x

Kendrick P and Stanley F (2002) Pilbara 4 (PIL4 ndash Roebourne synopsis)In lsquoA Biodiversity Audit of Western Australiarsquos 53 BiogeographicalSubregions in 2002rsquo (Eds J E May and N L McKenzie) pp 581ndash593(Department of Conservation and Land Management Perth)

Kennedy M Phillips B L Legge S Murphy S A and Faulkner R A(2012) Do dingoes suppress the activity of feral cats in northernAustralia Austral Ecology 37 134ndash139 doi101111j1442-9993201102256x

Kiesecker J M Copeland H Pocewicz A and McKenney B (2010)Development by design blending landscape-level planning with themitigation hierarchy Frontiers in Ecology and the Environment 8261ndash266 doi101890090005

King D R and Smith L A (1985) The distribution of the European red fox(Vulpes vulpes) in Western Australia Records of the Western AustralianMuseum 12 197ndash205

Knight A T Cowling R M Rouget M Balmford A Lombard A Tand Campbell B M (2008) Knowing but not doing selecting priorityconservation areas and the researchndashimplementation gap ConservationBiology 22 610ndash617 doi101111j1523-1739200800914x

Laurance W F Koster H Grooten M Anderson A B Zuidema P AZwick S Zagt R J Lynam A J Linkie M and Anten N P (2012)Making conservation research more relevant for conservationpractitioners Biological Conservation 153 164ndash168 doi101016jbiocon201205012

Legge S Murphy S Heathcote J Flaxman E Augusteyn J andCrossman M (2008) The short-term effects of an extensive and high-intensity fire on vertebrates in the tropical savannas of the centralKimberley northern Australia Wildlife Research 35 33ndash43 doi101071WR07016

Lester D (2015) Effective wildlife roadkill mitigation Journal of Trafficand Transportation Engineering 3 42ndash51 doi10172652328-2142201501005

Letnic M Tamayo B and Dickman C R (2005) The responsesof mammals to La Nintildea (El Nintildeo Southern Oscillation)-associatedrainfall predation and wildfire in central Australia Journal ofMammalogy 86 689ndash703 doi1016441545-1542(2005)086[0689TROMTL]20CO2

Letnic M Koch F Gordon C Crowther M S and Dickman C R(2009) Keystone effects of an alien top-predator stem extinctions ofnative mammals Proceedings of the Royal Society of London Series BBiological Sciences 276 3249ndash3256 doi101098rspb20090574

Lindenmayer D B (2015) Continental-level biodiversity collapseProceedings of the National Academy of Sciences of the United States ofAmerica 112 4514ndash4515 doi101073pnas1502766112

Lindenmayer D B Piggott M P and Wintle B A (2013) Counting thebooks while the library burns why conservation monitoring programs

need a plan for action Frontiers in Ecology and the Environment 11549ndash555 doi101890120220

Maron M Hobbs R J Moilanen A Matthews J W Christie KGardner T A Keith D A Lindenmayer D B and McAlpine C A(2012) Faustian bargains Restoration realities in the context ofbiodiversity offset policies Biological Conservation 155 141ndash148doi101016jbiocon201206003

McCall S C McCarthy M A van der Ree R Harper M J Cesarini Sand Soanes K (2010) Evidence that a highway reduces apparentsurvival rates of squirrel gliders Ecology and Society 15 27

McGregor R A Stokes V L and Craig M D (2014) Does forestrestoration in fragmented landscapes provide habitat for a wide-ranging carnivore Animal Conservation 17 467ndash475 doi101111acv12112

McKenzie N L Burbidge A A Baynes A Brereton R N DickmanC R Gordon G Gibson L A Menkhorst P W Robinson A CWilliams M R and Woinarski J C Z (2007) Analysis of factorsimplicated in the recent decline of Australiarsquos mammal fauna Journal ofBiogeography 34 597ndash611 doi101111j1365-2699200601639x

McKenzie N L van Leeuwen S and Pinder AM (2009) Introduction tothe Pilbara Biodiversity Survey 2002ndash2007 Records of the WesternAustralian Museum 78(Suppl) 3ndash89 doi1018195issn0313-122x78(1)2009003-089

Morris K Johnson B Orell P Gaikhorst G Wane A and MoroD (2003) Recovery of the threatened chuditch (Dasyurus geoffroii)a case study In lsquoPredators with Pouches The Biology of CarnivorousMarsupialsrsquo (Eds M Jones C Dickman and M Archer) pp 435ndash451(CSIRO Publishing Melbourne)

Nelson J L Scroggie M P and Belcher C A (2014) Developing acamera trap survey protocol to detect a rare marsupial carnivore thespotted-tailed quoll (Dasyurus maculatus) In lsquoCamera TrappingWildlife Management and Researchrsquo (Eds P D Meek and PJ Fleming) pp 271ndash279 (CSIRO Publishing Melbourne)

Newsome T M Dellinger J A Pavey C R Ripple W J Shores C RWirsing A J and Dickman C R (2015) The ecological effects ofproviding resource subsidies to predators Global Ecology andBiogeography 24 1ndash11 doi101111geb12236

OrsquoDonnell S Webb J K and Shine R (2010) Conditioned tasteaversion enhances the survival of an endangered predator imperilled bya toxic invader Journal of Applied Ecology 47 558ndash565 doi101111j1365-2664201001802x

Oakwood M (1997) The ecology of the northern quoll Dasyurushallucatus PhD Thesis Australian National University Canberra

Oakwood M (2000) Reproduction and demography of the northern quollDasyurus hallucatus in the lowland savanna of northern AustraliaAustralian Journal of Zoology 48 519ndash539 doi101071ZO00028

Oakwood M (2002) Spatial and social organization of a carnivorousmarsupial Dasyurus hallucatus (Marsupialia Dasyuridae) Journal ofZoology 257 237ndash248 doi101017S0952836902000833

Oakwood M and Foster P (2008) Monitoring extinction of the northernquoll Australian Academy of Science Newsletter 71 6

Palomares F and Caro T M (1999) Interspecific killing amongmammalian carnivores American Naturalist 153 492ndash508 doi101086303189

Pavey C R and Bastin G (2014) lsquoAustralian Rangelands and ClimateChange ndash Invasive Animalsrsquo (Ninti One Limited amp CSIRO AliceSprings)

Pierson J C Barton P S Lane PW and Lindenmayer D B (2015) Canhabitat surrogates predict the response of target species to landscapechange Biological Conservation 184 1ndash10 doi101016jbiocon201412017

Pollock A B (1999) Notes on status distribution and diet of northernquoll Dasyurus hallucatus in the MackayndashBowen area mideastern

Research priorities for the northern quoll Australian Mammalogy M

Queensland Australian Zoologist 31 388ndash395 doi107882AZ1999040

Ramanaidou E R and Morris R C (2010) A synopsis of the channeliron deposits of the Hamersley Province Western AustraliaTransactions of the Institutions of Mining and Metallurgy Section BApplied Earth Science 119 56ndash59 doi101179037174510X12853354810624

Schmitt L H Bradley A J Kemper C M Kitchener D J HumphreysW F and How R A (1989) Ecology and physiology of the northernquoll Dasyurus hallucatus (Marsupialia Dasyuridae) at MitchellPlateau Kimberley Western Australia Journal of Zoology 217539ndash558 doi101111j1469-79981989tb02510x

Short J and Smith A (1994) Mammal decline and recovery in AustraliaJournal of Mammalogy 75 288ndash297 doi1023071382547

Spencer P How R Hillyer M Cook A Morris K Stevenson C andUmbrello L (2013) Genetic analysis of northern quolls from the PilbaraRegion ofWestern Australia Year one ndash final report Unpublished reportDepartment of Sustainability Environment Water Population andCommunities Canberra and Department of Parks and Wildlife Perth

Sutherland W J Fleishman E Mascia M B Pretty J and Rudd M A(2011) Methods for collaboratively identifying research priorities andemerging issues in science and policyMethods in Ecology and Evolution2 238ndash247 doi101111j2041-210X201000083x

Sutherst R W Floyd R B and Maywald G F (1996) The potentialgeographical distribution of the cane toad Bufo marinus L in AustraliaConservation Biology 10 294ndash299 doi101046j1523-1739199610010294x

Taylor B D and Goldingay R L (2003) Cutting the carnage wildlifeusage of road culverts in north-eastern New South Wales WildlifeResearch 30 529ndash537 doi101071WR01062

Tingley R Phillips B L Letnic M Brown G P Shine R and BairdS J (2013) Identifying optimal barriers to halt the invasion of canetoads Rhinella marina in arid Australia Journal of Applied Ecology 50129ndash137 doi1011111365-266412021

Van Vreeswyk A M E Payne A L Leighton K A and HennigP (2004) An Inventory and Condition Survey of the Pilbara RegionWestern Australia Technical Bulletin No 92 Department of AgricultureWestern Australia Perth

Webb J Legge S Tuft K Cremona T and Austin C (2015) Can wemitigate cane toad impacts on northern quolls Final report National

Environmental Research Program Northern Australia Hub CharlesDarwin University Darwin

Wintle B A Walshe T V Parris K M and McCarthy M A (2012)Designing occupancy surveys and interpreting non-detection whenobservations are imperfect Diversity amp Distributions 18 417ndash424doi101111j1472-4642201100874x

Woinarski J Menkhorst K Gambold N and Braithwaite R W (1992)Mammals of the Bungle Bungle area In lsquoA Survey of the Wildlife andVegetation of Purnululu (Bungle Bungle) National Park and AdjacentArearsquo (Ed J C Z Woinarski) Research Bulletin No 6 pp 53ndash67(CSIRO Division of Wildlife and Ecology Darwin)

Woinarski J C Z Risler J and Kean L (2004) Response of vegetationand vertebrate fauna to 23 years of fire exclusion in a tropical Eucalyptusopen forest Northern Territory Australia Austral Ecology 29 156ndash176doi101111j1442-9993200401333x

Woinarski J C Z Oakwood M Winter J Burnett S Milne D FosterP Myles H and Holmes B (2008) Surviving the toads patterns ofpersistence of the northern quoll Dasyurus hallucatus in QueenslandReport submitted to the Natural Heritage Trust Strategic ReserveProgram as a component of project 2005162 Monitoring andManagement of Cane Toad Impact in theNorthern Territory Available athttplrmntgovauplants-and-animalsinformation-and-publicationspublications2008 [accessed 14 November 2013]

Woinarski J C Z Armstrong M Brennan K Fisher A Griffiths A DHill B Milne D J Palmer C Ward S Watson M Winderlich Sand Young S (2010) Monitoring indicates rapid and severe decline ofnative small mammals in Kakadu National Park northern AustraliaWildlife Research 37 116ndash126 doi101071WR09125

Woinarski J Burbidge A and Harrison P (2014) lsquoAction Plan forAustralian Mammals 2012rsquo (CSIRO Publishing Melbourne)

Woinarski J C Burbidge A A and Harrison P L (2015) Ongoingunraveling of a continental fauna decline and extinction of Australianmammals since European settlement Proceedings of the NationalAcademy of Sciences of the United States of America 112 4531ndash4540doi101073pnas1417301112

Woolley P A Krajewski C and Westerman M (2015) Phylogeneticrelationships within Dasyurus (Dasyuromorphia Dasyuridae) quollsystematics based on molecular evidence and male characteristicsJournal of Mammalogy 96 37ndash46 doi101093jmammalgyu028

N Australian Mammalogy V A Cramer et al

wwwpublishcsiroaujournalsam

Page 7: Research priorities for the northern quoll (Dasyurus ......The Pilbara population of the northern quoll (Dasyurus hallucatus) has been seldom studied, and the impacts of threats such

habitat complexity (eg diversity of boulder sizes) and thepresence of permanent water in the quarry which may haveincreased prey availability at these sites

Movement of animals through the landscape for dispersaland repopulation is another important consideration for a speciesin which both sexes tend towards semelparity and is thereforepredisposed to local extinction (Oakwood 1997) Improvedunderstanding of how the northern quoll (particularly dispersingmales) uses areas of lower habitat quality and the identificationof such habitat in habitat-suitability models will allow futurelandscape-scale strategic assessments to better predict thecombined impacts of disturbance on the ability of northern quolls

to move through the landscape Analysis of regional populationgenetics will also improve our understanding of the level ofdispersal occurring across the landscape

Priority 3 Population dynamics

Better understand the population dynamics of thenorthern quoll in the Pilbara

Little information is available on the number ofsubpopulations of northern quoll in the Pilbara how thesesubpopulations interact how stable they are in space and timeand how they respond to disturbance As no demographic

Table 2 Records of the northern quoll in each land system type in the PilbaraRecords were current as of 23 May 2014

Land system Area (km2) No of records Land system description

Augustus 61 1 Rugged ranges hills ridges and plateaux supporting mulga shrublands and hard spinifex grasslandsBoolaloo 1502 245 Granite hills domes and tor fields and sandy plains with shrubby spinifex grasslandsBoolgeeda 7748 29 Stony lower slopes and plains below hill systems supporting hard and soft spinifex grasslands andmulga

shrublandsCalcrete 1444 2 Low calcrete platforms and plains supporting shrubby hard spinifex grasslandsCallawa 1003 9 Highly dissected low hills mesas and gravelly plains of sandstone and conglomerate supporting soft and

hard spinifex grasslandsCapricorn 5296 257 Hills and ridges of sandstone and dolomite shrubby hard and soft spinifex grasslandsCoongimah 3244 12 Plateau surfaces low hills with steep slopes and undulating uplands supporting hard spinifex grasslandsDune 138 1 Dune fields supporting soft spinifex grasslandsGranitic 4020 1 Rugged granitic hills supporting shrubby hard and soft spinifex grasslandsHorseflat 1261 4 Gilgaied clay plains supporting tussock grasslands and minor grassy snakewood shrublandsLittoral 1577 3 Bare coastal mudflats with mangroves on seaward fringes samphire flats sandy islands coastal dunes

and beachesMacroy 13 095 94 Stony plains and occasional tor fields based on granite supporting hard and soft spinifex grasslandsMcKay 4202 21 Hills ridges plateau remnants and breakaways of metasedimentary rocks supporting hard spinifex

grasslandsMallina 2557 8 Sandy surfaced alluvial plains supporting soft spinifex (and occasionally hard spinifex) grasslandsMarandoo 459 2 Basalt hills and restricted stony plains supporting grassy mulga shrublandsMosquito 1840 3 Stony plains and prominent ridges of schist and other metamorphic rocks supporting hard spinifex

grasslandsNewman 14 580 81 Rugged jaspilite plateaux ridges and mountains supporting hard spinifex grasslandsOnslow 424 1 Sandplains dunes and clay plains supporting soft spinifex grasslands and minor tussock grasslandsParaburdoo 565 3 Basalt-derived stony gilgai plains and stony plains supporting snakewood and mulga shrublands with

spinifex and tussock grasslandsParadise 1479 8 Alluvial plains supporting soft spinifex grasslands and tussock grasslandsRiver 4088 37 Active flood plains major rivers supporting grassy eucalypt woodlands tussock grasslands and soft

spinifex grasslandsRobe 865 369 Low limonite mesas and buttes supporting soft spinifex (and occasionally hard spinifex) grasslandsRocklea 22 993 40 Basalt hills plateaux lower slopes and minor stony plains supporting hard spinifex (and occasionally

soft spinifex) grasslandsRuth 346 36 Hills and ridges of volcanic and other rocks supporting hard spinifex (occasionally soft spinifex)

grasslandsStuart 1794 3 Gently undulating stony plains supporting hard and soft spinifex grassland and snakewood shrublandsTaylor 129 3 Stony plains and isolated low hills of sedimentary rocks supporting hard and soft spinifex grasslandsTalga 2124 9 Hills and ridges of greenstone and chert and stony plains supporting hard and soft spinifex grasslandsTuree 581 2 Stony alluvial plains with gilgaied and non-gilgaied surfaces supporting tussock grasslands and grassy

shrublandsUaroo 7681 37 Broad sandy plains supporting shrubby hard and soft spinifex grasslandsUrandy 1311 6 Stony plains alluvial plains and drainage lines supporting shrubby soft spinifex grasslandsWhite Springs 266 4 Stony gilgai plains supporting tussock grasslands and hard spinifex grasslandsWona 1815 10 Basalt upland gilgai plains supporting tussock grasslands and minor hard spinifex grasslandsYamerina 1207 1 Floodplains and deltaic deposits supporting tussock grasslands grassy woodlands and minor halophytic

low shrublands

Research priorities for the northern quoll Australian Mammalogy G

studies have been undertaken in the region Cadenhead et al(2015) used demographic information from the NorthernTerritory and Kimberley populations in their PVA Uncertaintyabout data quality and the limited information available ondemographic parameters for the Pilbara constrained the level ofcertainty around the outcomes of the PVA The model doeshowever provide a framework into which new data can beincorporated as they become available For example preliminarydata from the Pilbara monitoring program suggest that theproportion of male die-off in the Pilbara population is lowerthan in the Kimberley and Northern Territory populations(Dunlop unpubl data) Cadenhead et al (2015) stress the needfor more demographic and occurrence data to be collectedbefore further development decisions are made (see lsquoDevelopappropriate and standardised survey and monitoring methodsrsquoand lsquoImprove our understanding of habitat requirementsrsquo above)

Investigate population structure and the interactionbetween populations

Analysis of the genetic diversity and population structure ofthe Pilbara population of the northern quoll is ongoingOnly small differences in genetic diversity as measured byheterozygosity were found between samples obtained at 16locations within the Pilbara with the exception of the isolatedpopulation on Dolphin Island (Spencer et al 2013) This resultindicates a high level of movement across the landscape byindividuals during mating and dispersal phases Spatial structureanalysis of samples collected from around Pannawonicasuggested a lsquoneighbour sizersquo of ~15 km (Spencer et al 2013)Current data show no evidence of genetic bottleneckssuggesting that if there has been a population decline in thePilbara that this has not affected genetic diversity A usefulapproach would be to combine genetic information with datafrom radio-tracking or GPS collars to better understand theuse of lsquolandscapersquo space and how this influences populationdynamics

Priority 4 Key threats and the interaction of these threats

Introduced predators

Introduced cats and foxes are considered to be one of themain causes of the decline and extinction of the critical-weight-range mammals (35 g to 55 kg) in Australia (Short and Smith1994 McKenzie et al 2007 Woinarski et al 2015) but theirimpacts on the northern quoll are less clear A recent review offeral cat diets across Australia found that northern quolls werepresent in the diet of feral cats (Doherty et al 2015a) andWoinarski et al (2014) consider the threat posed by cats to thenorthern quoll to be severe although the impacts of cats may bereduced in rugged refuge areas The impacts of cats on othermuch larger species of quoll appear to be inconsistent withsome evidence of coexistence between cats and quolls beforethe arrival of foxes (Glen and Dickman 2008 and referencescontained within) Due to the limited overlap in their distributionthroughout Australia the threat posed by foxes to northernquolls is less understood than for cats and Woinarski et al(2014) consider the threat posed by foxes to be moderate on anational level Foxes are common in coastal near-coastal and

riparian areas of the Pilbara and extend as far north as Broome(King and Smith 1985 Kendrick and Stanley 2002) They arerenowned for their aggression towards smaller carnivores(Palomares and Caro 1999) and neither the larger chuditch(Dasyurus geoffroii) nor spotted-tail quoll (Dasyurus maculatus)are known to coexist with large numbers of foxes (A Glen perscomm) The interactions between quolls cats and foxesare potentially more complex than for other medium-sizedmarsupials due to niche overlap in diet and competition for densites with the larger spotted-tailed quoll and chuditch bothexperiencing exploitation and interference competition fromfoxes (Morris et al 2003 Glen et al 2011) Competitionbetween feral cats and the northern quoll may potentially beintense during periods of low rainfall in the Pilbara when preyavailability is low Understanding whether foxes and cats aresignificant predators of or competitors with the northern quollin the Pilbara and how this affects their population viabilitywill clarify the level of risk posed by these predators Thesausage-style baits used for feral cats (eg Eradicatreg andCuriosityreg) are potentially attractive to the northern quoll withthe risk of mortality or reduced fitness Trials to assess themagnitude of the risks from Eradicatreg baits are currentlyunderway in the Pilbara

The role of dingoes and wild dogs in the mortality of andinteractions between mesopredators (both native and introduced)requires further investigation Dingoes are known to kill northernquolls butmay not eat these kills (Oakwood 1997) A recent studyin the Northern Territory found that camp dogs and dingoes werethe major source of mortality for northern quolls at the study site(Webb et al 2015) and roaming domestic dogs were identified asa cause of mortality for northern quolls in Queensland (Pollock1999) Human resource subsidies (eg stock carrion in rangelandareas) can artificially support predator populations thus indirectlyincreasing predation pressure on prey (Newsome et al 2015)A dynamic debate currently exists around the role of the dingo inregulating populations of feral cats and foxes and the implicationsof this for the conservation of the critical-weight-range mammalsin Australia (see Johnson et al 2007 Letnic et al 2009 Allenet al 2011 2013 Kennedy et al 2012) Understanding thepopulation density at which the role of the dingo as a regulator ofintroduced predators outweighs its own impact as a direct predatorof the northern quoll will inform whether dingowild dog controlwould benefit the northern quoll

Cane toads

The introduced cane toad now occupies more than half thehistoric range of the northern quoll and has caused an immediatecollapse or extirpation of northern quoll populations in theareas it has invaded (Burnett 1997 Oakwood and Foster2008 Woinarski et al 2008 2010) Earlier modelling work thatconsidered only climate parameters as well as more recentwork that incorporates future climate change suggest thatcane toads will not invade the Pilbara (Sutherst et al 1996Pavey and Bastin 2014) Other recent work that considers thefundamental niche of the cane toad and incorporates the locationof artificial watering points predicts that they will colonisethe Pilbara from the Kimberley via the coastal edge of theGreat Sandy Desert (Kearney et al 2008 Florance et al 2011

H Australian Mammalogy V A Cramer et al

Tingley et al 2013) This has created some level of uncertaintyabout the likely impact of cane toads on northern quolls in thePilbara Strategies for cane toad management were not rankedhighly by participants in the threat-management frameworkdeveloped by Carwardine et al (2014) which the authorsexplained as possibly due to uncertainty about how far canetoads will spread into the region

The model developed by Tingley et al (2013) suggested thatthe closure of 100 artificial water bodies at one of three potentiallocations along the coastal corridor between the Kimberley andthe Pilbara would prevent the invasion of cane toads into thePilbara While their model did account for uncharacteristicallywet years every three decades the dispersal of toads betweenwater bodies was determined by the number of rainy daysoccurring during the year (ie toads could only disperse betweenwater bodies for three days following rainfall) rather thanrainfall amount the associated likelihood of flooding and theavailability of free water across the landscape The cyclonic andthunderstorm nature of much of the rainfall in the north-west ofWestern Australia means that the region is prone to significantand sometimes extended flooding In April 1934 the countryaround Anna Plains (250 km south of Broome) was flooded forhundreds of miles when 27 inches (686mm) of rain fell infour days (Anon 1934) In March 2004 over 350mm of rain wasrecorded within a 48-h period at Telfer on the southern edge ofthe Great Sandy Desert due to Tropical Cyclone Fay Thiscaused the flooding of the De Grey and Oakover river systemsand flooded 140 km of the Telfer Access Road with floodwaterstaking several months to subside (wwwbomgovau) Extendingthe model of Tingley et al (2013) to account for such events willhelp clarify whether closing artificial water bodies is likely toprevent the invasion of cane toads into the Pilbara or may onlyassist in delaying their inevitable invasion

Comparison of SDMs (eg Cadenhead et al 2015) fornorthern quoll with fundamental niche models for cane toads(eg see Kearney et al 2008) within the Pilbara may identifypotential refuge areas for the northern quoll and these areasshould be prioritised for conservation actions Because theinvasion point of cane toads into the Pilbara is likely to be narrow(beginning around the mouth of the De Grey River Tingleyet al 2013) a proactive approach to training northern quollsto avoid cane toads may prove beneficial Conditioned taste-aversion (CTA) has been used successfully to train captive-reared northern quolls to avoid cane toads by offering themsmall dead toads infused with thiabendazole before theirrelease into the wild (OrsquoDonnell et al 2010)Monitoring of thesequolls after their release showed that some of their offspringhad learnt to avoid eating cane toads either through ingestionof small non-lethal toads or via social learning from theirmothers (Webb et al 2015) Fields trials of toad-aversion baits(either sausages or meatballs made from toad flesh andcontaining thiabendazole) are currently underway in the centralKimberley where cane toads were expected to invade in late2015 (Webb et al 2015) Further testing of CTA approachesunder a variety of field conditions is required The developmentof stable long-lasting baits suitable for aerial deployment isconsidered a pressing need as well as a captive colony ofnorthern quolls upon which to test the efficacy of toad-aversionbaits for inducing aversion to live toads (Webb et al 2015)

Infrastructure development and interactionswith humans

Infrastructure development may increase the likelihood ofnorthern quoll deaths due to motor vehicle strikes or beingcrushed in machinery Significant chuditch mortality wasobserved along mine access roads in the south-west of WesternAustralia (McGregor et al 2014) and the widening and sealingof an access road dramatically increased the number of roadkills and led to the extirpation of the resident population ofeastern quolls (D viverrinus) in Tasmania (Jones 2000)Infrastructure corridors such as high-tension power-lineeasements and ore-haulage railways are increasingly beinginstalled throughout the range of the northern quoll Althoughthese do not create the same mortality risk as roads they bisectand fragment habitat create large open spaces and potentiallyincrease predation risk

A recent small-scale radio-tracking study (see Dunlop et al2014a) around several quarries in the Pilbara found that northernquolls crossed roads but not railway lines The short-term natureof this work and the limited number of quolls able to be trackedmeans that the influence of roads railway lines and infrastructurecorridors on the movement and mortality of northern quollsrequires further assessment Using radio-tracking techniques todetermine crossing rates and road kill events may be difficultover large areas so population studies comparing groups ofquolls near major roads with those not affected by infrastructure(eg McCall et al 2010) may provide a more feasible meansof assessing the impacts of roads If vehicle strikes are foundto significantly contribute to the mortality of a local populationof the northern quoll then mitigation methods should betrialled on sections of road identified as road-kill hotspotsCulverts and underpasses have been shown to be used by avariety of small and medium-sized mammals and frogs (Taylorand Goldingay 2003) while installing rumble strips on roadshas led to decreased rates of road-kill of nocturnal animals(Lester 2015)

Interactions between threats

At present wildfire the subsequent loss of vegetation coverand increased predation of northern quolls is likely to be themost significant of the interactions between threats The directeffects of wildfire have been found to be less significant inthe subsequent decline of small to medium-sized mammalsparticularly for denning species such as the northern quoll thanthe indirect consequences of reduced resource availability andthe increased risk of predation due to lack of cover (Oakwood2000 Letnic et al 2005 McKenzie et al 2007 Legge et al2008 Doherty et al 2015b) Male northern quolls are perhapsmost vulnerable to such losses of cover during the breedingseason when their range can extend to over 100 ha (Oakwood2002) or where males survive after breeding their foragingrange is extended to gain the nutrition required to recover fromtheir poor condition (Cook 2010) Loss of cover due to late dry-season fires at East Alligator River in the Northern Territorymeant little cover was available for young quolls when theybegan foraging with dog and dingo predation a major source ofmortality (Webb et al 2015) The cumulative impacts of canetoad poisoning fire and predation are highly likely to lead to the

Research priorities for the northern quoll Australian Mammalogy I

extirpation of small and disjunct groups of northern quoll in thePilbara Population viability modelling of the East AlligatorRiver population suggested that high mortality from predationcombined with late dry-season burning was preventing therecovery of northern quoll populations after cane toad invasion

(Webb et al 2015) Improved knowledge of the severity of thesethreats to northern quolls in the Pilbara and their incorporationinto the SDM and PVA framework developed by Cadenheadet al (2015) will allow for better identification of priority areasfor integrated cane toad fire and predator management

Table 3 Expected timelines for the undertaking of research under each priority over the next 10 years and the funds currently allocated fromthe Pilbara Northern Quoll Offset Trust under each research priority

Rank Theme Research priority Current or proposed researchactivity

Time over next10 years

Current funding allocation(approximate)

1 Survey andmonitoring

Develop appropriate andstandardised survey andmonitoring methods

Develop monitoring protocol(traps and cameras) andinvestigate new methods

Ongoing survey and monitoringin conjunction with ResearchThemes 2 and 3 to better mapnorthern quoll distribution andto determine long-termpopulation trends

Years 1ndash2Years 3ndash10

$200 000 per annum over10 years for ResearchThemes 1 2 and 3(inclusive of salaries)

2 Habitatrequirements

Define areas of criticalhabitat for northernquoll in the Pilbara

Identify and describe criticalhabitat and undertake predictivemodelling of unsurveyed areasto inform further survey effort

Years 3ndash4 $50 000 for predictivemodelling (EdithCowan University)

Understand howdisturbance affectshabitat quality andconnectivity

Ongoing collection of data ondisturbance and habitat qualityin conjunction with ResearchThemes 1 and 4

Incorporation of new data intopredictive models

Years 4ndash10 See above

3 Populationdynamics

Better understand thepopulation dynamics ofthe northern quoll in thePilbara

Collect data on demographicparameters breeding dispersaland home-range use toincorporate into future PVA

Continued collectionof data inconjunction withpriorities 1 and 2during years 3ndash10

See above

Investigate populationstructure and theinteraction betweenpopulations

Analysis of existing tissuesamples and newly collectedsamples

Years 1ndash10 $120 000 for initial roundof genetic analysis(Years 3ndash5 MurdochUniversity)

4 Key threats Introduced predators Spatio-temporal interactionsbetween northern quolls andintroduced predators

Other projects as they arise

Years 2ndash4 (currentPhD projectUniversity ofQueensland)

$12 000 contribution tofield expenses

Cane toads Predictive modelling of canetoad distribution in the Pilbara

Years 3ndash4 Part of predictivemodelling by EdithCowan University

Infrastructuredevelopment andinteractions withhumans

Investigation of impacts oflinear infrastructure on northernquoll movement

Year 4 (currentHonours projectEdith CowanUniversity)

$15 000

Interactions betweenthreats

No on-ground projects currentlyplanned

Potential to be assessed throughPVA as data are collected underother research priorities

No funding currentlyallocated

5 Recolonisation ofrestored orartificial habitat

Determine the ability ofthe northern quoll torecolonise disturbedareas or coloniseartificial habitat

As projects arise Years 5ndash8 incollaboration withminesite restorationprograms

No funding currentlyallocated

J Australian Mammalogy V A Cramer et al

Priority 5 Recolonisation of restored andor artificial habitat

While anecdotal evidence suggests that northern quolls will denin waste rock dumps whether artificial or restored habitatprovides for long-term population viability is not known andlong-term assessment of created habitat is required Weacknowledge that a large gap exists between the desire to userestored or artificial habitat as an ecologically equivalent offsetand our limited ability to actually restore or re-create habitat thatis equivalent in structure composition and function particularlyon highly disturbed sites (Maron et al 2012) Yet there is a futureopportunity in the Pilbara to conduct lsquonaturalrsquo experiments onthe recolonisation of quolls after mining ceases including howto best design waste rock dumps so that habitat complexityand productivity is maximised by for example experimentingwith the size and positioning of boulders and their spatialarrangement in relation to surrounding landscape features Theuse of such artificial habitat should be tested in the first stagethrough direct monitoring of animals that have colonised orhave been translocated to the site Such experimentation isunderway at Atlas Ironrsquos Mt Dove operation (Atlas Iron 2012)Population modelling can then be used to predict futurepopulation sizes rates of growth and spread and the probabilityof long-term persistence in the context of various managementactions (eg predator control Armstrong and Reynolds 2012)

Resource allocation and timelines

While the allocation of resources and funding and the periodover which each of the research priorities was expected to beaddressed was not explicitly discussed during the workshopthe ranking of the priorities was taken to indicate the sequencein which the priorities should be addressed At the time of theworkshop some of the research questions identified werealready being investigated within various projects and a portionof the Pilbara Northern Quoll Offset Trust allocated to these InTable 3 we indicate the expected timelines over which someof the research programs are expected to be undertaken WhileAU$175M has been allocated to northern quoll research in thePilbara it is important to note that much of this amount is notcurrently available and will not become available until severalapproved mine developments have become operational InTable 3 we also indicate the proportion of offset funds expectedto be allocated to each research priority Research undertaken inother regions (eg research into CTA in the Kimberley regionahead of the cane toad invasion front) cannot be funded from thePilbara Northern Quoll Offset Trust but will help address someof the research questions identified here

Lessons learnt from the workshop process

The workshop approach used here was less formal than recentlarge-scale exercises used to elicit research priorities for emergingissues in science and policy at national or global levels (reviewedin Sutherland et al 2011) The focus in our (much smaller)workshop was on inclusivity rather than identifying participantsbased on purposive sampling of expertise or organisationalaffiliation We believe that the inclusive nature of the workshopwas a strength but we also recognise that this perhaps createdsome level of bias when ranking the research prioritiesAcknowledging both the experience and knowledge requirements

of those who work regularly in the Pilbara is an importantcomponent of improving the design of research programs that aimto improve conservation management in the region Yet theresearch priorities and their rankings are likely strongly reflectiveof the current information needs of most participants and maynot align with what conservation scientists believe to be themost critical areas of research for northern quoll conservation intothe future Baseline survey and monitoring are often the firstrequirements of a management or research plan for the northernquoll when offset conditions are prescribed and this was reflectedin the primary ranking given to the development of standardsurvey and monitoring methods We acknowledge the concernthat this could simply lead to offset funding being used to lsquomonitorthe extinctionrsquo of the northern quoll particularly after canetoads invade the Pilbara unless these data are used to triggermanagement actions (Oakwood and Foster 2008 Lindenmayeret al 2013) Clearly the currently limited information on habitatuse and demographic parameters for the northern quoll in thePilbara restricts the utility of more sophisticated researchapproaches such as developing robust SDM and PVA While thesevere threat that cane toads pose to the northern quoll wasrecognised this threat is not immediate in the region and mayexplain the less prominent ranking given to research to mitigatethe impacts of cane toads

The northern quoll has become emblematic of the dramaticand rapid species declines occurring in northern Australia andthe level of interest in this species was reflected in the highnumber of participants attending the workshop (one of six runby Parks and Wildlife) Given the number of attendees betterutilisation of the time available would have been gained byproviding participants with a brief before the workshopand asking them to contribute potential research questionsanonymously for interactive review by other participants Inthe exercises reviewed by Sutherland et al (2011) questionswere then discarded if they were not considered to meet criteriathat were previously agreed upon Some areas of researchthat gained some attention on the day may have been discardedif given greater consideration or interrogation before theworkshop For example there was some discussion duringthe workshop of investigating the use of habitat surrogates tomonitor the northern quoll yet research has shown that there islittle validity in using surrogates to monitor the abundance of asingle species (Pierson et al 2015)

Conclusions

Offset funds generated under the environmental approvalsprocess provide an opportunity to mobilise considerablefinancial resources for conservation (Kiesecker et al 2010) Yetthere remain significant conceptual and practical limitations tobe resolved to ensure that direct offsets provide measurableconservation outcomes that are ecologically equivalent to thelosses sustained by proceeding with the development (Maronet al 2012 Bull et al 2013) The collaborative identificationand ranking of research priorities undertaken here is the firststep in the process of developing and evaluating potentialconservation actions for the northern quoll in the Pilbara andserves as a mechanism to ensure that the research priorities ofstate and federal agencies reflects the knowledge needs of

Research priorities for the northern quoll Australian Mammalogy K

practitioners and proponents currently operating in the regionThe provision of offset funds over the next decade providesa unique opportunity to undertake research on a species ofconservation significance at a regional scale and over a longperiod through an integrated and strategic program of researchrather than the collection of ad hoc data While a morestructured approach to defining research questions before theworkshop may have led to greater critical reflection upon anddiscussion of some of the research priorities on the day theworkshop provided an opportunity for an open and diverse-ranging discussion in a collaborative environment amongstthe broader community of scientists environmental consultantsand mining proponents involved in the management andconservation of the northern quoll in the Pilbara

Acknowledgements

We gratefully acknowledge the financial support of Roy Hill who sponsoredthe workshop and partially funded VACrsquos salary We thank the Departmentof the Environmentrsquos Pilbara Taskforce especially Veronica Ritchie fortheir support of interested or affected parties The success of the workshopwas in great part due to the facilitation skills of Craig Salt of SustainableConsulting We thank all of the workshop participants for their open andcollaborative approach to the process Mike Bamford Eddie van EttenDavid Peacock and Peter Lyon provided useful comments on an earlier draftWe are grateful to John Woinarski for his constructive advice thatsubstantially improved the structure and clarity of the manuscript and ananonymous reviewer for careful and critical reading of the previousversion Lesley Gibson Russell Palmer and Al Glen generously discussedvarious aspects of the paper with VAC The map was prepared by CristinaRamalho

References

Allen B L Engeman R M and Allen L R (2011) Wild dogma anexamination of recent lsquoevidencersquo for dingo regulation of invasivemesopredator release in Australia Current Zoology 57 568ndash583

Allen B L Fleming P J Allen L R Engeman R M Ballard G andLeung L K-P (2013) As clear as mud a critical review of evidencefor the ecological roles of Australian dingoes Biological Conservation159 158ndash174 doi101016jbiocon201212004

Anon (1934) Floods in Western Australia Town isolated for week PerthMonday The Sydney Morning Herald 9

Armstrong D P and Reynolds M H (2012) Modelling reintroducedpopulations the state of the art and future directions In lsquoReintroductionBiology Integrating Science and Managementrsquo (Eds J G Ewen DP Armstrong K A Parker and P J Seddon) pp 165ndash222 (Wiley-Blackwell UK)

Atlas Iron (2012) Scope of works ndash Mount Dove artificial quoll habitatUnpublished report Atlas Iron Perth Available at httpwwwatlasironcomauIRMCompanyShowPageaspxCategoryId=190ampCPID=5424ampEID=94365410 [accessed 19 August 2015]

Austin C (2014) Can remote cameras accurately estimate populations ofthe endangered northern quoll BSc(Honours) Thesis University ofTechnology Sydney

Begg R J (1981) The small mammals of Little Nourlangie RockNT III Ecology ofDasyurus hallucatus the northern quoll (MarsupialiaDasyuridae) Wildlife Research 8 73ndash85 doi101071WR9810073

Braithwaite R W and Griffiths A D (1994) Demographic variationand range contraction in the northern quoll Dasyurus hallucatus(Marsupialia Dasyuridae)Wildlife Research 21 203ndash217 doi101071WR9940203

Bull J W Suttle K B Gordon A Singh N J and Milner-Gulland E J(2013) Biodiversity offsets in theory and practice Oryx 47 369ndash380doi101017S003060531200172X

Burnett S (1997) Colonizing cane toads cause population declines innative predators reliable anecdotal information and managementimplications Pacific Conservation Biology 3 65ndash72

Cadenhead N C R White A L and Wintle B A (2015) Identifyingconservation priorities and assessing impacts of proposed futuredevelopment in the Pilbara Bioregion in Western Australia Report bythe National Environment Research Program Decisions Hub to theDepartment of the Environment The University of MelbourneMelbourne

Carwardine J Nicol S Van Leeuwen S Walters B Firn J Reeson AMartin T G and Chades I (2014) Priority threat management forPilbara species of conservation significance CSIRO EcosystemSciences Brisbane

Cook A (2010) Habitat use and home-range of the northern quollDasyurushallucatus effects of fire MSc Thesis University ofWestern AustraliaPerth

Department of Sustainability Environment Water Population andCommunities (2012) Environment and Biodiversity Conservation Act1999 Environmental Offsets Policy Department of SustainabilityEnvironment Water Population and Communities Canberra

Doherty T S Davis R A Etten E J Algar D Collier N DickmanC R Edwards G Masters P Palmer R and Robinson S (2015a)A continental-scale analysis of feral cat diet in Australia Journal ofBiogeography 42 964ndash975 doi101111jbi12469

Doherty T S Dickman C R Nimmo D G and Ritchie E G (2015b)Multiple threats or multiplying the threats Interactions betweeninvasive predators and other ecological disturbances BiologicalConservation 190 60ndash68 doi101016jbiocon201505013

Dunlop J Cook A and Lees J (2014a) BHP Billiton Iron Ore Pilbaranorthern quoll monitoring project Final report May 2010ndashJune 2013Unpublished report prepared for BHP Billiton Iron Ore Department ofParks and Wildlife Perth

Dunlop J Cook A and Morris K (2014b) Pilbara northern quollproject surveying and monitoring Dasyurus hallucatus in the PilbaraWestern Australia Department of Parks and Wildlife Perth Availableat librarydpawwagovau

Florance D Webb J K Dempster T Kearney M R Worthing A andLetnic M (2011) Excluding access to invasion hubs can contain thespread of an invasive vertebrate Proceedings of the Royal Society ofLondon B Biological Sciences 278 2900ndash2908 doi101098rspb20110032

Gaillard J-M Hebblewhite M Loison A Fuller M Powell R BasilleM and Van Moorter B (2010) Habitatndashperformance relationshipsfinding the right metric at a given spatial scale PhilosophicalTransactions of the Royal Society of London Series B BiologicalSciences 365 2255ndash2265 doi101098rstb20100085

Glen A S and Dickman C R (2008) Niche overlap between marsupialand eutherian carnivores does competition threaten the endangeredspotted-tailed quoll Journal of Applied Ecology 45 700ndash707 doi101111j1365-2664200701449x

Glen A S Pennay M Dickman C R Wintle B A and Firestone K B(2011) Diets of sympatric native and introduced carnivores in theBarrington Tops eastern Australia Austral Ecology 36 290ndash296doi101111j1442-9993201002149x

Hennig P (2004) A brief land use history In lsquoAn Inventory and ConditionSurvey of the Pilbara Region Western Australiarsquo (Eds A M E VanVreeswyk A L Payne K A Leighton and P Hennig) TechnicalBulletin No 92 pp 13ndash18 (Western Australian Department ofAgriculture Perth)

Hohnen R Ashby J Tuft K and McGregor H (2013) Individualidentification of northern quolls (Dasyurus hallucatus) using remotecameras Australian Mammalogy 35 131ndash135 doi101071AM12015

L Australian Mammalogy V A Cramer et al

How R A Spencer P B S and Schmitt L H (2009) Island populationshave high conservation value for northern Australiarsquos top marsupialpredator ahead of a threatening process Journal of Zoology 278206ndash217 doi101111j1469-7998200900569x

Johnson C N Isaac J L and Fisher D O (2007) Rarity of a top predatortriggers continent-wide collapse of mammal prey dingoes andmarsupials in Australia Proceedings of the Royal Society of LondonSeries B Biological Sciences 274 341ndash346 doi101098rspb20063711

Jones M E (2000) Road upgrade road mortality and remedial measuresimpacts on a population of eastern quolls and Tasmanian devilsWildlifeResearch 27 289ndash296 doi101071WR98069

Kearney M Phillips B L Tracy C R Christian K A Betts G andPorter W P (2008) Modelling species distributions without usingspecies distributions the cane toad in Australia under current andfuture climates Ecography 31 423ndash434 doi101111j0906-7590200805457x

Kendrick P and Stanley F (2002) Pilbara 4 (PIL4 ndash Roebourne synopsis)In lsquoA Biodiversity Audit of Western Australiarsquos 53 BiogeographicalSubregions in 2002rsquo (Eds J E May and N L McKenzie) pp 581ndash593(Department of Conservation and Land Management Perth)

Kennedy M Phillips B L Legge S Murphy S A and Faulkner R A(2012) Do dingoes suppress the activity of feral cats in northernAustralia Austral Ecology 37 134ndash139 doi101111j1442-9993201102256x

Kiesecker J M Copeland H Pocewicz A and McKenney B (2010)Development by design blending landscape-level planning with themitigation hierarchy Frontiers in Ecology and the Environment 8261ndash266 doi101890090005

King D R and Smith L A (1985) The distribution of the European red fox(Vulpes vulpes) in Western Australia Records of the Western AustralianMuseum 12 197ndash205

Knight A T Cowling R M Rouget M Balmford A Lombard A Tand Campbell B M (2008) Knowing but not doing selecting priorityconservation areas and the researchndashimplementation gap ConservationBiology 22 610ndash617 doi101111j1523-1739200800914x

Laurance W F Koster H Grooten M Anderson A B Zuidema P AZwick S Zagt R J Lynam A J Linkie M and Anten N P (2012)Making conservation research more relevant for conservationpractitioners Biological Conservation 153 164ndash168 doi101016jbiocon201205012

Legge S Murphy S Heathcote J Flaxman E Augusteyn J andCrossman M (2008) The short-term effects of an extensive and high-intensity fire on vertebrates in the tropical savannas of the centralKimberley northern Australia Wildlife Research 35 33ndash43 doi101071WR07016

Lester D (2015) Effective wildlife roadkill mitigation Journal of Trafficand Transportation Engineering 3 42ndash51 doi10172652328-2142201501005

Letnic M Tamayo B and Dickman C R (2005) The responsesof mammals to La Nintildea (El Nintildeo Southern Oscillation)-associatedrainfall predation and wildfire in central Australia Journal ofMammalogy 86 689ndash703 doi1016441545-1542(2005)086[0689TROMTL]20CO2

Letnic M Koch F Gordon C Crowther M S and Dickman C R(2009) Keystone effects of an alien top-predator stem extinctions ofnative mammals Proceedings of the Royal Society of London Series BBiological Sciences 276 3249ndash3256 doi101098rspb20090574

Lindenmayer D B (2015) Continental-level biodiversity collapseProceedings of the National Academy of Sciences of the United States ofAmerica 112 4514ndash4515 doi101073pnas1502766112

Lindenmayer D B Piggott M P and Wintle B A (2013) Counting thebooks while the library burns why conservation monitoring programs

need a plan for action Frontiers in Ecology and the Environment 11549ndash555 doi101890120220

Maron M Hobbs R J Moilanen A Matthews J W Christie KGardner T A Keith D A Lindenmayer D B and McAlpine C A(2012) Faustian bargains Restoration realities in the context ofbiodiversity offset policies Biological Conservation 155 141ndash148doi101016jbiocon201206003

McCall S C McCarthy M A van der Ree R Harper M J Cesarini Sand Soanes K (2010) Evidence that a highway reduces apparentsurvival rates of squirrel gliders Ecology and Society 15 27

McGregor R A Stokes V L and Craig M D (2014) Does forestrestoration in fragmented landscapes provide habitat for a wide-ranging carnivore Animal Conservation 17 467ndash475 doi101111acv12112

McKenzie N L Burbidge A A Baynes A Brereton R N DickmanC R Gordon G Gibson L A Menkhorst P W Robinson A CWilliams M R and Woinarski J C Z (2007) Analysis of factorsimplicated in the recent decline of Australiarsquos mammal fauna Journal ofBiogeography 34 597ndash611 doi101111j1365-2699200601639x

McKenzie N L van Leeuwen S and Pinder AM (2009) Introduction tothe Pilbara Biodiversity Survey 2002ndash2007 Records of the WesternAustralian Museum 78(Suppl) 3ndash89 doi1018195issn0313-122x78(1)2009003-089

Morris K Johnson B Orell P Gaikhorst G Wane A and MoroD (2003) Recovery of the threatened chuditch (Dasyurus geoffroii)a case study In lsquoPredators with Pouches The Biology of CarnivorousMarsupialsrsquo (Eds M Jones C Dickman and M Archer) pp 435ndash451(CSIRO Publishing Melbourne)

Nelson J L Scroggie M P and Belcher C A (2014) Developing acamera trap survey protocol to detect a rare marsupial carnivore thespotted-tailed quoll (Dasyurus maculatus) In lsquoCamera TrappingWildlife Management and Researchrsquo (Eds P D Meek and PJ Fleming) pp 271ndash279 (CSIRO Publishing Melbourne)

Newsome T M Dellinger J A Pavey C R Ripple W J Shores C RWirsing A J and Dickman C R (2015) The ecological effects ofproviding resource subsidies to predators Global Ecology andBiogeography 24 1ndash11 doi101111geb12236

OrsquoDonnell S Webb J K and Shine R (2010) Conditioned tasteaversion enhances the survival of an endangered predator imperilled bya toxic invader Journal of Applied Ecology 47 558ndash565 doi101111j1365-2664201001802x

Oakwood M (1997) The ecology of the northern quoll Dasyurushallucatus PhD Thesis Australian National University Canberra

Oakwood M (2000) Reproduction and demography of the northern quollDasyurus hallucatus in the lowland savanna of northern AustraliaAustralian Journal of Zoology 48 519ndash539 doi101071ZO00028

Oakwood M (2002) Spatial and social organization of a carnivorousmarsupial Dasyurus hallucatus (Marsupialia Dasyuridae) Journal ofZoology 257 237ndash248 doi101017S0952836902000833

Oakwood M and Foster P (2008) Monitoring extinction of the northernquoll Australian Academy of Science Newsletter 71 6

Palomares F and Caro T M (1999) Interspecific killing amongmammalian carnivores American Naturalist 153 492ndash508 doi101086303189

Pavey C R and Bastin G (2014) lsquoAustralian Rangelands and ClimateChange ndash Invasive Animalsrsquo (Ninti One Limited amp CSIRO AliceSprings)

Pierson J C Barton P S Lane PW and Lindenmayer D B (2015) Canhabitat surrogates predict the response of target species to landscapechange Biological Conservation 184 1ndash10 doi101016jbiocon201412017

Pollock A B (1999) Notes on status distribution and diet of northernquoll Dasyurus hallucatus in the MackayndashBowen area mideastern

Research priorities for the northern quoll Australian Mammalogy M

Queensland Australian Zoologist 31 388ndash395 doi107882AZ1999040

Ramanaidou E R and Morris R C (2010) A synopsis of the channeliron deposits of the Hamersley Province Western AustraliaTransactions of the Institutions of Mining and Metallurgy Section BApplied Earth Science 119 56ndash59 doi101179037174510X12853354810624

Schmitt L H Bradley A J Kemper C M Kitchener D J HumphreysW F and How R A (1989) Ecology and physiology of the northernquoll Dasyurus hallucatus (Marsupialia Dasyuridae) at MitchellPlateau Kimberley Western Australia Journal of Zoology 217539ndash558 doi101111j1469-79981989tb02510x

Short J and Smith A (1994) Mammal decline and recovery in AustraliaJournal of Mammalogy 75 288ndash297 doi1023071382547

Spencer P How R Hillyer M Cook A Morris K Stevenson C andUmbrello L (2013) Genetic analysis of northern quolls from the PilbaraRegion ofWestern Australia Year one ndash final report Unpublished reportDepartment of Sustainability Environment Water Population andCommunities Canberra and Department of Parks and Wildlife Perth

Sutherland W J Fleishman E Mascia M B Pretty J and Rudd M A(2011) Methods for collaboratively identifying research priorities andemerging issues in science and policyMethods in Ecology and Evolution2 238ndash247 doi101111j2041-210X201000083x

Sutherst R W Floyd R B and Maywald G F (1996) The potentialgeographical distribution of the cane toad Bufo marinus L in AustraliaConservation Biology 10 294ndash299 doi101046j1523-1739199610010294x

Taylor B D and Goldingay R L (2003) Cutting the carnage wildlifeusage of road culverts in north-eastern New South Wales WildlifeResearch 30 529ndash537 doi101071WR01062

Tingley R Phillips B L Letnic M Brown G P Shine R and BairdS J (2013) Identifying optimal barriers to halt the invasion of canetoads Rhinella marina in arid Australia Journal of Applied Ecology 50129ndash137 doi1011111365-266412021

Van Vreeswyk A M E Payne A L Leighton K A and HennigP (2004) An Inventory and Condition Survey of the Pilbara RegionWestern Australia Technical Bulletin No 92 Department of AgricultureWestern Australia Perth

Webb J Legge S Tuft K Cremona T and Austin C (2015) Can wemitigate cane toad impacts on northern quolls Final report National

Environmental Research Program Northern Australia Hub CharlesDarwin University Darwin

Wintle B A Walshe T V Parris K M and McCarthy M A (2012)Designing occupancy surveys and interpreting non-detection whenobservations are imperfect Diversity amp Distributions 18 417ndash424doi101111j1472-4642201100874x

Woinarski J Menkhorst K Gambold N and Braithwaite R W (1992)Mammals of the Bungle Bungle area In lsquoA Survey of the Wildlife andVegetation of Purnululu (Bungle Bungle) National Park and AdjacentArearsquo (Ed J C Z Woinarski) Research Bulletin No 6 pp 53ndash67(CSIRO Division of Wildlife and Ecology Darwin)

Woinarski J C Z Risler J and Kean L (2004) Response of vegetationand vertebrate fauna to 23 years of fire exclusion in a tropical Eucalyptusopen forest Northern Territory Australia Austral Ecology 29 156ndash176doi101111j1442-9993200401333x

Woinarski J C Z Oakwood M Winter J Burnett S Milne D FosterP Myles H and Holmes B (2008) Surviving the toads patterns ofpersistence of the northern quoll Dasyurus hallucatus in QueenslandReport submitted to the Natural Heritage Trust Strategic ReserveProgram as a component of project 2005162 Monitoring andManagement of Cane Toad Impact in theNorthern Territory Available athttplrmntgovauplants-and-animalsinformation-and-publicationspublications2008 [accessed 14 November 2013]

Woinarski J C Z Armstrong M Brennan K Fisher A Griffiths A DHill B Milne D J Palmer C Ward S Watson M Winderlich Sand Young S (2010) Monitoring indicates rapid and severe decline ofnative small mammals in Kakadu National Park northern AustraliaWildlife Research 37 116ndash126 doi101071WR09125

Woinarski J Burbidge A and Harrison P (2014) lsquoAction Plan forAustralian Mammals 2012rsquo (CSIRO Publishing Melbourne)

Woinarski J C Burbidge A A and Harrison P L (2015) Ongoingunraveling of a continental fauna decline and extinction of Australianmammals since European settlement Proceedings of the NationalAcademy of Sciences of the United States of America 112 4531ndash4540doi101073pnas1417301112

Woolley P A Krajewski C and Westerman M (2015) Phylogeneticrelationships within Dasyurus (Dasyuromorphia Dasyuridae) quollsystematics based on molecular evidence and male characteristicsJournal of Mammalogy 96 37ndash46 doi101093jmammalgyu028

N Australian Mammalogy V A Cramer et al

wwwpublishcsiroaujournalsam

Page 8: Research priorities for the northern quoll (Dasyurus ......The Pilbara population of the northern quoll (Dasyurus hallucatus) has been seldom studied, and the impacts of threats such

studies have been undertaken in the region Cadenhead et al(2015) used demographic information from the NorthernTerritory and Kimberley populations in their PVA Uncertaintyabout data quality and the limited information available ondemographic parameters for the Pilbara constrained the level ofcertainty around the outcomes of the PVA The model doeshowever provide a framework into which new data can beincorporated as they become available For example preliminarydata from the Pilbara monitoring program suggest that theproportion of male die-off in the Pilbara population is lowerthan in the Kimberley and Northern Territory populations(Dunlop unpubl data) Cadenhead et al (2015) stress the needfor more demographic and occurrence data to be collectedbefore further development decisions are made (see lsquoDevelopappropriate and standardised survey and monitoring methodsrsquoand lsquoImprove our understanding of habitat requirementsrsquo above)

Investigate population structure and the interactionbetween populations

Analysis of the genetic diversity and population structure ofthe Pilbara population of the northern quoll is ongoingOnly small differences in genetic diversity as measured byheterozygosity were found between samples obtained at 16locations within the Pilbara with the exception of the isolatedpopulation on Dolphin Island (Spencer et al 2013) This resultindicates a high level of movement across the landscape byindividuals during mating and dispersal phases Spatial structureanalysis of samples collected from around Pannawonicasuggested a lsquoneighbour sizersquo of ~15 km (Spencer et al 2013)Current data show no evidence of genetic bottleneckssuggesting that if there has been a population decline in thePilbara that this has not affected genetic diversity A usefulapproach would be to combine genetic information with datafrom radio-tracking or GPS collars to better understand theuse of lsquolandscapersquo space and how this influences populationdynamics

Priority 4 Key threats and the interaction of these threats

Introduced predators

Introduced cats and foxes are considered to be one of themain causes of the decline and extinction of the critical-weight-range mammals (35 g to 55 kg) in Australia (Short and Smith1994 McKenzie et al 2007 Woinarski et al 2015) but theirimpacts on the northern quoll are less clear A recent review offeral cat diets across Australia found that northern quolls werepresent in the diet of feral cats (Doherty et al 2015a) andWoinarski et al (2014) consider the threat posed by cats to thenorthern quoll to be severe although the impacts of cats may bereduced in rugged refuge areas The impacts of cats on othermuch larger species of quoll appear to be inconsistent withsome evidence of coexistence between cats and quolls beforethe arrival of foxes (Glen and Dickman 2008 and referencescontained within) Due to the limited overlap in their distributionthroughout Australia the threat posed by foxes to northernquolls is less understood than for cats and Woinarski et al(2014) consider the threat posed by foxes to be moderate on anational level Foxes are common in coastal near-coastal and

riparian areas of the Pilbara and extend as far north as Broome(King and Smith 1985 Kendrick and Stanley 2002) They arerenowned for their aggression towards smaller carnivores(Palomares and Caro 1999) and neither the larger chuditch(Dasyurus geoffroii) nor spotted-tail quoll (Dasyurus maculatus)are known to coexist with large numbers of foxes (A Glen perscomm) The interactions between quolls cats and foxesare potentially more complex than for other medium-sizedmarsupials due to niche overlap in diet and competition for densites with the larger spotted-tailed quoll and chuditch bothexperiencing exploitation and interference competition fromfoxes (Morris et al 2003 Glen et al 2011) Competitionbetween feral cats and the northern quoll may potentially beintense during periods of low rainfall in the Pilbara when preyavailability is low Understanding whether foxes and cats aresignificant predators of or competitors with the northern quollin the Pilbara and how this affects their population viabilitywill clarify the level of risk posed by these predators Thesausage-style baits used for feral cats (eg Eradicatreg andCuriosityreg) are potentially attractive to the northern quoll withthe risk of mortality or reduced fitness Trials to assess themagnitude of the risks from Eradicatreg baits are currentlyunderway in the Pilbara

The role of dingoes and wild dogs in the mortality of andinteractions between mesopredators (both native and introduced)requires further investigation Dingoes are known to kill northernquolls butmay not eat these kills (Oakwood 1997) A recent studyin the Northern Territory found that camp dogs and dingoes werethe major source of mortality for northern quolls at the study site(Webb et al 2015) and roaming domestic dogs were identified asa cause of mortality for northern quolls in Queensland (Pollock1999) Human resource subsidies (eg stock carrion in rangelandareas) can artificially support predator populations thus indirectlyincreasing predation pressure on prey (Newsome et al 2015)A dynamic debate currently exists around the role of the dingo inregulating populations of feral cats and foxes and the implicationsof this for the conservation of the critical-weight-range mammalsin Australia (see Johnson et al 2007 Letnic et al 2009 Allenet al 2011 2013 Kennedy et al 2012) Understanding thepopulation density at which the role of the dingo as a regulator ofintroduced predators outweighs its own impact as a direct predatorof the northern quoll will inform whether dingowild dog controlwould benefit the northern quoll

Cane toads

The introduced cane toad now occupies more than half thehistoric range of the northern quoll and has caused an immediatecollapse or extirpation of northern quoll populations in theareas it has invaded (Burnett 1997 Oakwood and Foster2008 Woinarski et al 2008 2010) Earlier modelling work thatconsidered only climate parameters as well as more recentwork that incorporates future climate change suggest thatcane toads will not invade the Pilbara (Sutherst et al 1996Pavey and Bastin 2014) Other recent work that considers thefundamental niche of the cane toad and incorporates the locationof artificial watering points predicts that they will colonisethe Pilbara from the Kimberley via the coastal edge of theGreat Sandy Desert (Kearney et al 2008 Florance et al 2011

H Australian Mammalogy V A Cramer et al

Tingley et al 2013) This has created some level of uncertaintyabout the likely impact of cane toads on northern quolls in thePilbara Strategies for cane toad management were not rankedhighly by participants in the threat-management frameworkdeveloped by Carwardine et al (2014) which the authorsexplained as possibly due to uncertainty about how far canetoads will spread into the region

The model developed by Tingley et al (2013) suggested thatthe closure of 100 artificial water bodies at one of three potentiallocations along the coastal corridor between the Kimberley andthe Pilbara would prevent the invasion of cane toads into thePilbara While their model did account for uncharacteristicallywet years every three decades the dispersal of toads betweenwater bodies was determined by the number of rainy daysoccurring during the year (ie toads could only disperse betweenwater bodies for three days following rainfall) rather thanrainfall amount the associated likelihood of flooding and theavailability of free water across the landscape The cyclonic andthunderstorm nature of much of the rainfall in the north-west ofWestern Australia means that the region is prone to significantand sometimes extended flooding In April 1934 the countryaround Anna Plains (250 km south of Broome) was flooded forhundreds of miles when 27 inches (686mm) of rain fell infour days (Anon 1934) In March 2004 over 350mm of rain wasrecorded within a 48-h period at Telfer on the southern edge ofthe Great Sandy Desert due to Tropical Cyclone Fay Thiscaused the flooding of the De Grey and Oakover river systemsand flooded 140 km of the Telfer Access Road with floodwaterstaking several months to subside (wwwbomgovau) Extendingthe model of Tingley et al (2013) to account for such events willhelp clarify whether closing artificial water bodies is likely toprevent the invasion of cane toads into the Pilbara or may onlyassist in delaying their inevitable invasion

Comparison of SDMs (eg Cadenhead et al 2015) fornorthern quoll with fundamental niche models for cane toads(eg see Kearney et al 2008) within the Pilbara may identifypotential refuge areas for the northern quoll and these areasshould be prioritised for conservation actions Because theinvasion point of cane toads into the Pilbara is likely to be narrow(beginning around the mouth of the De Grey River Tingleyet al 2013) a proactive approach to training northern quollsto avoid cane toads may prove beneficial Conditioned taste-aversion (CTA) has been used successfully to train captive-reared northern quolls to avoid cane toads by offering themsmall dead toads infused with thiabendazole before theirrelease into the wild (OrsquoDonnell et al 2010)Monitoring of thesequolls after their release showed that some of their offspringhad learnt to avoid eating cane toads either through ingestionof small non-lethal toads or via social learning from theirmothers (Webb et al 2015) Fields trials of toad-aversion baits(either sausages or meatballs made from toad flesh andcontaining thiabendazole) are currently underway in the centralKimberley where cane toads were expected to invade in late2015 (Webb et al 2015) Further testing of CTA approachesunder a variety of field conditions is required The developmentof stable long-lasting baits suitable for aerial deployment isconsidered a pressing need as well as a captive colony ofnorthern quolls upon which to test the efficacy of toad-aversionbaits for inducing aversion to live toads (Webb et al 2015)

Infrastructure development and interactionswith humans

Infrastructure development may increase the likelihood ofnorthern quoll deaths due to motor vehicle strikes or beingcrushed in machinery Significant chuditch mortality wasobserved along mine access roads in the south-west of WesternAustralia (McGregor et al 2014) and the widening and sealingof an access road dramatically increased the number of roadkills and led to the extirpation of the resident population ofeastern quolls (D viverrinus) in Tasmania (Jones 2000)Infrastructure corridors such as high-tension power-lineeasements and ore-haulage railways are increasingly beinginstalled throughout the range of the northern quoll Althoughthese do not create the same mortality risk as roads they bisectand fragment habitat create large open spaces and potentiallyincrease predation risk

A recent small-scale radio-tracking study (see Dunlop et al2014a) around several quarries in the Pilbara found that northernquolls crossed roads but not railway lines The short-term natureof this work and the limited number of quolls able to be trackedmeans that the influence of roads railway lines and infrastructurecorridors on the movement and mortality of northern quollsrequires further assessment Using radio-tracking techniques todetermine crossing rates and road kill events may be difficultover large areas so population studies comparing groups ofquolls near major roads with those not affected by infrastructure(eg McCall et al 2010) may provide a more feasible meansof assessing the impacts of roads If vehicle strikes are foundto significantly contribute to the mortality of a local populationof the northern quoll then mitigation methods should betrialled on sections of road identified as road-kill hotspotsCulverts and underpasses have been shown to be used by avariety of small and medium-sized mammals and frogs (Taylorand Goldingay 2003) while installing rumble strips on roadshas led to decreased rates of road-kill of nocturnal animals(Lester 2015)

Interactions between threats

At present wildfire the subsequent loss of vegetation coverand increased predation of northern quolls is likely to be themost significant of the interactions between threats The directeffects of wildfire have been found to be less significant inthe subsequent decline of small to medium-sized mammalsparticularly for denning species such as the northern quoll thanthe indirect consequences of reduced resource availability andthe increased risk of predation due to lack of cover (Oakwood2000 Letnic et al 2005 McKenzie et al 2007 Legge et al2008 Doherty et al 2015b) Male northern quolls are perhapsmost vulnerable to such losses of cover during the breedingseason when their range can extend to over 100 ha (Oakwood2002) or where males survive after breeding their foragingrange is extended to gain the nutrition required to recover fromtheir poor condition (Cook 2010) Loss of cover due to late dry-season fires at East Alligator River in the Northern Territorymeant little cover was available for young quolls when theybegan foraging with dog and dingo predation a major source ofmortality (Webb et al 2015) The cumulative impacts of canetoad poisoning fire and predation are highly likely to lead to the

Research priorities for the northern quoll Australian Mammalogy I

extirpation of small and disjunct groups of northern quoll in thePilbara Population viability modelling of the East AlligatorRiver population suggested that high mortality from predationcombined with late dry-season burning was preventing therecovery of northern quoll populations after cane toad invasion

(Webb et al 2015) Improved knowledge of the severity of thesethreats to northern quolls in the Pilbara and their incorporationinto the SDM and PVA framework developed by Cadenheadet al (2015) will allow for better identification of priority areasfor integrated cane toad fire and predator management

Table 3 Expected timelines for the undertaking of research under each priority over the next 10 years and the funds currently allocated fromthe Pilbara Northern Quoll Offset Trust under each research priority

Rank Theme Research priority Current or proposed researchactivity

Time over next10 years

Current funding allocation(approximate)

1 Survey andmonitoring

Develop appropriate andstandardised survey andmonitoring methods

Develop monitoring protocol(traps and cameras) andinvestigate new methods

Ongoing survey and monitoringin conjunction with ResearchThemes 2 and 3 to better mapnorthern quoll distribution andto determine long-termpopulation trends

Years 1ndash2Years 3ndash10

$200 000 per annum over10 years for ResearchThemes 1 2 and 3(inclusive of salaries)

2 Habitatrequirements

Define areas of criticalhabitat for northernquoll in the Pilbara

Identify and describe criticalhabitat and undertake predictivemodelling of unsurveyed areasto inform further survey effort

Years 3ndash4 $50 000 for predictivemodelling (EdithCowan University)

Understand howdisturbance affectshabitat quality andconnectivity

Ongoing collection of data ondisturbance and habitat qualityin conjunction with ResearchThemes 1 and 4

Incorporation of new data intopredictive models

Years 4ndash10 See above

3 Populationdynamics

Better understand thepopulation dynamics ofthe northern quoll in thePilbara

Collect data on demographicparameters breeding dispersaland home-range use toincorporate into future PVA

Continued collectionof data inconjunction withpriorities 1 and 2during years 3ndash10

See above

Investigate populationstructure and theinteraction betweenpopulations

Analysis of existing tissuesamples and newly collectedsamples

Years 1ndash10 $120 000 for initial roundof genetic analysis(Years 3ndash5 MurdochUniversity)

4 Key threats Introduced predators Spatio-temporal interactionsbetween northern quolls andintroduced predators

Other projects as they arise

Years 2ndash4 (currentPhD projectUniversity ofQueensland)

$12 000 contribution tofield expenses

Cane toads Predictive modelling of canetoad distribution in the Pilbara

Years 3ndash4 Part of predictivemodelling by EdithCowan University

Infrastructuredevelopment andinteractions withhumans

Investigation of impacts oflinear infrastructure on northernquoll movement

Year 4 (currentHonours projectEdith CowanUniversity)

$15 000

Interactions betweenthreats

No on-ground projects currentlyplanned

Potential to be assessed throughPVA as data are collected underother research priorities

No funding currentlyallocated

5 Recolonisation ofrestored orartificial habitat

Determine the ability ofthe northern quoll torecolonise disturbedareas or coloniseartificial habitat

As projects arise Years 5ndash8 incollaboration withminesite restorationprograms

No funding currentlyallocated

J Australian Mammalogy V A Cramer et al

Priority 5 Recolonisation of restored andor artificial habitat

While anecdotal evidence suggests that northern quolls will denin waste rock dumps whether artificial or restored habitatprovides for long-term population viability is not known andlong-term assessment of created habitat is required Weacknowledge that a large gap exists between the desire to userestored or artificial habitat as an ecologically equivalent offsetand our limited ability to actually restore or re-create habitat thatis equivalent in structure composition and function particularlyon highly disturbed sites (Maron et al 2012) Yet there is a futureopportunity in the Pilbara to conduct lsquonaturalrsquo experiments onthe recolonisation of quolls after mining ceases including howto best design waste rock dumps so that habitat complexityand productivity is maximised by for example experimentingwith the size and positioning of boulders and their spatialarrangement in relation to surrounding landscape features Theuse of such artificial habitat should be tested in the first stagethrough direct monitoring of animals that have colonised orhave been translocated to the site Such experimentation isunderway at Atlas Ironrsquos Mt Dove operation (Atlas Iron 2012)Population modelling can then be used to predict futurepopulation sizes rates of growth and spread and the probabilityof long-term persistence in the context of various managementactions (eg predator control Armstrong and Reynolds 2012)

Resource allocation and timelines

While the allocation of resources and funding and the periodover which each of the research priorities was expected to beaddressed was not explicitly discussed during the workshopthe ranking of the priorities was taken to indicate the sequencein which the priorities should be addressed At the time of theworkshop some of the research questions identified werealready being investigated within various projects and a portionof the Pilbara Northern Quoll Offset Trust allocated to these InTable 3 we indicate the expected timelines over which someof the research programs are expected to be undertaken WhileAU$175M has been allocated to northern quoll research in thePilbara it is important to note that much of this amount is notcurrently available and will not become available until severalapproved mine developments have become operational InTable 3 we also indicate the proportion of offset funds expectedto be allocated to each research priority Research undertaken inother regions (eg research into CTA in the Kimberley regionahead of the cane toad invasion front) cannot be funded from thePilbara Northern Quoll Offset Trust but will help address someof the research questions identified here

Lessons learnt from the workshop process

The workshop approach used here was less formal than recentlarge-scale exercises used to elicit research priorities for emergingissues in science and policy at national or global levels (reviewedin Sutherland et al 2011) The focus in our (much smaller)workshop was on inclusivity rather than identifying participantsbased on purposive sampling of expertise or organisationalaffiliation We believe that the inclusive nature of the workshopwas a strength but we also recognise that this perhaps createdsome level of bias when ranking the research prioritiesAcknowledging both the experience and knowledge requirements

of those who work regularly in the Pilbara is an importantcomponent of improving the design of research programs that aimto improve conservation management in the region Yet theresearch priorities and their rankings are likely strongly reflectiveof the current information needs of most participants and maynot align with what conservation scientists believe to be themost critical areas of research for northern quoll conservation intothe future Baseline survey and monitoring are often the firstrequirements of a management or research plan for the northernquoll when offset conditions are prescribed and this was reflectedin the primary ranking given to the development of standardsurvey and monitoring methods We acknowledge the concernthat this could simply lead to offset funding being used to lsquomonitorthe extinctionrsquo of the northern quoll particularly after canetoads invade the Pilbara unless these data are used to triggermanagement actions (Oakwood and Foster 2008 Lindenmayeret al 2013) Clearly the currently limited information on habitatuse and demographic parameters for the northern quoll in thePilbara restricts the utility of more sophisticated researchapproaches such as developing robust SDM and PVA While thesevere threat that cane toads pose to the northern quoll wasrecognised this threat is not immediate in the region and mayexplain the less prominent ranking given to research to mitigatethe impacts of cane toads

The northern quoll has become emblematic of the dramaticand rapid species declines occurring in northern Australia andthe level of interest in this species was reflected in the highnumber of participants attending the workshop (one of six runby Parks and Wildlife) Given the number of attendees betterutilisation of the time available would have been gained byproviding participants with a brief before the workshopand asking them to contribute potential research questionsanonymously for interactive review by other participants Inthe exercises reviewed by Sutherland et al (2011) questionswere then discarded if they were not considered to meet criteriathat were previously agreed upon Some areas of researchthat gained some attention on the day may have been discardedif given greater consideration or interrogation before theworkshop For example there was some discussion duringthe workshop of investigating the use of habitat surrogates tomonitor the northern quoll yet research has shown that there islittle validity in using surrogates to monitor the abundance of asingle species (Pierson et al 2015)

Conclusions

Offset funds generated under the environmental approvalsprocess provide an opportunity to mobilise considerablefinancial resources for conservation (Kiesecker et al 2010) Yetthere remain significant conceptual and practical limitations tobe resolved to ensure that direct offsets provide measurableconservation outcomes that are ecologically equivalent to thelosses sustained by proceeding with the development (Maronet al 2012 Bull et al 2013) The collaborative identificationand ranking of research priorities undertaken here is the firststep in the process of developing and evaluating potentialconservation actions for the northern quoll in the Pilbara andserves as a mechanism to ensure that the research priorities ofstate and federal agencies reflects the knowledge needs of

Research priorities for the northern quoll Australian Mammalogy K

practitioners and proponents currently operating in the regionThe provision of offset funds over the next decade providesa unique opportunity to undertake research on a species ofconservation significance at a regional scale and over a longperiod through an integrated and strategic program of researchrather than the collection of ad hoc data While a morestructured approach to defining research questions before theworkshop may have led to greater critical reflection upon anddiscussion of some of the research priorities on the day theworkshop provided an opportunity for an open and diverse-ranging discussion in a collaborative environment amongstthe broader community of scientists environmental consultantsand mining proponents involved in the management andconservation of the northern quoll in the Pilbara

Acknowledgements

We gratefully acknowledge the financial support of Roy Hill who sponsoredthe workshop and partially funded VACrsquos salary We thank the Departmentof the Environmentrsquos Pilbara Taskforce especially Veronica Ritchie fortheir support of interested or affected parties The success of the workshopwas in great part due to the facilitation skills of Craig Salt of SustainableConsulting We thank all of the workshop participants for their open andcollaborative approach to the process Mike Bamford Eddie van EttenDavid Peacock and Peter Lyon provided useful comments on an earlier draftWe are grateful to John Woinarski for his constructive advice thatsubstantially improved the structure and clarity of the manuscript and ananonymous reviewer for careful and critical reading of the previousversion Lesley Gibson Russell Palmer and Al Glen generously discussedvarious aspects of the paper with VAC The map was prepared by CristinaRamalho

References

Allen B L Engeman R M and Allen L R (2011) Wild dogma anexamination of recent lsquoevidencersquo for dingo regulation of invasivemesopredator release in Australia Current Zoology 57 568ndash583

Allen B L Fleming P J Allen L R Engeman R M Ballard G andLeung L K-P (2013) As clear as mud a critical review of evidencefor the ecological roles of Australian dingoes Biological Conservation159 158ndash174 doi101016jbiocon201212004

Anon (1934) Floods in Western Australia Town isolated for week PerthMonday The Sydney Morning Herald 9

Armstrong D P and Reynolds M H (2012) Modelling reintroducedpopulations the state of the art and future directions In lsquoReintroductionBiology Integrating Science and Managementrsquo (Eds J G Ewen DP Armstrong K A Parker and P J Seddon) pp 165ndash222 (Wiley-Blackwell UK)

Atlas Iron (2012) Scope of works ndash Mount Dove artificial quoll habitatUnpublished report Atlas Iron Perth Available at httpwwwatlasironcomauIRMCompanyShowPageaspxCategoryId=190ampCPID=5424ampEID=94365410 [accessed 19 August 2015]

Austin C (2014) Can remote cameras accurately estimate populations ofthe endangered northern quoll BSc(Honours) Thesis University ofTechnology Sydney

Begg R J (1981) The small mammals of Little Nourlangie RockNT III Ecology ofDasyurus hallucatus the northern quoll (MarsupialiaDasyuridae) Wildlife Research 8 73ndash85 doi101071WR9810073

Braithwaite R W and Griffiths A D (1994) Demographic variationand range contraction in the northern quoll Dasyurus hallucatus(Marsupialia Dasyuridae)Wildlife Research 21 203ndash217 doi101071WR9940203

Bull J W Suttle K B Gordon A Singh N J and Milner-Gulland E J(2013) Biodiversity offsets in theory and practice Oryx 47 369ndash380doi101017S003060531200172X

Burnett S (1997) Colonizing cane toads cause population declines innative predators reliable anecdotal information and managementimplications Pacific Conservation Biology 3 65ndash72

Cadenhead N C R White A L and Wintle B A (2015) Identifyingconservation priorities and assessing impacts of proposed futuredevelopment in the Pilbara Bioregion in Western Australia Report bythe National Environment Research Program Decisions Hub to theDepartment of the Environment The University of MelbourneMelbourne

Carwardine J Nicol S Van Leeuwen S Walters B Firn J Reeson AMartin T G and Chades I (2014) Priority threat management forPilbara species of conservation significance CSIRO EcosystemSciences Brisbane

Cook A (2010) Habitat use and home-range of the northern quollDasyurushallucatus effects of fire MSc Thesis University ofWestern AustraliaPerth

Department of Sustainability Environment Water Population andCommunities (2012) Environment and Biodiversity Conservation Act1999 Environmental Offsets Policy Department of SustainabilityEnvironment Water Population and Communities Canberra

Doherty T S Davis R A Etten E J Algar D Collier N DickmanC R Edwards G Masters P Palmer R and Robinson S (2015a)A continental-scale analysis of feral cat diet in Australia Journal ofBiogeography 42 964ndash975 doi101111jbi12469

Doherty T S Dickman C R Nimmo D G and Ritchie E G (2015b)Multiple threats or multiplying the threats Interactions betweeninvasive predators and other ecological disturbances BiologicalConservation 190 60ndash68 doi101016jbiocon201505013

Dunlop J Cook A and Lees J (2014a) BHP Billiton Iron Ore Pilbaranorthern quoll monitoring project Final report May 2010ndashJune 2013Unpublished report prepared for BHP Billiton Iron Ore Department ofParks and Wildlife Perth

Dunlop J Cook A and Morris K (2014b) Pilbara northern quollproject surveying and monitoring Dasyurus hallucatus in the PilbaraWestern Australia Department of Parks and Wildlife Perth Availableat librarydpawwagovau

Florance D Webb J K Dempster T Kearney M R Worthing A andLetnic M (2011) Excluding access to invasion hubs can contain thespread of an invasive vertebrate Proceedings of the Royal Society ofLondon B Biological Sciences 278 2900ndash2908 doi101098rspb20110032

Gaillard J-M Hebblewhite M Loison A Fuller M Powell R BasilleM and Van Moorter B (2010) Habitatndashperformance relationshipsfinding the right metric at a given spatial scale PhilosophicalTransactions of the Royal Society of London Series B BiologicalSciences 365 2255ndash2265 doi101098rstb20100085

Glen A S and Dickman C R (2008) Niche overlap between marsupialand eutherian carnivores does competition threaten the endangeredspotted-tailed quoll Journal of Applied Ecology 45 700ndash707 doi101111j1365-2664200701449x

Glen A S Pennay M Dickman C R Wintle B A and Firestone K B(2011) Diets of sympatric native and introduced carnivores in theBarrington Tops eastern Australia Austral Ecology 36 290ndash296doi101111j1442-9993201002149x

Hennig P (2004) A brief land use history In lsquoAn Inventory and ConditionSurvey of the Pilbara Region Western Australiarsquo (Eds A M E VanVreeswyk A L Payne K A Leighton and P Hennig) TechnicalBulletin No 92 pp 13ndash18 (Western Australian Department ofAgriculture Perth)

Hohnen R Ashby J Tuft K and McGregor H (2013) Individualidentification of northern quolls (Dasyurus hallucatus) using remotecameras Australian Mammalogy 35 131ndash135 doi101071AM12015

L Australian Mammalogy V A Cramer et al

How R A Spencer P B S and Schmitt L H (2009) Island populationshave high conservation value for northern Australiarsquos top marsupialpredator ahead of a threatening process Journal of Zoology 278206ndash217 doi101111j1469-7998200900569x

Johnson C N Isaac J L and Fisher D O (2007) Rarity of a top predatortriggers continent-wide collapse of mammal prey dingoes andmarsupials in Australia Proceedings of the Royal Society of LondonSeries B Biological Sciences 274 341ndash346 doi101098rspb20063711

Jones M E (2000) Road upgrade road mortality and remedial measuresimpacts on a population of eastern quolls and Tasmanian devilsWildlifeResearch 27 289ndash296 doi101071WR98069

Kearney M Phillips B L Tracy C R Christian K A Betts G andPorter W P (2008) Modelling species distributions without usingspecies distributions the cane toad in Australia under current andfuture climates Ecography 31 423ndash434 doi101111j0906-7590200805457x

Kendrick P and Stanley F (2002) Pilbara 4 (PIL4 ndash Roebourne synopsis)In lsquoA Biodiversity Audit of Western Australiarsquos 53 BiogeographicalSubregions in 2002rsquo (Eds J E May and N L McKenzie) pp 581ndash593(Department of Conservation and Land Management Perth)

Kennedy M Phillips B L Legge S Murphy S A and Faulkner R A(2012) Do dingoes suppress the activity of feral cats in northernAustralia Austral Ecology 37 134ndash139 doi101111j1442-9993201102256x

Kiesecker J M Copeland H Pocewicz A and McKenney B (2010)Development by design blending landscape-level planning with themitigation hierarchy Frontiers in Ecology and the Environment 8261ndash266 doi101890090005

King D R and Smith L A (1985) The distribution of the European red fox(Vulpes vulpes) in Western Australia Records of the Western AustralianMuseum 12 197ndash205

Knight A T Cowling R M Rouget M Balmford A Lombard A Tand Campbell B M (2008) Knowing but not doing selecting priorityconservation areas and the researchndashimplementation gap ConservationBiology 22 610ndash617 doi101111j1523-1739200800914x

Laurance W F Koster H Grooten M Anderson A B Zuidema P AZwick S Zagt R J Lynam A J Linkie M and Anten N P (2012)Making conservation research more relevant for conservationpractitioners Biological Conservation 153 164ndash168 doi101016jbiocon201205012

Legge S Murphy S Heathcote J Flaxman E Augusteyn J andCrossman M (2008) The short-term effects of an extensive and high-intensity fire on vertebrates in the tropical savannas of the centralKimberley northern Australia Wildlife Research 35 33ndash43 doi101071WR07016

Lester D (2015) Effective wildlife roadkill mitigation Journal of Trafficand Transportation Engineering 3 42ndash51 doi10172652328-2142201501005

Letnic M Tamayo B and Dickman C R (2005) The responsesof mammals to La Nintildea (El Nintildeo Southern Oscillation)-associatedrainfall predation and wildfire in central Australia Journal ofMammalogy 86 689ndash703 doi1016441545-1542(2005)086[0689TROMTL]20CO2

Letnic M Koch F Gordon C Crowther M S and Dickman C R(2009) Keystone effects of an alien top-predator stem extinctions ofnative mammals Proceedings of the Royal Society of London Series BBiological Sciences 276 3249ndash3256 doi101098rspb20090574

Lindenmayer D B (2015) Continental-level biodiversity collapseProceedings of the National Academy of Sciences of the United States ofAmerica 112 4514ndash4515 doi101073pnas1502766112

Lindenmayer D B Piggott M P and Wintle B A (2013) Counting thebooks while the library burns why conservation monitoring programs

need a plan for action Frontiers in Ecology and the Environment 11549ndash555 doi101890120220

Maron M Hobbs R J Moilanen A Matthews J W Christie KGardner T A Keith D A Lindenmayer D B and McAlpine C A(2012) Faustian bargains Restoration realities in the context ofbiodiversity offset policies Biological Conservation 155 141ndash148doi101016jbiocon201206003

McCall S C McCarthy M A van der Ree R Harper M J Cesarini Sand Soanes K (2010) Evidence that a highway reduces apparentsurvival rates of squirrel gliders Ecology and Society 15 27

McGregor R A Stokes V L and Craig M D (2014) Does forestrestoration in fragmented landscapes provide habitat for a wide-ranging carnivore Animal Conservation 17 467ndash475 doi101111acv12112

McKenzie N L Burbidge A A Baynes A Brereton R N DickmanC R Gordon G Gibson L A Menkhorst P W Robinson A CWilliams M R and Woinarski J C Z (2007) Analysis of factorsimplicated in the recent decline of Australiarsquos mammal fauna Journal ofBiogeography 34 597ndash611 doi101111j1365-2699200601639x

McKenzie N L van Leeuwen S and Pinder AM (2009) Introduction tothe Pilbara Biodiversity Survey 2002ndash2007 Records of the WesternAustralian Museum 78(Suppl) 3ndash89 doi1018195issn0313-122x78(1)2009003-089

Morris K Johnson B Orell P Gaikhorst G Wane A and MoroD (2003) Recovery of the threatened chuditch (Dasyurus geoffroii)a case study In lsquoPredators with Pouches The Biology of CarnivorousMarsupialsrsquo (Eds M Jones C Dickman and M Archer) pp 435ndash451(CSIRO Publishing Melbourne)

Nelson J L Scroggie M P and Belcher C A (2014) Developing acamera trap survey protocol to detect a rare marsupial carnivore thespotted-tailed quoll (Dasyurus maculatus) In lsquoCamera TrappingWildlife Management and Researchrsquo (Eds P D Meek and PJ Fleming) pp 271ndash279 (CSIRO Publishing Melbourne)

Newsome T M Dellinger J A Pavey C R Ripple W J Shores C RWirsing A J and Dickman C R (2015) The ecological effects ofproviding resource subsidies to predators Global Ecology andBiogeography 24 1ndash11 doi101111geb12236

OrsquoDonnell S Webb J K and Shine R (2010) Conditioned tasteaversion enhances the survival of an endangered predator imperilled bya toxic invader Journal of Applied Ecology 47 558ndash565 doi101111j1365-2664201001802x

Oakwood M (1997) The ecology of the northern quoll Dasyurushallucatus PhD Thesis Australian National University Canberra

Oakwood M (2000) Reproduction and demography of the northern quollDasyurus hallucatus in the lowland savanna of northern AustraliaAustralian Journal of Zoology 48 519ndash539 doi101071ZO00028

Oakwood M (2002) Spatial and social organization of a carnivorousmarsupial Dasyurus hallucatus (Marsupialia Dasyuridae) Journal ofZoology 257 237ndash248 doi101017S0952836902000833

Oakwood M and Foster P (2008) Monitoring extinction of the northernquoll Australian Academy of Science Newsletter 71 6

Palomares F and Caro T M (1999) Interspecific killing amongmammalian carnivores American Naturalist 153 492ndash508 doi101086303189

Pavey C R and Bastin G (2014) lsquoAustralian Rangelands and ClimateChange ndash Invasive Animalsrsquo (Ninti One Limited amp CSIRO AliceSprings)

Pierson J C Barton P S Lane PW and Lindenmayer D B (2015) Canhabitat surrogates predict the response of target species to landscapechange Biological Conservation 184 1ndash10 doi101016jbiocon201412017

Pollock A B (1999) Notes on status distribution and diet of northernquoll Dasyurus hallucatus in the MackayndashBowen area mideastern

Research priorities for the northern quoll Australian Mammalogy M

Queensland Australian Zoologist 31 388ndash395 doi107882AZ1999040

Ramanaidou E R and Morris R C (2010) A synopsis of the channeliron deposits of the Hamersley Province Western AustraliaTransactions of the Institutions of Mining and Metallurgy Section BApplied Earth Science 119 56ndash59 doi101179037174510X12853354810624

Schmitt L H Bradley A J Kemper C M Kitchener D J HumphreysW F and How R A (1989) Ecology and physiology of the northernquoll Dasyurus hallucatus (Marsupialia Dasyuridae) at MitchellPlateau Kimberley Western Australia Journal of Zoology 217539ndash558 doi101111j1469-79981989tb02510x

Short J and Smith A (1994) Mammal decline and recovery in AustraliaJournal of Mammalogy 75 288ndash297 doi1023071382547

Spencer P How R Hillyer M Cook A Morris K Stevenson C andUmbrello L (2013) Genetic analysis of northern quolls from the PilbaraRegion ofWestern Australia Year one ndash final report Unpublished reportDepartment of Sustainability Environment Water Population andCommunities Canberra and Department of Parks and Wildlife Perth

Sutherland W J Fleishman E Mascia M B Pretty J and Rudd M A(2011) Methods for collaboratively identifying research priorities andemerging issues in science and policyMethods in Ecology and Evolution2 238ndash247 doi101111j2041-210X201000083x

Sutherst R W Floyd R B and Maywald G F (1996) The potentialgeographical distribution of the cane toad Bufo marinus L in AustraliaConservation Biology 10 294ndash299 doi101046j1523-1739199610010294x

Taylor B D and Goldingay R L (2003) Cutting the carnage wildlifeusage of road culverts in north-eastern New South Wales WildlifeResearch 30 529ndash537 doi101071WR01062

Tingley R Phillips B L Letnic M Brown G P Shine R and BairdS J (2013) Identifying optimal barriers to halt the invasion of canetoads Rhinella marina in arid Australia Journal of Applied Ecology 50129ndash137 doi1011111365-266412021

Van Vreeswyk A M E Payne A L Leighton K A and HennigP (2004) An Inventory and Condition Survey of the Pilbara RegionWestern Australia Technical Bulletin No 92 Department of AgricultureWestern Australia Perth

Webb J Legge S Tuft K Cremona T and Austin C (2015) Can wemitigate cane toad impacts on northern quolls Final report National

Environmental Research Program Northern Australia Hub CharlesDarwin University Darwin

Wintle B A Walshe T V Parris K M and McCarthy M A (2012)Designing occupancy surveys and interpreting non-detection whenobservations are imperfect Diversity amp Distributions 18 417ndash424doi101111j1472-4642201100874x

Woinarski J Menkhorst K Gambold N and Braithwaite R W (1992)Mammals of the Bungle Bungle area In lsquoA Survey of the Wildlife andVegetation of Purnululu (Bungle Bungle) National Park and AdjacentArearsquo (Ed J C Z Woinarski) Research Bulletin No 6 pp 53ndash67(CSIRO Division of Wildlife and Ecology Darwin)

Woinarski J C Z Risler J and Kean L (2004) Response of vegetationand vertebrate fauna to 23 years of fire exclusion in a tropical Eucalyptusopen forest Northern Territory Australia Austral Ecology 29 156ndash176doi101111j1442-9993200401333x

Woinarski J C Z Oakwood M Winter J Burnett S Milne D FosterP Myles H and Holmes B (2008) Surviving the toads patterns ofpersistence of the northern quoll Dasyurus hallucatus in QueenslandReport submitted to the Natural Heritage Trust Strategic ReserveProgram as a component of project 2005162 Monitoring andManagement of Cane Toad Impact in theNorthern Territory Available athttplrmntgovauplants-and-animalsinformation-and-publicationspublications2008 [accessed 14 November 2013]

Woinarski J C Z Armstrong M Brennan K Fisher A Griffiths A DHill B Milne D J Palmer C Ward S Watson M Winderlich Sand Young S (2010) Monitoring indicates rapid and severe decline ofnative small mammals in Kakadu National Park northern AustraliaWildlife Research 37 116ndash126 doi101071WR09125

Woinarski J Burbidge A and Harrison P (2014) lsquoAction Plan forAustralian Mammals 2012rsquo (CSIRO Publishing Melbourne)

Woinarski J C Burbidge A A and Harrison P L (2015) Ongoingunraveling of a continental fauna decline and extinction of Australianmammals since European settlement Proceedings of the NationalAcademy of Sciences of the United States of America 112 4531ndash4540doi101073pnas1417301112

Woolley P A Krajewski C and Westerman M (2015) Phylogeneticrelationships within Dasyurus (Dasyuromorphia Dasyuridae) quollsystematics based on molecular evidence and male characteristicsJournal of Mammalogy 96 37ndash46 doi101093jmammalgyu028

N Australian Mammalogy V A Cramer et al

wwwpublishcsiroaujournalsam

Page 9: Research priorities for the northern quoll (Dasyurus ......The Pilbara population of the northern quoll (Dasyurus hallucatus) has been seldom studied, and the impacts of threats such

Tingley et al 2013) This has created some level of uncertaintyabout the likely impact of cane toads on northern quolls in thePilbara Strategies for cane toad management were not rankedhighly by participants in the threat-management frameworkdeveloped by Carwardine et al (2014) which the authorsexplained as possibly due to uncertainty about how far canetoads will spread into the region

The model developed by Tingley et al (2013) suggested thatthe closure of 100 artificial water bodies at one of three potentiallocations along the coastal corridor between the Kimberley andthe Pilbara would prevent the invasion of cane toads into thePilbara While their model did account for uncharacteristicallywet years every three decades the dispersal of toads betweenwater bodies was determined by the number of rainy daysoccurring during the year (ie toads could only disperse betweenwater bodies for three days following rainfall) rather thanrainfall amount the associated likelihood of flooding and theavailability of free water across the landscape The cyclonic andthunderstorm nature of much of the rainfall in the north-west ofWestern Australia means that the region is prone to significantand sometimes extended flooding In April 1934 the countryaround Anna Plains (250 km south of Broome) was flooded forhundreds of miles when 27 inches (686mm) of rain fell infour days (Anon 1934) In March 2004 over 350mm of rain wasrecorded within a 48-h period at Telfer on the southern edge ofthe Great Sandy Desert due to Tropical Cyclone Fay Thiscaused the flooding of the De Grey and Oakover river systemsand flooded 140 km of the Telfer Access Road with floodwaterstaking several months to subside (wwwbomgovau) Extendingthe model of Tingley et al (2013) to account for such events willhelp clarify whether closing artificial water bodies is likely toprevent the invasion of cane toads into the Pilbara or may onlyassist in delaying their inevitable invasion

Comparison of SDMs (eg Cadenhead et al 2015) fornorthern quoll with fundamental niche models for cane toads(eg see Kearney et al 2008) within the Pilbara may identifypotential refuge areas for the northern quoll and these areasshould be prioritised for conservation actions Because theinvasion point of cane toads into the Pilbara is likely to be narrow(beginning around the mouth of the De Grey River Tingleyet al 2013) a proactive approach to training northern quollsto avoid cane toads may prove beneficial Conditioned taste-aversion (CTA) has been used successfully to train captive-reared northern quolls to avoid cane toads by offering themsmall dead toads infused with thiabendazole before theirrelease into the wild (OrsquoDonnell et al 2010)Monitoring of thesequolls after their release showed that some of their offspringhad learnt to avoid eating cane toads either through ingestionof small non-lethal toads or via social learning from theirmothers (Webb et al 2015) Fields trials of toad-aversion baits(either sausages or meatballs made from toad flesh andcontaining thiabendazole) are currently underway in the centralKimberley where cane toads were expected to invade in late2015 (Webb et al 2015) Further testing of CTA approachesunder a variety of field conditions is required The developmentof stable long-lasting baits suitable for aerial deployment isconsidered a pressing need as well as a captive colony ofnorthern quolls upon which to test the efficacy of toad-aversionbaits for inducing aversion to live toads (Webb et al 2015)

Infrastructure development and interactionswith humans

Infrastructure development may increase the likelihood ofnorthern quoll deaths due to motor vehicle strikes or beingcrushed in machinery Significant chuditch mortality wasobserved along mine access roads in the south-west of WesternAustralia (McGregor et al 2014) and the widening and sealingof an access road dramatically increased the number of roadkills and led to the extirpation of the resident population ofeastern quolls (D viverrinus) in Tasmania (Jones 2000)Infrastructure corridors such as high-tension power-lineeasements and ore-haulage railways are increasingly beinginstalled throughout the range of the northern quoll Althoughthese do not create the same mortality risk as roads they bisectand fragment habitat create large open spaces and potentiallyincrease predation risk

A recent small-scale radio-tracking study (see Dunlop et al2014a) around several quarries in the Pilbara found that northernquolls crossed roads but not railway lines The short-term natureof this work and the limited number of quolls able to be trackedmeans that the influence of roads railway lines and infrastructurecorridors on the movement and mortality of northern quollsrequires further assessment Using radio-tracking techniques todetermine crossing rates and road kill events may be difficultover large areas so population studies comparing groups ofquolls near major roads with those not affected by infrastructure(eg McCall et al 2010) may provide a more feasible meansof assessing the impacts of roads If vehicle strikes are foundto significantly contribute to the mortality of a local populationof the northern quoll then mitigation methods should betrialled on sections of road identified as road-kill hotspotsCulverts and underpasses have been shown to be used by avariety of small and medium-sized mammals and frogs (Taylorand Goldingay 2003) while installing rumble strips on roadshas led to decreased rates of road-kill of nocturnal animals(Lester 2015)

Interactions between threats

At present wildfire the subsequent loss of vegetation coverand increased predation of northern quolls is likely to be themost significant of the interactions between threats The directeffects of wildfire have been found to be less significant inthe subsequent decline of small to medium-sized mammalsparticularly for denning species such as the northern quoll thanthe indirect consequences of reduced resource availability andthe increased risk of predation due to lack of cover (Oakwood2000 Letnic et al 2005 McKenzie et al 2007 Legge et al2008 Doherty et al 2015b) Male northern quolls are perhapsmost vulnerable to such losses of cover during the breedingseason when their range can extend to over 100 ha (Oakwood2002) or where males survive after breeding their foragingrange is extended to gain the nutrition required to recover fromtheir poor condition (Cook 2010) Loss of cover due to late dry-season fires at East Alligator River in the Northern Territorymeant little cover was available for young quolls when theybegan foraging with dog and dingo predation a major source ofmortality (Webb et al 2015) The cumulative impacts of canetoad poisoning fire and predation are highly likely to lead to the

Research priorities for the northern quoll Australian Mammalogy I

extirpation of small and disjunct groups of northern quoll in thePilbara Population viability modelling of the East AlligatorRiver population suggested that high mortality from predationcombined with late dry-season burning was preventing therecovery of northern quoll populations after cane toad invasion

(Webb et al 2015) Improved knowledge of the severity of thesethreats to northern quolls in the Pilbara and their incorporationinto the SDM and PVA framework developed by Cadenheadet al (2015) will allow for better identification of priority areasfor integrated cane toad fire and predator management

Table 3 Expected timelines for the undertaking of research under each priority over the next 10 years and the funds currently allocated fromthe Pilbara Northern Quoll Offset Trust under each research priority

Rank Theme Research priority Current or proposed researchactivity

Time over next10 years

Current funding allocation(approximate)

1 Survey andmonitoring

Develop appropriate andstandardised survey andmonitoring methods

Develop monitoring protocol(traps and cameras) andinvestigate new methods

Ongoing survey and monitoringin conjunction with ResearchThemes 2 and 3 to better mapnorthern quoll distribution andto determine long-termpopulation trends

Years 1ndash2Years 3ndash10

$200 000 per annum over10 years for ResearchThemes 1 2 and 3(inclusive of salaries)

2 Habitatrequirements

Define areas of criticalhabitat for northernquoll in the Pilbara

Identify and describe criticalhabitat and undertake predictivemodelling of unsurveyed areasto inform further survey effort

Years 3ndash4 $50 000 for predictivemodelling (EdithCowan University)

Understand howdisturbance affectshabitat quality andconnectivity

Ongoing collection of data ondisturbance and habitat qualityin conjunction with ResearchThemes 1 and 4

Incorporation of new data intopredictive models

Years 4ndash10 See above

3 Populationdynamics

Better understand thepopulation dynamics ofthe northern quoll in thePilbara

Collect data on demographicparameters breeding dispersaland home-range use toincorporate into future PVA

Continued collectionof data inconjunction withpriorities 1 and 2during years 3ndash10

See above

Investigate populationstructure and theinteraction betweenpopulations

Analysis of existing tissuesamples and newly collectedsamples

Years 1ndash10 $120 000 for initial roundof genetic analysis(Years 3ndash5 MurdochUniversity)

4 Key threats Introduced predators Spatio-temporal interactionsbetween northern quolls andintroduced predators

Other projects as they arise

Years 2ndash4 (currentPhD projectUniversity ofQueensland)

$12 000 contribution tofield expenses

Cane toads Predictive modelling of canetoad distribution in the Pilbara

Years 3ndash4 Part of predictivemodelling by EdithCowan University

Infrastructuredevelopment andinteractions withhumans

Investigation of impacts oflinear infrastructure on northernquoll movement

Year 4 (currentHonours projectEdith CowanUniversity)

$15 000

Interactions betweenthreats

No on-ground projects currentlyplanned

Potential to be assessed throughPVA as data are collected underother research priorities

No funding currentlyallocated

5 Recolonisation ofrestored orartificial habitat

Determine the ability ofthe northern quoll torecolonise disturbedareas or coloniseartificial habitat

As projects arise Years 5ndash8 incollaboration withminesite restorationprograms

No funding currentlyallocated

J Australian Mammalogy V A Cramer et al

Priority 5 Recolonisation of restored andor artificial habitat

While anecdotal evidence suggests that northern quolls will denin waste rock dumps whether artificial or restored habitatprovides for long-term population viability is not known andlong-term assessment of created habitat is required Weacknowledge that a large gap exists between the desire to userestored or artificial habitat as an ecologically equivalent offsetand our limited ability to actually restore or re-create habitat thatis equivalent in structure composition and function particularlyon highly disturbed sites (Maron et al 2012) Yet there is a futureopportunity in the Pilbara to conduct lsquonaturalrsquo experiments onthe recolonisation of quolls after mining ceases including howto best design waste rock dumps so that habitat complexityand productivity is maximised by for example experimentingwith the size and positioning of boulders and their spatialarrangement in relation to surrounding landscape features Theuse of such artificial habitat should be tested in the first stagethrough direct monitoring of animals that have colonised orhave been translocated to the site Such experimentation isunderway at Atlas Ironrsquos Mt Dove operation (Atlas Iron 2012)Population modelling can then be used to predict futurepopulation sizes rates of growth and spread and the probabilityof long-term persistence in the context of various managementactions (eg predator control Armstrong and Reynolds 2012)

Resource allocation and timelines

While the allocation of resources and funding and the periodover which each of the research priorities was expected to beaddressed was not explicitly discussed during the workshopthe ranking of the priorities was taken to indicate the sequencein which the priorities should be addressed At the time of theworkshop some of the research questions identified werealready being investigated within various projects and a portionof the Pilbara Northern Quoll Offset Trust allocated to these InTable 3 we indicate the expected timelines over which someof the research programs are expected to be undertaken WhileAU$175M has been allocated to northern quoll research in thePilbara it is important to note that much of this amount is notcurrently available and will not become available until severalapproved mine developments have become operational InTable 3 we also indicate the proportion of offset funds expectedto be allocated to each research priority Research undertaken inother regions (eg research into CTA in the Kimberley regionahead of the cane toad invasion front) cannot be funded from thePilbara Northern Quoll Offset Trust but will help address someof the research questions identified here

Lessons learnt from the workshop process

The workshop approach used here was less formal than recentlarge-scale exercises used to elicit research priorities for emergingissues in science and policy at national or global levels (reviewedin Sutherland et al 2011) The focus in our (much smaller)workshop was on inclusivity rather than identifying participantsbased on purposive sampling of expertise or organisationalaffiliation We believe that the inclusive nature of the workshopwas a strength but we also recognise that this perhaps createdsome level of bias when ranking the research prioritiesAcknowledging both the experience and knowledge requirements

of those who work regularly in the Pilbara is an importantcomponent of improving the design of research programs that aimto improve conservation management in the region Yet theresearch priorities and their rankings are likely strongly reflectiveof the current information needs of most participants and maynot align with what conservation scientists believe to be themost critical areas of research for northern quoll conservation intothe future Baseline survey and monitoring are often the firstrequirements of a management or research plan for the northernquoll when offset conditions are prescribed and this was reflectedin the primary ranking given to the development of standardsurvey and monitoring methods We acknowledge the concernthat this could simply lead to offset funding being used to lsquomonitorthe extinctionrsquo of the northern quoll particularly after canetoads invade the Pilbara unless these data are used to triggermanagement actions (Oakwood and Foster 2008 Lindenmayeret al 2013) Clearly the currently limited information on habitatuse and demographic parameters for the northern quoll in thePilbara restricts the utility of more sophisticated researchapproaches such as developing robust SDM and PVA While thesevere threat that cane toads pose to the northern quoll wasrecognised this threat is not immediate in the region and mayexplain the less prominent ranking given to research to mitigatethe impacts of cane toads

The northern quoll has become emblematic of the dramaticand rapid species declines occurring in northern Australia andthe level of interest in this species was reflected in the highnumber of participants attending the workshop (one of six runby Parks and Wildlife) Given the number of attendees betterutilisation of the time available would have been gained byproviding participants with a brief before the workshopand asking them to contribute potential research questionsanonymously for interactive review by other participants Inthe exercises reviewed by Sutherland et al (2011) questionswere then discarded if they were not considered to meet criteriathat were previously agreed upon Some areas of researchthat gained some attention on the day may have been discardedif given greater consideration or interrogation before theworkshop For example there was some discussion duringthe workshop of investigating the use of habitat surrogates tomonitor the northern quoll yet research has shown that there islittle validity in using surrogates to monitor the abundance of asingle species (Pierson et al 2015)

Conclusions

Offset funds generated under the environmental approvalsprocess provide an opportunity to mobilise considerablefinancial resources for conservation (Kiesecker et al 2010) Yetthere remain significant conceptual and practical limitations tobe resolved to ensure that direct offsets provide measurableconservation outcomes that are ecologically equivalent to thelosses sustained by proceeding with the development (Maronet al 2012 Bull et al 2013) The collaborative identificationand ranking of research priorities undertaken here is the firststep in the process of developing and evaluating potentialconservation actions for the northern quoll in the Pilbara andserves as a mechanism to ensure that the research priorities ofstate and federal agencies reflects the knowledge needs of

Research priorities for the northern quoll Australian Mammalogy K

practitioners and proponents currently operating in the regionThe provision of offset funds over the next decade providesa unique opportunity to undertake research on a species ofconservation significance at a regional scale and over a longperiod through an integrated and strategic program of researchrather than the collection of ad hoc data While a morestructured approach to defining research questions before theworkshop may have led to greater critical reflection upon anddiscussion of some of the research priorities on the day theworkshop provided an opportunity for an open and diverse-ranging discussion in a collaborative environment amongstthe broader community of scientists environmental consultantsand mining proponents involved in the management andconservation of the northern quoll in the Pilbara

Acknowledgements

We gratefully acknowledge the financial support of Roy Hill who sponsoredthe workshop and partially funded VACrsquos salary We thank the Departmentof the Environmentrsquos Pilbara Taskforce especially Veronica Ritchie fortheir support of interested or affected parties The success of the workshopwas in great part due to the facilitation skills of Craig Salt of SustainableConsulting We thank all of the workshop participants for their open andcollaborative approach to the process Mike Bamford Eddie van EttenDavid Peacock and Peter Lyon provided useful comments on an earlier draftWe are grateful to John Woinarski for his constructive advice thatsubstantially improved the structure and clarity of the manuscript and ananonymous reviewer for careful and critical reading of the previousversion Lesley Gibson Russell Palmer and Al Glen generously discussedvarious aspects of the paper with VAC The map was prepared by CristinaRamalho

References

Allen B L Engeman R M and Allen L R (2011) Wild dogma anexamination of recent lsquoevidencersquo for dingo regulation of invasivemesopredator release in Australia Current Zoology 57 568ndash583

Allen B L Fleming P J Allen L R Engeman R M Ballard G andLeung L K-P (2013) As clear as mud a critical review of evidencefor the ecological roles of Australian dingoes Biological Conservation159 158ndash174 doi101016jbiocon201212004

Anon (1934) Floods in Western Australia Town isolated for week PerthMonday The Sydney Morning Herald 9

Armstrong D P and Reynolds M H (2012) Modelling reintroducedpopulations the state of the art and future directions In lsquoReintroductionBiology Integrating Science and Managementrsquo (Eds J G Ewen DP Armstrong K A Parker and P J Seddon) pp 165ndash222 (Wiley-Blackwell UK)

Atlas Iron (2012) Scope of works ndash Mount Dove artificial quoll habitatUnpublished report Atlas Iron Perth Available at httpwwwatlasironcomauIRMCompanyShowPageaspxCategoryId=190ampCPID=5424ampEID=94365410 [accessed 19 August 2015]

Austin C (2014) Can remote cameras accurately estimate populations ofthe endangered northern quoll BSc(Honours) Thesis University ofTechnology Sydney

Begg R J (1981) The small mammals of Little Nourlangie RockNT III Ecology ofDasyurus hallucatus the northern quoll (MarsupialiaDasyuridae) Wildlife Research 8 73ndash85 doi101071WR9810073

Braithwaite R W and Griffiths A D (1994) Demographic variationand range contraction in the northern quoll Dasyurus hallucatus(Marsupialia Dasyuridae)Wildlife Research 21 203ndash217 doi101071WR9940203

Bull J W Suttle K B Gordon A Singh N J and Milner-Gulland E J(2013) Biodiversity offsets in theory and practice Oryx 47 369ndash380doi101017S003060531200172X

Burnett S (1997) Colonizing cane toads cause population declines innative predators reliable anecdotal information and managementimplications Pacific Conservation Biology 3 65ndash72

Cadenhead N C R White A L and Wintle B A (2015) Identifyingconservation priorities and assessing impacts of proposed futuredevelopment in the Pilbara Bioregion in Western Australia Report bythe National Environment Research Program Decisions Hub to theDepartment of the Environment The University of MelbourneMelbourne

Carwardine J Nicol S Van Leeuwen S Walters B Firn J Reeson AMartin T G and Chades I (2014) Priority threat management forPilbara species of conservation significance CSIRO EcosystemSciences Brisbane

Cook A (2010) Habitat use and home-range of the northern quollDasyurushallucatus effects of fire MSc Thesis University ofWestern AustraliaPerth

Department of Sustainability Environment Water Population andCommunities (2012) Environment and Biodiversity Conservation Act1999 Environmental Offsets Policy Department of SustainabilityEnvironment Water Population and Communities Canberra

Doherty T S Davis R A Etten E J Algar D Collier N DickmanC R Edwards G Masters P Palmer R and Robinson S (2015a)A continental-scale analysis of feral cat diet in Australia Journal ofBiogeography 42 964ndash975 doi101111jbi12469

Doherty T S Dickman C R Nimmo D G and Ritchie E G (2015b)Multiple threats or multiplying the threats Interactions betweeninvasive predators and other ecological disturbances BiologicalConservation 190 60ndash68 doi101016jbiocon201505013

Dunlop J Cook A and Lees J (2014a) BHP Billiton Iron Ore Pilbaranorthern quoll monitoring project Final report May 2010ndashJune 2013Unpublished report prepared for BHP Billiton Iron Ore Department ofParks and Wildlife Perth

Dunlop J Cook A and Morris K (2014b) Pilbara northern quollproject surveying and monitoring Dasyurus hallucatus in the PilbaraWestern Australia Department of Parks and Wildlife Perth Availableat librarydpawwagovau

Florance D Webb J K Dempster T Kearney M R Worthing A andLetnic M (2011) Excluding access to invasion hubs can contain thespread of an invasive vertebrate Proceedings of the Royal Society ofLondon B Biological Sciences 278 2900ndash2908 doi101098rspb20110032

Gaillard J-M Hebblewhite M Loison A Fuller M Powell R BasilleM and Van Moorter B (2010) Habitatndashperformance relationshipsfinding the right metric at a given spatial scale PhilosophicalTransactions of the Royal Society of London Series B BiologicalSciences 365 2255ndash2265 doi101098rstb20100085

Glen A S and Dickman C R (2008) Niche overlap between marsupialand eutherian carnivores does competition threaten the endangeredspotted-tailed quoll Journal of Applied Ecology 45 700ndash707 doi101111j1365-2664200701449x

Glen A S Pennay M Dickman C R Wintle B A and Firestone K B(2011) Diets of sympatric native and introduced carnivores in theBarrington Tops eastern Australia Austral Ecology 36 290ndash296doi101111j1442-9993201002149x

Hennig P (2004) A brief land use history In lsquoAn Inventory and ConditionSurvey of the Pilbara Region Western Australiarsquo (Eds A M E VanVreeswyk A L Payne K A Leighton and P Hennig) TechnicalBulletin No 92 pp 13ndash18 (Western Australian Department ofAgriculture Perth)

Hohnen R Ashby J Tuft K and McGregor H (2013) Individualidentification of northern quolls (Dasyurus hallucatus) using remotecameras Australian Mammalogy 35 131ndash135 doi101071AM12015

L Australian Mammalogy V A Cramer et al

How R A Spencer P B S and Schmitt L H (2009) Island populationshave high conservation value for northern Australiarsquos top marsupialpredator ahead of a threatening process Journal of Zoology 278206ndash217 doi101111j1469-7998200900569x

Johnson C N Isaac J L and Fisher D O (2007) Rarity of a top predatortriggers continent-wide collapse of mammal prey dingoes andmarsupials in Australia Proceedings of the Royal Society of LondonSeries B Biological Sciences 274 341ndash346 doi101098rspb20063711

Jones M E (2000) Road upgrade road mortality and remedial measuresimpacts on a population of eastern quolls and Tasmanian devilsWildlifeResearch 27 289ndash296 doi101071WR98069

Kearney M Phillips B L Tracy C R Christian K A Betts G andPorter W P (2008) Modelling species distributions without usingspecies distributions the cane toad in Australia under current andfuture climates Ecography 31 423ndash434 doi101111j0906-7590200805457x

Kendrick P and Stanley F (2002) Pilbara 4 (PIL4 ndash Roebourne synopsis)In lsquoA Biodiversity Audit of Western Australiarsquos 53 BiogeographicalSubregions in 2002rsquo (Eds J E May and N L McKenzie) pp 581ndash593(Department of Conservation and Land Management Perth)

Kennedy M Phillips B L Legge S Murphy S A and Faulkner R A(2012) Do dingoes suppress the activity of feral cats in northernAustralia Austral Ecology 37 134ndash139 doi101111j1442-9993201102256x

Kiesecker J M Copeland H Pocewicz A and McKenney B (2010)Development by design blending landscape-level planning with themitigation hierarchy Frontiers in Ecology and the Environment 8261ndash266 doi101890090005

King D R and Smith L A (1985) The distribution of the European red fox(Vulpes vulpes) in Western Australia Records of the Western AustralianMuseum 12 197ndash205

Knight A T Cowling R M Rouget M Balmford A Lombard A Tand Campbell B M (2008) Knowing but not doing selecting priorityconservation areas and the researchndashimplementation gap ConservationBiology 22 610ndash617 doi101111j1523-1739200800914x

Laurance W F Koster H Grooten M Anderson A B Zuidema P AZwick S Zagt R J Lynam A J Linkie M and Anten N P (2012)Making conservation research more relevant for conservationpractitioners Biological Conservation 153 164ndash168 doi101016jbiocon201205012

Legge S Murphy S Heathcote J Flaxman E Augusteyn J andCrossman M (2008) The short-term effects of an extensive and high-intensity fire on vertebrates in the tropical savannas of the centralKimberley northern Australia Wildlife Research 35 33ndash43 doi101071WR07016

Lester D (2015) Effective wildlife roadkill mitigation Journal of Trafficand Transportation Engineering 3 42ndash51 doi10172652328-2142201501005

Letnic M Tamayo B and Dickman C R (2005) The responsesof mammals to La Nintildea (El Nintildeo Southern Oscillation)-associatedrainfall predation and wildfire in central Australia Journal ofMammalogy 86 689ndash703 doi1016441545-1542(2005)086[0689TROMTL]20CO2

Letnic M Koch F Gordon C Crowther M S and Dickman C R(2009) Keystone effects of an alien top-predator stem extinctions ofnative mammals Proceedings of the Royal Society of London Series BBiological Sciences 276 3249ndash3256 doi101098rspb20090574

Lindenmayer D B (2015) Continental-level biodiversity collapseProceedings of the National Academy of Sciences of the United States ofAmerica 112 4514ndash4515 doi101073pnas1502766112

Lindenmayer D B Piggott M P and Wintle B A (2013) Counting thebooks while the library burns why conservation monitoring programs

need a plan for action Frontiers in Ecology and the Environment 11549ndash555 doi101890120220

Maron M Hobbs R J Moilanen A Matthews J W Christie KGardner T A Keith D A Lindenmayer D B and McAlpine C A(2012) Faustian bargains Restoration realities in the context ofbiodiversity offset policies Biological Conservation 155 141ndash148doi101016jbiocon201206003

McCall S C McCarthy M A van der Ree R Harper M J Cesarini Sand Soanes K (2010) Evidence that a highway reduces apparentsurvival rates of squirrel gliders Ecology and Society 15 27

McGregor R A Stokes V L and Craig M D (2014) Does forestrestoration in fragmented landscapes provide habitat for a wide-ranging carnivore Animal Conservation 17 467ndash475 doi101111acv12112

McKenzie N L Burbidge A A Baynes A Brereton R N DickmanC R Gordon G Gibson L A Menkhorst P W Robinson A CWilliams M R and Woinarski J C Z (2007) Analysis of factorsimplicated in the recent decline of Australiarsquos mammal fauna Journal ofBiogeography 34 597ndash611 doi101111j1365-2699200601639x

McKenzie N L van Leeuwen S and Pinder AM (2009) Introduction tothe Pilbara Biodiversity Survey 2002ndash2007 Records of the WesternAustralian Museum 78(Suppl) 3ndash89 doi1018195issn0313-122x78(1)2009003-089

Morris K Johnson B Orell P Gaikhorst G Wane A and MoroD (2003) Recovery of the threatened chuditch (Dasyurus geoffroii)a case study In lsquoPredators with Pouches The Biology of CarnivorousMarsupialsrsquo (Eds M Jones C Dickman and M Archer) pp 435ndash451(CSIRO Publishing Melbourne)

Nelson J L Scroggie M P and Belcher C A (2014) Developing acamera trap survey protocol to detect a rare marsupial carnivore thespotted-tailed quoll (Dasyurus maculatus) In lsquoCamera TrappingWildlife Management and Researchrsquo (Eds P D Meek and PJ Fleming) pp 271ndash279 (CSIRO Publishing Melbourne)

Newsome T M Dellinger J A Pavey C R Ripple W J Shores C RWirsing A J and Dickman C R (2015) The ecological effects ofproviding resource subsidies to predators Global Ecology andBiogeography 24 1ndash11 doi101111geb12236

OrsquoDonnell S Webb J K and Shine R (2010) Conditioned tasteaversion enhances the survival of an endangered predator imperilled bya toxic invader Journal of Applied Ecology 47 558ndash565 doi101111j1365-2664201001802x

Oakwood M (1997) The ecology of the northern quoll Dasyurushallucatus PhD Thesis Australian National University Canberra

Oakwood M (2000) Reproduction and demography of the northern quollDasyurus hallucatus in the lowland savanna of northern AustraliaAustralian Journal of Zoology 48 519ndash539 doi101071ZO00028

Oakwood M (2002) Spatial and social organization of a carnivorousmarsupial Dasyurus hallucatus (Marsupialia Dasyuridae) Journal ofZoology 257 237ndash248 doi101017S0952836902000833

Oakwood M and Foster P (2008) Monitoring extinction of the northernquoll Australian Academy of Science Newsletter 71 6

Palomares F and Caro T M (1999) Interspecific killing amongmammalian carnivores American Naturalist 153 492ndash508 doi101086303189

Pavey C R and Bastin G (2014) lsquoAustralian Rangelands and ClimateChange ndash Invasive Animalsrsquo (Ninti One Limited amp CSIRO AliceSprings)

Pierson J C Barton P S Lane PW and Lindenmayer D B (2015) Canhabitat surrogates predict the response of target species to landscapechange Biological Conservation 184 1ndash10 doi101016jbiocon201412017

Pollock A B (1999) Notes on status distribution and diet of northernquoll Dasyurus hallucatus in the MackayndashBowen area mideastern

Research priorities for the northern quoll Australian Mammalogy M

Queensland Australian Zoologist 31 388ndash395 doi107882AZ1999040

Ramanaidou E R and Morris R C (2010) A synopsis of the channeliron deposits of the Hamersley Province Western AustraliaTransactions of the Institutions of Mining and Metallurgy Section BApplied Earth Science 119 56ndash59 doi101179037174510X12853354810624

Schmitt L H Bradley A J Kemper C M Kitchener D J HumphreysW F and How R A (1989) Ecology and physiology of the northernquoll Dasyurus hallucatus (Marsupialia Dasyuridae) at MitchellPlateau Kimberley Western Australia Journal of Zoology 217539ndash558 doi101111j1469-79981989tb02510x

Short J and Smith A (1994) Mammal decline and recovery in AustraliaJournal of Mammalogy 75 288ndash297 doi1023071382547

Spencer P How R Hillyer M Cook A Morris K Stevenson C andUmbrello L (2013) Genetic analysis of northern quolls from the PilbaraRegion ofWestern Australia Year one ndash final report Unpublished reportDepartment of Sustainability Environment Water Population andCommunities Canberra and Department of Parks and Wildlife Perth

Sutherland W J Fleishman E Mascia M B Pretty J and Rudd M A(2011) Methods for collaboratively identifying research priorities andemerging issues in science and policyMethods in Ecology and Evolution2 238ndash247 doi101111j2041-210X201000083x

Sutherst R W Floyd R B and Maywald G F (1996) The potentialgeographical distribution of the cane toad Bufo marinus L in AustraliaConservation Biology 10 294ndash299 doi101046j1523-1739199610010294x

Taylor B D and Goldingay R L (2003) Cutting the carnage wildlifeusage of road culverts in north-eastern New South Wales WildlifeResearch 30 529ndash537 doi101071WR01062

Tingley R Phillips B L Letnic M Brown G P Shine R and BairdS J (2013) Identifying optimal barriers to halt the invasion of canetoads Rhinella marina in arid Australia Journal of Applied Ecology 50129ndash137 doi1011111365-266412021

Van Vreeswyk A M E Payne A L Leighton K A and HennigP (2004) An Inventory and Condition Survey of the Pilbara RegionWestern Australia Technical Bulletin No 92 Department of AgricultureWestern Australia Perth

Webb J Legge S Tuft K Cremona T and Austin C (2015) Can wemitigate cane toad impacts on northern quolls Final report National

Environmental Research Program Northern Australia Hub CharlesDarwin University Darwin

Wintle B A Walshe T V Parris K M and McCarthy M A (2012)Designing occupancy surveys and interpreting non-detection whenobservations are imperfect Diversity amp Distributions 18 417ndash424doi101111j1472-4642201100874x

Woinarski J Menkhorst K Gambold N and Braithwaite R W (1992)Mammals of the Bungle Bungle area In lsquoA Survey of the Wildlife andVegetation of Purnululu (Bungle Bungle) National Park and AdjacentArearsquo (Ed J C Z Woinarski) Research Bulletin No 6 pp 53ndash67(CSIRO Division of Wildlife and Ecology Darwin)

Woinarski J C Z Risler J and Kean L (2004) Response of vegetationand vertebrate fauna to 23 years of fire exclusion in a tropical Eucalyptusopen forest Northern Territory Australia Austral Ecology 29 156ndash176doi101111j1442-9993200401333x

Woinarski J C Z Oakwood M Winter J Burnett S Milne D FosterP Myles H and Holmes B (2008) Surviving the toads patterns ofpersistence of the northern quoll Dasyurus hallucatus in QueenslandReport submitted to the Natural Heritage Trust Strategic ReserveProgram as a component of project 2005162 Monitoring andManagement of Cane Toad Impact in theNorthern Territory Available athttplrmntgovauplants-and-animalsinformation-and-publicationspublications2008 [accessed 14 November 2013]

Woinarski J C Z Armstrong M Brennan K Fisher A Griffiths A DHill B Milne D J Palmer C Ward S Watson M Winderlich Sand Young S (2010) Monitoring indicates rapid and severe decline ofnative small mammals in Kakadu National Park northern AustraliaWildlife Research 37 116ndash126 doi101071WR09125

Woinarski J Burbidge A and Harrison P (2014) lsquoAction Plan forAustralian Mammals 2012rsquo (CSIRO Publishing Melbourne)

Woinarski J C Burbidge A A and Harrison P L (2015) Ongoingunraveling of a continental fauna decline and extinction of Australianmammals since European settlement Proceedings of the NationalAcademy of Sciences of the United States of America 112 4531ndash4540doi101073pnas1417301112

Woolley P A Krajewski C and Westerman M (2015) Phylogeneticrelationships within Dasyurus (Dasyuromorphia Dasyuridae) quollsystematics based on molecular evidence and male characteristicsJournal of Mammalogy 96 37ndash46 doi101093jmammalgyu028

N Australian Mammalogy V A Cramer et al

wwwpublishcsiroaujournalsam

Page 10: Research priorities for the northern quoll (Dasyurus ......The Pilbara population of the northern quoll (Dasyurus hallucatus) has been seldom studied, and the impacts of threats such

extirpation of small and disjunct groups of northern quoll in thePilbara Population viability modelling of the East AlligatorRiver population suggested that high mortality from predationcombined with late dry-season burning was preventing therecovery of northern quoll populations after cane toad invasion

(Webb et al 2015) Improved knowledge of the severity of thesethreats to northern quolls in the Pilbara and their incorporationinto the SDM and PVA framework developed by Cadenheadet al (2015) will allow for better identification of priority areasfor integrated cane toad fire and predator management

Table 3 Expected timelines for the undertaking of research under each priority over the next 10 years and the funds currently allocated fromthe Pilbara Northern Quoll Offset Trust under each research priority

Rank Theme Research priority Current or proposed researchactivity

Time over next10 years

Current funding allocation(approximate)

1 Survey andmonitoring

Develop appropriate andstandardised survey andmonitoring methods

Develop monitoring protocol(traps and cameras) andinvestigate new methods

Ongoing survey and monitoringin conjunction with ResearchThemes 2 and 3 to better mapnorthern quoll distribution andto determine long-termpopulation trends

Years 1ndash2Years 3ndash10

$200 000 per annum over10 years for ResearchThemes 1 2 and 3(inclusive of salaries)

2 Habitatrequirements

Define areas of criticalhabitat for northernquoll in the Pilbara

Identify and describe criticalhabitat and undertake predictivemodelling of unsurveyed areasto inform further survey effort

Years 3ndash4 $50 000 for predictivemodelling (EdithCowan University)

Understand howdisturbance affectshabitat quality andconnectivity

Ongoing collection of data ondisturbance and habitat qualityin conjunction with ResearchThemes 1 and 4

Incorporation of new data intopredictive models

Years 4ndash10 See above

3 Populationdynamics

Better understand thepopulation dynamics ofthe northern quoll in thePilbara

Collect data on demographicparameters breeding dispersaland home-range use toincorporate into future PVA

Continued collectionof data inconjunction withpriorities 1 and 2during years 3ndash10

See above

Investigate populationstructure and theinteraction betweenpopulations

Analysis of existing tissuesamples and newly collectedsamples

Years 1ndash10 $120 000 for initial roundof genetic analysis(Years 3ndash5 MurdochUniversity)

4 Key threats Introduced predators Spatio-temporal interactionsbetween northern quolls andintroduced predators

Other projects as they arise

Years 2ndash4 (currentPhD projectUniversity ofQueensland)

$12 000 contribution tofield expenses

Cane toads Predictive modelling of canetoad distribution in the Pilbara

Years 3ndash4 Part of predictivemodelling by EdithCowan University

Infrastructuredevelopment andinteractions withhumans

Investigation of impacts oflinear infrastructure on northernquoll movement

Year 4 (currentHonours projectEdith CowanUniversity)

$15 000

Interactions betweenthreats

No on-ground projects currentlyplanned

Potential to be assessed throughPVA as data are collected underother research priorities

No funding currentlyallocated

5 Recolonisation ofrestored orartificial habitat

Determine the ability ofthe northern quoll torecolonise disturbedareas or coloniseartificial habitat

As projects arise Years 5ndash8 incollaboration withminesite restorationprograms

No funding currentlyallocated

J Australian Mammalogy V A Cramer et al

Priority 5 Recolonisation of restored andor artificial habitat

While anecdotal evidence suggests that northern quolls will denin waste rock dumps whether artificial or restored habitatprovides for long-term population viability is not known andlong-term assessment of created habitat is required Weacknowledge that a large gap exists between the desire to userestored or artificial habitat as an ecologically equivalent offsetand our limited ability to actually restore or re-create habitat thatis equivalent in structure composition and function particularlyon highly disturbed sites (Maron et al 2012) Yet there is a futureopportunity in the Pilbara to conduct lsquonaturalrsquo experiments onthe recolonisation of quolls after mining ceases including howto best design waste rock dumps so that habitat complexityand productivity is maximised by for example experimentingwith the size and positioning of boulders and their spatialarrangement in relation to surrounding landscape features Theuse of such artificial habitat should be tested in the first stagethrough direct monitoring of animals that have colonised orhave been translocated to the site Such experimentation isunderway at Atlas Ironrsquos Mt Dove operation (Atlas Iron 2012)Population modelling can then be used to predict futurepopulation sizes rates of growth and spread and the probabilityof long-term persistence in the context of various managementactions (eg predator control Armstrong and Reynolds 2012)

Resource allocation and timelines

While the allocation of resources and funding and the periodover which each of the research priorities was expected to beaddressed was not explicitly discussed during the workshopthe ranking of the priorities was taken to indicate the sequencein which the priorities should be addressed At the time of theworkshop some of the research questions identified werealready being investigated within various projects and a portionof the Pilbara Northern Quoll Offset Trust allocated to these InTable 3 we indicate the expected timelines over which someof the research programs are expected to be undertaken WhileAU$175M has been allocated to northern quoll research in thePilbara it is important to note that much of this amount is notcurrently available and will not become available until severalapproved mine developments have become operational InTable 3 we also indicate the proportion of offset funds expectedto be allocated to each research priority Research undertaken inother regions (eg research into CTA in the Kimberley regionahead of the cane toad invasion front) cannot be funded from thePilbara Northern Quoll Offset Trust but will help address someof the research questions identified here

Lessons learnt from the workshop process

The workshop approach used here was less formal than recentlarge-scale exercises used to elicit research priorities for emergingissues in science and policy at national or global levels (reviewedin Sutherland et al 2011) The focus in our (much smaller)workshop was on inclusivity rather than identifying participantsbased on purposive sampling of expertise or organisationalaffiliation We believe that the inclusive nature of the workshopwas a strength but we also recognise that this perhaps createdsome level of bias when ranking the research prioritiesAcknowledging both the experience and knowledge requirements

of those who work regularly in the Pilbara is an importantcomponent of improving the design of research programs that aimto improve conservation management in the region Yet theresearch priorities and their rankings are likely strongly reflectiveof the current information needs of most participants and maynot align with what conservation scientists believe to be themost critical areas of research for northern quoll conservation intothe future Baseline survey and monitoring are often the firstrequirements of a management or research plan for the northernquoll when offset conditions are prescribed and this was reflectedin the primary ranking given to the development of standardsurvey and monitoring methods We acknowledge the concernthat this could simply lead to offset funding being used to lsquomonitorthe extinctionrsquo of the northern quoll particularly after canetoads invade the Pilbara unless these data are used to triggermanagement actions (Oakwood and Foster 2008 Lindenmayeret al 2013) Clearly the currently limited information on habitatuse and demographic parameters for the northern quoll in thePilbara restricts the utility of more sophisticated researchapproaches such as developing robust SDM and PVA While thesevere threat that cane toads pose to the northern quoll wasrecognised this threat is not immediate in the region and mayexplain the less prominent ranking given to research to mitigatethe impacts of cane toads

The northern quoll has become emblematic of the dramaticand rapid species declines occurring in northern Australia andthe level of interest in this species was reflected in the highnumber of participants attending the workshop (one of six runby Parks and Wildlife) Given the number of attendees betterutilisation of the time available would have been gained byproviding participants with a brief before the workshopand asking them to contribute potential research questionsanonymously for interactive review by other participants Inthe exercises reviewed by Sutherland et al (2011) questionswere then discarded if they were not considered to meet criteriathat were previously agreed upon Some areas of researchthat gained some attention on the day may have been discardedif given greater consideration or interrogation before theworkshop For example there was some discussion duringthe workshop of investigating the use of habitat surrogates tomonitor the northern quoll yet research has shown that there islittle validity in using surrogates to monitor the abundance of asingle species (Pierson et al 2015)

Conclusions

Offset funds generated under the environmental approvalsprocess provide an opportunity to mobilise considerablefinancial resources for conservation (Kiesecker et al 2010) Yetthere remain significant conceptual and practical limitations tobe resolved to ensure that direct offsets provide measurableconservation outcomes that are ecologically equivalent to thelosses sustained by proceeding with the development (Maronet al 2012 Bull et al 2013) The collaborative identificationand ranking of research priorities undertaken here is the firststep in the process of developing and evaluating potentialconservation actions for the northern quoll in the Pilbara andserves as a mechanism to ensure that the research priorities ofstate and federal agencies reflects the knowledge needs of

Research priorities for the northern quoll Australian Mammalogy K

practitioners and proponents currently operating in the regionThe provision of offset funds over the next decade providesa unique opportunity to undertake research on a species ofconservation significance at a regional scale and over a longperiod through an integrated and strategic program of researchrather than the collection of ad hoc data While a morestructured approach to defining research questions before theworkshop may have led to greater critical reflection upon anddiscussion of some of the research priorities on the day theworkshop provided an opportunity for an open and diverse-ranging discussion in a collaborative environment amongstthe broader community of scientists environmental consultantsand mining proponents involved in the management andconservation of the northern quoll in the Pilbara

Acknowledgements

We gratefully acknowledge the financial support of Roy Hill who sponsoredthe workshop and partially funded VACrsquos salary We thank the Departmentof the Environmentrsquos Pilbara Taskforce especially Veronica Ritchie fortheir support of interested or affected parties The success of the workshopwas in great part due to the facilitation skills of Craig Salt of SustainableConsulting We thank all of the workshop participants for their open andcollaborative approach to the process Mike Bamford Eddie van EttenDavid Peacock and Peter Lyon provided useful comments on an earlier draftWe are grateful to John Woinarski for his constructive advice thatsubstantially improved the structure and clarity of the manuscript and ananonymous reviewer for careful and critical reading of the previousversion Lesley Gibson Russell Palmer and Al Glen generously discussedvarious aspects of the paper with VAC The map was prepared by CristinaRamalho

References

Allen B L Engeman R M and Allen L R (2011) Wild dogma anexamination of recent lsquoevidencersquo for dingo regulation of invasivemesopredator release in Australia Current Zoology 57 568ndash583

Allen B L Fleming P J Allen L R Engeman R M Ballard G andLeung L K-P (2013) As clear as mud a critical review of evidencefor the ecological roles of Australian dingoes Biological Conservation159 158ndash174 doi101016jbiocon201212004

Anon (1934) Floods in Western Australia Town isolated for week PerthMonday The Sydney Morning Herald 9

Armstrong D P and Reynolds M H (2012) Modelling reintroducedpopulations the state of the art and future directions In lsquoReintroductionBiology Integrating Science and Managementrsquo (Eds J G Ewen DP Armstrong K A Parker and P J Seddon) pp 165ndash222 (Wiley-Blackwell UK)

Atlas Iron (2012) Scope of works ndash Mount Dove artificial quoll habitatUnpublished report Atlas Iron Perth Available at httpwwwatlasironcomauIRMCompanyShowPageaspxCategoryId=190ampCPID=5424ampEID=94365410 [accessed 19 August 2015]

Austin C (2014) Can remote cameras accurately estimate populations ofthe endangered northern quoll BSc(Honours) Thesis University ofTechnology Sydney

Begg R J (1981) The small mammals of Little Nourlangie RockNT III Ecology ofDasyurus hallucatus the northern quoll (MarsupialiaDasyuridae) Wildlife Research 8 73ndash85 doi101071WR9810073

Braithwaite R W and Griffiths A D (1994) Demographic variationand range contraction in the northern quoll Dasyurus hallucatus(Marsupialia Dasyuridae)Wildlife Research 21 203ndash217 doi101071WR9940203

Bull J W Suttle K B Gordon A Singh N J and Milner-Gulland E J(2013) Biodiversity offsets in theory and practice Oryx 47 369ndash380doi101017S003060531200172X

Burnett S (1997) Colonizing cane toads cause population declines innative predators reliable anecdotal information and managementimplications Pacific Conservation Biology 3 65ndash72

Cadenhead N C R White A L and Wintle B A (2015) Identifyingconservation priorities and assessing impacts of proposed futuredevelopment in the Pilbara Bioregion in Western Australia Report bythe National Environment Research Program Decisions Hub to theDepartment of the Environment The University of MelbourneMelbourne

Carwardine J Nicol S Van Leeuwen S Walters B Firn J Reeson AMartin T G and Chades I (2014) Priority threat management forPilbara species of conservation significance CSIRO EcosystemSciences Brisbane

Cook A (2010) Habitat use and home-range of the northern quollDasyurushallucatus effects of fire MSc Thesis University ofWestern AustraliaPerth

Department of Sustainability Environment Water Population andCommunities (2012) Environment and Biodiversity Conservation Act1999 Environmental Offsets Policy Department of SustainabilityEnvironment Water Population and Communities Canberra

Doherty T S Davis R A Etten E J Algar D Collier N DickmanC R Edwards G Masters P Palmer R and Robinson S (2015a)A continental-scale analysis of feral cat diet in Australia Journal ofBiogeography 42 964ndash975 doi101111jbi12469

Doherty T S Dickman C R Nimmo D G and Ritchie E G (2015b)Multiple threats or multiplying the threats Interactions betweeninvasive predators and other ecological disturbances BiologicalConservation 190 60ndash68 doi101016jbiocon201505013

Dunlop J Cook A and Lees J (2014a) BHP Billiton Iron Ore Pilbaranorthern quoll monitoring project Final report May 2010ndashJune 2013Unpublished report prepared for BHP Billiton Iron Ore Department ofParks and Wildlife Perth

Dunlop J Cook A and Morris K (2014b) Pilbara northern quollproject surveying and monitoring Dasyurus hallucatus in the PilbaraWestern Australia Department of Parks and Wildlife Perth Availableat librarydpawwagovau

Florance D Webb J K Dempster T Kearney M R Worthing A andLetnic M (2011) Excluding access to invasion hubs can contain thespread of an invasive vertebrate Proceedings of the Royal Society ofLondon B Biological Sciences 278 2900ndash2908 doi101098rspb20110032

Gaillard J-M Hebblewhite M Loison A Fuller M Powell R BasilleM and Van Moorter B (2010) Habitatndashperformance relationshipsfinding the right metric at a given spatial scale PhilosophicalTransactions of the Royal Society of London Series B BiologicalSciences 365 2255ndash2265 doi101098rstb20100085

Glen A S and Dickman C R (2008) Niche overlap between marsupialand eutherian carnivores does competition threaten the endangeredspotted-tailed quoll Journal of Applied Ecology 45 700ndash707 doi101111j1365-2664200701449x

Glen A S Pennay M Dickman C R Wintle B A and Firestone K B(2011) Diets of sympatric native and introduced carnivores in theBarrington Tops eastern Australia Austral Ecology 36 290ndash296doi101111j1442-9993201002149x

Hennig P (2004) A brief land use history In lsquoAn Inventory and ConditionSurvey of the Pilbara Region Western Australiarsquo (Eds A M E VanVreeswyk A L Payne K A Leighton and P Hennig) TechnicalBulletin No 92 pp 13ndash18 (Western Australian Department ofAgriculture Perth)

Hohnen R Ashby J Tuft K and McGregor H (2013) Individualidentification of northern quolls (Dasyurus hallucatus) using remotecameras Australian Mammalogy 35 131ndash135 doi101071AM12015

L Australian Mammalogy V A Cramer et al

How R A Spencer P B S and Schmitt L H (2009) Island populationshave high conservation value for northern Australiarsquos top marsupialpredator ahead of a threatening process Journal of Zoology 278206ndash217 doi101111j1469-7998200900569x

Johnson C N Isaac J L and Fisher D O (2007) Rarity of a top predatortriggers continent-wide collapse of mammal prey dingoes andmarsupials in Australia Proceedings of the Royal Society of LondonSeries B Biological Sciences 274 341ndash346 doi101098rspb20063711

Jones M E (2000) Road upgrade road mortality and remedial measuresimpacts on a population of eastern quolls and Tasmanian devilsWildlifeResearch 27 289ndash296 doi101071WR98069

Kearney M Phillips B L Tracy C R Christian K A Betts G andPorter W P (2008) Modelling species distributions without usingspecies distributions the cane toad in Australia under current andfuture climates Ecography 31 423ndash434 doi101111j0906-7590200805457x

Kendrick P and Stanley F (2002) Pilbara 4 (PIL4 ndash Roebourne synopsis)In lsquoA Biodiversity Audit of Western Australiarsquos 53 BiogeographicalSubregions in 2002rsquo (Eds J E May and N L McKenzie) pp 581ndash593(Department of Conservation and Land Management Perth)

Kennedy M Phillips B L Legge S Murphy S A and Faulkner R A(2012) Do dingoes suppress the activity of feral cats in northernAustralia Austral Ecology 37 134ndash139 doi101111j1442-9993201102256x

Kiesecker J M Copeland H Pocewicz A and McKenney B (2010)Development by design blending landscape-level planning with themitigation hierarchy Frontiers in Ecology and the Environment 8261ndash266 doi101890090005

King D R and Smith L A (1985) The distribution of the European red fox(Vulpes vulpes) in Western Australia Records of the Western AustralianMuseum 12 197ndash205

Knight A T Cowling R M Rouget M Balmford A Lombard A Tand Campbell B M (2008) Knowing but not doing selecting priorityconservation areas and the researchndashimplementation gap ConservationBiology 22 610ndash617 doi101111j1523-1739200800914x

Laurance W F Koster H Grooten M Anderson A B Zuidema P AZwick S Zagt R J Lynam A J Linkie M and Anten N P (2012)Making conservation research more relevant for conservationpractitioners Biological Conservation 153 164ndash168 doi101016jbiocon201205012

Legge S Murphy S Heathcote J Flaxman E Augusteyn J andCrossman M (2008) The short-term effects of an extensive and high-intensity fire on vertebrates in the tropical savannas of the centralKimberley northern Australia Wildlife Research 35 33ndash43 doi101071WR07016

Lester D (2015) Effective wildlife roadkill mitigation Journal of Trafficand Transportation Engineering 3 42ndash51 doi10172652328-2142201501005

Letnic M Tamayo B and Dickman C R (2005) The responsesof mammals to La Nintildea (El Nintildeo Southern Oscillation)-associatedrainfall predation and wildfire in central Australia Journal ofMammalogy 86 689ndash703 doi1016441545-1542(2005)086[0689TROMTL]20CO2

Letnic M Koch F Gordon C Crowther M S and Dickman C R(2009) Keystone effects of an alien top-predator stem extinctions ofnative mammals Proceedings of the Royal Society of London Series BBiological Sciences 276 3249ndash3256 doi101098rspb20090574

Lindenmayer D B (2015) Continental-level biodiversity collapseProceedings of the National Academy of Sciences of the United States ofAmerica 112 4514ndash4515 doi101073pnas1502766112

Lindenmayer D B Piggott M P and Wintle B A (2013) Counting thebooks while the library burns why conservation monitoring programs

need a plan for action Frontiers in Ecology and the Environment 11549ndash555 doi101890120220

Maron M Hobbs R J Moilanen A Matthews J W Christie KGardner T A Keith D A Lindenmayer D B and McAlpine C A(2012) Faustian bargains Restoration realities in the context ofbiodiversity offset policies Biological Conservation 155 141ndash148doi101016jbiocon201206003

McCall S C McCarthy M A van der Ree R Harper M J Cesarini Sand Soanes K (2010) Evidence that a highway reduces apparentsurvival rates of squirrel gliders Ecology and Society 15 27

McGregor R A Stokes V L and Craig M D (2014) Does forestrestoration in fragmented landscapes provide habitat for a wide-ranging carnivore Animal Conservation 17 467ndash475 doi101111acv12112

McKenzie N L Burbidge A A Baynes A Brereton R N DickmanC R Gordon G Gibson L A Menkhorst P W Robinson A CWilliams M R and Woinarski J C Z (2007) Analysis of factorsimplicated in the recent decline of Australiarsquos mammal fauna Journal ofBiogeography 34 597ndash611 doi101111j1365-2699200601639x

McKenzie N L van Leeuwen S and Pinder AM (2009) Introduction tothe Pilbara Biodiversity Survey 2002ndash2007 Records of the WesternAustralian Museum 78(Suppl) 3ndash89 doi1018195issn0313-122x78(1)2009003-089

Morris K Johnson B Orell P Gaikhorst G Wane A and MoroD (2003) Recovery of the threatened chuditch (Dasyurus geoffroii)a case study In lsquoPredators with Pouches The Biology of CarnivorousMarsupialsrsquo (Eds M Jones C Dickman and M Archer) pp 435ndash451(CSIRO Publishing Melbourne)

Nelson J L Scroggie M P and Belcher C A (2014) Developing acamera trap survey protocol to detect a rare marsupial carnivore thespotted-tailed quoll (Dasyurus maculatus) In lsquoCamera TrappingWildlife Management and Researchrsquo (Eds P D Meek and PJ Fleming) pp 271ndash279 (CSIRO Publishing Melbourne)

Newsome T M Dellinger J A Pavey C R Ripple W J Shores C RWirsing A J and Dickman C R (2015) The ecological effects ofproviding resource subsidies to predators Global Ecology andBiogeography 24 1ndash11 doi101111geb12236

OrsquoDonnell S Webb J K and Shine R (2010) Conditioned tasteaversion enhances the survival of an endangered predator imperilled bya toxic invader Journal of Applied Ecology 47 558ndash565 doi101111j1365-2664201001802x

Oakwood M (1997) The ecology of the northern quoll Dasyurushallucatus PhD Thesis Australian National University Canberra

Oakwood M (2000) Reproduction and demography of the northern quollDasyurus hallucatus in the lowland savanna of northern AustraliaAustralian Journal of Zoology 48 519ndash539 doi101071ZO00028

Oakwood M (2002) Spatial and social organization of a carnivorousmarsupial Dasyurus hallucatus (Marsupialia Dasyuridae) Journal ofZoology 257 237ndash248 doi101017S0952836902000833

Oakwood M and Foster P (2008) Monitoring extinction of the northernquoll Australian Academy of Science Newsletter 71 6

Palomares F and Caro T M (1999) Interspecific killing amongmammalian carnivores American Naturalist 153 492ndash508 doi101086303189

Pavey C R and Bastin G (2014) lsquoAustralian Rangelands and ClimateChange ndash Invasive Animalsrsquo (Ninti One Limited amp CSIRO AliceSprings)

Pierson J C Barton P S Lane PW and Lindenmayer D B (2015) Canhabitat surrogates predict the response of target species to landscapechange Biological Conservation 184 1ndash10 doi101016jbiocon201412017

Pollock A B (1999) Notes on status distribution and diet of northernquoll Dasyurus hallucatus in the MackayndashBowen area mideastern

Research priorities for the northern quoll Australian Mammalogy M

Queensland Australian Zoologist 31 388ndash395 doi107882AZ1999040

Ramanaidou E R and Morris R C (2010) A synopsis of the channeliron deposits of the Hamersley Province Western AustraliaTransactions of the Institutions of Mining and Metallurgy Section BApplied Earth Science 119 56ndash59 doi101179037174510X12853354810624

Schmitt L H Bradley A J Kemper C M Kitchener D J HumphreysW F and How R A (1989) Ecology and physiology of the northernquoll Dasyurus hallucatus (Marsupialia Dasyuridae) at MitchellPlateau Kimberley Western Australia Journal of Zoology 217539ndash558 doi101111j1469-79981989tb02510x

Short J and Smith A (1994) Mammal decline and recovery in AustraliaJournal of Mammalogy 75 288ndash297 doi1023071382547

Spencer P How R Hillyer M Cook A Morris K Stevenson C andUmbrello L (2013) Genetic analysis of northern quolls from the PilbaraRegion ofWestern Australia Year one ndash final report Unpublished reportDepartment of Sustainability Environment Water Population andCommunities Canberra and Department of Parks and Wildlife Perth

Sutherland W J Fleishman E Mascia M B Pretty J and Rudd M A(2011) Methods for collaboratively identifying research priorities andemerging issues in science and policyMethods in Ecology and Evolution2 238ndash247 doi101111j2041-210X201000083x

Sutherst R W Floyd R B and Maywald G F (1996) The potentialgeographical distribution of the cane toad Bufo marinus L in AustraliaConservation Biology 10 294ndash299 doi101046j1523-1739199610010294x

Taylor B D and Goldingay R L (2003) Cutting the carnage wildlifeusage of road culverts in north-eastern New South Wales WildlifeResearch 30 529ndash537 doi101071WR01062

Tingley R Phillips B L Letnic M Brown G P Shine R and BairdS J (2013) Identifying optimal barriers to halt the invasion of canetoads Rhinella marina in arid Australia Journal of Applied Ecology 50129ndash137 doi1011111365-266412021

Van Vreeswyk A M E Payne A L Leighton K A and HennigP (2004) An Inventory and Condition Survey of the Pilbara RegionWestern Australia Technical Bulletin No 92 Department of AgricultureWestern Australia Perth

Webb J Legge S Tuft K Cremona T and Austin C (2015) Can wemitigate cane toad impacts on northern quolls Final report National

Environmental Research Program Northern Australia Hub CharlesDarwin University Darwin

Wintle B A Walshe T V Parris K M and McCarthy M A (2012)Designing occupancy surveys and interpreting non-detection whenobservations are imperfect Diversity amp Distributions 18 417ndash424doi101111j1472-4642201100874x

Woinarski J Menkhorst K Gambold N and Braithwaite R W (1992)Mammals of the Bungle Bungle area In lsquoA Survey of the Wildlife andVegetation of Purnululu (Bungle Bungle) National Park and AdjacentArearsquo (Ed J C Z Woinarski) Research Bulletin No 6 pp 53ndash67(CSIRO Division of Wildlife and Ecology Darwin)

Woinarski J C Z Risler J and Kean L (2004) Response of vegetationand vertebrate fauna to 23 years of fire exclusion in a tropical Eucalyptusopen forest Northern Territory Australia Austral Ecology 29 156ndash176doi101111j1442-9993200401333x

Woinarski J C Z Oakwood M Winter J Burnett S Milne D FosterP Myles H and Holmes B (2008) Surviving the toads patterns ofpersistence of the northern quoll Dasyurus hallucatus in QueenslandReport submitted to the Natural Heritage Trust Strategic ReserveProgram as a component of project 2005162 Monitoring andManagement of Cane Toad Impact in theNorthern Territory Available athttplrmntgovauplants-and-animalsinformation-and-publicationspublications2008 [accessed 14 November 2013]

Woinarski J C Z Armstrong M Brennan K Fisher A Griffiths A DHill B Milne D J Palmer C Ward S Watson M Winderlich Sand Young S (2010) Monitoring indicates rapid and severe decline ofnative small mammals in Kakadu National Park northern AustraliaWildlife Research 37 116ndash126 doi101071WR09125

Woinarski J Burbidge A and Harrison P (2014) lsquoAction Plan forAustralian Mammals 2012rsquo (CSIRO Publishing Melbourne)

Woinarski J C Burbidge A A and Harrison P L (2015) Ongoingunraveling of a continental fauna decline and extinction of Australianmammals since European settlement Proceedings of the NationalAcademy of Sciences of the United States of America 112 4531ndash4540doi101073pnas1417301112

Woolley P A Krajewski C and Westerman M (2015) Phylogeneticrelationships within Dasyurus (Dasyuromorphia Dasyuridae) quollsystematics based on molecular evidence and male characteristicsJournal of Mammalogy 96 37ndash46 doi101093jmammalgyu028

N Australian Mammalogy V A Cramer et al

wwwpublishcsiroaujournalsam

Page 11: Research priorities for the northern quoll (Dasyurus ......The Pilbara population of the northern quoll (Dasyurus hallucatus) has been seldom studied, and the impacts of threats such

Priority 5 Recolonisation of restored andor artificial habitat

While anecdotal evidence suggests that northern quolls will denin waste rock dumps whether artificial or restored habitatprovides for long-term population viability is not known andlong-term assessment of created habitat is required Weacknowledge that a large gap exists between the desire to userestored or artificial habitat as an ecologically equivalent offsetand our limited ability to actually restore or re-create habitat thatis equivalent in structure composition and function particularlyon highly disturbed sites (Maron et al 2012) Yet there is a futureopportunity in the Pilbara to conduct lsquonaturalrsquo experiments onthe recolonisation of quolls after mining ceases including howto best design waste rock dumps so that habitat complexityand productivity is maximised by for example experimentingwith the size and positioning of boulders and their spatialarrangement in relation to surrounding landscape features Theuse of such artificial habitat should be tested in the first stagethrough direct monitoring of animals that have colonised orhave been translocated to the site Such experimentation isunderway at Atlas Ironrsquos Mt Dove operation (Atlas Iron 2012)Population modelling can then be used to predict futurepopulation sizes rates of growth and spread and the probabilityof long-term persistence in the context of various managementactions (eg predator control Armstrong and Reynolds 2012)

Resource allocation and timelines

While the allocation of resources and funding and the periodover which each of the research priorities was expected to beaddressed was not explicitly discussed during the workshopthe ranking of the priorities was taken to indicate the sequencein which the priorities should be addressed At the time of theworkshop some of the research questions identified werealready being investigated within various projects and a portionof the Pilbara Northern Quoll Offset Trust allocated to these InTable 3 we indicate the expected timelines over which someof the research programs are expected to be undertaken WhileAU$175M has been allocated to northern quoll research in thePilbara it is important to note that much of this amount is notcurrently available and will not become available until severalapproved mine developments have become operational InTable 3 we also indicate the proportion of offset funds expectedto be allocated to each research priority Research undertaken inother regions (eg research into CTA in the Kimberley regionahead of the cane toad invasion front) cannot be funded from thePilbara Northern Quoll Offset Trust but will help address someof the research questions identified here

Lessons learnt from the workshop process

The workshop approach used here was less formal than recentlarge-scale exercises used to elicit research priorities for emergingissues in science and policy at national or global levels (reviewedin Sutherland et al 2011) The focus in our (much smaller)workshop was on inclusivity rather than identifying participantsbased on purposive sampling of expertise or organisationalaffiliation We believe that the inclusive nature of the workshopwas a strength but we also recognise that this perhaps createdsome level of bias when ranking the research prioritiesAcknowledging both the experience and knowledge requirements

of those who work regularly in the Pilbara is an importantcomponent of improving the design of research programs that aimto improve conservation management in the region Yet theresearch priorities and their rankings are likely strongly reflectiveof the current information needs of most participants and maynot align with what conservation scientists believe to be themost critical areas of research for northern quoll conservation intothe future Baseline survey and monitoring are often the firstrequirements of a management or research plan for the northernquoll when offset conditions are prescribed and this was reflectedin the primary ranking given to the development of standardsurvey and monitoring methods We acknowledge the concernthat this could simply lead to offset funding being used to lsquomonitorthe extinctionrsquo of the northern quoll particularly after canetoads invade the Pilbara unless these data are used to triggermanagement actions (Oakwood and Foster 2008 Lindenmayeret al 2013) Clearly the currently limited information on habitatuse and demographic parameters for the northern quoll in thePilbara restricts the utility of more sophisticated researchapproaches such as developing robust SDM and PVA While thesevere threat that cane toads pose to the northern quoll wasrecognised this threat is not immediate in the region and mayexplain the less prominent ranking given to research to mitigatethe impacts of cane toads

The northern quoll has become emblematic of the dramaticand rapid species declines occurring in northern Australia andthe level of interest in this species was reflected in the highnumber of participants attending the workshop (one of six runby Parks and Wildlife) Given the number of attendees betterutilisation of the time available would have been gained byproviding participants with a brief before the workshopand asking them to contribute potential research questionsanonymously for interactive review by other participants Inthe exercises reviewed by Sutherland et al (2011) questionswere then discarded if they were not considered to meet criteriathat were previously agreed upon Some areas of researchthat gained some attention on the day may have been discardedif given greater consideration or interrogation before theworkshop For example there was some discussion duringthe workshop of investigating the use of habitat surrogates tomonitor the northern quoll yet research has shown that there islittle validity in using surrogates to monitor the abundance of asingle species (Pierson et al 2015)

Conclusions

Offset funds generated under the environmental approvalsprocess provide an opportunity to mobilise considerablefinancial resources for conservation (Kiesecker et al 2010) Yetthere remain significant conceptual and practical limitations tobe resolved to ensure that direct offsets provide measurableconservation outcomes that are ecologically equivalent to thelosses sustained by proceeding with the development (Maronet al 2012 Bull et al 2013) The collaborative identificationand ranking of research priorities undertaken here is the firststep in the process of developing and evaluating potentialconservation actions for the northern quoll in the Pilbara andserves as a mechanism to ensure that the research priorities ofstate and federal agencies reflects the knowledge needs of

Research priorities for the northern quoll Australian Mammalogy K

practitioners and proponents currently operating in the regionThe provision of offset funds over the next decade providesa unique opportunity to undertake research on a species ofconservation significance at a regional scale and over a longperiod through an integrated and strategic program of researchrather than the collection of ad hoc data While a morestructured approach to defining research questions before theworkshop may have led to greater critical reflection upon anddiscussion of some of the research priorities on the day theworkshop provided an opportunity for an open and diverse-ranging discussion in a collaborative environment amongstthe broader community of scientists environmental consultantsand mining proponents involved in the management andconservation of the northern quoll in the Pilbara

Acknowledgements

We gratefully acknowledge the financial support of Roy Hill who sponsoredthe workshop and partially funded VACrsquos salary We thank the Departmentof the Environmentrsquos Pilbara Taskforce especially Veronica Ritchie fortheir support of interested or affected parties The success of the workshopwas in great part due to the facilitation skills of Craig Salt of SustainableConsulting We thank all of the workshop participants for their open andcollaborative approach to the process Mike Bamford Eddie van EttenDavid Peacock and Peter Lyon provided useful comments on an earlier draftWe are grateful to John Woinarski for his constructive advice thatsubstantially improved the structure and clarity of the manuscript and ananonymous reviewer for careful and critical reading of the previousversion Lesley Gibson Russell Palmer and Al Glen generously discussedvarious aspects of the paper with VAC The map was prepared by CristinaRamalho

References

Allen B L Engeman R M and Allen L R (2011) Wild dogma anexamination of recent lsquoevidencersquo for dingo regulation of invasivemesopredator release in Australia Current Zoology 57 568ndash583

Allen B L Fleming P J Allen L R Engeman R M Ballard G andLeung L K-P (2013) As clear as mud a critical review of evidencefor the ecological roles of Australian dingoes Biological Conservation159 158ndash174 doi101016jbiocon201212004

Anon (1934) Floods in Western Australia Town isolated for week PerthMonday The Sydney Morning Herald 9

Armstrong D P and Reynolds M H (2012) Modelling reintroducedpopulations the state of the art and future directions In lsquoReintroductionBiology Integrating Science and Managementrsquo (Eds J G Ewen DP Armstrong K A Parker and P J Seddon) pp 165ndash222 (Wiley-Blackwell UK)

Atlas Iron (2012) Scope of works ndash Mount Dove artificial quoll habitatUnpublished report Atlas Iron Perth Available at httpwwwatlasironcomauIRMCompanyShowPageaspxCategoryId=190ampCPID=5424ampEID=94365410 [accessed 19 August 2015]

Austin C (2014) Can remote cameras accurately estimate populations ofthe endangered northern quoll BSc(Honours) Thesis University ofTechnology Sydney

Begg R J (1981) The small mammals of Little Nourlangie RockNT III Ecology ofDasyurus hallucatus the northern quoll (MarsupialiaDasyuridae) Wildlife Research 8 73ndash85 doi101071WR9810073

Braithwaite R W and Griffiths A D (1994) Demographic variationand range contraction in the northern quoll Dasyurus hallucatus(Marsupialia Dasyuridae)Wildlife Research 21 203ndash217 doi101071WR9940203

Bull J W Suttle K B Gordon A Singh N J and Milner-Gulland E J(2013) Biodiversity offsets in theory and practice Oryx 47 369ndash380doi101017S003060531200172X

Burnett S (1997) Colonizing cane toads cause population declines innative predators reliable anecdotal information and managementimplications Pacific Conservation Biology 3 65ndash72

Cadenhead N C R White A L and Wintle B A (2015) Identifyingconservation priorities and assessing impacts of proposed futuredevelopment in the Pilbara Bioregion in Western Australia Report bythe National Environment Research Program Decisions Hub to theDepartment of the Environment The University of MelbourneMelbourne

Carwardine J Nicol S Van Leeuwen S Walters B Firn J Reeson AMartin T G and Chades I (2014) Priority threat management forPilbara species of conservation significance CSIRO EcosystemSciences Brisbane

Cook A (2010) Habitat use and home-range of the northern quollDasyurushallucatus effects of fire MSc Thesis University ofWestern AustraliaPerth

Department of Sustainability Environment Water Population andCommunities (2012) Environment and Biodiversity Conservation Act1999 Environmental Offsets Policy Department of SustainabilityEnvironment Water Population and Communities Canberra

Doherty T S Davis R A Etten E J Algar D Collier N DickmanC R Edwards G Masters P Palmer R and Robinson S (2015a)A continental-scale analysis of feral cat diet in Australia Journal ofBiogeography 42 964ndash975 doi101111jbi12469

Doherty T S Dickman C R Nimmo D G and Ritchie E G (2015b)Multiple threats or multiplying the threats Interactions betweeninvasive predators and other ecological disturbances BiologicalConservation 190 60ndash68 doi101016jbiocon201505013

Dunlop J Cook A and Lees J (2014a) BHP Billiton Iron Ore Pilbaranorthern quoll monitoring project Final report May 2010ndashJune 2013Unpublished report prepared for BHP Billiton Iron Ore Department ofParks and Wildlife Perth

Dunlop J Cook A and Morris K (2014b) Pilbara northern quollproject surveying and monitoring Dasyurus hallucatus in the PilbaraWestern Australia Department of Parks and Wildlife Perth Availableat librarydpawwagovau

Florance D Webb J K Dempster T Kearney M R Worthing A andLetnic M (2011) Excluding access to invasion hubs can contain thespread of an invasive vertebrate Proceedings of the Royal Society ofLondon B Biological Sciences 278 2900ndash2908 doi101098rspb20110032

Gaillard J-M Hebblewhite M Loison A Fuller M Powell R BasilleM and Van Moorter B (2010) Habitatndashperformance relationshipsfinding the right metric at a given spatial scale PhilosophicalTransactions of the Royal Society of London Series B BiologicalSciences 365 2255ndash2265 doi101098rstb20100085

Glen A S and Dickman C R (2008) Niche overlap between marsupialand eutherian carnivores does competition threaten the endangeredspotted-tailed quoll Journal of Applied Ecology 45 700ndash707 doi101111j1365-2664200701449x

Glen A S Pennay M Dickman C R Wintle B A and Firestone K B(2011) Diets of sympatric native and introduced carnivores in theBarrington Tops eastern Australia Austral Ecology 36 290ndash296doi101111j1442-9993201002149x

Hennig P (2004) A brief land use history In lsquoAn Inventory and ConditionSurvey of the Pilbara Region Western Australiarsquo (Eds A M E VanVreeswyk A L Payne K A Leighton and P Hennig) TechnicalBulletin No 92 pp 13ndash18 (Western Australian Department ofAgriculture Perth)

Hohnen R Ashby J Tuft K and McGregor H (2013) Individualidentification of northern quolls (Dasyurus hallucatus) using remotecameras Australian Mammalogy 35 131ndash135 doi101071AM12015

L Australian Mammalogy V A Cramer et al

How R A Spencer P B S and Schmitt L H (2009) Island populationshave high conservation value for northern Australiarsquos top marsupialpredator ahead of a threatening process Journal of Zoology 278206ndash217 doi101111j1469-7998200900569x

Johnson C N Isaac J L and Fisher D O (2007) Rarity of a top predatortriggers continent-wide collapse of mammal prey dingoes andmarsupials in Australia Proceedings of the Royal Society of LondonSeries B Biological Sciences 274 341ndash346 doi101098rspb20063711

Jones M E (2000) Road upgrade road mortality and remedial measuresimpacts on a population of eastern quolls and Tasmanian devilsWildlifeResearch 27 289ndash296 doi101071WR98069

Kearney M Phillips B L Tracy C R Christian K A Betts G andPorter W P (2008) Modelling species distributions without usingspecies distributions the cane toad in Australia under current andfuture climates Ecography 31 423ndash434 doi101111j0906-7590200805457x

Kendrick P and Stanley F (2002) Pilbara 4 (PIL4 ndash Roebourne synopsis)In lsquoA Biodiversity Audit of Western Australiarsquos 53 BiogeographicalSubregions in 2002rsquo (Eds J E May and N L McKenzie) pp 581ndash593(Department of Conservation and Land Management Perth)

Kennedy M Phillips B L Legge S Murphy S A and Faulkner R A(2012) Do dingoes suppress the activity of feral cats in northernAustralia Austral Ecology 37 134ndash139 doi101111j1442-9993201102256x

Kiesecker J M Copeland H Pocewicz A and McKenney B (2010)Development by design blending landscape-level planning with themitigation hierarchy Frontiers in Ecology and the Environment 8261ndash266 doi101890090005

King D R and Smith L A (1985) The distribution of the European red fox(Vulpes vulpes) in Western Australia Records of the Western AustralianMuseum 12 197ndash205

Knight A T Cowling R M Rouget M Balmford A Lombard A Tand Campbell B M (2008) Knowing but not doing selecting priorityconservation areas and the researchndashimplementation gap ConservationBiology 22 610ndash617 doi101111j1523-1739200800914x

Laurance W F Koster H Grooten M Anderson A B Zuidema P AZwick S Zagt R J Lynam A J Linkie M and Anten N P (2012)Making conservation research more relevant for conservationpractitioners Biological Conservation 153 164ndash168 doi101016jbiocon201205012

Legge S Murphy S Heathcote J Flaxman E Augusteyn J andCrossman M (2008) The short-term effects of an extensive and high-intensity fire on vertebrates in the tropical savannas of the centralKimberley northern Australia Wildlife Research 35 33ndash43 doi101071WR07016

Lester D (2015) Effective wildlife roadkill mitigation Journal of Trafficand Transportation Engineering 3 42ndash51 doi10172652328-2142201501005

Letnic M Tamayo B and Dickman C R (2005) The responsesof mammals to La Nintildea (El Nintildeo Southern Oscillation)-associatedrainfall predation and wildfire in central Australia Journal ofMammalogy 86 689ndash703 doi1016441545-1542(2005)086[0689TROMTL]20CO2

Letnic M Koch F Gordon C Crowther M S and Dickman C R(2009) Keystone effects of an alien top-predator stem extinctions ofnative mammals Proceedings of the Royal Society of London Series BBiological Sciences 276 3249ndash3256 doi101098rspb20090574

Lindenmayer D B (2015) Continental-level biodiversity collapseProceedings of the National Academy of Sciences of the United States ofAmerica 112 4514ndash4515 doi101073pnas1502766112

Lindenmayer D B Piggott M P and Wintle B A (2013) Counting thebooks while the library burns why conservation monitoring programs

need a plan for action Frontiers in Ecology and the Environment 11549ndash555 doi101890120220

Maron M Hobbs R J Moilanen A Matthews J W Christie KGardner T A Keith D A Lindenmayer D B and McAlpine C A(2012) Faustian bargains Restoration realities in the context ofbiodiversity offset policies Biological Conservation 155 141ndash148doi101016jbiocon201206003

McCall S C McCarthy M A van der Ree R Harper M J Cesarini Sand Soanes K (2010) Evidence that a highway reduces apparentsurvival rates of squirrel gliders Ecology and Society 15 27

McGregor R A Stokes V L and Craig M D (2014) Does forestrestoration in fragmented landscapes provide habitat for a wide-ranging carnivore Animal Conservation 17 467ndash475 doi101111acv12112

McKenzie N L Burbidge A A Baynes A Brereton R N DickmanC R Gordon G Gibson L A Menkhorst P W Robinson A CWilliams M R and Woinarski J C Z (2007) Analysis of factorsimplicated in the recent decline of Australiarsquos mammal fauna Journal ofBiogeography 34 597ndash611 doi101111j1365-2699200601639x

McKenzie N L van Leeuwen S and Pinder AM (2009) Introduction tothe Pilbara Biodiversity Survey 2002ndash2007 Records of the WesternAustralian Museum 78(Suppl) 3ndash89 doi1018195issn0313-122x78(1)2009003-089

Morris K Johnson B Orell P Gaikhorst G Wane A and MoroD (2003) Recovery of the threatened chuditch (Dasyurus geoffroii)a case study In lsquoPredators with Pouches The Biology of CarnivorousMarsupialsrsquo (Eds M Jones C Dickman and M Archer) pp 435ndash451(CSIRO Publishing Melbourne)

Nelson J L Scroggie M P and Belcher C A (2014) Developing acamera trap survey protocol to detect a rare marsupial carnivore thespotted-tailed quoll (Dasyurus maculatus) In lsquoCamera TrappingWildlife Management and Researchrsquo (Eds P D Meek and PJ Fleming) pp 271ndash279 (CSIRO Publishing Melbourne)

Newsome T M Dellinger J A Pavey C R Ripple W J Shores C RWirsing A J and Dickman C R (2015) The ecological effects ofproviding resource subsidies to predators Global Ecology andBiogeography 24 1ndash11 doi101111geb12236

OrsquoDonnell S Webb J K and Shine R (2010) Conditioned tasteaversion enhances the survival of an endangered predator imperilled bya toxic invader Journal of Applied Ecology 47 558ndash565 doi101111j1365-2664201001802x

Oakwood M (1997) The ecology of the northern quoll Dasyurushallucatus PhD Thesis Australian National University Canberra

Oakwood M (2000) Reproduction and demography of the northern quollDasyurus hallucatus in the lowland savanna of northern AustraliaAustralian Journal of Zoology 48 519ndash539 doi101071ZO00028

Oakwood M (2002) Spatial and social organization of a carnivorousmarsupial Dasyurus hallucatus (Marsupialia Dasyuridae) Journal ofZoology 257 237ndash248 doi101017S0952836902000833

Oakwood M and Foster P (2008) Monitoring extinction of the northernquoll Australian Academy of Science Newsletter 71 6

Palomares F and Caro T M (1999) Interspecific killing amongmammalian carnivores American Naturalist 153 492ndash508 doi101086303189

Pavey C R and Bastin G (2014) lsquoAustralian Rangelands and ClimateChange ndash Invasive Animalsrsquo (Ninti One Limited amp CSIRO AliceSprings)

Pierson J C Barton P S Lane PW and Lindenmayer D B (2015) Canhabitat surrogates predict the response of target species to landscapechange Biological Conservation 184 1ndash10 doi101016jbiocon201412017

Pollock A B (1999) Notes on status distribution and diet of northernquoll Dasyurus hallucatus in the MackayndashBowen area mideastern

Research priorities for the northern quoll Australian Mammalogy M

Queensland Australian Zoologist 31 388ndash395 doi107882AZ1999040

Ramanaidou E R and Morris R C (2010) A synopsis of the channeliron deposits of the Hamersley Province Western AustraliaTransactions of the Institutions of Mining and Metallurgy Section BApplied Earth Science 119 56ndash59 doi101179037174510X12853354810624

Schmitt L H Bradley A J Kemper C M Kitchener D J HumphreysW F and How R A (1989) Ecology and physiology of the northernquoll Dasyurus hallucatus (Marsupialia Dasyuridae) at MitchellPlateau Kimberley Western Australia Journal of Zoology 217539ndash558 doi101111j1469-79981989tb02510x

Short J and Smith A (1994) Mammal decline and recovery in AustraliaJournal of Mammalogy 75 288ndash297 doi1023071382547

Spencer P How R Hillyer M Cook A Morris K Stevenson C andUmbrello L (2013) Genetic analysis of northern quolls from the PilbaraRegion ofWestern Australia Year one ndash final report Unpublished reportDepartment of Sustainability Environment Water Population andCommunities Canberra and Department of Parks and Wildlife Perth

Sutherland W J Fleishman E Mascia M B Pretty J and Rudd M A(2011) Methods for collaboratively identifying research priorities andemerging issues in science and policyMethods in Ecology and Evolution2 238ndash247 doi101111j2041-210X201000083x

Sutherst R W Floyd R B and Maywald G F (1996) The potentialgeographical distribution of the cane toad Bufo marinus L in AustraliaConservation Biology 10 294ndash299 doi101046j1523-1739199610010294x

Taylor B D and Goldingay R L (2003) Cutting the carnage wildlifeusage of road culverts in north-eastern New South Wales WildlifeResearch 30 529ndash537 doi101071WR01062

Tingley R Phillips B L Letnic M Brown G P Shine R and BairdS J (2013) Identifying optimal barriers to halt the invasion of canetoads Rhinella marina in arid Australia Journal of Applied Ecology 50129ndash137 doi1011111365-266412021

Van Vreeswyk A M E Payne A L Leighton K A and HennigP (2004) An Inventory and Condition Survey of the Pilbara RegionWestern Australia Technical Bulletin No 92 Department of AgricultureWestern Australia Perth

Webb J Legge S Tuft K Cremona T and Austin C (2015) Can wemitigate cane toad impacts on northern quolls Final report National

Environmental Research Program Northern Australia Hub CharlesDarwin University Darwin

Wintle B A Walshe T V Parris K M and McCarthy M A (2012)Designing occupancy surveys and interpreting non-detection whenobservations are imperfect Diversity amp Distributions 18 417ndash424doi101111j1472-4642201100874x

Woinarski J Menkhorst K Gambold N and Braithwaite R W (1992)Mammals of the Bungle Bungle area In lsquoA Survey of the Wildlife andVegetation of Purnululu (Bungle Bungle) National Park and AdjacentArearsquo (Ed J C Z Woinarski) Research Bulletin No 6 pp 53ndash67(CSIRO Division of Wildlife and Ecology Darwin)

Woinarski J C Z Risler J and Kean L (2004) Response of vegetationand vertebrate fauna to 23 years of fire exclusion in a tropical Eucalyptusopen forest Northern Territory Australia Austral Ecology 29 156ndash176doi101111j1442-9993200401333x

Woinarski J C Z Oakwood M Winter J Burnett S Milne D FosterP Myles H and Holmes B (2008) Surviving the toads patterns ofpersistence of the northern quoll Dasyurus hallucatus in QueenslandReport submitted to the Natural Heritage Trust Strategic ReserveProgram as a component of project 2005162 Monitoring andManagement of Cane Toad Impact in theNorthern Territory Available athttplrmntgovauplants-and-animalsinformation-and-publicationspublications2008 [accessed 14 November 2013]

Woinarski J C Z Armstrong M Brennan K Fisher A Griffiths A DHill B Milne D J Palmer C Ward S Watson M Winderlich Sand Young S (2010) Monitoring indicates rapid and severe decline ofnative small mammals in Kakadu National Park northern AustraliaWildlife Research 37 116ndash126 doi101071WR09125

Woinarski J Burbidge A and Harrison P (2014) lsquoAction Plan forAustralian Mammals 2012rsquo (CSIRO Publishing Melbourne)

Woinarski J C Burbidge A A and Harrison P L (2015) Ongoingunraveling of a continental fauna decline and extinction of Australianmammals since European settlement Proceedings of the NationalAcademy of Sciences of the United States of America 112 4531ndash4540doi101073pnas1417301112

Woolley P A Krajewski C and Westerman M (2015) Phylogeneticrelationships within Dasyurus (Dasyuromorphia Dasyuridae) quollsystematics based on molecular evidence and male characteristicsJournal of Mammalogy 96 37ndash46 doi101093jmammalgyu028

N Australian Mammalogy V A Cramer et al

wwwpublishcsiroaujournalsam

Page 12: Research priorities for the northern quoll (Dasyurus ......The Pilbara population of the northern quoll (Dasyurus hallucatus) has been seldom studied, and the impacts of threats such

practitioners and proponents currently operating in the regionThe provision of offset funds over the next decade providesa unique opportunity to undertake research on a species ofconservation significance at a regional scale and over a longperiod through an integrated and strategic program of researchrather than the collection of ad hoc data While a morestructured approach to defining research questions before theworkshop may have led to greater critical reflection upon anddiscussion of some of the research priorities on the day theworkshop provided an opportunity for an open and diverse-ranging discussion in a collaborative environment amongstthe broader community of scientists environmental consultantsand mining proponents involved in the management andconservation of the northern quoll in the Pilbara

Acknowledgements

We gratefully acknowledge the financial support of Roy Hill who sponsoredthe workshop and partially funded VACrsquos salary We thank the Departmentof the Environmentrsquos Pilbara Taskforce especially Veronica Ritchie fortheir support of interested or affected parties The success of the workshopwas in great part due to the facilitation skills of Craig Salt of SustainableConsulting We thank all of the workshop participants for their open andcollaborative approach to the process Mike Bamford Eddie van EttenDavid Peacock and Peter Lyon provided useful comments on an earlier draftWe are grateful to John Woinarski for his constructive advice thatsubstantially improved the structure and clarity of the manuscript and ananonymous reviewer for careful and critical reading of the previousversion Lesley Gibson Russell Palmer and Al Glen generously discussedvarious aspects of the paper with VAC The map was prepared by CristinaRamalho

References

Allen B L Engeman R M and Allen L R (2011) Wild dogma anexamination of recent lsquoevidencersquo for dingo regulation of invasivemesopredator release in Australia Current Zoology 57 568ndash583

Allen B L Fleming P J Allen L R Engeman R M Ballard G andLeung L K-P (2013) As clear as mud a critical review of evidencefor the ecological roles of Australian dingoes Biological Conservation159 158ndash174 doi101016jbiocon201212004

Anon (1934) Floods in Western Australia Town isolated for week PerthMonday The Sydney Morning Herald 9

Armstrong D P and Reynolds M H (2012) Modelling reintroducedpopulations the state of the art and future directions In lsquoReintroductionBiology Integrating Science and Managementrsquo (Eds J G Ewen DP Armstrong K A Parker and P J Seddon) pp 165ndash222 (Wiley-Blackwell UK)

Atlas Iron (2012) Scope of works ndash Mount Dove artificial quoll habitatUnpublished report Atlas Iron Perth Available at httpwwwatlasironcomauIRMCompanyShowPageaspxCategoryId=190ampCPID=5424ampEID=94365410 [accessed 19 August 2015]

Austin C (2014) Can remote cameras accurately estimate populations ofthe endangered northern quoll BSc(Honours) Thesis University ofTechnology Sydney

Begg R J (1981) The small mammals of Little Nourlangie RockNT III Ecology ofDasyurus hallucatus the northern quoll (MarsupialiaDasyuridae) Wildlife Research 8 73ndash85 doi101071WR9810073

Braithwaite R W and Griffiths A D (1994) Demographic variationand range contraction in the northern quoll Dasyurus hallucatus(Marsupialia Dasyuridae)Wildlife Research 21 203ndash217 doi101071WR9940203

Bull J W Suttle K B Gordon A Singh N J and Milner-Gulland E J(2013) Biodiversity offsets in theory and practice Oryx 47 369ndash380doi101017S003060531200172X

Burnett S (1997) Colonizing cane toads cause population declines innative predators reliable anecdotal information and managementimplications Pacific Conservation Biology 3 65ndash72

Cadenhead N C R White A L and Wintle B A (2015) Identifyingconservation priorities and assessing impacts of proposed futuredevelopment in the Pilbara Bioregion in Western Australia Report bythe National Environment Research Program Decisions Hub to theDepartment of the Environment The University of MelbourneMelbourne

Carwardine J Nicol S Van Leeuwen S Walters B Firn J Reeson AMartin T G and Chades I (2014) Priority threat management forPilbara species of conservation significance CSIRO EcosystemSciences Brisbane

Cook A (2010) Habitat use and home-range of the northern quollDasyurushallucatus effects of fire MSc Thesis University ofWestern AustraliaPerth

Department of Sustainability Environment Water Population andCommunities (2012) Environment and Biodiversity Conservation Act1999 Environmental Offsets Policy Department of SustainabilityEnvironment Water Population and Communities Canberra

Doherty T S Davis R A Etten E J Algar D Collier N DickmanC R Edwards G Masters P Palmer R and Robinson S (2015a)A continental-scale analysis of feral cat diet in Australia Journal ofBiogeography 42 964ndash975 doi101111jbi12469

Doherty T S Dickman C R Nimmo D G and Ritchie E G (2015b)Multiple threats or multiplying the threats Interactions betweeninvasive predators and other ecological disturbances BiologicalConservation 190 60ndash68 doi101016jbiocon201505013

Dunlop J Cook A and Lees J (2014a) BHP Billiton Iron Ore Pilbaranorthern quoll monitoring project Final report May 2010ndashJune 2013Unpublished report prepared for BHP Billiton Iron Ore Department ofParks and Wildlife Perth

Dunlop J Cook A and Morris K (2014b) Pilbara northern quollproject surveying and monitoring Dasyurus hallucatus in the PilbaraWestern Australia Department of Parks and Wildlife Perth Availableat librarydpawwagovau

Florance D Webb J K Dempster T Kearney M R Worthing A andLetnic M (2011) Excluding access to invasion hubs can contain thespread of an invasive vertebrate Proceedings of the Royal Society ofLondon B Biological Sciences 278 2900ndash2908 doi101098rspb20110032

Gaillard J-M Hebblewhite M Loison A Fuller M Powell R BasilleM and Van Moorter B (2010) Habitatndashperformance relationshipsfinding the right metric at a given spatial scale PhilosophicalTransactions of the Royal Society of London Series B BiologicalSciences 365 2255ndash2265 doi101098rstb20100085

Glen A S and Dickman C R (2008) Niche overlap between marsupialand eutherian carnivores does competition threaten the endangeredspotted-tailed quoll Journal of Applied Ecology 45 700ndash707 doi101111j1365-2664200701449x

Glen A S Pennay M Dickman C R Wintle B A and Firestone K B(2011) Diets of sympatric native and introduced carnivores in theBarrington Tops eastern Australia Austral Ecology 36 290ndash296doi101111j1442-9993201002149x

Hennig P (2004) A brief land use history In lsquoAn Inventory and ConditionSurvey of the Pilbara Region Western Australiarsquo (Eds A M E VanVreeswyk A L Payne K A Leighton and P Hennig) TechnicalBulletin No 92 pp 13ndash18 (Western Australian Department ofAgriculture Perth)

Hohnen R Ashby J Tuft K and McGregor H (2013) Individualidentification of northern quolls (Dasyurus hallucatus) using remotecameras Australian Mammalogy 35 131ndash135 doi101071AM12015

L Australian Mammalogy V A Cramer et al

How R A Spencer P B S and Schmitt L H (2009) Island populationshave high conservation value for northern Australiarsquos top marsupialpredator ahead of a threatening process Journal of Zoology 278206ndash217 doi101111j1469-7998200900569x

Johnson C N Isaac J L and Fisher D O (2007) Rarity of a top predatortriggers continent-wide collapse of mammal prey dingoes andmarsupials in Australia Proceedings of the Royal Society of LondonSeries B Biological Sciences 274 341ndash346 doi101098rspb20063711

Jones M E (2000) Road upgrade road mortality and remedial measuresimpacts on a population of eastern quolls and Tasmanian devilsWildlifeResearch 27 289ndash296 doi101071WR98069

Kearney M Phillips B L Tracy C R Christian K A Betts G andPorter W P (2008) Modelling species distributions without usingspecies distributions the cane toad in Australia under current andfuture climates Ecography 31 423ndash434 doi101111j0906-7590200805457x

Kendrick P and Stanley F (2002) Pilbara 4 (PIL4 ndash Roebourne synopsis)In lsquoA Biodiversity Audit of Western Australiarsquos 53 BiogeographicalSubregions in 2002rsquo (Eds J E May and N L McKenzie) pp 581ndash593(Department of Conservation and Land Management Perth)

Kennedy M Phillips B L Legge S Murphy S A and Faulkner R A(2012) Do dingoes suppress the activity of feral cats in northernAustralia Austral Ecology 37 134ndash139 doi101111j1442-9993201102256x

Kiesecker J M Copeland H Pocewicz A and McKenney B (2010)Development by design blending landscape-level planning with themitigation hierarchy Frontiers in Ecology and the Environment 8261ndash266 doi101890090005

King D R and Smith L A (1985) The distribution of the European red fox(Vulpes vulpes) in Western Australia Records of the Western AustralianMuseum 12 197ndash205

Knight A T Cowling R M Rouget M Balmford A Lombard A Tand Campbell B M (2008) Knowing but not doing selecting priorityconservation areas and the researchndashimplementation gap ConservationBiology 22 610ndash617 doi101111j1523-1739200800914x

Laurance W F Koster H Grooten M Anderson A B Zuidema P AZwick S Zagt R J Lynam A J Linkie M and Anten N P (2012)Making conservation research more relevant for conservationpractitioners Biological Conservation 153 164ndash168 doi101016jbiocon201205012

Legge S Murphy S Heathcote J Flaxman E Augusteyn J andCrossman M (2008) The short-term effects of an extensive and high-intensity fire on vertebrates in the tropical savannas of the centralKimberley northern Australia Wildlife Research 35 33ndash43 doi101071WR07016

Lester D (2015) Effective wildlife roadkill mitigation Journal of Trafficand Transportation Engineering 3 42ndash51 doi10172652328-2142201501005

Letnic M Tamayo B and Dickman C R (2005) The responsesof mammals to La Nintildea (El Nintildeo Southern Oscillation)-associatedrainfall predation and wildfire in central Australia Journal ofMammalogy 86 689ndash703 doi1016441545-1542(2005)086[0689TROMTL]20CO2

Letnic M Koch F Gordon C Crowther M S and Dickman C R(2009) Keystone effects of an alien top-predator stem extinctions ofnative mammals Proceedings of the Royal Society of London Series BBiological Sciences 276 3249ndash3256 doi101098rspb20090574

Lindenmayer D B (2015) Continental-level biodiversity collapseProceedings of the National Academy of Sciences of the United States ofAmerica 112 4514ndash4515 doi101073pnas1502766112

Lindenmayer D B Piggott M P and Wintle B A (2013) Counting thebooks while the library burns why conservation monitoring programs

need a plan for action Frontiers in Ecology and the Environment 11549ndash555 doi101890120220

Maron M Hobbs R J Moilanen A Matthews J W Christie KGardner T A Keith D A Lindenmayer D B and McAlpine C A(2012) Faustian bargains Restoration realities in the context ofbiodiversity offset policies Biological Conservation 155 141ndash148doi101016jbiocon201206003

McCall S C McCarthy M A van der Ree R Harper M J Cesarini Sand Soanes K (2010) Evidence that a highway reduces apparentsurvival rates of squirrel gliders Ecology and Society 15 27

McGregor R A Stokes V L and Craig M D (2014) Does forestrestoration in fragmented landscapes provide habitat for a wide-ranging carnivore Animal Conservation 17 467ndash475 doi101111acv12112

McKenzie N L Burbidge A A Baynes A Brereton R N DickmanC R Gordon G Gibson L A Menkhorst P W Robinson A CWilliams M R and Woinarski J C Z (2007) Analysis of factorsimplicated in the recent decline of Australiarsquos mammal fauna Journal ofBiogeography 34 597ndash611 doi101111j1365-2699200601639x

McKenzie N L van Leeuwen S and Pinder AM (2009) Introduction tothe Pilbara Biodiversity Survey 2002ndash2007 Records of the WesternAustralian Museum 78(Suppl) 3ndash89 doi1018195issn0313-122x78(1)2009003-089

Morris K Johnson B Orell P Gaikhorst G Wane A and MoroD (2003) Recovery of the threatened chuditch (Dasyurus geoffroii)a case study In lsquoPredators with Pouches The Biology of CarnivorousMarsupialsrsquo (Eds M Jones C Dickman and M Archer) pp 435ndash451(CSIRO Publishing Melbourne)

Nelson J L Scroggie M P and Belcher C A (2014) Developing acamera trap survey protocol to detect a rare marsupial carnivore thespotted-tailed quoll (Dasyurus maculatus) In lsquoCamera TrappingWildlife Management and Researchrsquo (Eds P D Meek and PJ Fleming) pp 271ndash279 (CSIRO Publishing Melbourne)

Newsome T M Dellinger J A Pavey C R Ripple W J Shores C RWirsing A J and Dickman C R (2015) The ecological effects ofproviding resource subsidies to predators Global Ecology andBiogeography 24 1ndash11 doi101111geb12236

OrsquoDonnell S Webb J K and Shine R (2010) Conditioned tasteaversion enhances the survival of an endangered predator imperilled bya toxic invader Journal of Applied Ecology 47 558ndash565 doi101111j1365-2664201001802x

Oakwood M (1997) The ecology of the northern quoll Dasyurushallucatus PhD Thesis Australian National University Canberra

Oakwood M (2000) Reproduction and demography of the northern quollDasyurus hallucatus in the lowland savanna of northern AustraliaAustralian Journal of Zoology 48 519ndash539 doi101071ZO00028

Oakwood M (2002) Spatial and social organization of a carnivorousmarsupial Dasyurus hallucatus (Marsupialia Dasyuridae) Journal ofZoology 257 237ndash248 doi101017S0952836902000833

Oakwood M and Foster P (2008) Monitoring extinction of the northernquoll Australian Academy of Science Newsletter 71 6

Palomares F and Caro T M (1999) Interspecific killing amongmammalian carnivores American Naturalist 153 492ndash508 doi101086303189

Pavey C R and Bastin G (2014) lsquoAustralian Rangelands and ClimateChange ndash Invasive Animalsrsquo (Ninti One Limited amp CSIRO AliceSprings)

Pierson J C Barton P S Lane PW and Lindenmayer D B (2015) Canhabitat surrogates predict the response of target species to landscapechange Biological Conservation 184 1ndash10 doi101016jbiocon201412017

Pollock A B (1999) Notes on status distribution and diet of northernquoll Dasyurus hallucatus in the MackayndashBowen area mideastern

Research priorities for the northern quoll Australian Mammalogy M

Queensland Australian Zoologist 31 388ndash395 doi107882AZ1999040

Ramanaidou E R and Morris R C (2010) A synopsis of the channeliron deposits of the Hamersley Province Western AustraliaTransactions of the Institutions of Mining and Metallurgy Section BApplied Earth Science 119 56ndash59 doi101179037174510X12853354810624

Schmitt L H Bradley A J Kemper C M Kitchener D J HumphreysW F and How R A (1989) Ecology and physiology of the northernquoll Dasyurus hallucatus (Marsupialia Dasyuridae) at MitchellPlateau Kimberley Western Australia Journal of Zoology 217539ndash558 doi101111j1469-79981989tb02510x

Short J and Smith A (1994) Mammal decline and recovery in AustraliaJournal of Mammalogy 75 288ndash297 doi1023071382547

Spencer P How R Hillyer M Cook A Morris K Stevenson C andUmbrello L (2013) Genetic analysis of northern quolls from the PilbaraRegion ofWestern Australia Year one ndash final report Unpublished reportDepartment of Sustainability Environment Water Population andCommunities Canberra and Department of Parks and Wildlife Perth

Sutherland W J Fleishman E Mascia M B Pretty J and Rudd M A(2011) Methods for collaboratively identifying research priorities andemerging issues in science and policyMethods in Ecology and Evolution2 238ndash247 doi101111j2041-210X201000083x

Sutherst R W Floyd R B and Maywald G F (1996) The potentialgeographical distribution of the cane toad Bufo marinus L in AustraliaConservation Biology 10 294ndash299 doi101046j1523-1739199610010294x

Taylor B D and Goldingay R L (2003) Cutting the carnage wildlifeusage of road culverts in north-eastern New South Wales WildlifeResearch 30 529ndash537 doi101071WR01062

Tingley R Phillips B L Letnic M Brown G P Shine R and BairdS J (2013) Identifying optimal barriers to halt the invasion of canetoads Rhinella marina in arid Australia Journal of Applied Ecology 50129ndash137 doi1011111365-266412021

Van Vreeswyk A M E Payne A L Leighton K A and HennigP (2004) An Inventory and Condition Survey of the Pilbara RegionWestern Australia Technical Bulletin No 92 Department of AgricultureWestern Australia Perth

Webb J Legge S Tuft K Cremona T and Austin C (2015) Can wemitigate cane toad impacts on northern quolls Final report National

Environmental Research Program Northern Australia Hub CharlesDarwin University Darwin

Wintle B A Walshe T V Parris K M and McCarthy M A (2012)Designing occupancy surveys and interpreting non-detection whenobservations are imperfect Diversity amp Distributions 18 417ndash424doi101111j1472-4642201100874x

Woinarski J Menkhorst K Gambold N and Braithwaite R W (1992)Mammals of the Bungle Bungle area In lsquoA Survey of the Wildlife andVegetation of Purnululu (Bungle Bungle) National Park and AdjacentArearsquo (Ed J C Z Woinarski) Research Bulletin No 6 pp 53ndash67(CSIRO Division of Wildlife and Ecology Darwin)

Woinarski J C Z Risler J and Kean L (2004) Response of vegetationand vertebrate fauna to 23 years of fire exclusion in a tropical Eucalyptusopen forest Northern Territory Australia Austral Ecology 29 156ndash176doi101111j1442-9993200401333x

Woinarski J C Z Oakwood M Winter J Burnett S Milne D FosterP Myles H and Holmes B (2008) Surviving the toads patterns ofpersistence of the northern quoll Dasyurus hallucatus in QueenslandReport submitted to the Natural Heritage Trust Strategic ReserveProgram as a component of project 2005162 Monitoring andManagement of Cane Toad Impact in theNorthern Territory Available athttplrmntgovauplants-and-animalsinformation-and-publicationspublications2008 [accessed 14 November 2013]

Woinarski J C Z Armstrong M Brennan K Fisher A Griffiths A DHill B Milne D J Palmer C Ward S Watson M Winderlich Sand Young S (2010) Monitoring indicates rapid and severe decline ofnative small mammals in Kakadu National Park northern AustraliaWildlife Research 37 116ndash126 doi101071WR09125

Woinarski J Burbidge A and Harrison P (2014) lsquoAction Plan forAustralian Mammals 2012rsquo (CSIRO Publishing Melbourne)

Woinarski J C Burbidge A A and Harrison P L (2015) Ongoingunraveling of a continental fauna decline and extinction of Australianmammals since European settlement Proceedings of the NationalAcademy of Sciences of the United States of America 112 4531ndash4540doi101073pnas1417301112

Woolley P A Krajewski C and Westerman M (2015) Phylogeneticrelationships within Dasyurus (Dasyuromorphia Dasyuridae) quollsystematics based on molecular evidence and male characteristicsJournal of Mammalogy 96 37ndash46 doi101093jmammalgyu028

N Australian Mammalogy V A Cramer et al

wwwpublishcsiroaujournalsam

Page 13: Research priorities for the northern quoll (Dasyurus ......The Pilbara population of the northern quoll (Dasyurus hallucatus) has been seldom studied, and the impacts of threats such

How R A Spencer P B S and Schmitt L H (2009) Island populationshave high conservation value for northern Australiarsquos top marsupialpredator ahead of a threatening process Journal of Zoology 278206ndash217 doi101111j1469-7998200900569x

Johnson C N Isaac J L and Fisher D O (2007) Rarity of a top predatortriggers continent-wide collapse of mammal prey dingoes andmarsupials in Australia Proceedings of the Royal Society of LondonSeries B Biological Sciences 274 341ndash346 doi101098rspb20063711

Jones M E (2000) Road upgrade road mortality and remedial measuresimpacts on a population of eastern quolls and Tasmanian devilsWildlifeResearch 27 289ndash296 doi101071WR98069

Kearney M Phillips B L Tracy C R Christian K A Betts G andPorter W P (2008) Modelling species distributions without usingspecies distributions the cane toad in Australia under current andfuture climates Ecography 31 423ndash434 doi101111j0906-7590200805457x

Kendrick P and Stanley F (2002) Pilbara 4 (PIL4 ndash Roebourne synopsis)In lsquoA Biodiversity Audit of Western Australiarsquos 53 BiogeographicalSubregions in 2002rsquo (Eds J E May and N L McKenzie) pp 581ndash593(Department of Conservation and Land Management Perth)

Kennedy M Phillips B L Legge S Murphy S A and Faulkner R A(2012) Do dingoes suppress the activity of feral cats in northernAustralia Austral Ecology 37 134ndash139 doi101111j1442-9993201102256x

Kiesecker J M Copeland H Pocewicz A and McKenney B (2010)Development by design blending landscape-level planning with themitigation hierarchy Frontiers in Ecology and the Environment 8261ndash266 doi101890090005

King D R and Smith L A (1985) The distribution of the European red fox(Vulpes vulpes) in Western Australia Records of the Western AustralianMuseum 12 197ndash205

Knight A T Cowling R M Rouget M Balmford A Lombard A Tand Campbell B M (2008) Knowing but not doing selecting priorityconservation areas and the researchndashimplementation gap ConservationBiology 22 610ndash617 doi101111j1523-1739200800914x

Laurance W F Koster H Grooten M Anderson A B Zuidema P AZwick S Zagt R J Lynam A J Linkie M and Anten N P (2012)Making conservation research more relevant for conservationpractitioners Biological Conservation 153 164ndash168 doi101016jbiocon201205012

Legge S Murphy S Heathcote J Flaxman E Augusteyn J andCrossman M (2008) The short-term effects of an extensive and high-intensity fire on vertebrates in the tropical savannas of the centralKimberley northern Australia Wildlife Research 35 33ndash43 doi101071WR07016

Lester D (2015) Effective wildlife roadkill mitigation Journal of Trafficand Transportation Engineering 3 42ndash51 doi10172652328-2142201501005

Letnic M Tamayo B and Dickman C R (2005) The responsesof mammals to La Nintildea (El Nintildeo Southern Oscillation)-associatedrainfall predation and wildfire in central Australia Journal ofMammalogy 86 689ndash703 doi1016441545-1542(2005)086[0689TROMTL]20CO2

Letnic M Koch F Gordon C Crowther M S and Dickman C R(2009) Keystone effects of an alien top-predator stem extinctions ofnative mammals Proceedings of the Royal Society of London Series BBiological Sciences 276 3249ndash3256 doi101098rspb20090574

Lindenmayer D B (2015) Continental-level biodiversity collapseProceedings of the National Academy of Sciences of the United States ofAmerica 112 4514ndash4515 doi101073pnas1502766112

Lindenmayer D B Piggott M P and Wintle B A (2013) Counting thebooks while the library burns why conservation monitoring programs

need a plan for action Frontiers in Ecology and the Environment 11549ndash555 doi101890120220

Maron M Hobbs R J Moilanen A Matthews J W Christie KGardner T A Keith D A Lindenmayer D B and McAlpine C A(2012) Faustian bargains Restoration realities in the context ofbiodiversity offset policies Biological Conservation 155 141ndash148doi101016jbiocon201206003

McCall S C McCarthy M A van der Ree R Harper M J Cesarini Sand Soanes K (2010) Evidence that a highway reduces apparentsurvival rates of squirrel gliders Ecology and Society 15 27

McGregor R A Stokes V L and Craig M D (2014) Does forestrestoration in fragmented landscapes provide habitat for a wide-ranging carnivore Animal Conservation 17 467ndash475 doi101111acv12112

McKenzie N L Burbidge A A Baynes A Brereton R N DickmanC R Gordon G Gibson L A Menkhorst P W Robinson A CWilliams M R and Woinarski J C Z (2007) Analysis of factorsimplicated in the recent decline of Australiarsquos mammal fauna Journal ofBiogeography 34 597ndash611 doi101111j1365-2699200601639x

McKenzie N L van Leeuwen S and Pinder AM (2009) Introduction tothe Pilbara Biodiversity Survey 2002ndash2007 Records of the WesternAustralian Museum 78(Suppl) 3ndash89 doi1018195issn0313-122x78(1)2009003-089

Morris K Johnson B Orell P Gaikhorst G Wane A and MoroD (2003) Recovery of the threatened chuditch (Dasyurus geoffroii)a case study In lsquoPredators with Pouches The Biology of CarnivorousMarsupialsrsquo (Eds M Jones C Dickman and M Archer) pp 435ndash451(CSIRO Publishing Melbourne)

Nelson J L Scroggie M P and Belcher C A (2014) Developing acamera trap survey protocol to detect a rare marsupial carnivore thespotted-tailed quoll (Dasyurus maculatus) In lsquoCamera TrappingWildlife Management and Researchrsquo (Eds P D Meek and PJ Fleming) pp 271ndash279 (CSIRO Publishing Melbourne)

Newsome T M Dellinger J A Pavey C R Ripple W J Shores C RWirsing A J and Dickman C R (2015) The ecological effects ofproviding resource subsidies to predators Global Ecology andBiogeography 24 1ndash11 doi101111geb12236

OrsquoDonnell S Webb J K and Shine R (2010) Conditioned tasteaversion enhances the survival of an endangered predator imperilled bya toxic invader Journal of Applied Ecology 47 558ndash565 doi101111j1365-2664201001802x

Oakwood M (1997) The ecology of the northern quoll Dasyurushallucatus PhD Thesis Australian National University Canberra

Oakwood M (2000) Reproduction and demography of the northern quollDasyurus hallucatus in the lowland savanna of northern AustraliaAustralian Journal of Zoology 48 519ndash539 doi101071ZO00028

Oakwood M (2002) Spatial and social organization of a carnivorousmarsupial Dasyurus hallucatus (Marsupialia Dasyuridae) Journal ofZoology 257 237ndash248 doi101017S0952836902000833

Oakwood M and Foster P (2008) Monitoring extinction of the northernquoll Australian Academy of Science Newsletter 71 6

Palomares F and Caro T M (1999) Interspecific killing amongmammalian carnivores American Naturalist 153 492ndash508 doi101086303189

Pavey C R and Bastin G (2014) lsquoAustralian Rangelands and ClimateChange ndash Invasive Animalsrsquo (Ninti One Limited amp CSIRO AliceSprings)

Pierson J C Barton P S Lane PW and Lindenmayer D B (2015) Canhabitat surrogates predict the response of target species to landscapechange Biological Conservation 184 1ndash10 doi101016jbiocon201412017

Pollock A B (1999) Notes on status distribution and diet of northernquoll Dasyurus hallucatus in the MackayndashBowen area mideastern

Research priorities for the northern quoll Australian Mammalogy M

Queensland Australian Zoologist 31 388ndash395 doi107882AZ1999040

Ramanaidou E R and Morris R C (2010) A synopsis of the channeliron deposits of the Hamersley Province Western AustraliaTransactions of the Institutions of Mining and Metallurgy Section BApplied Earth Science 119 56ndash59 doi101179037174510X12853354810624

Schmitt L H Bradley A J Kemper C M Kitchener D J HumphreysW F and How R A (1989) Ecology and physiology of the northernquoll Dasyurus hallucatus (Marsupialia Dasyuridae) at MitchellPlateau Kimberley Western Australia Journal of Zoology 217539ndash558 doi101111j1469-79981989tb02510x

Short J and Smith A (1994) Mammal decline and recovery in AustraliaJournal of Mammalogy 75 288ndash297 doi1023071382547

Spencer P How R Hillyer M Cook A Morris K Stevenson C andUmbrello L (2013) Genetic analysis of northern quolls from the PilbaraRegion ofWestern Australia Year one ndash final report Unpublished reportDepartment of Sustainability Environment Water Population andCommunities Canberra and Department of Parks and Wildlife Perth

Sutherland W J Fleishman E Mascia M B Pretty J and Rudd M A(2011) Methods for collaboratively identifying research priorities andemerging issues in science and policyMethods in Ecology and Evolution2 238ndash247 doi101111j2041-210X201000083x

Sutherst R W Floyd R B and Maywald G F (1996) The potentialgeographical distribution of the cane toad Bufo marinus L in AustraliaConservation Biology 10 294ndash299 doi101046j1523-1739199610010294x

Taylor B D and Goldingay R L (2003) Cutting the carnage wildlifeusage of road culverts in north-eastern New South Wales WildlifeResearch 30 529ndash537 doi101071WR01062

Tingley R Phillips B L Letnic M Brown G P Shine R and BairdS J (2013) Identifying optimal barriers to halt the invasion of canetoads Rhinella marina in arid Australia Journal of Applied Ecology 50129ndash137 doi1011111365-266412021

Van Vreeswyk A M E Payne A L Leighton K A and HennigP (2004) An Inventory and Condition Survey of the Pilbara RegionWestern Australia Technical Bulletin No 92 Department of AgricultureWestern Australia Perth

Webb J Legge S Tuft K Cremona T and Austin C (2015) Can wemitigate cane toad impacts on northern quolls Final report National

Environmental Research Program Northern Australia Hub CharlesDarwin University Darwin

Wintle B A Walshe T V Parris K M and McCarthy M A (2012)Designing occupancy surveys and interpreting non-detection whenobservations are imperfect Diversity amp Distributions 18 417ndash424doi101111j1472-4642201100874x

Woinarski J Menkhorst K Gambold N and Braithwaite R W (1992)Mammals of the Bungle Bungle area In lsquoA Survey of the Wildlife andVegetation of Purnululu (Bungle Bungle) National Park and AdjacentArearsquo (Ed J C Z Woinarski) Research Bulletin No 6 pp 53ndash67(CSIRO Division of Wildlife and Ecology Darwin)

Woinarski J C Z Risler J and Kean L (2004) Response of vegetationand vertebrate fauna to 23 years of fire exclusion in a tropical Eucalyptusopen forest Northern Territory Australia Austral Ecology 29 156ndash176doi101111j1442-9993200401333x

Woinarski J C Z Oakwood M Winter J Burnett S Milne D FosterP Myles H and Holmes B (2008) Surviving the toads patterns ofpersistence of the northern quoll Dasyurus hallucatus in QueenslandReport submitted to the Natural Heritage Trust Strategic ReserveProgram as a component of project 2005162 Monitoring andManagement of Cane Toad Impact in theNorthern Territory Available athttplrmntgovauplants-and-animalsinformation-and-publicationspublications2008 [accessed 14 November 2013]

Woinarski J C Z Armstrong M Brennan K Fisher A Griffiths A DHill B Milne D J Palmer C Ward S Watson M Winderlich Sand Young S (2010) Monitoring indicates rapid and severe decline ofnative small mammals in Kakadu National Park northern AustraliaWildlife Research 37 116ndash126 doi101071WR09125

Woinarski J Burbidge A and Harrison P (2014) lsquoAction Plan forAustralian Mammals 2012rsquo (CSIRO Publishing Melbourne)

Woinarski J C Burbidge A A and Harrison P L (2015) Ongoingunraveling of a continental fauna decline and extinction of Australianmammals since European settlement Proceedings of the NationalAcademy of Sciences of the United States of America 112 4531ndash4540doi101073pnas1417301112

Woolley P A Krajewski C and Westerman M (2015) Phylogeneticrelationships within Dasyurus (Dasyuromorphia Dasyuridae) quollsystematics based on molecular evidence and male characteristicsJournal of Mammalogy 96 37ndash46 doi101093jmammalgyu028

N Australian Mammalogy V A Cramer et al

wwwpublishcsiroaujournalsam

Page 14: Research priorities for the northern quoll (Dasyurus ......The Pilbara population of the northern quoll (Dasyurus hallucatus) has been seldom studied, and the impacts of threats such

Queensland Australian Zoologist 31 388ndash395 doi107882AZ1999040

Ramanaidou E R and Morris R C (2010) A synopsis of the channeliron deposits of the Hamersley Province Western AustraliaTransactions of the Institutions of Mining and Metallurgy Section BApplied Earth Science 119 56ndash59 doi101179037174510X12853354810624

Schmitt L H Bradley A J Kemper C M Kitchener D J HumphreysW F and How R A (1989) Ecology and physiology of the northernquoll Dasyurus hallucatus (Marsupialia Dasyuridae) at MitchellPlateau Kimberley Western Australia Journal of Zoology 217539ndash558 doi101111j1469-79981989tb02510x

Short J and Smith A (1994) Mammal decline and recovery in AustraliaJournal of Mammalogy 75 288ndash297 doi1023071382547

Spencer P How R Hillyer M Cook A Morris K Stevenson C andUmbrello L (2013) Genetic analysis of northern quolls from the PilbaraRegion ofWestern Australia Year one ndash final report Unpublished reportDepartment of Sustainability Environment Water Population andCommunities Canberra and Department of Parks and Wildlife Perth

Sutherland W J Fleishman E Mascia M B Pretty J and Rudd M A(2011) Methods for collaboratively identifying research priorities andemerging issues in science and policyMethods in Ecology and Evolution2 238ndash247 doi101111j2041-210X201000083x

Sutherst R W Floyd R B and Maywald G F (1996) The potentialgeographical distribution of the cane toad Bufo marinus L in AustraliaConservation Biology 10 294ndash299 doi101046j1523-1739199610010294x

Taylor B D and Goldingay R L (2003) Cutting the carnage wildlifeusage of road culverts in north-eastern New South Wales WildlifeResearch 30 529ndash537 doi101071WR01062

Tingley R Phillips B L Letnic M Brown G P Shine R and BairdS J (2013) Identifying optimal barriers to halt the invasion of canetoads Rhinella marina in arid Australia Journal of Applied Ecology 50129ndash137 doi1011111365-266412021

Van Vreeswyk A M E Payne A L Leighton K A and HennigP (2004) An Inventory and Condition Survey of the Pilbara RegionWestern Australia Technical Bulletin No 92 Department of AgricultureWestern Australia Perth

Webb J Legge S Tuft K Cremona T and Austin C (2015) Can wemitigate cane toad impacts on northern quolls Final report National

Environmental Research Program Northern Australia Hub CharlesDarwin University Darwin

Wintle B A Walshe T V Parris K M and McCarthy M A (2012)Designing occupancy surveys and interpreting non-detection whenobservations are imperfect Diversity amp Distributions 18 417ndash424doi101111j1472-4642201100874x

Woinarski J Menkhorst K Gambold N and Braithwaite R W (1992)Mammals of the Bungle Bungle area In lsquoA Survey of the Wildlife andVegetation of Purnululu (Bungle Bungle) National Park and AdjacentArearsquo (Ed J C Z Woinarski) Research Bulletin No 6 pp 53ndash67(CSIRO Division of Wildlife and Ecology Darwin)

Woinarski J C Z Risler J and Kean L (2004) Response of vegetationand vertebrate fauna to 23 years of fire exclusion in a tropical Eucalyptusopen forest Northern Territory Australia Austral Ecology 29 156ndash176doi101111j1442-9993200401333x

Woinarski J C Z Oakwood M Winter J Burnett S Milne D FosterP Myles H and Holmes B (2008) Surviving the toads patterns ofpersistence of the northern quoll Dasyurus hallucatus in QueenslandReport submitted to the Natural Heritage Trust Strategic ReserveProgram as a component of project 2005162 Monitoring andManagement of Cane Toad Impact in theNorthern Territory Available athttplrmntgovauplants-and-animalsinformation-and-publicationspublications2008 [accessed 14 November 2013]

Woinarski J C Z Armstrong M Brennan K Fisher A Griffiths A DHill B Milne D J Palmer C Ward S Watson M Winderlich Sand Young S (2010) Monitoring indicates rapid and severe decline ofnative small mammals in Kakadu National Park northern AustraliaWildlife Research 37 116ndash126 doi101071WR09125

Woinarski J Burbidge A and Harrison P (2014) lsquoAction Plan forAustralian Mammals 2012rsquo (CSIRO Publishing Melbourne)

Woinarski J C Burbidge A A and Harrison P L (2015) Ongoingunraveling of a continental fauna decline and extinction of Australianmammals since European settlement Proceedings of the NationalAcademy of Sciences of the United States of America 112 4531ndash4540doi101073pnas1417301112

Woolley P A Krajewski C and Westerman M (2015) Phylogeneticrelationships within Dasyurus (Dasyuromorphia Dasyuridae) quollsystematics based on molecular evidence and male characteristicsJournal of Mammalogy 96 37ndash46 doi101093jmammalgyu028

N Australian Mammalogy V A Cramer et al

wwwpublishcsiroaujournalsam


GIL ADAMSON In weiter Ferne die Hunde - bilder.buecher.de · Über dem Mädchen im Wassergraben hing der Mond, klein und hart. Blasser Schaum quoll hoch, wo ihre Schuhe im Schlamm

GIL ADAMSON In weiter Ferne die Hunde - bilder.buecher.de · Über dem Mädchen im Wassergraben hing der Mond, klein und hart. Blasser Schaum quoll hoch, wo ihre Schuhe im Schlamm

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THE BRONZE QUOLL, DASYURUS SPARTACUS (MARSUPIALIA: …bionames.org/bionames-archive/issn/0310-0049/11/145.pdf · spartacus (spirit specimen PM22004) 2.1 mm, D. geoffroii (spirit specimen

THE BRONZE QUOLL, DASYURUS SPARTACUS (MARSUPIALIA: …bionames.org/bionames-archive/issn/0310-0049/11/145.pdf · spartacus (spirit specimen PM22004) 2.1 mm, D. geoffroii (spirit specimen

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Comparative physiology of Australian quolls (Dasyurus

Comparative physiology of Australian quolls (Dasyurus

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The Ability of Spotted-tailed Quolls to Locate and Consume ...Dasyurus maculatus, is . the last remaining quoll species extant in southeastern Australia. It is a forest-dependent carnivorous

The Ability of Spotted-tailed Quolls to Locate and Consume ...Dasyurus maculatus, is . the last remaining quoll species extant in southeastern Australia. It is a forest-dependent carnivorous

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Featherdale Endangered Species Spotted-tail quoll · Endangered Species Spotted-tail quoll By Kaizen Earl-Sawa 4M Elderslie Public School. Quolls There are four types of quolls in

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Offset Plan. The northern quoll cat bait uptake and ...A composite map of male and female quoll movement areas, and aerial bait drops, and the suspected spread of baits on the ground,

Offset Plan. The northern quoll cat bait uptake and ...A composite map of male and female quoll movement areas, and aerial bait drops, and the suspected spread of baits on the ground,

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Impact of fox baiting on tiger quoll populations - Final ...environment.gov.au/system/files/resources/321120e5-e421-4740-af6… · The potential for fox baiting to impact on tiger

Impact of fox baiting on tiger quoll populations - Final ...environment.gov.au/system/files/resources/321120e5-e421-4740-af6… · The potential for fox baiting to impact on tiger

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Chuditch (Dasyurus geoffroii) - Department of the Environment

Chuditch (Dasyurus geoffroii) - Department of the Environment

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Northern Quoll Regional Analysis Report v3

Northern Quoll Regional Analysis Report v3

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Habitat use and home-range of the northern quoll, · Habitat use and home-range of the northern quoll, Dasyurus hallucatus: effects of fire . This thesis is submitted in partial fulfilment

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Documents
Spotted Quoll Open Pit Nickel Mine€¦ · 5 4. Key environmental factors It is the EPA’s opinion that the following key environmental factors relevant to the proposal require evaluation

Spotted Quoll Open Pit Nickel Mine€¦ · 5 4. Key environmental factors It is the EPA’s opinion that the following key environmental factors relevant to the proposal require evaluation

Documents
SHINGLEBACK PADEMELON MU(24) STREET QUOLL PLACE … · mu(06) mu(06) street street glider street close street avenue crescent proposed environment education centre * dragon marron

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Name tags wordpress  · Web viewMt Evelyn Caterpillar Guides Enchanted Camp 2012. royal guild. Quoll. Mt Evelyn Caterpillar Guides Enchanted Camp 2012. elf guild. Author: Em Meades

Name tags wordpress  · Web viewMt Evelyn Caterpillar Guides Enchanted Camp 2012. royal guild. Quoll. Mt Evelyn Caterpillar Guides Enchanted Camp 2012. elf guild. Author: Em Meades

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The Effects of Mining Infrastructure on Northern quoll ... · Honours Thesis by Melinda Henderson Bachelor of Science (Conservation and Wildlife Biology) This document is confidential

The Effects of Mining Infrastructure on Northern quoll ... · Honours Thesis by Melinda Henderson Bachelor of Science (Conservation and Wildlife Biology) This document is confidential

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Activity Report - ASX · Activity Report For the period ending 31st March 2013 generation and reduced unit cash costs of production from the previous quarter. in place. Spotted Quoll

Activity Report - ASX · Activity Report For the period ending 31st March 2013 generation and reduced unit cash costs of production from the previous quarter. in place. Spotted Quoll

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Rainfall and topography predict gene flow among ...northern quoll (Dasyurus hallucatus) is a carnivorous marsupial that has experienced dramatic population decli-nes in recent decades

Rainfall and topography predict gene flow among ...northern quoll (Dasyurus hallucatus) is a carnivorous marsupial that has experienced dramatic population decli-nes in recent decades

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Iron Ore Holdings Ltd Buckland Project & Northern Quoll: adaptive mine design and management 30 July 2013 0

Iron Ore Holdings Ltd Buckland Project & Northern Quoll: adaptive mine design and management 30 July 2013 0

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Northern Quoll Workshop TalkBook · Vanja Sekizovic Calibre Global Veronica Ritchie DSEWPaC Viki Cramer DPaW Zoe Stokes Atlas Iron. Setting the scene Electronic whiteboard 01

Northern Quoll Workshop TalkBook · Vanja Sekizovic Calibre Global Veronica Ritchie DSEWPaC Viki Cramer DPaW Zoe Stokes Atlas Iron. Setting the scene Electronic whiteboard 01

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Northern Quoll (Dasyurus hallucatus)...(Dasyurus hallucatus) This small, omnivorous (eats both animals and plants) marsupial is one of the four Australian species of quoll. The Northern

Northern Quoll (Dasyurus hallucatus)...(Dasyurus hallucatus) This small, omnivorous (eats both animals and plants) marsupial is one of the four Australian species of quoll. The Northern

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Comparative Postnatal Ontogeny of the Skull in the ... · Comparative Postnatal Ontogeny of the Skull in the Australidelphian Metatherian Dasyurus albopunctatus (Marsupialia: Dasyuromorpha:

Comparative Postnatal Ontogeny of the Skull in the ... · Comparative Postnatal Ontogeny of the Skull in the Australidelphian Metatherian Dasyurus albopunctatus (Marsupialia: Dasyuromorpha:

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The Sturt's Desert Rose and Spotted Quoll

The Sturt's Desert Rose and Spotted Quoll

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2018 northern quoll annual report

2018 northern quoll annual report

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Quoll product datasheet FINAL 30-05 - Home - … ltd 7/36 Campbell Ave | Cromer NSW 2099 | Australia | Tel +61 403 606 187 | info@nocturnallighting.net.au | Code LED(/(Watt(s) Beam(Angle

Quoll product datasheet FINAL 30-05 - Home - … ltd 7/36 Campbell Ave | Cromer NSW 2099 | Australia | Tel +61 403 606 187 | [email protected] | Code LED(/(Watt(s) Beam(Angle

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Burnett, Scott Edward (2001) Ecology and conservation ...researchonline.jcu.edu.au/17435/3/03_References_and_Appendices.pdf · conservation status of the northern spot-tailed quoll,

Burnett, Scott Edward (2001) Ecology and conservation ...researchonline.jcu.edu.au/17435/3/03_References_and_Appendices.pdf · conservation status of the northern spot-tailed quoll,

Documents
Copyright Act 1968 · Draft Action Plan - Spotted-tailed Quoll 3 Occasional sightings of . D. maculatus . continue to occur across the ACT and surrounding region and there have been

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National Recovery Plan for the Spotted-tailed Quoll … · Web viewSpotted-tailed Quoll Dasyurus maculatus Department of Environment, Land, Water and Planning Prepared by the Victorian

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Koolan Island quoll demographics & genetics

Koolan Island quoll demographics & genetics

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The Reproductive Cycl in the Marsupiae l Dasyurus viverrinus.Dasyurus has only one breeding season1 in the year, which extends over the winter months, i.e. from the end of May to the

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Ecology and management of the northern quoll Dasyurus ... · Ecology and management of the northern quoll Dasyurus hallucatus in the Pilbara. Progress Report Judy Dunlop, Annette

Ecology and management of the northern quoll Dasyurus ... · Ecology and management of the northern quoll Dasyurus hallucatus in the Pilbara. Progress Report Judy Dunlop, Annette

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Drought, disease or devil declines? Identifying the cause ... whole thesis.pdf · Drought, disease or devil declines? Identifying the cause of decline of the eastern quoll, Dasyurus

Drought, disease or devil declines? Identifying the cause ... whole thesis.pdf · Drought, disease or devil declines? Identifying the cause of decline of the eastern quoll, Dasyurus

Documents