Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is...

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Review of the distribution, causes for the decline and recommendations for management of the quokka, Setonix brachyurus (Macropodidae: Marsupialia), an endemic macropodid marsupial from south-west Western Australia PAUL J. DE TORES 1* , MATT W. HAYWARD 2,3,4 , MICHAEL J. DILLON 3 AND ROBERT I. BRAZELL 5 1 Department of Environment and Conservation, Wildlife Research Centre, P.O. Box 51 Wanneroo, Western Australia 6946 2 School of Biological, Earth and Environmental Science, University of New South Wales, Sydney, New South Wales 2052 3 Department of Environment and Conservation, Dwellingup Research Centre, Banksiadale Road, Dwellingup, Western Australia 6213 4 Current address: Mammal Research Institute, Polish Academy of Science, 17-230 Biglowieja, Poland 5 Department of Environment and Conservation, Wellington District, P.O. Box 809 Collie, Western Australia 6225 * Contact person for correspondence: Paul de Tores, email: [email protected] ABSTRACT The former and current distribution of the quokka, Setonix brachyurus, was mapped from published and all available unpublished records. At the time of European settlement the quokka was widespread and abundant and its distribution encompassed an area of approximately 41 200 km 2 of south-west Western Australia inclusive of two offshore islands, Bald Island and Rottnest Island. Historical reports indicated an extensive population decline occurred in the 1930s. The decline continued, with a previously undocumented decline apparent in the period from 1980 to 1992. However, this decline may be an artefact of the time scales used for mapping and may well equate with a previously reported decline for a suite of south-west mammals in the 1970s. By 1992 the quokka’s distribution had been reduced to an area of approximately 17 800 km 2 . An increased awareness of the presence of the quokka on the mainland has resulted in numerous reportings of quokka presence since 1992, has confirmed the existence of several populations at the northern extent of the quokka’s known geographic range and indicated the current, 2005, distribution to be similar to that in 1992. However, survey and population estimates at six of these mainland locations from the northern jarrah forest indicated low abundance. There have been no population estimates elsewhere on the mainland. Two populations have been reported from the Swan Coastal Plain, but neither has been confirmed extant. Predation by the introduced fox, Vulpes vulpes , is implicated as a major cause of the quokka’s initial decline, while ongoing predation, habitat destruction and modification through altered fire regimes have contributed to the continued decline. Specific conservation management actions are recommended, namely: (i) Implementing an active adaptive management program in the northern jarrah forest to determine quokka population response to habitat manipulation through the use of fire, fox baiting and pig control; (ii) Surveying the Stirling Range and Green Range populations with emphasis placed on determining population size and population genetic structure; (iii) Surveying the reported occurrences from the Swan Coastal Plain, with emphasis on unambiguously determining presence. If confirmed, priority should be directed to assessing population size and determining the management requirements to ensure persistence of the population; (iv) Surveying southern forest and south coast populations to assess quokka population size, the extent of movement between subpopulations and assessment of the range of habitat types used by quokkas. The latter should be combined with spatial analyses of known extant populations and suitable and potentially suitable habitat; (v) Determining the role of fire in establishing and maintaining preferred habitat of southern forest and south coast populations; and (vi) Establishing a program to assess the potential effects from management operations. Keywords: quokka, Setonix brachyurus, distribution, fox predation, fire, adaptive management INTRODUCTION The quokka, Setonix brachyurus (Macropodidae: Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west Western Australia and two offshore islands – Conservation Science W. Aust. 6 (1) : 13–73 (2007)

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Page 1: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Review of the distribution, causes for the decline andrecommendations for management of the quokka,Setonix brachyurus (Macropodidae: Marsupialia), anendemic macropodid marsupial from south-westWestern Australia

PAUL J. DE TORES1*, MATT W. HAYWARD2,3,4, MICHAEL J. DILLON3 AND ROBERT I. BRAZELL5

1 Department of Environment and Conservation, Wildlife Research Centre,P.O. Box 51 Wanneroo, Western Australia 6946

2 School of Biological, Earth and Environmental Science, University of New South Wales,Sydney, New South Wales 2052

3 Department of Environment and Conservation, Dwellingup Research Centre,Banksiadale Road, Dwellingup, Western Australia 6213

4 Current address: Mammal Research Institute, Polish Academy of Science,17-230 Biglowieja, Poland

5 Department of Environment and Conservation, Wellington District, P.O. Box 809 Collie, Western Australia 6225* Contact person for correspondence: Paul de Tores, email: [email protected]

ABSTRACT

The former and current distribution of the quokka,Setonix brachyurus, was mapped from published and allavailable unpublished records. At the time of Europeansettlement the quokka was widespread and abundant andits distribution encompassed an area of approximately41 200 km2 of south-west Western Australia inclusive oftwo offshore islands, Bald Island and Rottnest Island.Historical reports indicated an extensive populationdecline occurred in the 1930s. The decline continued,with a previously undocumented decline apparent in theperiod from 1980 to 1992. However, this decline maybe an artefact of the time scales used for mapping andmay well equate with a previously reported decline for asuite of south-west mammals in the 1970s. By 1992 thequokka’s distribution had been reduced to an area ofapproximately 17 800 km2. An increased awareness ofthe presence of the quokka on the mainland has resultedin numerous reportings of quokka presence since 1992,has confirmed the existence of several populations at thenorthern extent of the quokka’s known geographic rangeand indicated the current, 2005, distribution to be similarto that in 1992. However, survey and populationestimates at six of these mainland locations from thenorthern jarrah forest indicated low abundance. Therehave been no population estimates elsewhere on themainland. Two populations have been reported fromthe Swan Coastal Plain, but neither has been confirmedextant.Predation by the introduced fox, Vulpes vulpes, isimplicated as a major cause of the quokka’s initial decline,while ongoing predation, habitat destruction andmodification through altered fire regimes havecontributed to the continued decline.

Specific conservation management actions arerecommended, namely: (i) Implementing an activeadaptive management program in the northern jarrahforest to determine quokka population response tohabitat manipulation through the use of fire, fox baitingand pig control; (ii) Surveying the Stirling Range andGreen Range populations with emphasis placed ondetermining population size and population geneticstructure; (iii) Surveying the reported occurrences fromthe Swan Coastal Plain, with emphasis on unambiguouslydetermining presence. If confirmed, priority should bedirected to assessing population size and determiningthe management requirements to ensure persistence ofthe population; (iv) Surveying southern forest and southcoast populations to assess quokka population size, theextent of movement between subpopulations andassessment of the range of habitat types used by quokkas.The latter should be combined with spatial analyses ofknown extant populations and suitable and potentiallysuitable habitat; (v) Determining the role of fire inestablishing and maintaining preferred habitat of southernforest and south coast populations; and (vi) Establishinga program to assess the potential effects frommanagement operations.

Keywords: quokka, Setonix brachyurus, distribution, foxpredation, fire, adaptive management

INTRODUCTION

The quokka, Setonix brachyurus (Macropodidae:Marsupialia) (Quoy & Gaimard 1830) is a small to mediumsized macropodid marsupial, endemic to the mainland ofsouth-west Western Australia and two offshore islands –

Conservation Science W. Aust. 6 (1) : 13–73 (2007)

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Bald Island, east of Albany and Rottnest Island, 20kmwest of Perth (Fig. 1). Adult males range in weight from2.7 to 4.2 kg and adult females from 1.6 to 3.5kg(Hayward et al. 2003; Kitchener 1995). The quokka isknown to the Aboriginal, or Noongar, people of south-west Western Australia by a range of names including‘Ban-gup’, ‘Bungeup’, ‘Quak –a’ (Gould 1863;Shortridge 1909), ‘kwoka’ and ‘bangop’ (Abbott 2001a).

The quokka was the second Australian marsupialrecorded by Europeans, the first appears to have beenFrancis Pelsart’s description of the Tammar Wallaby fromthe Houtman Abrolhos on 1629. The first Europeanrecord of a quokka is attributed to Samuel Volckertzoon(or Volckersen) who, in 1658, when visiting the thenun-named Rottnest Island off the coast near Perth,described the quokka as ‘a wild cat resembling a civet-cat but with browner hair’ (Alexander 1914). In 1696when Willem de Vlamingh visited the island he describedthe quokka as ‘a kind of rat as big as a common cat’(Alexander 1914). He named the island Rottenest (nowRottnest), meaning rat’s nest. The quokka is oftenerroneously reported by local media as occurring onlyon Rottnest Island. Its presence on the mainland wasdescribed in 1957 as largely unreported (Barker et al.1957) and it is still widely thought of as only occurringon Rottnest Island.

At the time of European settlement of south-westWestern Australia, the mainland distribution of thequokka was thought to extend from the Moore Riverdistrict 80–100 km north of Perth (Baynes 1979), toareas south and south-east of Perth at locations withinthe Swan Coastal Plain, the jarrah, Eucalyptus marginata,and karri, E. diversicolor, forests south-east of Perth, theCape Leeuwin–Cape Naturaliste region, the south coastand east to the Hunter River east of Bremer Bay (Baynes1979; Baynes et al. 1975; Glauert 1933; Kitchener 1995;Ride 1970) (Fig. 1). In addition to the Rottnest Islandand Bald Island populations, quokkas were reported fromBreaksea Island, near Albany (Western AustralianMuseum records), ‘Twin Peak’ Island and other off shoreislands near Esperance (Shortridge 1909). Subsequentreferences to the quokka’s historic distribution on themainland, at or prior to the time of European settlement,have included the south coast of Western Australia as fareast as Esperance (Shortridge 1909; Troughton 1965).

This inferred former distribution was based on theabove reports and location records. All location recordswere within four biogeographical regions, or bioregions,as identified in the Interim Biogeographical Regionalisationfor Australia (IBRA) (Thackway & Cresswell 1995),namely the Swan Coastal Plain, the Jarrah Forest, theWarren and the Esperance Plains bioregions (Fig. 1).

Figure 1. The four biogeographical regions, as defined by the Interim Biogeographical Regionalisation for Australia (IBRA)(Thackway and Cresswell, 1995), within south-west Western Australia which encompass all records of occurrence of the quokka,Setonix brachyurus. Previously unpublished trapping locations are shown as + for sites trapped in the period 1992 to 2002 toconfirm presence and/or to derive an estimate of abundance

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Distribution of the quokka 15

Various authors have expressed concern at the extentof the quokka’s decline from this inferred historicdistribution and attributed different levels of threat to itsconservation status. Clarke (1948) suggested the quokkawas almost confined to Rottnest Island. White (1952),although noting there was a misleading impression ofrarity on the mainland, conceded the quokka wasbecoming less abundant there. Loaring (in Serventy etal. 1954) believed quokkas had vanished from their ‘gullyhaunts’ in the Darling Plateau by the 1920s. Sharman(1954) noted the quokka appeared common only onRottnest and Bald Islands with isolated colonies at somemainland sites.

Barker et al. (1957) cited a 1957 newspaper articlewhich suggested the quokka was extinct on the mainland.That report generated interest in quokka populations,and in the same year, led to the confirmation of thepresence of a small colony of quokkas near Byford,immediately south of the Perth metropolitan area (Barkeret al. 1957). Hart et al. (1986) and similarly Perry (1973)also noted there was a widely held, yet incorrect, beliefthe quokka had become extinct on the mainland afterthe population crash of the 1930s. This collapse of quokkapopulations on the mainland in the 1930s is widelyreported and Ride (1970) recorded, prior to this collapse,that the quokka was common in the south-west andpopulations were sufficiently abundant for quokkashooting to be a ‘familiar sport’.

Serventy (in Serventy et al. 1954) reported anapparent increase in quokka numbers in the DarlingRange and Manjimup areas in the early 1950s. However,this seems to be contradicted by reports from Loaringand Glauert (also reporting in Serventy et al. 1954).Dell (1983) believed Serventy’s (1954) reported increasein numbers of the quokka and other species could notbe substantiated and was an artefact of an increasedawareness by observers. How et al. (1987), whenreporting on a vertebrate fauna survey for the areabetween Busselton and Albany, believed quokkapopulations had persisted on the mainland but wererapidly diminishing.

In an attempt to quantify the conservation status andcauses of decline for a suite of macropod species, Johnsonet al. (1989) concluded the quokka had experienced asubstantial (85–90%) decline in its geographic range onthe mainland. These authors identified the quokka aswarranting a high priority for conservation management.

The quokka is currently listed as a Threatened Species,in the sub-category Vulnerable, pursuant to the WorldConservation Union (IUCN) Red List of ThreatenedSpecies (Hilton-Taylor 2000). It is listed nationally asThreatened Fauna, in the sub-category Vulnerable,pursuant to the Commonwealth of Australia’sEnvironment Protection and Biodiversity Conservation Act1999 (EPBC Act) criteria for Vulnerable. In 1996 itwas placed on the Western Australian list of ‘fauna whichis rare or likely to become extinct’ pursuant to Section14(2)(ba) of the Western Australian WildlifeConservation Act 1950.

This paper reviews the distribution of the quokka,

assesses the probable causes of the species decline andproposes specific conservation managementrecommendations. The conservation status is reviewedin a forthcoming publication.

METHODS

Distribution maps and terminology

Location records for the quokka were obtained frompublished and unpublished accounts of fossil, subfossiland surface cave deposits, published and unpublishedrecords of distribution/presence, database records ofthe Western Australian Museum (WAM), databaserecords of the Australian Museum, database records ofthe Western Australian Department of Environment andConservation (DEC, formerly the Department ofConservation and Land Management, CALM) (CALMunpublished; Gilfillan unpublished). Additionalinformation on WAM records was supplied by NorahCooper (pers. comm. to PJdeT)1 and additional sub-fossil records were obtained from searches of theuncatalogued collection of the Palaeontology andAnthropology Sections of the Western AustralianMuseum (E. Jefferys pers. comm. to MWH)2 .

Records of presence were provided from ourpreviously unpublished field survey and trapping datafrom the early 1970s to 2002 with additional detaileddata from the period 1992 to 2002 from trapping at sixlocations within the northern jarrah forest – GervasseForest Block, Rosella Road, Albany Highway (Travellers’Arms Site), Holyoake Swamp, Kesners Swamp and Lewis(Wild Pig) Swamp (Fig. 1.). Field survey consideredquokkas to be present at a site if confirmed throughtrapping, recent roadkills or by the presence of scatswithin characteristic runways in densely vegetated areasconsidered typical of quokka habitat (see Barker et al.1957; Christensen et al. 1985; Christensen & Kimber1975; Dillon 1993; Kitchener 1995; Maxwell et al. 1996;Sinclair 1999). The presence of scats in open areas andaway from characteristic runways, was consideredinsufficient to infer presence (Alacs et al. 2003).

Locations from all sources were mapped using thegeographic information system (GIS) software ArcGIS(ESRI 1999-2004). Location records were excludedwhen they were unable to be validated and/or if notsupported by location co-ordinates, or if location co-ordinates could not be determined. The inferred pre-European distribution (inclusive of sub-fossil records)was mapped from these records, as was the distributionat six subsequent times, namely at the time of Europeansettlement, at 1920, 1950, 1980, 1992 and 2005. Theinferred distribution at the time of European settlementand at 1920 pre-dates arrival of the fox and the firstevidence of fox predation in south-west Western Australia.The 1950 distribution includes records from 1921 to

1 Norah Cooper: Curator of Mammals, Western Australian Museum, Perth2 Elizabeth Jefferys: University of New South Wales

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1950. The 1980 distribution includes records from 1951to 1980. The 1992 distribution includes records from1981 to 1992. The 2005 distribution is inferred fromrecords post 1992, where populations were verified orwhere there was no evidence to indicate previously knownpopulations had not persisted. These time frames allowquantitative and visual assessment of the decline fromthe pre-European distribution. Our use of the term ‘pre-European distribution’ is synonymous with the term‘original distribution’ as used by Baynes (1987). Thecurrent (2005) distribution was also mapped to depictdensity of sighting and location records. Densitycategories were derived through kernel analysis inArcGIS. Densities reflect the density of sighting/locationrecords and do not necessarily equate with populationdensities.

Areas surveyed, population estimates andsurvivorship

Estimates of population size were obtained frompublished accounts and our previously unpublishedsurvey and trapping records at locations within thenorthern and southern jarrah forests, where the northernjarrah forest is broadly defined as the jarrah, marri andwandoo forests north of the Preston River (Fig. 1). Thesouthern forest is broadly defined as the jarrah, marri,

karri, tingle and wandoo forests south of the PrestonRiver, north and west of the Frankland River. The southcoast refers to the area east of the Frankland River (Fig.1).

The Gervasse site (Fig. 1) is subject to an ongoingmonitoring program (capture-mark-release-recapture)and capture data were analysed for eleven trappingsessions conducted in June/July 1992, March 1993,June/July 1993, January 1994, August 1994, April 1995,January 1996, January 1997, February 1998, March1999 and February 2000. To enable comparison withpopulation estimates from five other northern jarrahforest sites (Hayward et al. 2003), estimates of populationsize from the Gervasse site were derived from the ‘deathbut no immigration’ Jolly-Seber (JS) model using thesoftware interface modified to accommodate unequaltime periods between trapping sessions (Barker &McGlinchy 2001).

‘Known to be alive’ (KTBA) estimates were alsoderived for the Gervasse site, where an animal trappedat trapping session t(n-i), not trapped at t(n) andsubsequently trapped at t(n+i) is assumed to be alive att(n). The number of captures at the other five sitesreported from the northern jarrah forest was too low toderive an estimate of, or index to, population size. Wherepresence was confirmed, data from these sites are reportedas the total number of individuals trapped.

Figure 2. The southwest of Western Australia showing location records for the quokka, Setonix brachyurus. The full source for eachrecord is listed in tables 1 to 8.

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Distribution of the quokka 17

RESULTS

Records of occurrenceFossil and subfossil records are shown in Table 1,Australian Museum records in Table 2, published recordsin Table 3, unpublished records (from various sources)in Table 4, unpublished records from the Departmentof Conservation and Land Management databases, nowDepartment of Environment and Conservation (DEC)(CALM unpublished; Gilfillan unpublished) in Table 5.Our previously unpublished location data for extantpopulations are shown in Tables 6 and 7. All recordswere mapped and shown in Fig. 2. Data from theWestern Australian Museum (Kitchener & Vicker 1981)and additional WAM records (WA Museum databaserecords; pers. comm. from E. Jefferys; pers. comm.from N. Cooper) are presented in mapped form only.Five of the records unable to be validated or unable tobe confirmed are shown in Table 8. These records arealso mapped in Fig. 2, but excluded when inferringdistribution. The population translocated to a predator-proof fenced sanctuary (Karakamia Sanctuary, east ofPerth) is also listed in Table 8 and mapped in Fig. 2,but also excluded from inferred distribution, as is thecaptive colony formerly housed at the University ofWestern Australia and the location of a failedtranslocation – the Harry Waring Marsupial Reserve,Jandakot (Short et al. 1992). Two unconfirmed records

(Muddy Lake, south of Bunbury and a WaterCorporation reserve near Dunsborough) are also listedin Table 8 and mapped in Fig. 2, but excluded frominferred distribution mapping.

Estimates of population size andsurvivorship

The ‘deaths but no immigration’ Jolly-Seber estimate ofpopulation size for Gervasse was 49 ±7.9 and is shownin Table 7, as are results from opportunistic trappingprograms initiated to determine presence at five othersites. Known to be alive (KTBA) estimates for the period1992 to 2000 for the Gervasse population are shown inTable 9.

DISTRIBUTION

Distribution pre European settlement

There is no evidence to suggest the quokka has extendedits distribution since European settlement and allpublished and unpublished accounts imply a progressivecontraction of this distribution. Therefore, distributionpre-European settlement is inferred from current andhistorically known location records and from subfossilrecords and is shown in Fig. 3. The age, i.e. sub-fossil,of the Hasting’s Cave deposits east of Jurien Bay (Table 1and Fig. 2) indicate this to be the northern extent of the

Figure 3. The inferred distribution of the quokka, Setonix brachyurus, pre European settlement and at five subsequent timeperiods, 1920, 1950, 1980, 1992 and 2005. The distribution at all time periods includes Rottnest Island and Bald Island. The brokenline indicates the 700mm rainfall isohyet, the solid line indicates the 1000mm rainfall isohyet.

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quokka’s pre-European distribution, as concluded byBaynes (1979). Sub fossil deposits at Yanchep (Merrilees1965) and the series of deposits from Perth to JurienBay (E. Jefferys pers. comm. to MWH) indicate thisdistribution was continuous. The Hunter River record(Kitchener & Vicker 1981), east of Albany is the easternmost record. Therefore, the inferred pre-Europeanextent of occurrence on the mainland extends from JurienBay, southward through the Swan Coastal Plain, thejarrah and karri forest areas, the Leeuwin-Naturalisteregion, the south coast and east to the Hunter River,east of Albany. This distribution encompassed an areaof approximately 44 300 km2, inclusive of Rottnest Island(15.5 km2) and Bald Island (7.8 km2). This area isexclusive of Breaksea Island, North Twin Peak Islandand South Twin Peak Island and exclusive of the southcoast, east of the Hunter River, considered by Shortridge(1909) to be within the quokka’s original distribution.

The two records from Breaksea Island are from skullscollected by Ian Abbott3 in 1975 (Kitchener & Vicker1981). There have been no further records fromBreaksea Island and the true origin of these skulls isunclear. The skulls may be sub-fossil specimensrepresenting a natural population which subsequentlydied out after it became isolated on the Breaksea peninsulawhen it was separated from the mainland approximately7 000 years ago (Ian Abbott pers. comm. to PJdeT).Alternatively, explanations (as proposed by Ian Abbott,pers. comm. to PJdeT) include: (i) quokkas still occuron Breaksea Island but have been overlooked; (ii) sealersknown to have been operating in the area from the 1790s,and/or Noongar women held captive on the Island,caught quokkas on the mainland and left their remainson Breaksea Island. This is supported by historicalrecords which refer to sealers marooning Noongarwomen on islands; and (iii) residents on the Island,including a lighthouse keeper present there from 1858,or other visitors, attempted and failed to establish aquokka population on the Island.

Recent work examining the effect of weeds on nestingseabirds has involved considerable time traversing allvegetation types on Breaksea Island and has failed todetect any sign of quokka activity or presence and studiesexamining sub-fossil remains of seabirds have notidentified fossil or subfossil quokka remains (PeterCollins, pers. comm. to PJdeT)4 . Although quokkaremains may have been overlooked (Peter Collins, pers.comm. to PJdeT), there seems to be insufficient evidenceto include Breaksea Island within the original distributionof the quokka.

Shortridge (1909) provided the sole reference toquokka presence on ‘Twin Peak Island’, presumablyreferring to either North Twin Peak Island or SouthTwin Peak Island in the Recherche Archipelago, off the

coast at Esperance. Glauert (in Serventy et al. 1954)described the quokka as plentiful on Rottnest Island and‘some islands off the south coast’. However, there are noother records of occurrence of the quokka from theislands off Esperance and numerous authors (Abbott &Burbidge 1995; Burbidge 1977; Calaby 1971; Glauert1933; Maxwell et al. 1996; Sharman 1954; Waring 1956)list the quokka as present on only Rottnest Island andBald Island. Quokka presence was not recorded on anyof the islands in the Recherche group by Serventy(1953), however, the tammar wallaby, Macropus eugenii,was recorded as plentiful on Middle Island and NorthTwin Peak Island. Similarly, Kabay and Start (1976) wereunable to confirm presence of quokkas on either Northor South Twin Peak Island when they surveyed bothislands in 1976. The tammar and quokka do not occursympatrically on any island and various hypotheses havebeen forwarded to explain this (see Clarke 1948; Main1961; Serventy 1951; Serventy 1953). The mostparsimonious explanation for the lack of sympatricoccurrence is that the small to medium size insularmacropod fauna is determined by the climatic conditionsat the time of separation from the mainland, withpersistence determined by continued availability ofsuitable habitat. Where islands are small, inter-specificcompetition operates and one species only will persist.Main (1961) hypothesised an island area in excess of sixsquare miles was necessary to support a single speciesof small macropod. This hypothesis is consistent withthe occurrence of tammars only on Garden Island andquokkas only on Rottnest Island. It is also consistentwith the presence of quokkas only on Bald Island.

Given the size of North and South Twin Peak Islandand the presence of the tammar on North Twin PeakIsland, it is unlikely the quokka also occurred there. The‘Twin Peak Island’ record of Shortridge (1909) thereforeappears to be spurious and we believe the record is areference to the tammar wallaby, presumably from NorthTwin Peak Island. Thus, we have interpreted the pre-European distribution of the quokka to be exclusive ofthe islands of the Recherche and exclusive of the southcoast, east of the Hunter River. The absence of quokkasfrom the fossil and sub fossil records from this region isconsistent with this inferred distribution.

The inferred pre-historic distribution is not differentfrom the inferred distribution, pre-European settlement.Although abundant in the fossil record dating to thePleistocene (Balme et al. 1978) (Table 1), the quokkaappears to have been restricted to the south-west cornerof Australia. The mainland population was presumablysplit by rising sea levels, with the Rottnest Islandpopulation separated between 6 000 and 8 000 yearsbefore present (BP) (Churchill 1959). The Bald Islandpopulation was separated almost 10 000 years BP (Storr1965).

Quokka fossil deposits from Devil’s Lair were datedto 35 000 years BP (Balme et al. 1978), although morerecent techniques suggest an age in excess of 40 000years (Turney et al. 2001) (Table 1). Balme et al. (1978)concluded quokkas became increasingly abundant from

3 Ian Abbott: Science Advisor and Senior Principal Research Scientist,Western Australian Department of Environment and Conservation, ScienceDivision, Kensington

4 Peter Collins: Fauna Conservation Officer, Western Australian Departmentof Environment and Conservation, South Coast Region, Albany

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Distribution of the quokka 19

32 000 years BP, then showed a slow decline, followedby an increase in abundance. The peak in abundancecoincided with a wetter but cooler climate and the periodof decline coincided with a period of intense aridity,relative to present conditions. Despite these variations,the quokka was still one of the most abundant speciesthroughout the Devil’s Lair deposits (Balme et al. 1978).Balme et al. (1978) concluded the persistence of thequokka throughout this period of climatic variationimplies a continuous presence of thickly vegetatedwatercourses and forest, although the forest speciescomposition may have changed. Importantly, the post-glacial period of aridity experienced in other parts ofsouthern Australia was not as severe in the south-westcorner of Australia, thus allowing forest dwellingmammals to persist (Balme et al. 1978).

Distribution at the time of Europeansettlement

The inferred distribution at the time of Europeansettlement is shown in Fig. 3. Published and unpublishedreports (Table 3 and 4) indicate the quokka was locallyabundant and the distribution encompassed an area ofapproximately 41 200 km2. This indicates a relativelyminor southern contraction from the pre-Europeandistribution.

With the exception of records north and north-eastof Albany, including the Stirling Range National Park,the distribution follows the pattern of rainfall, with recordsof occurrence confined to areas now receiving an averageannual rainfall of 700 mm or more (Fig. 3). Althoughthe distribution is shown as continuous, the Stirling Rangeand Green Range populations are isolated from all othersouth coast populations. The Stirling Range populationmay be in an area of higher rainfall (see climaticinfluences, below).

Distribution at 1920

The inferred distribution at 1920 shows no contractionfrom that at the time of European settlement (Fig. 3).

Distribution at 1950

The inferred distribution at 1950 is shown in Fig. 3encompasses an area of occurrence of approximately37 800 km2 and shows a contraction from that at 1920.

Reports of quokka abundance prior to the mid 1930scontrast to later reports (Tables 3 and 4). Prince (1984)included the quokka amongst the species of kangaroosand wallabies commercially exploited in the first 30 yearsof the 20th century. Records of the specific number ofquokkas harvested were not kept and harvests of thequokka and the tammar wallaby were combined withother small macropods. Potential commercial exploitationof the quokka ceased in 1952, with proclamation of theFauna Protection Act of 1950 (Prince 1984). As none

of these harvests was from Rottnest Island (R.I.T. Prince,pers. comm. to PJdeT)5 , the implication is the quokkawas sufficiently abundant on the mainland in the earlypart of the 20th century to enable it to be commerciallyexploited. These harvested animals were in addition tothe large number of quokkas shot in locally organized‘quokka shoots’ which occurred in the 1940s (Les Wilson,pers. comm. to PJdeT)6 and it seems the quokka waswidespread and abundant prior to the 1930s, albeit inrestricted habitat (Gould 1863; Perry 1971; Perry 1973;Shortridge 1909; White 1952). Long-term residents fromthe Northcliffe area (pers. comm. to PJdeT) reportedthat in the 1930s, and perhaps into the 1940s, quokkasregularly browsed in pasture well away from areas ofrestricted creekline habitat. Further evidence indicatingthe quokka was at high density prior to the decline inthe 1930s is provided by the 1933 proclamation of thequokka as ‘vermin’ pursuant to the Vermin Act, 1918(Government Gazette of Western Australia 1933).

The subsequent decline is reported anecdotally only.This decline is also only partially reflected by the declinein inferred distribution at 1950 (Fig. 3) which showsonly a moderate southward contraction from the inferreddistribution at 1920. Populations were still present, orinferred by subsequent ‘rediscoveries’ to have been present,as far north as the Perth metropolitan area. No westwardcontraction in range is apparent and similarly, populationspersisted on the Swan Coastal Plain. Although thedistribution at 1950 shows only a moderate contractionfrom 1920, the number of known locations clearlydeclined and abundance at each also appears to havedeclined. Quokkas may have persisted as far east as theHunter River area, east of Albany. However, supportiveevidence is provided by one WAM registered specimenrecord only, collected in 1970 (Kitchener & Vicker1981). The specimen is a part of a mandible (NorahCooper, pers. comm. to PJdeT), is undated and may beconsiderably older than the collection date implies. Thus,the westward contraction may have commenced priorto the collection date of the Hunter River specimen andthe eastern extent of the distribution at 1950 (and 1980,see below) may have extended no farther than theWaychinicup Bay/Waychinicup National Park and GreenRange area (Fig. 3).

Distribution at 1980

The inferred distribution at 1980 is shown in Fig. 3 andshows no contraction from that at 1950. However, asfor the pattern of decline described at 1950, recordsindicate the number of known populations had decreasedand most published accounts referred to the quokka asrare on the mainland (Table 3).

Distribution at 1992

The inferred distribution at 1992 is shown in Fig. 3encompasses an area of approximately 17 800 km2 and

5 R.I.T. (Bob) Prince: Senior Research Scientist, Western Australian Department of Environment and Conservation, Science Division, Woodvale6 Les Wilson: long-term resident of the Northcliffe area.

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20 P.J. de Torres et al.

shows a marked contraction from the distribution at1980. With the exception of a recently reportedrediscovery at Muddy Lake, south of Bunbury (Dell &Hyder-Griffiths 2002) (Table 4) and an unconfirmedrecord from the Dunsborough area (Table 8), the lastrecords from the Swan Coastal Plain are from 1961 fromBibra Lake, at the southern outskirts of the Perthmetropolitan area (Kitchener & Vicker 1981) and from1975 and 1976 from the Bunbury/Muddy Lake area(Table 4) (Hart et al. 1986; Kitchener & Vicker 1981).The last confirmed record from the Cape Leeuwin–CapeNaturaliste area is from 1979 from Wardanup Hill, southof Dunsborough (Kitchener & Vicker 1981).

Populations were present, or inferred as still presentby subsequent ‘rediscoveries’, at Churchman Forest Block,within 10 km of the Perth metropolitan area. Exclusiveof Bald Island, the eastern most records are from theWaychinicup National Park area east of Albany and GreenRange, northeast of Albany. The Stirling Rangepopulation(s) is shown as isolated from other south coastpopulations and may have been isolated prior to 1992.

Current distribution

The inferred current distribution is shown in Figs 3 and4 and is unchanged from 1992.

Trapping in 1995–96 in the Dwellingup andManjimup areas at sites known to support quokkapopulations in the 1990s failed to confirm presence atnumerous sites (Dillon 1996) (Table 10). With theexception of the unconfirmed records from Muddy Lakeand Dunsborough, quokkas are now considered absentfrom the Swan Coastal Plain. The skull collected at MuddyLake (Dell & Hyder-Griffiths 2002) in September 2002has no associated soft tissue and is not aged (NorahCooper, pers. comm. to PJdeT). Its collection date doesnot necessarily infer presence of an extant population atthe time of collection. Quokka presence is otherwiseonly inferred at this location by the presence of runways.There is an unconfirmed report of quokka occurrencefrom a Water Corporation managed reserve in theDunsborough area and unsubstantiated anecdotal reportsof presence in the Cape Leeuwin–Cape Naturaliste area.

Figure 4. The inferred current (2005) distribution of the quokka, Setonix brachyurus, showing density of location / sightingrecords. The solid line indicates the 1000mm rainfall isohyet.

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Distribution of the quokka 21

Presence of suitable habitat and anecdotal reports suggestquokkas may also still occur in the lower catchment ofthe Blackwood River, north-east of Augusta (RogerHearn, pers. comm. to PJdeT)7 . Data unavailable forthis review and referred to by Liddelow (2006) indicatedseveral populations may occur west of Nannup. Thecurrent inferred distribution (Fig. 3) includes theserecords, however the pattern of these records ofoccurrence combined with sites also examined byLiddelow (2006) where quokkas were previously knownto occur and are considered to no longer occur, suggestthis western forest margin may be experiencing the samedecline detected in the northern jarrah forest.

The most northerly records from the Darling Plateauare from Churchman Forest Block, approximately10 km south-east of the Perth metropolitan area (Table4). The confirmed sites from the northern jarrah forestextend in a narrow band along the Darling Plateau andare bounded by the 1 000 mm annual rainfall isohyet(Fig. 3). There are no confirmed records of extantpopulations between Collie (Davis Forest Block) andNannup (Boronia, Gregory and Helms forest blocks)(Table 6, Fig. 3). With the exception of the recordsfrom Boronia Forest Block and records east of Nannup(Liddelow 2006), the Nannup population(s) and forestpopulations farther south are also almost entirely withinthe 1 000 mm annual rainfall isohyet (Fig. 3). The mosteasterly mainland records are from the south coastnear Waychinicup National Park, east of Albany, frompresence inferred by collection of hair from GreenRange north-north-east of Albany (J.A. Friend, pers.comm. to PJdeT)8 and from the Stirling RangeNational Park, northeast of Albany. As depicted bythe inferred distribution at 1992, the Stirling Rangeand Green Range population(s) appear to be, and mayalways have been, isolated from other south coastpopulations.

Figure 4 shows the current inferred distribution basedon records of occurrence from 1992 to present. Thedensity gradient depicted (Fig. 4) reflects the density ofsighting/location records. It does not necessarily reflectpopulation densities and the nodes of inferred highpopulation density are, to some extent, likely to be anartefact of proximity to townsites and local interest inreporting sightings.

DISCUSSION

Distribution and population estimates

The contraction of distribution has been greatest fromthe northern extent of the geographic range and fromthe Swan Coastal Plain. The northern-most known extantpopulation is approximately 10 km south-east of Perth.With the exception of the unconfirmed records from

Muddy Lake and the Water Corporation reserve nearDunsborough, the quokka is now considered absent fromthe Swan Coastal Plain and the Cape Leeuwin–CapeNaturaliste area and occurs only as far east asWaychinicup National Park (Figs 2 and 3).

Surveys aimed at determining presence in forest areassince the early 1970s have confirmed quokka populationsfrom the northern (Dwellingup) and southern(Manjimup) forest areas are patchy and discontinuous.The known northern limit of the present distributionwas extended in 1995 by discovery of a population nearJarrahdale (Rosella Road) (Tables 6 and 7), in 1996 byre-discovery of a population north-east of Jarrahdale(Albany Highway/Travellers’ Arms) (Tables 6 and 7)and in 2001 and 2002 by discovery of populations withinChurchman Forest Block, 10 km south-east of the Perthmetropolitan area (John Liddington, pers. comm. toPJdeT, Table 4)9 . Although there are in excess of 40records of occurrence in the northern jarrah forest post1992 (Tables 3, 4, 5, 6 and 7), several of these recordsof occurrence may constitute a single population. Equallyimportantly, confirmation of presence does not equatewith persistence of a population. Figure 4 shouldtherefore be interpreted cautiously, as each sighting/location record does not necessarily reflect a separatepopulation and the density gradient depicted reflects thedensity of location/sighting records, not populationdensity. Although the nodes which depict highest densityin the southern forest, near Manjimup, and the southcoast, near Walpole-Nornalup National Park, concur withour observational and unquantified survey data whichsuggest these areas support the highest densitypopulations (exclusive of Rottenest Island), these nodesalso reflect proximity to townsites or areas where thereis local interest in reporting sightings. Therefore, thehigh density nodes are also likely to reflect the level oflocal survey effort and the level of local interest inreporting sighting records. They should not be seen asquantified estimates of abundance.

Estimates of abundance were available from six sitesonly, rarely reported more than twenty to thirty individualsand often comprised considerably fewer (Table 3 and7). The Gervasse population is the only knownpopulation shown to support in excess of forty individuals(Table 7). The northern-most sites appear to be atcritically low density. The Rosella Road population isthought to be at risk of local extinction (Hayward et al.2003). The Wild Pig Swamp (Lewis Swamp) sitesupported quokkas in the early 1990s (Dillon 1993),however despite the evidence of fresh activity, trappingfailed to confirm quokka presence when trapped in 1995(de Tores, Dillon, Tomkinson and Buehrig, Tables 6and 7) and in 1998–2000 (Hayward et al. 2003). TheHolyoake site is thought to be locally extinct (our dataand Hayward et al. 2003).

Results from published accounts (Christensen et al.

7 Roger Hearn: Regional Ecologist, Western Australian Department of Environment and Conservation, Manjimup8 J.A. Friend: Principal Research Scientist, Western Australian Department of Environment and Conservation, Science Division, Albany9 John Liddington: Water Authority of Western Australia, Armadale

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22 P.J. de Torres et al.

1985) and our trapping surveys (Tables 6 and 7) haveconsistently returned low capture rates from these forestareas. Capture rates of 0.3 (Dillon 1993) and 0.07 to0.99 (Hayward et al. 2003) quokkas per 100 trap nightsfor sites in the Dwellingup area and 1.2 per 100 trapnights over three sites north of Dwellingup (Sinclair1999), are representative of trapping success at theselow density sites.

Hayward et al. (2003) believed the northern jarrahforest population formerly constituted a functionalmetapopulation which is now under threat of collapse.Collapse of this metapopulation, or collapse of theremaining catchment-confined populations, would resultin a considerable contraction of the quokkas’ distribution.Given the lag time required to detect a decline in apopulation, we believe there is an urgent need for acoordinated and site-prioritised monitoring program atknown quokka sites and appropriate quantitative analysesof monitoring results (see section on managementrecommendations).

An additional series of surveys has been conductedpost collation of the data reported here (GraemeLiddelow10 and Alan Wright11 , pers. com. to PJdeT)and indicate presence of quokkas at numerous additionallocations within the southern forest and northern jarrahforest. However, no data were collected on populationsize or movement between populations. The likelihoodof persistence of these populations is unknown.

Habitat use

Hayward (unpublished, reported in summary in de Toreset al. 2004) concluded all extant northern jarrah forestpopulations were associated with creeklines characterisedand dominated by the ti-tree, Taxandria linearifolia(formerly Agonis linearifolia), and further concludedquokka populations from the northern jarrah forest arerestricted to areas supporting a structural mosaic of burntand unburnt vegetation. These findings concur withrecords from the mid 1970s to 2002 (our records,various unpublished accounts and confirmed sightings).

Hayward (unpublished, reported in summary in deTores et al. 2004) also examined 66 sites in the northernjarrah forest to assess quokka presence/absence. GeneralLinear Modeling (GLM) and model selection throughuse of a mixed approach of stepwise removal of variablesand the Information-Theoretic approach and AkaikeInformation Criterion (AIC) (Burnham & Anderson2002) was used to identify the ‘best’ model to describethe preferred habitat of the quokka. We do not advocatethis mixed approach and recommend use of theInformation-Theoretic approach and selection from aset of a priori candidate models for any further analyses.

From the mixed approach, three models equally well

described the preferred habitat. The explanatory variablesof the preferred models were:(i) the number of 1080 meat baits delivered per hectare

(an increased number of baits was correlated withquokka presence);

(ii) the average age of the swamp in terms of yearssince last burn (there was a positive correlationbetween years since last burn and quokka presence);

(iii) a habitat factor score (NJF4) (characterised bypossessing large areas of Taxandria swamp burntfive to nine years previously – positively correlatedwith quokka presence);

(iv) a habitat factor score (NJF2) (characterized by ahigh proportion of jarrah – marri open forest andTaxandria swamp burnt 15–19 years previously –this variable was negatively correlated with quokkapresence); and

(v) increasing distance to anthropogenic disturbance.Hayward (unpublished, reported in summary in de

Tores et al. 2004) concluded these features highlightthe importance of a mosaic of age classes within theswamp. This mosaic needs to support early (< 10 yearspost-fire) and late (long unburnt) seral stages. Theintermediate seral stage (15–19 years post fire) is avoided.Therefore, this is a mosaic of specific age classes (youngand old) rather than simply a mosaic of mixed age classes.We recommend adopting an active adaptive managementprogram to determine whether conservationmanagement of the quokka can be improved, throughthe use of fire, to create and maintain this preferredmosaic (see management recommendations section).

The relative importance to the quokka of plant speciesadapted to different fire intervals also needs to beassessed. For example, species such as Taxandrialinearifolia will rapidly generate from root stocks andprovide a dense cover within one year post a spring fire(Kimber 1974), whereas many other species fromriparian zones generate from seed and are adapted tolonger intervals between fire (Abbott 1999; Burrows &Wardell-Johnson 2003). Potential conflict with otherconservation management priorities must also beaddressed. For example, riparian systems supportingquokka populations may also support fire sensitiverelictual plant taxa and communities (Burrows & Wardell-Johnson 2003). Therefore, we caution that any imposedfire regime and adaptive management approach shouldbe supported by an appropriate level of monitoring ofall species potentially affected by the regime.

Although present in the northern jarrah forest, thehabitat occupied there suggests the quokka is not aforest-dwelling species, or more specifically, is not aspecies of the more open northern jarrah forest. Historicaccounts of the appearance of the south-west forestsand the pattern of burning by Noongar people innorthern jarrah forest have been interpreted by Hallam(1975) as indicating the forests ‘comprised tall, straight,mature trees, all frequently scorched but clear of undergrowthand easy to move through’. If the forest structure was asdescribed by Hallam (1975), it would not have been

10 Graeme Liddelow: Senior Technical Officer, Western Australian Departmentof Environment and Conservation, Science Division, Manjimup

11 Alan Wright: Nature Conservation Officer, Western Australian Departmentof Environment and Conservation, Swan Region, Jarrahdale

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Distribution of the quokka 23

conducive to a contiguous distribution of a speciesdependent upon swamp and creekline vegetation andmay have restricted the quokka to discrete catchments,with minimal dispersal and movement of individualsbetween catchments.

In southern forest areas, populations appear to beless discrete and the quokka inhabits a broader range ofhabitats including ti-tree thickets in the upper reaches ofcreek systems, dense streamside beds of rushes, karriregrowth, ridges supporting karri and tingle, Eucalyptusguilfoylei and E. jacksonii, forests (Christensen et al.1985) and occurs widely within forest areas supportingan understorey of spreading sword-sedge, Lepidospermaeffusum, and Anarthria scabra (Greg Freebury, pers.comm. to PJdeT)12 . Populations from the south coastalso appear to be less discrete than the northern jarrahforest populations and occur from Walpole-NornalupNational Park to the Mt Manypeaks area, northeast ofAlbany (Fig. 3). The extent of mixing (dispersal,immigration and emigration) between sub-populationsfrom the southern forest and the extent of mixingbetween sub-populations from the south coast areas isnot known.

Population genetics – the northern jarrahforest metapopulation

Hayward et al. (2003) hypothesised the northern jarrahforest formerly supported a metapopulation which is nowin a state of ‘terminal collapse’. Conversely, results froma study on the genetic structure (Alacs 2001) of thesame populations revealed these northern jarrah forestpopulations showed no evidence of mixing. The latterstudy suggests these populations may never haveconstituted a single metapopulation. However, the twofindings are not mutually exclusive. Alacs (2001) andHayward et al. (2003) examined the same geographicallyseparated populations (different catchments) from withinthe northern jarrah forest. We hypothesise there mayhave been mixing (gene flow) within catchments, butlittle or no mixing between catchments. Each catchmentmay have constituted a functional metapopulation.

Historical evidence from the Swan Coastal Plain isconsistent with this hypothesis and suggests thepopulations from the Coastal Plain were discrete andconfined to individual swamps and/or catchments orsub-catchments. We further hypothesise the populationsfrom the northern jarrah forest, although confined tocatchments or sub-catchments, mixed with populationsfrom the same catchment on the Swan Coastal Plain.Hence metapopulations may have existed, but under thisscenario, each catchment or subcatchment would havesupported a metapopulation. With progressivefragmentation of suitable habitat in the jarrah forest andon the coastal plain (draining of swamps and increaseddistances between suitable habitat patches), movement

between habitat fragments would have been less frequentand eventually ceased.

Factors influencing decline of the quokka

Numerous studies have attempted to determine the causesof the widespread decline in Australia’s mammalian faunasince European settlement (see for example Burbidge& McKenzie 1989; Calaby & Grigg 1989; Calver &Dell 1998a; Calver & Dell 1998b; Morton 1990; Recher& Lim 1990; Short et al. 2002; Short & Calaby 2001;Smith & Quinn 1996; Wilson & Friend 1999). Thesereviews have been selective and addressed some, butnot all of the plausible causes for decline. In acomprehensive analysis, Abbott (2001b) reviewed therole of fifteen possible causes for the decline of the bilby,Macrotis lagotis. His conclusion concurred with Watts(1969) i.e. that the fox was ‘the necessary and sufficientfactor associated with regional declines’ of the bilby.

Different studies have attributed the decline indistribution and abundance of the quokka to specific,but varying causes. A marked decline of the quokkawas recorded during the 1930s (White 1952). Christensen(1978; Christensen 1980b) presented evidence for adecline in a range of mammal species from south-westWestern Australia in the period from 1973 to 1978 andbelieved this decline was the result of an increased levelof predation by foxes. Christensen (1978) furthersuggested this increase in fox predation coincided with acessation of widespread 1080 baiting for rabbits, whichin turn led to a reduction of secondary poisoning offoxes. King et al. (1981) concurred with this belief. Thephenomenon of secondary poisoning of foxes wassubsequently demonstrated by Algar and Kinnear (1996).Our data suggest a period of marked decline in the extentof occurrence between 1980 and 1992 (Fig. 3).However, this period of decline may be an artefact ofthe time periods used for mapping. The mapped declinemay reflect the decline identified by Christensen (1978).

A widespread decline of many species, particularly inmore arid areas, was recorded in the 1880s (Shortridge1909). However, Shortridge (1909) also noted themammals of the south-west, to as far north as the MooreRiver, had not ‘disappeared in the same extraordinaryway’. Burbidge and McKenzie (1989) attributed themodern (post 1900) decline of Western Australia’s non-volant terrestrial, critical weight range (35 g–5.5 kg)mammalian fauna to environmental changes sinceEuropean settlement. They believed these changesreduced available productivity by diverting resources tohumans and introduced species and reduced vegetativecover through grazing and altered fire regimes. Criticalweight range mammals suffered the greatest declinesdue to their limited mobility and relatively high metabolicrequirements (Burbidge & McKenzie 1989). Theyfurther identified factors likely to ameliorate susceptibilityto decline and believed critical weight range mammalspecies with the ability to use, or with a requirement forrockpiles were afforded additional protection. Kinnearet al. (1988) had previously identified the predation

12 Greg Freebury: Operations Officer – Nature Conservation, WesternAustralian Department of Environment and Conservation, South CoastRegion, Albany

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24 P.J. de Torres et al.

refuge value of rockpiles and subsequently (Kinnear etal. 1998) provided more empirical evidence for this.Burbidge and McKenzie (1989) believed species usingburrows, sheltering in hollow logs or to some extent,those with an arboreal habit, would also be affordedprotection from predation. We believe the quokka, witha requirement for a vegetation mosaic which includesareas difficult to penetrate, represents a further groupof critical weight range mammal fauna benefiting fromuse of a predation refuge, analogous to the rock wallabyuse of rockpiles. However, we caution that althoughrefuges may provide temporal respite from predation,long-term persistence of predation may still lead to localand regional extinctions, despite the presence of refuges.

Burbidge and McKenzie (1989) claimed the arid zonefauna were most at risk and fauna from the mesic zonewere somewhat buffered from the changes which resultedin reduced resource availability. The more mesic south-west Western Australian environment may have bufferedthe fauna from this initial decline. Recent researchsuggests fauna from mesic areas are no longer secure(Woinarski et al. 2001). Mesic areas may have moreavailable environmental productivity and may toleratelonger periods of the various disturbances responsiblefor the decline observed in arid areas. However,continued depletion of environmental resources and areduction in refugia may be contributing to a moremodern decline. The decline of the quokka is consistentwith this pattern, with the distribution contracting fromthe more arid northern and west coast areas, whilepopulations from the more mesic southern forest andsouth coast areas have persisted (Figs 3 and 4).

Predation by the dingo, European red fox andferal cat and interaction with other species

Three eutherian predators, the dingo, Canis lupus dingo,the European red fox, Vulpes vulpes, and the cat, Feliscatus, have been introduced to Australia since the arrivalof humans. These eutherian predators acted on apredator-naïve marsupial prey which had not interactedwith medium-sized cursorial predators for the preceding20 000 to 30 000 years (Johnson et al. 1989).

The Dingo

The dingo was introduced to Australia by Asian seafarersapproximately 3 500 to 4 000 years ago (Corbett 1995b).Where present, the primary prey of the dingo is usuallymedium sized mammals (Corbett 1995a) and largemacropods (Corbett & Newsome 1987). However,dingoes will prey upon a wide range of species (Corbett& Newsome 1987; Newsome et al. 1983; Robertshaw& Harden 1985; Thomson 1992; Whitehouse 1977)and the preferred prey in any particular area appears tobe determined not only by prey abundance, but by preyavailability (Corbett 1995a; Corbett & Newsome 1987;Vernes et al. 2001). In north-west Western Australia,Thomson (1992) found large prey items, mostlykangaroos, featured prominently, with smaller prey items

taken less often. Taking smaller prey was associated witha breakdown of dingo packs and an increase in thenumber of solitary dingoes (Thomson 1992).

In south-west Western Australia, the quokka wascommon prior to the 1930s and may have been anoccasional prey item for the dingo, although the quokka’spreference for densely-vegetated habitat may have offeredrefuge from predation. However, the established presenceof the dingo within the region prior to the modern declineof the quokka, implies dingo predation was not the majorcause of that decline. Our own observations indicatedingoes are now exceedingly rare in the jarrah forest andare thus unlikely to be generating predation pressure ofany consequence on quokka populations. Historically, thedingo is believed to have had little major impact on themajority of Australian fauna with the exception of theThylacine, Thylacinus cynocephalus, and the Tasmaniandevil, Sarcophilus harrisii, (see Corbett 1995b; Jones 1995;Rounsevell & Mooney 1995).

The impact of the dingo on Australian native faunamay have been minimised through competitive inhibitionby the sympatric mesopredator, humans. Similarly,dingoes may exhibit an indirect effect as a mesopredator(Newsome 2001). Dingoes are thought to prey on theEuropean red fox and the feral cat where they occursympatrically in the Nullarbor region of Western Australia(Marsack & Campbell 1990), however Marsack andCampbell (1990) implied the evidence for this wascircumstantial. Thomson (1992), as part of an extensivestudy in north-west Western Australia, found the feralcat was present as a prey item only once, and the foxwas not present at all in the stomach contents from asample of 95 dingoes. Newsome et al. (1983) recordedthe fox and the cat present in 1.6 and 0.3 percentrespectively from the stomach contents of 530 dingoesin a trapping and dietary analysis study in south-easternAustralia. The lack of any other comprehensive datasupporting the assertion of intraguild predation suggestsdingoes are more likely to competitively inhibit foxesand cats than prey directly on them.

The European red fox

The red fox was introduced to eastern Australia in thelate 1860s to 1870s (Rolls 1969; Troughton 1965) andhad spread to the south-west of Western Australia bythe early 1930s (Gooding 1955; Jarman 1986; King &Smith 1985; Long 1988). The arrival of the fox in south-west Western Australia coincides with the reporteddecline of the quokka (White 1952) and numerous smallto medium size terrestrial native mammals from mainlandAustralia (Burbidge & McKenzie 1989; Daubney et al.undated; Friend 1990; Jenkins 1974; Richards & Short1996). Numerous authors have suggested the fox wasresponsible for this fauna decline, see for example thereview by Abbott (2001b). Although it is only relativelyrecently that the fox has been empirically linked to thesuppression of native mammal populations (see Kinnearet al. 1988; Kinnear et al. 1998), it seems very likelyfoxes were responsible for the initial decline of the quokka

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Distribution of the quokka 25

on the mainland and have contributed to its continueddecline. The abundance of quokkas on Rottnest Islandin the absence of foxes (Dunnet 1963; Holsworth 1964;Holsworth 1967; Kitchener 1970; Shield 1959) isconsistent with this hypothesis. Further evidence of theeffect of foxes on quokkas comes from a series ofreintroductions, and ultimate failure, of nearly 700Rottnest Island quokkas between 1972 and 1983 to afenced, but not fox-proof, enclosure at Jandakot, nearPerth (Short et al. 1992) (Table 8). By 1988 only ninequokkas remained in the 254 ha enclosure. The lowsurvivorship was attributed to predation by foxes andferal cats (Short et al. 1992).

The fox is now generally accepted as a significantpredator of medium size terrestrial native Australianmammals. Recognition has been at a national and statelevel, as demonstrated by the Commonwealth ofAustralia’s Threat Abatement Plan for Predation by theEuropean Red Fox (Biodiversity Group EnvironmentAustralia 1999) and two recently published state plansfor control of the red fox – the Victorian PestManagement. A framework for Action. Fox ManagementStrategy (DNRE 2002) and the New South Wales ThreatAbatement Plan for Predation by the Red Fox (Vulpesvulpes) (NSW NPWS 2001). Fauna managementprograms in Western Australia have recognised theimportance of fox control and the Western AustralianDepartment of Environment and Conservation (DEC)has implemented a broad scale fox control and faunarecovery program, Western Shield. The program isfocused on recovery of threatened fauna througheffective fox control and translocation of threatened faunaspecies in the presence of fox control.

The predecessor to Western Shield was OperationFoxglove, a large scale experimental 1080 baitingprogram within the northern jarrah forest of south-westWestern Australia (de Tores 1999). The areas baitedwithin the 550 000 ha study area of Operation Foxglove,combined with local baiting at the Gervasse site, includeor abutted all but one of the known extant occurrencesof the quokka in the northern jarrah forest. The Foxglovebaiting program commenced in 1994 and, despite thefact that quokka populations have not shown an increasein abundance since this baiting program commenced(Hayward et al. 2003), we hypothesise the quokka haspersisted in the northern jarrah forest as a result of thisbaiting and further believe effective long termconservation of the quokka requires implementing andmaintaining fox control programs (see managementrecommendations).

The feral cat

Aborigines from the desert regions of central Australiaand central Western Australia believe cats to have eitheralways been present or to have arrived ‘from the west’(Burbidge & McKenzie 1989). Although far fromclaiming the assertion as definitive, Gaynor (2000)believed this arrival from the west may have been a resultof cats escaping from Dutch shipwrecks off the west

coast of Australia. Dickman (1996) noted cats couldhave been introduced to north-western Australia as earlyas the 16th Century. However, Abbott (2002), from adetailed search of historical records, concluded cats werenot present on mainland Australia prior to Europeansettlement. Abbott (2002) presented a compellingargument and conceptual model to support his beliefthat cats spread from multiple points of introduction atcoastal locations in the period 1824 to 1886. Abbott(2002) and others before him (see Morton 1990) alsoargued there is insufficient evidence to support theassertion that cats were responsible for any mammalextinctions from arid Australia. However, there is generalconsensus in the literature (see Dickman 1996) tosuggest the cat has contributed to the decline of manyspecies. Although there is some empirical evidencecorrelating the presence of cats with extinctions of criticalweight range mammals from islands (Burbidge & Manly2002) and empirical evidence indicating predation byferal cats led to reduced population sizes of small nativevertebrates (Risbey et al. 2000), we caution againstinferring a causal relationship between cat predation andfauna extinctions on the mainland generally, and betweencat predation and quokka decline specifically.

Abbott (2002), citing Catling , Coman (1991) andDickman (1996) noted the optimal prey size for catswas approximately 200g. The weight of adult quokkasranges from 1.6 to 3.5kg for females and 2.7 to 4.2 kgfor males (Hayward et al. 2003; Kitchener 1995), whichsuggests if cats prey on quokkas, predation may berestricted to juveniles, or will favour predation ofjuveniles. Circumstantial evidence suggests feral cats preyon the young of bridled nailtail wallabies, Onychogaleafraenata, (Horsup & Evans 1993) and the brush-tailedrock-wallaby, Petrogale penicillata, (Short et al. 1992).More convincing evidence was presented implicating thecat as a significant predator of juvenile and adult alliedrock-wallabies, Petrogale assimilis, (Spencer 1991). Allthree species are larger than the quokka (Eldridge &Close 1995; Evans & Gordon 1995; Prince 1995),suggesting cats may well prey on juvenile and/or adultquokkas.

Cats have also been identified as predators of severalspecies of Australian mammals with adult body weightsgreater than 200 g. For example, cats were responsiblefor between 25 and 32% of predation events of adultwoylies, Bettongia penicillata, at translocation release sitesin the northern jarrah forest (de Tores 1999) andresponsible for predation of adult brushtail possums,Trichosurus vulpecula, in the same study (de Tores,Himbeck, MacArthur, Maxwell, White and Rosier,unpublished radio telemetry data). Cats have also beenreported as preying on adult rufous hare-wallabies,Lagorchestes hirsutus, (Gibson et al. 1994), the burrowingbettong, Bettongia lesueur, (Christensen & Burrows1994), the numbat, Myrmecobius fasciatus, (Friend &Thomas 1994) and the western ringtail possum (de Tores2005). Short et al. (1992) concluded cat predationcontributed to the failure of a translocation of the bandedhare-wallaby, Lagostrophus fasciatus. The above suggests

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26 P.J. de Torres et al.

cats prey on a suite of native mammal fauna, inclusive ofspecies larger than 200g and have the potential to preyon quokkas. Short et al. (1992) further concluded directpredation by foxes and cats was a more plausibleexplanation, than overgrazing by rabbits and othermacropods, for the failure of the quokka translocationsto Jandakot in the 1970s.

However, the failure of the Jandakot translocationsand the suspicion that cat predation contributed to thosefailures are insufficient to conclude cat predation wasresponsible for the initial and continued decline of quokkapopulations on the mainland. Equally confounding is theco-existence of cats and quokkas on Rottnest Island withno discernable reduction in quokka abundance. Similarlyconfounding is the presence of cats in Tasmania with nodiscernable reduction in quokka-size prey species.

Molsher (1999) concluded, from an experimentalstudy at Lake Burrendong, New South Wales,interspecific competition between foxes and cats was themost likely mechanism limiting feral cats. Interspecificcompetition between cats and foxes was also thought tobe a possible explanation for the observed increase incat numbers when fox density was experimentally reducedat Heirisson Prong, north-west Western Australia (Risbeyet al. 2000). Such a response (mesopredator release) isa potential outcome of DEC’s large-scale fox controland fauna recovery program, Western Shield. However,the lack of records of cat predation from Rottnest Islandand the absence of evidence to support the assertionthat the cat was responsible for other mammal extinctions(Abbott 2002), suggests cats are unlikely to be solely orprimarily responsible for the widespread decline of thequokka.

Other predators and interactions

Other predators are unlikely to have been responsible forthe decline in quokka populations. Nocturnal birds of preyare reported to have been responsible for fossil depositsfrom several cave sites (see Table 1 – owl accumulatedfossil deposits). Two owl species from the south-west ofWestern Australia, the barking owl, Ninox connivens, andthe masked owl, Tyto novaehollandiae, have been recordedtaking prey as large as young rabbits (Schodde &Tidemann 1982), so could presumably prey on juvenileor sub-adult quokkas. Although Johnstone (cited as apers. comm. by Abbott 1999) regarded the barking owlas a species favouring swamps and edges of rivers, it isnot a forest species (Abbott 1999) and was not recordedat any of the seventy forest sites surveyed by Liddelow etal. (2002). The masked owl is infrequently recorded inforest (Abbott 1999) and is more common in woodlandor at the interface of agricultural land and forest (Liddelowet al. 2002). Therefore, neither species is likely to havebeen responsible for the decline of the quokka, particularlywithin forest areas. The presence of quokka bonesrecovered from a wedge-tail eagle’s eyrie on Bald Island(Storr 1965) indicates the quokka may be an occasionalprey item of the wedge-tail eagle. However, we cautionagainst inferring presence of a prey species in the diet of a

predator equates with a predation induced decline of thedetected prey species. The quokka’s diurnal use of denselyvegetated swamps within forest areas on the mainland(Christensen et al. 1985; Storr 1964b; White 1952), useof Spreading Sword-sedge, Lepidosperma effusum, andAnarthria scabra habitat within other forest areas anduse of heath habitat on the south coast further minimisesthe risk of predation by eagles.

Large predatory snakes, e.g. the carpet python,Morelia spilota, and to a lesser extent goannas, Varanusspp., are able to prey upon species such as the southernbrown bandicoot, Isoodon obesulus, the common brushtailpossum, Trichosurus vulpecula , the brush-tailed bettong,Bettongia penicillata, (de Tores, Himbeck, MacArthur,Maxwell, White and Rosier, unpublished radio telemetrydata), the western ringtail possum, Pseudocheirusoccidentalis, (de Tores 2005) and the tammar wallaby,Macropus eugenii, (David Pearson, pers. comm. toMWH)13 and may also pose a predation threat to sub-adult quokkas. However, these predators have co-existedwith the quokka in the south-west of Western Australiathroughout the quokkas existence and there is noevidence to suggest they initiated the decline in quokkaabundance. The co-existence of high density populationsof the carpet python and the tammar wallaby on GardenIsland is consistent with this hypothesis.

The feral pig, Sus scrofa, an introduced opportunisticomnivore, is present in forest areas of south-west WesternAustralia and most climatic regions in Australia (Pavlov1995). In addition to the commercial health risks posed,where investigated, pigs have been found to pose asignificant environmental and management problem(Pavlov et al. 1992). Ecological damage associated withpig activity (rooting) in wet tropical forests in Queenslandwas shown to vary with forest type, with wet sclerophyllforest sites showing the greatest disturbance (Laurance& Harrington 1997). Our observations (MJD inparticular) have confirmed the presence of pigs atnumerous quokka sites and, although not supported byany quantitative data, there appears to be a trend ofincreasing occurrence of pig activity and pig abundanceat quokka sites in the south-west forests (MJD, personalobservations; Graeme Liddelow, pers. comm. to PJdeT).Non-target captures of quokkas at sites trapped for pigcontrol purposes (see records by Liddington and Stainesin Table 4 for the years 2001 and 2002) may indicate anoverlap of quokka and pig preferred habitat within thesouth-west forests. The relatively sedentary nature ofpigs (Caley 1997; Saunders & Kay 1996) and thepotential for pigs to significantly disturb quokka habitatby creating large openings and easier access for foxes,may pose an additional threat to the conservation statusof the quokka.

Climatic influencesThe mainland quokka populations appear to have beenhistorically limited to areas of the south-west of Western

13 David Pearson: Principal Research Scientist, Western AustralianDepartment of Environment and Conservation, Science Division, Woodvale

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Distribution of the quokka 27

Australia with an annual average rainfall in excess of700 mm. The current distribution of the quokka closelyfollows the limit of the 1 000 mm rainfall isohyet and,for all but the eastern extent of the distribution near theStirling Range National Park, appears to be limited toareas with an annual average rainfall of 700 mm. or more(Figs 3 and 4). This may reflect the quokka’s relativelyhigh water requirements (Main & Yadav 1971).

Populations in the vicinity of Stirling Range and GreenRange appear to be isolated from all other south coastpopulations and also appear to be within a lower rainfallzone (Figs 3 and 4). The locations of rainfall recordingstations provided by the Australian Bureau ofMeteorology (BoM) indicate stations are located to thenorth and south of the Stirling Range, with no stationsin the immediate vicinity of the known extant quokkapopulations. The Stirling Range may be in a pocket ofhigher, orographically generated rainfall, not detectedby the BoM rainfall recording stations and not reflectedin the pattern of rainfall isohyets. This hypothesis issupported by the presence of an isolated population ofthe red flowering gum, Corymbia ficifolia, (Brown et al.1998) and the occurrence several wet region bird species(Ian Abbott, pers. comm. to PJdeT) in the StirlingRange. The Corymbia ficifolia population is 100 kmnorth of its main distribution and the species is thoughtto be a relic from a past wetter climate (Brown et al.1998). Further evidence of this orographic effect wasprovided by Courtney (1993) who prepared a rainfallisohyet map depicting the highest peaks of the StirlingRange as receiving more than 700mm of rainfall annually.The Stirling Range quokka population(s) may have beentraditionally isolated or may represent a remnant of aonce contiguous population. In either case, the statusand security of these isolated populations and the isolatedoccurrence at Green Range warrant further investigation.

The Rottnest Island population suffers seasonalmortality over summer and this has been attributedindirectly to dehydration (Barker 1961; Holsworth 1964;Main et al. 1959; Packer 1968; Storr 1964b).Nonetheless, quokkas persisted in the fossil record fromsouth-west Western Australia throughout periods ofsignificant climatic variation from warm and wet to glacialaridity. Cook (1960) suggested climate change was thecause of the modern decline of the quokka. There seemsto be insufficient evidence to attribute climate changealone as the major cause of the decline of the quokkasince the 1930s. However, Balme et al. noted thepersistence of the quokka in the fossil record throughperiods of aridity may have been because the south-westcorner of Australia did not experience the extremes inaridity experienced by other parts of southern Australia.Therefore, the consequences of increasing aridityassociated with current patterns of climate change shouldbe seen as a potential threat to the continued persistenceof the quokka.

European colonisation and development

Reports from naturalists in the early 1900s indicated the

quokka was common prior to and until the 1930s (Gould1863; Shortridge 1909; White 1952) and, with very fewexceptions, most reports suggest quokkas were restrictedto specific habitats (see Holsworth 1967; Storr 1964b).Much of the coastal heath and shrubland habitats in thesouth-west of Western Australia where quokkas onceoccurred (Shortridge 1909; White 1952) have beencleared for urban development. The coastal plain, fromnorth of Perth to Busselton (Fig. 1) has few remainingpockets of undisturbed vegetation and the remainingfragments are small and easily invaded by introducedpredators. The areas known to have historicallysupported quokka populations from the Swan CoastalPlain were swamps or low lying seasonally waterloggedareas. These coastal plain swamps have been progressivelydrained, with very few retaining their original drainagepatterns. Similarly, the upper reaches of many creeksystems in the forested areas of the Darling Range havebeen cleared for agriculture, dammed to supply waterto Perth or split from connecting habitat by roads tosuch an extent that the remaining quokka habitat is highlyfragmented and in places may be too small to supportviable populations.

Other factors associated with European colonisationmay have similarly affected the quokka. Calver and Dell(1998b) believed all mammal species from the south-west forests of Western Australia historically had a widerdistribution and suggested the extant mammal fauna maybe resilient to changes in forest structure. However, theycautioned this resilience may not be the case andemphasised the importance of experimentallydemonstrating that there are no direct and indirect linksbetween forestry practices and declines in distributionand abundance on the suite of resident native fauna fromthe south-west of Western Australia. Increases inpredation rates (Wayne et al. 2000), increases in thearea of edge affected habitat (Wilson & Friend 1999),increases in interpatch distances (Hayward et al. 2003)and increases in roading disturbance (Calver & Dell1998a) may be indirect effects from management of thejarrah forest. The effect of these activities on quokkapopulations has not been quantified. Roading and otherdisturbances associated with mining and harvesting,resulting in removal of habitat and alteration of drainagepatterns, also have the potential to contribute to increasesin interpatch distances. Construction of logging accessroads in 2001–2002 in Nairn Forest Block in the southernforest region of south-west Western Australia has beenimplicated as an example of the detrimental effects fromharvesting and the associated roading activities. Postcommencement of roading there was an observedincrease in the number of reported quokka roadkills. Aminimum of 10 quokka roadkills was reported by a singleobserver within a six month period in close proximityto, and coinciding with roading activity at Nairn ForestBlock (John Austin, pers. comm. to PJdeT)14 . However,there are alternative and equally plausible explanations

14 John Austin: Long-term local resident, Northcliffe, Western Australia

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28 P.J. de Torres et al.

for this increase in observed roadkills. Quokkas mayhave been attracted to areas of new growth resultingfrom recent burns of roadside verges. This explanationis also consistent with the quokkas’ preference for avegetation mosaic which includes recently burnt areas(de Tores et al. 2004). Nonetheless, this uncertaintyhighlights the need to identity the cause of the largenumber of roadkills and to quantify the direct and indirecteffects from anthropogenic disturbance.

Fire

The Noongar people have occupied the south-westcorner of Australia for at least the last 40 000 years(Balme et al. 1978; Merrilees et al. 1973) and possiblylonger (Turney et al. 2001). In the period prior toEuropean settlement, the Noongar people of the south-west are thought to have burnt the jarrah forest, withlow intensity fires, with a minimum fire interval of twoto three years in the moist to high rainfall areas, and twoto five years in the forest areas with lower annual rainfalland lower fuel accumulation rates (Burrows et al. 1995).Burrows et al. (1995) concluded this pattern of burningwould have led to a vegetation mosaic which would haveincluded patches of unburnt forest, with most patches‘being less than 6 years since last fire’ (see also Wilson &Friend 1999). Riparian environments may haveexperienced longer intervals between fires (Abbott 2000;Burrows & Friend 1998).

Ward et al. (2001) (see also Ward & Sneeuwjagt1999) believed examining fire scars on grass trees,Xanthorrhoea spp., provided a more sensitive techniquefor determining fire history. From examination of grasstrees, Ward et al. (2001) believed Noongar burning, orthe pre-European frequency of burning in the jarrahforest, was once every three to four years. Lamont et al.(2003) cautioned that the susceptibility of grass trees tofire, and therefore their ability to reflect fire history,depends on their location in the landscape. Burrows andWardell-Johnson (2003) further cautioned againstinterpreting historic patterns of fire on a regional scaleon the basis of patterns observed among individual grasstrees. The findings of Ward et al. (2001) suggested ashorter interval between fires than concluded by Burrowset al. (1995). However, both studies indicated thefrequency of burning in forest areas of south-westWestern Australia has changed from the frequency usedby the Noongar people. Ward and Sneeuwjagt (1999)further implied these changes also apply to coastal areas.Abbott (2003) suggested the interval between fires setby the Noongar people in coastal areas may have beenas short as two to four years.

Alterations to fire regimes following Europeansettlement have been implicated with declines in faunaabundance and range (see review by Wilson & Friend1999). Burrows et al. (1995) believed three distinct fireregime periods or eras could be defined for the forestsof south-west Western Australia post Europeansettlement, namely (i) the first European era (1855–1920) where there was a cessation of Noongar burning

practices which led to longer intervals between fires andmore intense fires; (ii) the second European era (1920–1965) when forestry practices adopted a policy of fireexclusion. Burning was restricted to strategic buffers andfire suppression purposes. Much of the forest remainedunburnt and large high intensity wildfires occurred,fuelled by logging debris and naturally occurring highfuel loads; (iii) the third European era (1965–1995)where fuel reduction burning was based on a six to tenyear rotation. Although fire frequency for this periodwas comparable to the pre-European regimes, fireintensity was higher, fuelled primarily by logging debris(Burrows et al. 1995). Towards the end of this thirdera, and from 1962 in particular, the area burnt byprescribed fire greatly increased (Lachie McCaw15 pers.comm. to PJdeT and unpublished data). This was areflection of the then Forest Department’s expandedprogram of broad-scale burning which commenced after1954 (McCaw et al. 2005; Wallace 1966).

Although this third ‘era’ encompasses the period 1980to 1992, i.e. the period within which our data indicate alarge contraction in the distribution of the quokka, nodirect causal relationship can be drawn, particularly asthe areas of greatest quokka decline are outside forestareas subject to this fire regime. Fire regimes in forestareas post 1995 have been similar to the third Europeanera described by Burrows et al. (1995). Discussions withDepartmental operational staff involved in prescriptionburning, combined with the reported reduction in theextent of the area burnt annually in the south-west ofWestern Australia, indicated a trend of progressivelydecreasing prescribed-burn fire frequency in the south-west. The 1998–99 prescribed burn program achievedless than 50% of the planned prescribed burning programand was the lowest achieved since 1961 (CALM 1999).The areas ‘prescription burnt’ in the south-west forestregions in 1999–2000, 2000–2001 and 2001–2002 were134 308, 87 866 and 74 739 ha. respectively (CALM2000; CALM 2001; CALM 2002). From 2002 on thistrend of decreasing area burnt annually has been reversed,with 144 835 ha burnt as part of prescribed burns in2002–2003 and 192 119 ha in 2003–2004 (CALM2003; CALM 2004). We caution that increasing the area‘prescription burnt’ alone does not equate with burningto provide the preferred habitat mosaic. Our personalobservations (PJdeT) suggest broad scale burning has atendency to use natural barriers as fire boundaries. TheTaxandria linearifolia creeklines often act as suchnatural barriers and the long term effect has been toencroach upon these barriers and progressively reducethe size and extent of the T. linearifolia creeklines. Thistype of encroachment does not equate with providingthe burn conducive to maintaining the preferred habitatof the quokka. However, effective use of prescribed burnsdoes provide the opportunity to use fire to create thequokka’s preferred habitat mosaic as described by

15 Lachie McCaw: Senior Research Scientist, Western Australian Departmentof Environment and Conservation, Science Division, Manjimup

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Distribution of the quokka 29

Hayward (2002) and Hayward (unpublished, reportedin summary in de Tores et al. 2004).

The finding by Hayward (unpublished, reported insummary in de Tores et al. 2004) is only partly consistentwith findings from a small-scale study (Christensen &Kimber 1975) specifically examining the effect of fire onthe quokka in forest sites near Dwellingup in the 1970s.Christensen and Kimber (1975) concluded quokkasreturned to swamps to forage almost immediately aftera burn, showed an influx of new individuals (i.e.previously un-trapped animals) and became resident 18months after the fire. There appeared to be no residentquokka population where vegetative cover was entirelyremoved by the fire (Christensen & Kimber 1975). Anolder (unburnt for 15 years) site trapped in that studyled the authors to conclude quokkas desert sites unburntfor 15 years and have a preference for a spatial mosaicor patchy burn which provide areas of refuge and areasof foraging habitat (Christensen & Kimber 1975).Hayward (unpublished, reported in summary in de Toreset al. 2004) came to a different conclusion and suggestedthis 15 to 19 year post-fire component of the mosaic didnot represent the upper fuel age of the quokkas preferredmosaic. Hayward (unpublished, reported in summaryin de Tores et al. 2004) showed this component of themosaic was negatively correlated with quokka presence(avoided by quokkas) and the preferred mosaic includedan additional component which was long unburnt (seesection on habitat use). We therefore suggest a burningregime which will create and maintain the mosaicidentified by Hayward (unpublished, reported insummary in de Tores et al. 2004) is required in thenorthern jarrah forest. This would necessitate differentburning regimes from those currently used at quokkasites in the northern jarrah forest (see section onmanagement recommendations).

Small, scattered populations are also likely to besusceptible to stochastic events such as wildfires whichmay result in localised and more extensive extinctions.Kirke (1983) reported quokkas fleeing from wildfires atGreen Range, north of Albany. Similarly, in theNorthcliffe area, quokkas observed in large numbers inthe 1940s and earlier, and known to feed in paddocksaway from vegetated creeklines, were reported as lastseen in number at this location when fleeing from a firein the 1940s (Laurie Wilson, pers. comm. to PJdeT)16 .Numerous quokkas were also observed in an openpaddock after fleeing a wildfire in the Allen Road/HilltopRoad area in Walpole-Nornalup National Park, circa1987 (John Asher, pers. comm. to PJdeT)17 . Quokkasare now reported to have repopulated this area and arethought to be in large numbers (Greg Freebury, pers.comm. to PJdeT).

The only known record of occurrence from KarnetForest Block, west of Jarrahdale, is from quokkas

observed fleeing a fire in 1991 (unpublished records fromA.N. Start, Table 4). A high-intensity wildfire burnt alarge portion of the Stirling Range National Park inNovember 2000 (pers. obs. of MWH) and numerousquokka deaths were reported. A similar fire in the StirlingRange National Park in 1991 was thought to have beenequally damaging to quokkas. However, quokkas werereported to have repopulated burnt areas in subsequentyears (Sinclair 1999). A large number of quokka deathswas also recorded as a result of a wildfire in the NuytsWilderness area, near Walpole in 2001 (Middleton 2001).Quokkas have subsequently been detected in unburntpatches within the boundary of the Nuyts fire and quokkapresence has been inferred at three locations immediatelyoutside the burn boundary. Presence was inferred bydetection of scats within typical quokka runways in densepatches of Spreading Sword-sedge, Lepidospermaeffusum, and Anarthria scabra (Greg Freebury, pers.comm. to PJdeT). With the exception of the programestablished to monitor recovery from the Nuyts fire, theextent of documenting any recovery from these stochasticevents has been largely anecdotal.

The quokka appears to be capable of persisting in afire-prone environment and the absence of low intensityfire from many sites may be a contributing factor to thecollapse of the northern jarrah forest metapopulation asdescribed by Hayward et al. (2003). However, there issufficient evidence from the northern jarrah forest tosuggest quokka populations there are dependent on thepresence of a structural mosaic which incorporates areasburnt within the previous nine years, but also has a largeoverall fuel age (i.e. must also include areas longunburnt) (Hayward, unpublished, reported in summaryin de Tores et al. 2004).

Further support for the requirement of a mosaic,and not simply a requirement for the presence of firewithin the past nine years, is provided by the results fromsurveys in 1995–1996 (Dillon 1996). From 28 northernand southern forest locations previously known to supportquokka populations, nine no longer supported quokkasand seven of these nine sites had been burnt within theprevious 10 years (Table 10) (Dillon 1996). Althoughthese areas provide the component of the mosaic whichhas been burnt within the last 9 to 10 years, they may nolonger support areas of long unburnt habitat.

A fire regime of low intensity burns and with afrequency comparable to the third European era ofBurrows et al. (1995) may be appropriate to generatethe vegetation mosaic preferred by the quokka. However,no single fire regime will benefit all taxa. Of particularconcern is the threat to the Noisy Scrub-bird, Atrichornisclamosus. The noisy scrub-bird has a preferred habitatof densely vegetated creekline and gully vegetation(Abbott 1999; Burbidge 2003). A too frequent burningregime of creeklines where it and the quokka aresympatric may compromise noisy scrub-bird habitat and‘even a mild fire can render an area unsuitable for manyyears’ (Burbidge 2003). Although the only knownlocations where extant populations of the noisy scrub-bird and quokka occur sympatrically are at Two Peoples

16 Laurie Wilson: Long-term local resident, Northcliffe, Western Australia17 John Asher: Environmental Officer, Western Australian Department of

Environment and Conservation, Nature Conservation Division, Bunbury

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30 P.J. de Torres et al.

Bay, on the south coast, the noisy scrub-bird has recentlybeen translocated to sites within the northern jarrah forestwhere quokkas may also occur. Therefore, there is thepotential for conflicting fire management requirementsat these translocation release sites.

There are also indirect detrimental effects from fire.Christensen (1980a) recorded increases in the predationrate on the brush-tailed bettong following fire. This typeof indirect effect may also result in localised increasedpredation risk for other mammal populations. Kinnearet al. (Kinnear et al. 1988) suggested this type of indirectthreat may increase the risk of localised extinctions.

Hunting by Aboriginal people

Gardner (1957), citing John Lort Stokes, referred toobservations of Aboriginal people burning the bush forthe purpose of catching snakes, lizards and wallabies.These prey were speared as they fled the burningvegetation. Gould (1863) and Evans (undated) weremore specific and noted quokkas were eaten byAboriginal people. Evans (undated) noted the Aboriginalpeople from the Northcliffe area supplemented their dietwith quokka, kangaroo and marron. Gould (1863) notedquokkas were killed in great numbers at the end of theseason by Aboriginal people. Gould’s (1863) descriptionof Aboriginal people burning the bush to flush out theirprey is consistent with Gardner’s (1957) description. Theend of the season referred to by Gould (1863) is nodoubt a reference to the end of summer, as Meagher(1974), citing numerous historical records, noted towardsthe end of summer, Aboriginal people set fire to thebush to drive the wallabies from their retreats. Green(1989) referred to use of fire at Bald Head in January(mid summer) to ‘burn off the land for wallaby’ and theJanuary reference of ‘brought home … three wallaby fromBald Head where the Aborigines had fired the land to hunt’indicates burning for this purpose was not restricted tothe end of the summer season. At other times of theyear dogs were used to drive out prey (Meagher 1974).

Although Aboriginal people may have beenresponsible for exterminating some island populationsof macropods (Abbott 1980), there is no evidence tosuggest Aboriginal hunting of the quokka contributedto the species’ decline.

Disease

Reports from studies of colonies of captive quokkas showthe susceptibility of the species to disease (see Bradshaw1991). Anecdotal accounts in 1967 suggested a fatalherpes epidemic transmitted by a handler affected acaptive quokka colony (Burnet 1968). Salmonellainfections are common in the Rottnest Island and BaldIsland quokka populations but far less so on the mainland(Hart 1977; Hart et al. 1986). Hart (1981) foundcaptured mainland quokkas died within 24 hours ofexposure to bags in which Rottnest Island quokkas hadbeen held. These deaths were attributed to infectionsfrom Salmonella meunchen. Hart (1981) hypothesisedthese deaths may have been the result of cross infection

from use of the Rottenest Island bags, but conceded S.meunchen may have been present at an undetectablylow level in the mainland quokka population and becamelethal only when the animals were stressed throughtrapping.

Barker et al. (1957) recorded the presence of theparasite Ixodes australiensis in mainland populations ofthe quokka and several other parasites and diseases havebeen isolated from quokkas.

Mass mortality of quokkas was reported from theNorthcliffe area where swamps were reportedly ‘full ofquokka bodies’ in the early 1920s (George Gardner citedas a pers. comm. by How et al. 1987). How et al. (1987)attributed this to disease. Similarly, there are records ofan epidemic within the quokka population in the WarrenRiver area, near Manjimup in 1921 (Aldrich 1921; Lane-Poole 1921; Weston 1921). Unexplained quokka deathswere also reported from Crown reserves in the vicinityof Yallingup in 1933 (Aldrich 1933). Other references(Cook 1960; Perry 1973; Waring 1956; White 1952)also indicated mass deaths occurred as a result of diseasein the 1930s. Mass deaths have recently been attributedto surplus killing by foxes (Short et al. 2002).

The decline in arid zone mammals of the late 1800snoted by Shortridge (1909) and reported from theNullarbor Plain (Richards & Short 1996), has beensuggested to be a result of a ‘strange virus’ whichRichards and Short (1996) suggest, albeit based onanecdotal accounts, may have been the protozoanparasitic disease toxoplasmosis, passed on from feral cats.Toxoplasmosis was first recorded in the Rottnest Islandquokkas in 1961 (Gibb et al. 1966) and has beenreported in other marsupials including the eastern barredbandicoot, Perameles gunnii, (Lenghaus et al. 1990),the western ringtail possum, Pseudocheirus occidentalis,(de Tores 2005) and the chuditch, Dasyurus geoffroii,(Haigh et al. 1994). Berdoy et al. (2000) notedtoxoplasmosis may alter the behaviour of its intermediatehosts and increase its susceptibility to predation.

White (1952) believed fox predation, competition withrabbits, destruction of habitat through clearing andbushfires were supplementary to disease as the causalfactor for the quokkas’ virtual disappearance on themainland. Despite the references to quokkas dying fromdisease in the 1920s (Aldrich 1921; Lane-Poole 1921;Weston 1921) and the 1930s (Cook 1960; Perry 1973;Waring 1956; White 1952), the persistence of quokkason Rottnest Island and Bald Island during these periodssuggests disease was not the major contributor to theirdecline, or alternatively, if disease was responsible forthe decline on the mainland, it did not have the sameeffect on, or did not reach, Rottnest Island or Bald Island.It was during this period of decline on the mainland inthe 1930s when the quokka was no doubt at high densityon Rottnest Island, as it was in this period when it wasfirst referred to as a pest on Rottnest Island (Storr 1963).

Although White (1952) believed a decline caused bypredation, competition and habitat destruction wassupplementary to that caused by disease and Cook (1960)linked the decline of the quokka with disease, Recher

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Distribution of the quokka 31

and Lim (1990) considered disease to be a contributing,not causal, factor to the decline of the Australia’s mammalfauna. Johnson et al. (1989) discounted disease as a factorassociated with the decline of critical weight rangemacropods altogether. Dickman (1992) noted there wasno evidence to indicate widespread disease as the causeof the decline of mammals within Australasia. We concurwith these authors and suggest there is insufficientevidence to conclude disease alone was the cause of thedecline of the quokka. However, we caution againsttrivializing the potential effects from disease. The 1826King George’s Sound record of Quoy and Giamard notedthe quokka specimen described ‘was recently dead whenwe found it, probably from disease’ (Alexander 1916). Ifdisease was the cause of death, it is reasonable tohypothesise disease was brought to King George’s Soundby Europeans. Although the Rottnest Island populationappears to be secure, its possible exposure to disease asa result of human contact should be seen as a potentialthreat. Introduction of a disease, or a single catastrophicevent, could result in a chance extinction of this insularpopulation.

Management recommendations

The northern jarrah forest

We recommend an active adaptive managementapproach for conservation management of the quokkawithin the northern jarrah forest. The active adaptivemanagement approach requires implementing an agreedexperimental approach (i.e. agreed to by allstakeholders), whereby a set of models and managementactions have been formulated and appropriatemonitoring protocols established. We believe monitoringshould be focused on examining the response of thequokka to a variety of management practices.

Quantified data has shown the northern jarrah forestpopulations are at low density. Despite the presence offox baiting, these populations have not responded andpredation is still potentially limiting population response.Based on the rapid rate of fox re-invasion of the northernjarrah forest post aerial baiting events (de Tores 1999),the lower probability of survivorship of the woylie,Bettongia penicillata, in treatments baited four times peryear compared with six times per year, and thesignificantly lower levels of survivorship in areas abuttingagricultural land (de Tores 1999), we recommend theactive adaptive management program incorporates anassessment of the effectiveness of an increased frequencyof 1080 baiting.

The northern jarrah forest populations are highlyfragmented, the populations are not mixing and thepreferred habitat within the Taxandria swamps is acomplex mosaic of recently burnt and long unburntareas. An additional requirement of the preferred mosaicis to have a minimal area burnt 15 to 19 years previously.The preferred spatial configuration of these seral stagesis not known – specifically, the relative proportion ofeach required seral stage is not known and the upper

limit of ‘time since last fire’ , i.e. the maximum period oftime without fire within the long unburnt component ofthe mosaic, is not known. Fire management practiceswhich selectively burn long unburnt components of themosaic may be detrimental to the long-term conservationof quokkas in the northern jarrah forest. This practicehas the potential to increase fragmentation of quokkahabitat by removing a component of the preferredmosaic. This practice should not be seen as a methodfor creation of quokka habitat, and, at best, constitutesa program more akin to a trial and error approach thanan adaptive management program.

Therefore, we recommend the active adaptivemanagement program should assign high priority tospatial analyses of existing, historic and potential quokkasites in the northern jarrah forest, with the objective ofstratifying the existing Taxandria mosaic (with stratabased on the number of years post fire) and determiningwhether any of these swamps can be better managedthrough the use of fire. Under this scenario, fire wouldbe used to create the preferred mosaic. We alsorecommend monitoring should incorporate geneticanalyses to test the northern jarrah forest metapopulationhypothesis. We further advocate any monitoring programassociated with quokka conservation needs to incorporatea component to enable quantitative assessment of pigdamage to quokka habitat. Although we advocate useof conventional trapping techniques to determine quokkaabundance, we caution against the overuse of invasivetrapping and strongly recommend use of alternativemethods where applicable. These methods includemolecular techniques (Alacs et al. 2003). We alsorecommend development of techniques to quantifyabundance based on the extent of activity in quokka-likerunways. Currently assessment of activity in runways canmeasure activity levels only and should not be extrapolatedto infer abundance. Further development of thetechnique adopted by Hayward et al. (2005) isrecommended.

The active adaptive management program shouldspecify quantifiable conservation outcomes. Suchmeasurable long-term conservation outcomes includedetermining:

• the number of known extant quokka sites where thepreferred structural mosaic has been establishedthrough the use of fire;

• the number of new sites where the preferred structuralmosaic has been created through the use of fire;

• the number of sites where quokka populations remainstable or show an increase in abundance (asmeasured through survival analyses and populationestimates, respectively);

• the number of sites currently thought to be supportingpotentially suitable habitat, where quokkapresence has not been detected/confirmed, andwhere, post habitat manipulation, quokkapresence is confirmed and a populationestablished;

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32 P.J. de Torres et al.

• the number of sites where population mixing has beenconfirmed (i.e. sites where animals have beenknown to disperse to and where the source site isstill viable [the source sites can be identifiedthrough the use of molecular techniques, andpopulation viability can be determined byconventional monitoring [trapping] andpopulation modeling]).

The Swan Coastal Plain

High priority is recommended to unambiguouslydetermine quokka presence at Muddy Lake and the waterauthority reserve near Dunsborough. If presence isconfirmed, we recommend a monitoring program beimplemented to ensure any trend of increase or decreasein population can be detected. The requirement forintroduced predator control and habitat manipulationshould also be assessed. Minimal data were available onquokka presence elsewhere on the Swan Coastal Plain.We recommend a review of the former known locationsfrom the Swan Coastal Plain (historically these wereswamps), assessment of quokka presence at these sitesand examination of the effects from draining theseswamps. If appropriate, and where possible, theconservation value of re-instating former drainagepatterns should be examined.

The southern forest and south coast

The dearth of information on the size of each population/sub-population from the southern forest and south coastareas should be addressed. We recommend an initialapproach of confirming presence, through trapping, atsites where quokkas are thought to occur. We furtherrecommend undertaking spatial analysis of these extantpopulations and locations of known and potentiallysuitable habitat in conjunction with a survey andmonitoring program to assess population size. The extentof dispersal/immigration/emigration between habitatpatches should be quantified to determine whether thesub-populations constitute a functional metapopulation,discrete sub-populations or a panmictic population. Thisprocess would enable populations of high conservationvalue to be identified, where conservation value is assessedin terms of the population’s strategic value locally,regionally and globally as determined by its geographiclocation, demographics, genetic structure and importanceas a source population for re-stocking other populations/subpopulations.

As a matter of urgency, we recommend a morerigorous and strategic approach be implemented to assessthe potential effect from timber harvesting and associatedoperational activities. Assessment should identify theextent of quokka habitat to be modified, destroyed orretained by each proposed operation, the size of thepopulation(s) affected by the proposed operation, theconservation significance of the population, the potentialfor dispersal, numbers likely to disperse and dispersalpatterns, availability of suitable habitat within dispersaldistances, population size within areas of suitable habitat

within dispersal distance and the potential effect on thesepopulations.

We recommend investigation of the population(s)from the Stirling Range to assess the security of thesepopulations. Survey is recommended to assesspopulation size, habitat used and the security of thishabitat from stochastic events, in particular from wildfire.We also recommend examination of the genetic structureto determine whether the Stirling Range and GreenRange population(s) was, or is still, contiguous with othersouth coast populations.

CONCLUSION

The distribution of the quokka appears to have beentraditionally limited by climate and by rainfall in particular.Reduction in availability of suitable habitat, in conjunctionwith predation and changed fire regimes, appear to havelimited this distribution, or more specifically has furtherconfined the quokka to specific habitats within the limitsof its geographic range.

For the quokka, like numerous other native mammalspecies, the arrival of Europeans to Australia coincidedwith a slow but continual decline in abundance and range(Fig. 3). From 1900 in particular, the increasing humanpopulation in the south-west of Western Australia resultedin anthropogenic disturbances including vegetationclearance, logging, mining, hunting and changed fireregimes. Introduced predators have been implicated in40% of historic extinctions (Caughley & Gunn 1996)and it seems likely that predation pressure from theintroduced red fox, in conjunction with continued habitatalteration through fire exclusion and colonisation, furthercompromised the conservation status of the quokka.

However, the quokka, like many Australian nativemammals (Wilson & Friend 1999) appears to be resilientto individual disturbance factors but also appears to beincreasingly susceptible to the cumulative effect ofmultiple factors. None of these factors has occurred inisolation. Many commenced more or less in synchronywith the arrival of the fox (Recher & Lim 1990) and allcontinue to operate.

Although there is considerable agreement thatmultiple factors combined to contribute to the decline,there is also considerable disagreement as to the ultimatecause of the decline of critical weight range mammalssince European arrival (see Burbidge & McKenzie 1989;Lunney et al. 2001; Morton 1990; Recher & Lim 1990;Short et al. 2002; Short & Calaby 2001; Smith & Quinn1996; Wilson & Friend 1999). Abbott (2001b) concludedthe fox was the primary agent responsible for the declineof the bilby. However, a single explanatory hypothesisto account for past and continuing declines is unlikely tobe applicable to all species and it would seem logical thateach species is affected to a differing degree by eachfactor. Similarly, the relative importance of any factorwill vary spatially for each species. Kinnear et al. (2002)proposed two hypotheses, niche loss/damage andpredation, to account for the initial and continued decline

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Distribution of the quokka 33

of Australia’s critical weight range mammal fauna. Thehypotheses are not necessarily mutually exclusive (Kinnearet al. 2002). There is a wealth of evidence linking twofactors to the decline of the quokka; predation and aloss of the quokkas’ preferred habitat mosaic. The latterequates with the niche loss/damage hypothesis proposedby Kinnear et al. (2002). We believe the two hypotheses,niche loss/damage and predation, when acting inconcert, are sufficient to account for the pattern of declineof the quokka.

Islands, although not immune from disturbancefactors, often act as refuges for threatened species largelybecause the factors responsible for the decline on themainland are often absent from islands (Dickman 1992).The quokka population on Rottnest Island may provideclues to the significance of the various disturbance factorswhich led to its decline on the mainland. Compared tothe mainland, the Rottnest Island population has a highersusceptibility to disease (Salmonella) (Hart et al. 1986),encounters seasonal aridity leading to summer mortality(Hodgkin & Sheard 1959; Shield 1964) and a period ofanoestrus (Shield 1964). Rottnest Island and mainlandsouth-west Western Australia have experienced the effectsof development and have been subject to extensive habitatfragmentation. The success of the quokka populationon Rottnest Island, despite these disturbances, suggeststhe species is quite resilient. With respect to disturbancefactors, the most significant difference between RottnestIsland and the south-west mainland appears to be thepresence of the fox on the mainland. Recent interestingadvances in our understanding of the effect of the foxon native fauna reinforce this conclusion. The predationefficiency of foxes combined with the inadequate anti-predator defences of a predator-naïve fauna may lead to‘high and unsustainable’ levels of predation (Short et al.2002). Short et al. (2002) concluded this is the case forthe Australian mammal fauna which has evolved in relativeisolation since the break-up of Gondwana 50-60 millionyears BP (Heatwole 1987). Short et al. (2002) furtherconcluded surplus killing may be an outward sign of thismismatch and provided case studies to support theirassertion that the declines of many prey species, and inparticular those species with limited refugia, may be aresult of surplus killing by foxes.

The hypothesis that species with little or no refugewould have suffered most from this predation (Short etal. 2002) is consistent with the pattern of decline of criticalweight range mammals in the arid zone described byBurbidge and McKenzie (1989). The quokka is nowrestricted to areas of refuge (islands and dense vegetationon the mainland). The presence of this latter refuge seemslikely to have been the sole factor staving off extinctionresulting from fox predation and the combined effectsof altered fire regimes and habitat loss.

With the continued depletion of resources (nutrients,water, refuge), which have previously buffered the faunadecline in the more mesic areas of the mainland (Recher& Lim 1990; Woinarski et al. 2001), the bufferingcapability may be in the process of being compromised.Further extinctions may result (Recher & Lim 1990;

Woinarski et al. 2001) and the assumption that criticalweight range mammals in mesic areas are secure(Burbidge & McKenzie 1989) may no longer be valid.

ACKNOWLEDGEMENTS

Staf f from the Western Australia Department ofEnvironment and Conservation (DEC, formerly theDepartment of Conservation and Land Management,CALM) provided numerous records of the occurrenceof the quokka, and in particular we thank Sandra Gilfillan,Peter Orell and Peter Mawson for supplying databaserecords and Peter Collins, Greg Freebury, Ian Wilsonand Graeme Liddelow for additional information onsouthern forest and south coast populations. Thanks toRalph Staines for providing additional trapping andlocation records, Antoinette Tomkinson, Rob Buehrig,Kathy Himbeck, Cathy Lambert, Kaylene Parker andSuzanne Rosier for providing field assistance at varioustrapping sites and to John Liddington, from the WaterCorporation of Western Australia for providing additionaltrapping and location records. Thanks especially to LizJef ferys for collating quokka records from theuncatalogued records of the Palaeontology Departmentof the Museum of Western Australia, Norah Cooper forclarifying WAM records, Tony Start for providing manyunpublished records, Sandy Ingleby for providing accessto the database records of the Australian Museum, ClaireWright from the Rottnest Island Authority for providingadditional references and Tony Start and Jim Lane foralerting the authors to the issues relevant to quokkahabitat on the Swan Coastal Plain. Special thanks toMark Sheehan, Northcliffe, for his commitment toensuring all quokka records were forwarded to theauthors. We thank Robyn Wilson, Shane French andDavid Robertson from the Western AustralianDepartment of Environment and Conservation,Information Management Branch for GIS support. Inparticular we thank Lisa Wright, Wildlife ResearchLibrarian, DEC, for the tracking down requests forobscure references and Ian Abbott, Neil Burrows, R.I.T.(Bob) Prince, Lachie McCaw, Tony Start, AndrewBurbidge and Suzanne Rosier for advice on preliminarydrafts. We also thank Per Christensen for his constructivecriticism as a formal referee and for his valuable inputand support generally. We thank Don Bradshaw andDarrell Kitchener for comment as referees of a previousdraft and two anonymous referees for valuable commenton an earlier draft which encompassed a conservationstatus review. The work was funded by the WesternAustralian Department of Conservation and LandManagement and Alcoa World Alumina, Australia.

REFERENCES

Abbott I (1980). Aboriginal man as an exterminator ofkangaroo and wallaby populations on islands roundAustralia. Oecologia 44:347–354.

Page 22: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

34 P.J. de Torres et al.

Abbott I (1998). Avifauna of the Irwin Inlet-Broke Inlet-Mt Frankland region of south-west Western Australia,1912-1913. CALMScience 2(4):373–386.

Abbott I (1999). The avifauna of the forests of south-west Western Australia: changes in speciescomposition, distribution, and abundance followinganthropogenic disturbance. CALMScienceSupplement 5:1–175.

Abbott I (2000). Impact of agricultural developmentand changed fire regimes on species composition ofthe avifauna in the Denmark region of south-westWestern Australia, 1889–1999. CALMScience3(3):279–308.

Abbott I (2001a). Aboriginal names of mammal speciesin south-west Western Australia. CALMScience3(4):433–486.

Abbott I (2001b). The Bilby Macrotis lagotis(Marsupialia: Peramelidae) in south-western Australia:original range limits, subsequent decline, andpresumed regional extinction. Records of the WesternAustralian Museum 20:271–305.

Abbott I (2002). Origin and spread of the cat, Feliscatus, on mainland Australia, with a discussion ofthe magnitude of its early impact on native fauna.Wildlife Research 29(1):51–74.

Abbott I. (2003). Aboriginal fire regimes in south-westWestern Australia: evidence from historicaldocuments. In: Fire in Ecosystems of south-west WesternAustralia: impacts and management. (I AbbottNBurrows, Eds). Backhuys Publishing, Leiden, theNetherlands. Perth, Western Australia, pp. 119–146.

Abbott I, Burbidge AA (1995). The occurrence ofmammal species on the islands of Australia: asummary of existing knowledge. CALMScience1(3):259–324.

Alacs E, Aplers D, de Tores PJ et al. (2003). Identifyingthe presence of quokkas (Setonix brachyurus) andother macropods using cytochrome b analyses fromfaeces. Wildlife Research 30(1):41–47.

Alacs EA (2001). Conservation genetics of the Quokka,Setonix brachyurus. Honours thesis. VeterinaryBiology. Murdoch University. Perth, Western Australia

Aldrich J (1921). File note dated 9 May 1921 in WesternAustralian Government State Records OfficeArchives: 56/21 AN 108 Acc 477. Game Act –Diseases among Quaggas (Wallibies) in South West.

Aldrich J (1933). File note dated 9 September 1933 inWestern Australian Government State Records OfficeArchives: 56/21 AN 108 Acc 477. Game Act –Diseases among Quaggas (Wallibies) in South West.

Alexander WB (1914). The history of zoology in WesternAustralia. Part I. – Discoveries in the 17th Century.The Journal of the Natural History and Science Societyof Western Australia 5:49–64.

Alexander WB (1916). History of zoology in WesternAustralia. Part II. – 1797–1829. Journal andProceedings of the Royal Society of Western Australia1:83–149.

Alexander WB (1918). History of zoology in WesternAustralia. Part 3. – 1829–1840. Journal and

Proceedings of the Royal Society of Western AustraliaIII:37–69.

Algar D, Kinnear JE (1996). Secondary poisoning offoxes following a routine 1080 rabbit-baitingcampaign in the Western Australian wheatbelt.CALMScience 2(2):149–152.

Archer M (1974a). New information about thequaternary distribution of the Thylacine (Marsupialia,Thylacinidae) in Australia. Journal of the Royal Societyof Western Australia 57(2):43–50.

Archer M (1974b). Excavations in the Orchestra ShellCave, Wanneroo, Western Australia. Archaeology andPhysical Anthropology in Oceania 9(2):156–162.

Archer M, Baynes A (1972). Prehistoric mammal faunasfrom two small caves in the extreme south-west ofWestern Australia. Journal of the Royal Society ofWestern Australia 55(3):80–90.

Austin C (1979). The Well-Being of the JandakotQuokka Population. Population Analysis. Thesissubmitted in partial fulfillment of Bachelor of Science(Honours). Zoology Department. University ofWestern Australia. Perth. Western Australia. Perth

Balme J, Merrilees D, Porter JK (1978). Late Quaternarymammal remains, spanning about 30 000 years, fromexcavations in Devil’s Lair, Western Australia. Journalof the Royal Society of Western Australia 61(2):33–65.

Bannister JL (1970). Report on Biological Survey, TwoPeoples Bay Nature Reserve (No. 27956) February3 to 17, 1970. Unpublished Report for the WesternAustralian Museum. Perth, Western Australia.

Barker R, McGlinchy A. (2001). M-R-I. MarkRecapture Interface for Programs Capture and Jolly.Version 2. Landcare Research. Manaaki Whenua.New Zealand. http://www.landcareresearch.co.nz

Barker S (1961). Studies on marsupial nutrition. III.The copper-molybdenum-inorganic sulphateinteraction in the Rottnest quokka, Setonix brachyurus(Quoy & Gaimard). Australian Journal of BiologicalScience 14:646–658.

Barker S, Main AR, Sadleir RM (1957). Recent captureof the quokka (Setonix brachyurus) on the mainland.The Western Australian Naturalist 6:53–55.

Barrett S (1996). A Biological Survey of Mountains inSouthern Western Australia. Unpublished reportprepared for the Western Australian Department ofConservation and Land Management and theAustralian Nature Conservation Agency.

Baynes A (1979). The Analysis of a Late QuarternaryMammal Fauna from Hastings Cave, Jurien. PhDThesis. University of Western Australia. Perth

Baynes A. (1987). The original mammal fauna of theNullarbor and southern peripheral regions: evidencefrom skeletal remains in the superficial cave deposits.In: A Biological Survey of the Nullarbor Region Southand Western Australia in 1984. (NL McKenzieACRobinson, Eds). South Australian Department ofEnvironment and Planning. Adelaide, pp. 139–152.

Baynes A, Merrilees D, Porter JK (1975). Mammalremains from the upper levels of a late Pleistocene

Page 23: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka 35

deposit in Devil’s Lair, Western Australia. Journal ofthe Royal Society of Western Australia 58(4):97–126.

Berdoy M, Webster JP, MacDonald DW (2000). Fatalattraction in rats infected with Toxoplasma gondii.Proceedings of the Royal Society of London (Series B)267:1591–1594.

Biodiversity Group Environment Australia (1999). ThreatAbatement Plan for Predation by the European RedFox. Environment Australia under the NaturalHeritage Trust. Canberra, ACT, Australia.

Bradshaw SD. (1991). Rottnest Quokka Workshop. Aninquiry into the current status of the quokka, held in theRottnest Research Station on 9–10 March, 1991.Unpublished report prepared for the Rottnest IslandResearch Committee.

Brown KL, Hickman EJ, Lally TR (1998). Discoveryof the red flowering Corymbia ficifolia in the StirlingRange. The Western Australian Naturalist 22(1):23–24.

Burbidge AA (Ed.).) (1977). The Status of Kangaroosand Wallabies in Australia. Report of the WorkingGroup on Macropod Habitat of the Standing Committeeof the Council of Nature Conservation Ministers.Department of Environment, Housing andCommunity Development. Australian GovernmentPublishing Service, Canberra.

Burbidge AA, McKenzie NL (1989). Patterns in themodern decline of Western Australia’s vertebratefauna: causes and conservation implications.Biological Conservation 50:143–198.

Burbidge AA, Manly BFJ (2002). Mammal extinctionson Australian islands: causes and conservationimplications. Journal of Biogeography 29:465–473.

Burbidge AH. (2003). Birds and fire in theMediterranean climate of south-west WesternAustralia. In: Fire in Ecosystems of south-west WesternAustralia: impacts and management. (I AbbottNBurrows, Eds). Backhuys Publishing, Leiden, theNetherlands. Perth, Western Australia, pp. 321–348.

Burnet SM (1968). Changing Patterns: an atypicalautobiography. Sun Books. Melbourne.

Burnham KP, Anderson DR (2002). Model Selection andMultimodel Inference. A Practical Information-Theoretic Approach. Second Edition. Springer-Verlag.New York.

Burrows N, Wardell-Johnson G. (2003). Fire and plantinteractions in forested ecosystems of south-westAustralia. In: Fire in Ecosystems of south-west WesternAustralia: impacts and management. (I AbbottNBurrows, Eds). Backhuys Publishing, Leiden, theNetherlands. Perth, Western Australia, pp. 225–268.

Burrows ND, Friend GR. (1998). Biological indicatorsof appropriate fire regimes in southwest Australianecosystems. In: Fire in Ecosystem Management:Shifting the Paradigm from Suppression to Prescription.(TL PrudenLA Brennan, Eds). Proceedings of theTall Timbers Fire Ecology Conference No. 20. TallTimbers Research Station, Tallahassee, Florida. 413–421.

Burrows ND, Ward B, Robinson AD (1995). Jarrahforest fire history from stem analysis andanthropological evidence. Australian Forestry58(1):7–16.

Butler WH (1969). Remains of Sarcophilus the‘Tasmanian’ Devil (Marsupialia, Dasyuridae) fromcoastal dunes south of Scott River, Western Australia.The Western Australian Naturalist 11(4):87–88.

Calaby JH (1971). The current status of the Australianmacropodidae. Australian Journal of Zoology16(1):17–28.

Calaby JH, Grigg GC. (1989). Changes in macropodcommunities and populations in the past 200 years,and the future. In: Kangaroos, Wallabies and Rat-Kangaroos. Vol. 2 (G Grigg, P JarmanI Hume, Eds).Surrey Beatty & Sons Pty Limited. Chipping Norton,New South Wales, pp. 813–820.

Caley P (1997). Movements, activity patterns and habitatuse of feral pigs (Sus scrofa) in a tropical habitat.Wildlife Research 24(1):77–88.

CALM (1999). Department of Conservation and LandManagement Annual Report 1998–1999. Perth,Western Australia.

CALM (2000). Department of Conservation and LandManagement Annual Report 1999–2000. Perth,Western Australia.

CALM (2001). Department of Conservation and LandManagement Annual Report 2000–2001. Perth,Western Australia.

CALM (2002). Department of Conservation and LandManagement Annual Report 2001–2002. Perth,Western Australia.

CALM (2003). Department of Conservation and LandManagement Annual Report 2002–2003. Perth,Western Australia.

CALM (2004). Department of Conservation and LandManagement Annual Report 2003–2004. Perth,Western Australia.

CALM. (unpublished). Unpublished Department ofConservation and Land Management databaserecords of fauna occurrence in Western Australia.

Calver MC, Dell J (1998a). Conservation status ofmammals and birds in the southwestern Australianforests. II. Are there unstudied, indirect or long-termlinks between forestry practices and species declineand extinction? Pacific Conservation Biology 4:315–325.

Calver MC, Dell J (1998b). Conservation status ofmammals and birds in the southwestern Australianforests. I. Is there evidence of direct links betweenforestry practices and species decline and extinction?Pacific Conservation Biology 4:296–314.

Catling PC (1988). Similarities and contrasts in the dietsof foxes, Vulpes vulpes, and Cats, Felis catus, relativeto fluctuating prey populations and drought.Australian Wildlife Research 15:307–317.

Caughley G, Gunn A (1996). Conservation biology intheory and practice. Blackwell Science. Carlton.

Page 24: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

36 P.J. de Torres et al.

Christensen P (1978). The introduced european red fox(Vulpes vulpes): a serious threat to native mammalsof the south-west. Forests Department of WesternAustralia. Perth. November 1978.

Christensen P, Burrows N. (1994). Project desertdreaming: experimental reintroduction of mammalsto the Gibson Desert, Western Australia. In:Reintroduction Biology of Australian and NewZealand Fauna. (M Serena, Ed.). Surrey Beatty andSons. Sydney, pp. 199–207.

Christensen P, Annels A, Liddelow G et al. (1985).Vertebrate fauna in the southern forests of WesternAustralia: a survey. Bulletin 94. Forests Departmentof Western Australia. Perth. 1985.

Christensen PE (undated). Effect of Prescribed Burningon One Population of Quokka (Setonix brachyurus).Local Expt. 117. Unpublished report for the ForestsDepartment of Western Australia.

Christensen PE, Kimber PC. (1975). Effects ofprescribed burning on the flora and fauna of south-west Australian forests. In: Managing TerrestrialEcosystems. Brisbane. (J KikkawaHA Nix, Eds).Ecological Society of Australia. Proceedings of theEcological Society of Australia, Vol 9: 85–106.

Christensen PES (1980a). The biology of Bettongiapenicillata Gray, 1837, and Macropus eugenii(Desmarest, 1817) in relation to fire. Bulletin 91.Forests Department of Western Australia. Perth.1980.

Christensen PES (1980b). A sad day for native fauna.Forest Focus 23:3–12.

Churchill DM (1959). Late Quarternery eustaticchanges in the Swan River District. Journal of theRoyal Society of Western Australia 42:53–55.

Clarke JR (1948). Anatomy of the Quokka. Part 1. –External morphology and large intestine. Journal ofthe Royal Society of Western Australia 33:59–150.

Coman BJ. (1991). The ecology and control of feralcats in Australia. In: The Impact of Cats on NativeWildlife. (C Potter, Ed.). National EndangeredSpecies Program, Australian National Parks andWildlife Service, Canberra. 70–74.

Cook DL (1960). Some mammal remains found in cavesnear Margaret River. The Western AustralianNaturalist 7(4):107–108.

Cook DL (1963). The fossil vertebrate fauna of Strongs’Cave, Boranup, Western Australia. The WesternAustralian Naturalist 8(7):153–162.

Corbett LK (1995a). The Dingo in Australia and Asia.University of New South Wales Press. Sydney.

Corbett LK. (1995b). Dingo, Canis lupus dingo. In:The Mammals of Australia. (R Strahan, Ed.). ReedBooks. Sydney, pp. 696–698.

Corbett LK, Newsome AE (1987). The feeding ecologyof the dingo. III. Dietary relationships with widelyfluctuating prey populations in arid Australia: anhypothesis of alternation of predation. Oecologia74:215–227.

Courtney J. (1993). Climate. In: Mountains of Mystery.A Natural History Of The Striling Range. (CThomson, G HallG Friend, Eds). Western AustralianDepartment of Conservation and Land Management.Perth, Western Australia.

Crabb D (1973). Quokka on the Albany Highway. TheWestern Australian Naturalist 12(6):141.

Daubney ME, Gardner G, Moore EM et al. (Eds.)(undated). Northcliffe Remembers. NorthcliffePioneer Museum, Northcliffe, Western Australia.

de Tores P (1999). 1997–1998 Report to EnvironmentAustralia National Feral Animal Control ProgramISF06638. Control and Ecology of the Red Fox inWestern Australia – Prey Response to 1080 BaitingOver Large Areas. Unpublished report prepared forEnvironment Australia, National Feral AnimalControl Program. ISF06638. Department ofConservation and Land Management. Perth.

de Tores P (2005). A proposal for translocation of thewestern ringtail possum, Pseudocheirus occidentalis,an arboreal marsupial endemic to the south-west ofWestern Australia. Unpublished report prepared forthe Western Australian Department of Conservationand Land Management. Perth, Western Australia.

de Tores PJ, Hayward MW, Rosier SM (2004). Thewestern ringtail possum, Pseudocheirus occidentalis,and the quokka, Setonix brachyurus, case studies:Western Shield review – February 2003. ConservationScience Western Australia 5(2):235–257.

Dell J. (1983). The Importance of the Darling Scarp toFauna. In: Scarp Symposium. A one day symposium toreview land use and conflicts on the Darling Scarp.Perth. Western Australian Institute of Technology.WAIT Environmental Studies Group. Report No.10:

Dell J, Hyder-Griffiths B (2002). A PreliminaryAssessment of the Faunal Values of the Muddy LakesPortion of the South Bunbury to Capel CoastalCorridor. Unpublished Draft Report. Departmentof Environmental Protection.

Dickman CR. (1992). Conservation of Mammals in theAustralasian Region: the Importance of Islands. In:Australia and the Global Environmental Crisis. (JNColesJM Drew, Eds). Academy Press. Canberra, pp.175–214.

Dickman CR (1996). Overview of the impacts of feralcats on Australian native fauna. Australian NatureConservation Agency. Canberra.

Dillon M (1993). Summary of Studies Involving theMainland Quokka in the Dwellingup District Sincethe Early 1970s. Unpublished Report for theDepartment of Conservation and Land Management.Perth, Western Australia.

Dillon MJ (1996). Summary of all Sites Assessed forQuokka Abundance. Unpublished report preparedfor the Western Australian Department ofConservation and Land Management. Perth. WesternAustralia.

DNRE (2002). Victorian Pest Management. A

Page 25: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka 37

framework for Action. Fox Management Strategy. TheState of Victoria, Department of Natural Resourcesand Environment.

Dortch C, Merrilees D (1971). A Salvage Excavation inDevil’s Lair, Western Australia. Journal of the RoyalSociety of Western Australia 54(4):103–113.

Dortch CE, Merrilees D (1973). Human occupation ofDevil’s Lair, Western Australia during the Pleistocene.Archaeology & Physical Anthropology in OceaniaVIII(2):89–115.

Dunnet GM (1962). A population study of the quokka,Setonix brachyurus Quoy and Gaimard (Marsupialia).II. Habitat, movements, breeding and growth.CSIRO Wildlife Research 7(1):13–32.

Dunnet GM (1963). A population study of the quokka,Setonix brachyurus Quoy and Gaimard (Marsupialia).III. The estimation of population parameters bymeans of the recapture technique. CSIRO WildlifeResearch 8:78–117.

Eldridge MDB, Close RL. (1995). Allied rock-wallaby,Petrogale assimilis. In: The Mammals of Australia.(R Strahan, Ed.). Reed Books. Sydney, pp. 365–366.

ESRI. (1999–2004). ArcGIS 9. Environmental SystemsResearch Institute, Inc.

Evans HD (undated). Pioneering in the Karri Country.Pemberton-Northcliffe Tourist Bureau.

Evans M, Gordon G. (1995). Bridled Nailtail Wallaby,Onychogalea fraenata. In: The Mammals of Australia.(R Strahan, Ed.). Reed Books. Sydney, pp. 356–358.

Friend JA (1990). The Numbat Myrmecobius fasciatus(Myrmecobiidae): history of decline and potential forrecovery. Proceedings of the Ecological Society ofAustralia 16:369–377.

Friend JA, Thomas ND. (1994). Reintroduction andthe numbat recovery programme. In: ReintroductionBiology of Australian and New Zealand Fauna. (MSerena, Ed.). Surrey Beatty and Sons. Sydney, pp.189–198.

Friend JA, Butler T (nominally 1998). Gilbert’s PotorooRecovery Team Annual Report 1998. Unpublishedreport for the Western Australian Department ofConservation and Land Management. Albany,Western Australia.

Friend JA, Butler T (nominally 1999). Gilbert’s PotorooRecovery Team Annual Report 1999. Unpublishedreport for the Western Australian Department ofConservation and Land Management. Albany,Western Australia.

Gardner CA (1957). The fire factor in relation to thevegetation of Western Australia. The WesternAustralian Naturalist 5(7):166–173.

Gaynor A (2000). Report on the history of the arrival ofthe feral cat population in Western Australia.CALMScience 3(2):149–179.

Gibb DGA, Kakulas BA, Perrett DH et al. (1966).Toxoplasmosis in the Rottnest quokka (Setonixbrachyurus). Australian Journal of ExperimentalBiology and Medical Science 44:665–672.

Gibson DF, Lundie-Jenkins G, Langford DG et al.(1994). Predation by Feral Cats, Felis catus, on theRufous Hare-wallaby, Lagorchestes hirsutus, in theTanami Desert. Australian Mammalogy 17:103–108.

Gilfillan S. (unpublished). Unpublished database recordsof fauna occurrence from the south coast region ofWestern Australia.

Glauert L (1926). A List of Western Australian Fossils.Geological Survey of Western Australia Bulletin 88:36–71.

Glauert L (1933). The distribution of the marsupials inWestern Australia. Journal of the Royal Society ofWestern Australia 19:17–32.

Glauert L (1948). The cave fossils of the south-west.The Western Australian Naturalist 1:100–104.

Glauert L (1950). The development of our knowledgeof the marsupials of Western Australia. Journal of theRoyal Society of Western Australia:115–134.

Gooding CD (1955). The vermin bonus system inWestern Australia. Part 1 – The distribution ofpayments. The Journal of Agriculture of WesternAustralia 4(4):433–449.

Gould J (1863). The Mammals of Australia. Volume ll.The author. 26 Charlotte Street, BedfordSquare,London.

Government Gazette of Western Australia. (1933). TheVermin Act, 1918. Proclamation. Perth, WesternAustralia.

Green N. (1989). Aborigines of the Albany Region 1821–1898. The Bicentennial Dictionary of WesternAustralians. Volume VI. The University of WesternAustralia Press, Perth, Western Australia.

Haigh SA, Gaynor WT, Morris KD. (1994). A healthmonitoring program for captive, wild and translocatedchuditch (Dasyurus geoffroii). In: Proceedings of the1994 Conference of the Australian Association ofVeterinary Conservation Biologists. Canberra. (AWEnglishGR Phelps, Eds). Australian Association ofVeterinary Conservation Biologists. 52–66.

Hallam SJ (1975). Fire and Hearth a study of Aboriginalusage and European usurpation in south-westernAustralia. Australian Institue of Aboriginal Studies.Canberra.

Hart R (1977). Report on trapping program andassessment of health of quokkas at Holyoake Site,near Dwellingup. Unpublished report.

Hart RP (1981). The ecology of Salmonella in a wildmarsupial, the Quokka (Setonix brachyurus). PhDthesis. Department of Zoology. University of WesternAustralia. Perth

Hart RP, Bradshaw SD, Iveson JB (1986). Salmonellainfections and animal condition in the mainland andBald Island populations of the quokka (Setonixbrachyurus: Marsupialia). Journal of the Royal Societyof Western Australia 69(1):7–11.

Hayward MW (2002). The ecology of the quokka(Setonix brachyurus) (Macropodidae: Marsupialia) inthe northern jarrah forest of Australia. PhD thesis.

Page 26: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

38 P.J. de Torres et al.

School of Biological, Earth and EnvironmentalScience. University of New South Wales. Sydney

Hayward MW, de Tores PJ, Dillon MJ et al. (2003).Local population structure of a naturally occurringmetapopulation of the quokka (Setonix brachyurusMacropodidae: Marsupialia). Biological Conservation110:343–355.

Hayward MW, de Tores PJ, Dillon MJ et al. (2005).Using faecal pellet counts along transects to estimatequokka (Setonix brachyurus) population density.Wildlife Research 32(6):503–507.

Heatwole H. (1987). Major components and distributionof the terrestrial fauna. In: Fauna of Australia.General Articles. Vol. 1A (GR DyneDW Walton,Eds). Australian Government Publishing Service.Canberra, pp. 101–135.

Herrick EH (1961). Some preliminary experiments onadrenal function during seasonal stresses in a wildmarsupial (Setonix brachyurus). Journal of the RoyalSociety of Western Australia 44(2):61–64.

Hilton-Taylor C. (2000). 2000 IUCN Red List ofThreatened Species. IUCN, Gland, Switzerland andCambridge, UK.

Hodgkin EP, Sheard K (1959). Rottnest Island: theRottnest Biological Station and recent scientificresearch. 42(3):65–95.

Holsworth WN (1964). Marsupial Behaviour with specialreference to Population Homeostasis in the Quokkason the West End of Rottnest Island. PhD Thesis.Department of Zoology. University of WesternAustralia. Perth, Western Australia

Holsworth WN (1967). Population dynamics of thequokka, Setonix brachyurus, on the west end ofRottnest I., Western Australia. I. Habitat anddistribution of the quokka. Australian Journal ofZoology 15:29–46.

Horsup A, Evans M (1993). Predation by feral cats,Felis catus, on an endangered marsupial, the bridlednailtail wallaby, Onychogalea franenata. AustralianMammalogy 16(1):83–84.

How RA, Dell J, Humphreys WF (1987). The groundvertebrate fauna of coastal areas between Busseltonand Albany, Western Australia. Records of the WesternAustralian Museum 13(4):553–574.

Jarman P. (1986). The red fox – an exotic, large predator.In: The ecology of exotic animals and plants: someAustralian case histories. (RL Kitching, Ed.). JohnWiley and Sons. Brisbane, pp. 45–61.

Jenkins CFH (1974). The decline of the Dalgite (Macrotislagotis) and other wild life in the Avon Valley. TheWestern Australian Naturalist 12(8):169–172.

Johnson KA, Burbidge AA, McKenzie NL. (1989).Australian Macropodoidea: Status, causes of declineand future research and management. In: Kangaroos,Wallabies and Rat-Kangaroos. Vol. 2 (G Grigg, PJarmanI Hume, Eds). Surrey Beatty & Sons PtyLimited. Chipping Norton, New South Wales, pp.641–657.

Jones M. (1995). Tasmanina Devil, Sarcophilus harrisii.

In: The Mammals of Australia. (R Strahan, Ed.).Reed Books. Sydney, pp. 82–84.

Kabay ED, Start AN (1976). Results of the Search forthe Potoroo in South West and South Coast ofWestern Australia 1975/76. Unpublished reportprepared for the Western Australian Department ofFisheries and Wildlife. Perth, Western Australia.

Kimber PC. (1974). Some effects of prescribed burningon jarrah forest birds. In: 3rd Fire Ecology Symposium.Monash University, Melbourne. 49–57.

King DR, Smith LA (1985). The distribution of theEuropean red fox (Vulpes vulpes) in Western Australia.Records of the Western Australian Museum 12(2):197–205.

King DR, Oliver AJ, Mead RJ (1981). Bettongia andfluoroacetate: a role for 1080 in fauna management.Australian Wildlife Research 8:529–536.

Kinnear JE, Onus ML, Bromilow RN (1988). Foxcontrol and rock-wallaby population dynamics.Australian Wildlife Research 15:435–450.

Kinnear JE, Onus ML, Sumner NR (1998). Fox Controland rock-wallaby population dynamics – II. Anupdate. Wildlife Research 25(1):81–88.

Kinnear JE, Sumner NR, Onus ML (2002). The redfox in Australia – an exotic predator turned biocontrolagent. Biological Conservation 108:335–359.

Kirke A (1983). Quokka, Setonix brachyurus, at GreenRange. The Western Australian Naturalist15(6):146.

Kitchener DJ (1970). Aspects of the Response of theQuokka to Environmental Stress. PhD Thesis.University of Western Australia. Perth, WesternAustralia

Kitchener DJ (1972). The importance of shelter to thequokka, Setonix brachyurus (Marsupialia), on RottnestIsland. Australian Journal of Zoology 20:281–299.

Kitchener DJ (1973). Notes on home range andmovement in two small macropods, the potoroo(Potorous apicalis) and the quokka (Setonixbrachyurus). Mammalia 37:231–240.

Kitchener DJ. (1995). Quokka, Setonix brachyurus. In:The Mammals of Australia. (R Strahan, Ed.). ReedBooks. Sydney, pp. 401–403.

Kitchener DJ, Vicker E (1981). Catalogue of modernmammals in the Western Australian Museum 1895to 1981. Western Australian Museum. Perth.

Kitchener DJ, Chapman A, Barron G. (1978). Mammalsof the Northern Swan Coastal Plain. In: FaunalStudies of the Northern Swan Coastal Plain. AConsideration of Past and Future Changes. TheWestern Australian Museum, for the Department ofConservation and Environment. Perth, WesternAustralia.

Krefft G (1867). List of specimens collected by MrGeorge Masters, Assistant Curator, during the year1866. Annual Report. Australian Museum. pp: 6–7.

Krefft G (1869). Specimens collected by Mr GeorgeMasters, Assistant Curator, in Western Australia.Annual Report. Australian Museum. pp: 7–8.

Page 27: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka 39

Lamont BB, Ward DJ, Eldridge J et al. (2003). Believingthe Balga: a new method for gauging the fire historyof vegetation using grass trees. In: Fire in Ecosystemsof south-west Western Australia: impacts andmanagement. (I AbbottN Burrows, Eds). BackhuysPublishing, Leiden, the Netherlands. Perth, WesternAustralia, pp. 147–169.

Lane-Poole CE (1921). File note dated 17 May 1921 inWestern Australian Government State Records OfficeArchives: 56/21 AN 108 Acc 477. Game Act –Diseases among Quaggas (Wallibies) in South West.

Laurance WF, Harrington GN (1997). Ecologicalassociations of feeding sites of feral pigs in theQueensland wet tropics. Wildlife Research 24(5):579–590.

Lenghaus C, Obendorf DL, Wright FH. (1990).Veterinary aspects of Perameles gunnii biology withspecial reference to species conservation. In:Management and Conservation of Small Populations.Proceedings of a conference held in Melbourne,Australia. September 26–27, 1989. (TW ClarkJHSeebeck, Eds). Chicago Zoological Society.Brookfield, Illinois.

Liddelow GL (2006). Rapid survey of quokka (Setonixbrachyurus) in the southern forests. In AnnualResearch Activity Report. July 2005 – June 2006.Science Division. Discovering the nature of WA.Department of Consrevation and Land Management.Perth, Western Australia. pp: 71–72.

Liddelow GL, Wheeler IB, Kavanagh RP. (2002). Owlsin the Southwest Forests of Western Australia. In:Ecology and Conservation of Owls. (I Newton, RKavanagh, J Olsen et al., Eds). CSIRO. Melbourne,pp. 233–241.

Lilley I (1993). Recent research in southwestern WesternAustralia: a summary of initial findings. AustralianArchaeology 39:34–41.

Long JL (1988). Introduced birds and mammals inWestern Australia. Technical Series 1 (2nd Edn).Agricultural Protection Board, Perth, WesternAustralia.

Lowry DC. (1967). Geological Survey of WesternAustralia. Bureau of Mineral Resources, Geologyand Geophysics, Perth, Western Australia.

Lundelius E (1957). Additions to knowledge of theranges of Western Australian mammals. The WesternAustralian Naturalist 5:173–182.

Lundelius EL (1960). Post Pleistocene Faunal Successionin Western Australia and its Climatic Interpretation.Report of the International Geological Congress. XXISession. Part IV Chronology and Climatology of theQuaternary. Norden. 1960. pp: 142–153.

Lundelius EL (1966). Marsupial carnivore dens inAustralian caves. Studies in Speleology 1(4):174–181.

Lunney D, Matthews A, Grigg J (2001). The diet ofAntechinus agilis and A. swainsonii in unlogged andregenerating sites in Mumbula State Forest, south-eastern New South Wales. Wildlife Research28(5):459–464.

Main AR (1961). The occurrence of macropodidae onislands and its climatic and ecological implications.Journal of the Royal Society of Western Australia44(3):84–89.

Main AR, Yadav M (1971). Conservation of macropodsin reserves in Western Australia. BiologicalConservation 3(2):123–133.

Main AR, Shield JW, Waring H. (1959). Recent studieson marsupial ecology. In: Biogeography and Ecologyin Australia. (RL Keast, RL Crocker, CS Christian,Eds). The Hague, Netherlands, pp. 315–331.

Marsack P, Campbell G (1990). Feeding behaviour anddiet of dingoes in the Nullarbor Region, WesternAustralia. Australian Wildlife Research 17:349–357.

Maxwell S, Burbidge AA, Morris K (Eds.) (1996). The1996 Action Plan For Australian Marsupials andMonotremes. Endangered Species Program. ProjectNumber 500. Wildlife Australia, Canberra, ACT.

McCaw L, Hamilton T, Rumley C. (2005). Applicationof fire hstory records to contemporary managementissues in south-west Australian forests. In: A forestconscienceness: proceedings 6th National Conferenceof the Australian Forest History Society Inc, 12–17September 2004. Augusta, Western Australia. (MCalver, H Bigler-Cole, G Bolton et al., Eds).Millpress, Rotterdam.

Mead RJ, Twigg LE, King DR et al. (1985). Thetolerance to fluoroacetate of geographically separatedpopulations of the quokka (Setonix brachyurus).Australian Journal of Zoology 21(6):503–511.

Meagher SJ (1974). The food resources of the aboriginesof the south-west of Western Australia. Records of theWestern Australian Museum 3:14–65.

Merrilees D (1965). Quokka at Yanchep in historic time.The Western Australian Naturalist 10(1):18.

Merrilees D (1968). Man the destroyer: late Quaternarychanges in the Australian marsupial fauna. Journalof the Royal Society of Western Australia 51(1):1–24.

Merrilees D, Dix WC, Hallam SJ et al. (1973).Aboriginal man in southwest Australia. Journal of theRoyal Society of Western Australia 56:44–55.

Middleton E (2001). Nuyts Wilderness Wildfire ImpactReport. 7–11 March 2001, Walpole, WesternAustralia. Unpublished report prepared for theDepartment of Conservation and Land Management,Perth, Western Australia.

Molsher RL (1999). The ecology of feral cats, Felis catus,in the open forest in New South Wales: interactionswith food resources and foxes. PhD Thesis. Schoolof Biological Sciences. University of Sydney. Sydney

Morton SR. (1990). The impact of European settlementon the vertebrate animals of arid Australia: aconceptual model. In: Australian Ecosystems: 200years of Utilization, Degradation and Reconstruction.Geraldton, Western Australia. (DA Saunders, AJMHopkinsRA How, Eds). Ecological Society ofAustralia. Proceedings of the Ecological Society ofAustralia, Vol 16: 201–213.

Newsome AE. (2001). The biology and ecology of thedingo. In: A Symposium on the Dingo. (CR Dickman,

Page 28: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

40 P.J. de Torres et al.

D Lunney, Eds). Royal Zoological Society of New SouthWales. Mosman, New South Wales, pp. 20–33.

Newsome AE, Corbett LK, Catling PC et al. (1983).The feeding ecology of the dingo. I. Stomach contentsfrom trapping in southeastern Australia, and the non-target wildlife also caught in dingo traps. AustralianWildlife Research 10:477–486.

Nicholls DG (1971). Daily and seasonal movements ofthe quokka, Setonix brachyurus (Marsupialia), onRottnest Island. Australian Journal of Zoology 19:215–226.

NSW NPWS (2001). Threat Abatement Plan forPredation by the red fox (Vulpes vulpes). NSW NationalParks and Wildlife Service. Hurstville, Sydney.

Packer WC (1965). Environmental influences on dailyand seasonal activity in Setonix brachyurus (Quoy andGaimard) (Marsupialia). Animal Behaviour 13:270–283.

Packer WC (1968). Eosinophils and population densityin the marsupial Setonix. Journal of Mammalogy49(1):124–126.

Pavlov PM. (1995). Pig, Sus scrofa. In: The Mammals ofAustralia. (R Strahan, Ed.). Reed Books. Sydney,pp. 698–699.

Pavlov PM, Crome FHJ, Moore LA (1992). Feral pigs,rainforest conservation and exotic disease in northQueensland. Wildlife Research 19(2):179–194.

Pearce AT (1969). File note dated 24 February 1969 inWestern Australian Government State Records OfficeArchives: 65/50 Acc 477. Folio 120. F.C. Act –Idnetification of Fauna – General File.

Perry DH (1971). Observations of a young naturalistin the Lower Blackwood Valley in the year 1919. TheWestern Australian Naturalist 12(2):46–47.

Perry DH (1973). Some notes on the decline andsubsequent recovery of mammal populations in thesouth-west. The Western Australian Naturalist12(6):128–130.

Porter JK (1979). Vertebrate remains from a stratifiedHolocene deposit in Skull Cave, Western Australia,and a review of their significance. Journal of the RoyalSociety of Western Australia 61(4):109–117.

Prince RIT (1984). Exploitation of Kangaroos andWallabies in Western Australia I. A Review to 1970,with special reference on Red and Western GreyKangaroos. Wildlife Research Bulletin Number 13Western Australia. Department of Fisheries andWildlife, Perth, Western Australia.

Prince RIT. (1995). Banded hare-wallaby, Lagostrophusfasciatus. In: The Mammals of Australia. (R Strahan,Ed.). Reed Books. Sydney, pp. 406–408.

Recher HF, Lim L. (1990). A review of current ideas ofthe extinction, conservation and management ofAustralia’s terrestrial vertebrate fauna. In: AustralianEcosystems: 200 years of Utilization, Degradation andReconstruction. Geraldton, Western Australia. (DASaunders, AJM HopkinsRA How, Eds). EcologicalSociety of Australia. Proceedings of the EcologicalSociety of Australia, Vol.16: 287–301.

Richards JD, Short J (1996). History of the disappearanceof native fauna from the Nullarbor Plain through theeyes of long term resident Amy Crocker. The WesternAustralian Naturalist 21(2):89–96.

Ride WDL (1970). A Guide to the Native Mammals ofAustralia. Oxford University Press. Melbourne.

Risbey DA, Calver MC, Short JC et al. (2000). Theimpact of cats and foxes on the small vertebrate faunaof Heirisson Prong, Western Australia. WildlifeResearch 27(3):223–236.

Robertshaw JD, Harden RH (1985). The ecology ofthe dingo in north-eastern New South Wales III.Analysis of macropod bone fragments found in dingoscats. Australian Wildlife Research 12:163–171.

Roe R (1971). Trial excavation in a small cave, Gingin.The Western Australian Naturalist 11:183–184.

Rolls EC (1969). They All Ran Wild: The enthrallingstory of pests on the land in Australia. Angus andRobertson Ltd. Sydney.

Rounsevell DE, Mooney N. (1995). Thylacine,Thylacinus cynocephalus. In: The Mammals ofAustralia. (R Strahan, Ed.). Reed Books. Sydney,pp. 164–165.

Sadleir RMFS (1959). Comparative Aspects of theEcology and Physiology of Rottnest and ByfordPopulations of the Quokka (Setonix brachyurus Quoyand Gaimard). Honours thesis. University of WesternAustralia. Perth. Western Australia

Saunders G, Kay B (1996). Movements and home rangesof feral pigs (Sus scrofa) in Kosciusko National Park,New South Wales. Wildlife Research 23(6):711–720.

Schmidt W, Mason M (1973). Fire and fauna in thenorthern jarrah forest of Western Australia. TheWestern Australian Naturalist 12(6):162–164.

Schodde R, Tidemann SC (Eds.) (1982). The Reader’sDigest Complete Book of Australian Birds. Reader’sDigest Services Pty Ltd, Sydney.

Serventy DL (1951). Inter-specific competion on smallislands. The Western Australian Naturalist 3(1):59–60.

Serventy DL, Glauert L, Carnaby IC et al. (1954). Therecent increase of the rarer native mammals. TheWestern Australian Naturalist 4:128–141.

Serventy VN (1953). The archipelago of the Recherche.4. Mammals. Australian Geographical Society Report1:40–48.

Sharman GB (1954). The relationships of the quokka(Setonix brachyurus). The Western AustralianNaturalist 4:159–168.

Shield JW (1959). Rottnest Island: the Rottnest BiologicalStation and recent scientific research. 15. – Rottnestfield studies concerned with the Quokka. Journal ofthe Royal Society of Western Australia 42(3):76–82.

Shield JW (1964). A breeding season difference in twopopulations of the Australian macropod marsupial(Setonix brachyurus). Journal of Mammalogy45(4):616–625.

Short J, Kinnear JE, Robley A (2002). Surplus killing

Page 29: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka 41

by introduced predators in Australia – evidence forineffective anti-predator adaptions in native preyspecies? Biological Conservation 103:283–301.

Short J, Bradshaw SD, Giles J et al. (1992).Reintroduction of macropods (Marsupialia:Macropodoidea) in Australia – a review. BiologicalConservation 62:189–204.

Short JC, Calaby JH (2001). The status of Australianmammals in 1922 – collections and field notes ofmuseum collector Charles Hoy. Australian Zoologist31(4):533–562.

Shortridge GC (1909). An account of the geographicaldistribution of marsupials and monotremes of south-western Australia having special reference to thespecimens collected during the Balston Expeditionof 1904–1907. Proceedings of the Zoological Society(London) 1909:803–848.

Sinclair EA (1999). Genetic variation in two endangeredmarsupials, the quokka, Setonix brachyurus, andGilbert’s potoroo, Potorous gilbertii. PhD thesis.University of Western Australia. Perth, WesternAustralia

Smith AP, Quinn DG (1996). Patterns and causes ofextinction and decline in Australian conilurinerodents. Biological Conservation 77:243–267.

Smitz A (1998). Progress report on fate of quokkastranslocated to Karakamia Sanctuary, east of Perth.Unpublished report for Karakamia Sanctuary for theDepartment of Conservation and Land Management,Perth Western Australia.

Spencer PBS (1991). Evidence of predation by a feralcat Felis catus (Carnivora: Felidae) on an isolatedrock-wallaby colony in tropical Queensland.Australian Mammalogy 14:143–144.

Stewart DWR (1936). Notes on marsupial damage inpine plantations. Australian Forestry 1(2):41–42.

Storr GM (1963). Some factors inducing change in thevegetation of Rottnest Island. The Western AustralianNaturalist 9(1):15–22.

Storr GM (1964a). Studies on marsupial nutrition. IV.Diet of the quokka, Setonix brachyurus (Quoy &Gaimard), on Rottnest Island, Western Australia.Australian Journal of Biological Science 17(2):469–481.

Storr GM (1964b). The environment of the quokka(Setonix brachyurus) in the Darling Range, WesternAustralia. Journal of the Royal Society of WesternAustralia 47(1):1–2.

Storr GM (1965). Notes on Bald Island and the adjacentmainland. The Western Australian Naturalist9(8):187–196.

Thackway R, Cresswell ID (Eds.) (1995). An InterimBiogeographical Regionalisation for Australia: aFramework for Setting Priorities in the NationalReserves Cooperative Program. Version 4.0. AustralianNature Conservation Agency, Canberra.

Thomas O (1906). On mammals collected in South-west Australia for Mr. W. E. Balston. Proceedings ofthe Zoological Society of London 1906:468–478.

Thomson PC (1992). The behavioural ecology ofdingoes in north-western Australia. III. Hunting andfeeding behaviour, and diet. Wildlife Research19(5):531–542.

Troughton E (1965). Furred Animals of Australia.Angus and Robertson. Sydney, London, Melbourne.

Turney CSM, Bird MI, Fifield LK et al. (2001). Earlyhuman occupation at Devil’s Lair, southwesternAustralia 50,000 years ago. Quaternary Research55:3–13.

Vernes K, Dennis A, Winter J (2001). Mammalian dietand broad hunting strategy of the dingo (Canisfamiliaris dingo) in the wet tropical rain forests ofnortheastern Australia. Biotropica 33(2):339–345.

Wallace WR (1966). Fire in the jarrah forest. Journal ofthe Royal Society of Western Australia 49(2):33–44.

Walton DW (Ed.).) (1988). Zoological catalogue ofAustralia. 5 Mammalia. AGPS, Canberra.

Ward D, Sneeuwjagt R (1999). Believing the balga. Atime capsule of fire management. Landscope14(3):10–16.

Ward DJ, Lamont BB, Burrows CL (2001). Grasstreesreveal contrasting fire regimes in eucalypt forestsbefore and after European settlement in southwesternAustralia. Forest Ecology and Management 150:323–329.

Waring H (1956). Marsupial studies in Western Australia.The Australian Journal of Science 18:66–73.

Watts CHS (1969). Distribution and habits of the rabbitbandicoot. Transactions of the Royal Society of SouthAustralia 93:135–141.

Wayne A, Ward C, Rooney J et al. (2000). The immediateimpacts of timber harvesting and associated activitieson the Ngwayir (Pseudocheirus occidentalis) in theJarrah forest of Kingston State Forest Block. ProgressReport December 2000. Unpublished reportprepared for the Western Australian Department ofConservation and Land Management.

Weston LN (1921). File note dated 18 March 1921 inWestern Australian Government State Records OfficeArchives: 56/21 AN 108 Acc 477. Game Act –Diseases among Quaggas (Wallibies) in South West.

White SR (1952). The occurrence of the quokka in thesouth-west. The Western Australian Naturalist3(5):101–103.

Whitehouse SJO (1977). The diet of the Dingo inWestern Australia. Australian Wildlife Research4:145–150.

Wilson BA, Friend GR (1999). Responses of Australianmammals to disturbance factors. AustralianMammalogy 21(1):87–105.

Woinarski JCZ, Milne DJ, Wanganeen G (2001).Changes in mammal populations in relatively intactlandscapes of Kakadu National Park, NorthernTerritory, Australia. Austral Ecology 26:360–370.

Page 30: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

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Table 1

Fossil and sub-fossil records of Setonix brachyurus. The site names are listed in a north to south latitudinal cline and are mapped in Figure 2. Bolding and an asterisk indicatethe deposits at that site are considered to be part of the original (modern) mammal fauna as noted by the source for the record or as inferred by the age of the deposit. Boldingonly indicates a more subjective assessment of age, where deposits were implied as part of the original (modern) mammal fauna. The abbreviation BP means before present.

Site name Location Description Comment Age Source Comment on Setonix record

Hastings Cave / 8 km north east of Jurien, Described by (Lundelius Hastings Cave / Drover’s Cave listed 5 900 ± 140 years BP Baynes (1979) Listed from 1 layer only,Drovers Cave * approx 210 km north 1957; Lundelius 1960) as by (Lundelius 1957) as representative (Lundelius 1960), however mammal occurrences were from

of Perth entrance formed by collapse of Jurien Bay caves. Bone deposits Lundelius (1957) reported top deposits considered to lessof roof. Cave formed from were concentrated near cave mouth most foot of surface deposits to than 1,000 years old andarches developing (Lundelius 1957) and thought to be represent fauna prior to effect considered to be part of the(coherence) over dissolved derived from owl pellets. Concurs with of introduced species. Also original faunaaeolian calcarenite Baynes (1979). Fragmentation of listed presence of rabbits and(Baynes 1979). bones of larger mammals considered house mouse and interpreted Lundelius (1960) Lower levels only

to be indicative of mammalian this as indicating youngerpredator accumulation (Baynes deposits. Baynes (1979) listed Merrilees (1968) Merrilees noted the only1979), with a minor contribution as a ages (Hastings Cave) as 400 ± records of Setonix from theresult of human occupation. 70 years BP at a depth of Moore River – Dongara region

10 mm to 11 400 ± 200 years are from lower parts of HastingsBP at 2.98m. Cave deposit. However,

Jefferys (pers comm. to MWH)subsequently listed Setonixoccurrence at Echidna Cave(Dongara), an un-named cavenear the northern boundary ofNambung National Park, HouseCave (Moore – Dongara riverarea), Greensand Cliffs (Gingin)and McIntyre Gully (Gingin)

Yanchep * North of Perth A complete quokka skeleton The skull was penetrated by a (Merrilees 1965) Skeleton with iron rod throughrecovered in 1965 registered small iron stake implanted skullwith the WAM as part of the during the animal’s life time,fossil collection. thereby inferring persistence of

quokkas in Perth area inmodern times

Orchestra Shell Wanneroo, North of 4 stratigraphic levels, Lair, deposits presumed to be not specified Archer (1974b) Lower two levelsCave Perth. Location noted as progressive faunal accumulated by Sarcophilus.

approx 16km south of impoverishment towards top.Yanchep cave describedby Lundelius (1957)

Murray Cave Approximately 40km Considered to represent a 3 090 ± 90 years BP. Contains (Archer 1974a)north of Perth carnivore’s accumulation, youngest dated occurrence

typical of Sarcophilus (from charcoal at 1–7cm depth)of thylacine

Rainbow Cave * 1 km inland, 1.5km south Collapsed limestone cave, Lilley (1993) aged at 340 ± 45 Lilley (1993) At depths of 10,15, 25, 35, 50of the mouth of the 30m deep, 15m wide years BP at depth of 5–10cm; and 75cm. Deposits aboveMargaret river, south- 790 ± 50 years BP at depth of 30cm are therefore presumablywest Western Australia 25–30cm; 8340 ± 45 years BP at from the modern fauna

depth of 35–40cm; 4150 ± 70years BP at depth of 70–75cm;and a second age of 1030 ± 50

Page 31: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka 43ye

ars

BP

was

det

erm

ined

at

dept

h of

25–

30cm

Mam

mot

h C

ave

Mar

gare

t R

iver

are

a,W

as t

he f

irst

cave

inD

epos

its c

onsi

dere

d by

Gla

uert

Lund

eliu

s (1

960)

dat

ed d

epos

itsG

laue

rt (

1948

)so

uth

wes

t W

este

rnW

este

rn A

ustr

alia

to

yiel

d(1

948)

to

be w

ater

lain

. M

erril

ees

from

the

upp

er s

and

laye

r as

Aus

tral

iafo

ssil

vert

ebra

te r

emai

ns &

(196

8) c

onsi

dere

d de

posi

ts w

ere

grea

ter

than

37

000

year

s B

P.C

ook

(196

3)de

scrib

ed b

y G

laue

rt (

1948

).m

ore

cons

iste

nt w

ith t

alus

and

not

Mer

rilee

s (1

968)

dat

ed u

pper

Orig

inal

ly r

ecog

nise

d as

wat

er t

rans

port

ed a

s su

gges

ted

part

s of

dep

osit

as 3

1,00

0M

erril

ees

(196

8)ha

ving

tw

o st

ratig

raph

icby

Gla

uert

year

s B

Pun

its:

a ye

llow

-red

san

dLu

ndel

ius

(196

0)la

yer

abov

e co

arse

red

san

d.B

oth

laye

rs c

appe

d w

ithtr

aver

tine

(Lun

deliu

s 19

60).

Rem

nant

s of

a p

revi

ousl

ylo

nger

tun

nel

in c

alca

reou

sae

olia

nite

s

Mar

gare

t R

iver

not

desc

ribed

Gla

uert

(19

26)

refe

rred

to

colle

ctio

nsP

leis

toce

neG

laue

rt (

1926

)ca

ves

from

‘cer

tain

cav

es n

ear

the

Mar

gare

tR

iver

in

the

extr

eme

Sou

th-W

est’.

Loca

tion

take

n no

min

ally

her

e as

Mam

mot

h C

ave

Har

ley’

s C

ave

Cap

e Le

euw

in –

Cap

eno

t de

scrib

edLi

sted

by

Mer

rilee

s (1

968)

as

List

ed a

s fa

una

char

acte

ristic

of

Mer

rilee

s (1

968)

Nat

ural

iste

reg

ion,

sou

thre

pres

enta

tive

of s

peci

men

s co

llect

edth

e re

gion

for

som

e hu

ndre

d to

wes

t W

este

rn A

ustr

alia

by L

owry

(19

67)

from

sur

face

litte

ra

few

tho

usan

d ye

ars

prio

r to

from

cav

es i

n C

ape

Leeu

win

–Cap

eE

urop

ean

settl

emen

tN

atur

alis

te r

egio

n. H

owev

er,

noin

dica

tion

of t

hese

col

lect

ions

bei

ngm

ade

by L

owry

(19

67)

who

dis

cuss

edge

omor

phol

ogy

Lake

Cav

eM

arga

ret

Riv

er a

rea,

not

desc

ribed

not

give

nG

laue

rt (

1948

)so

uth

wes

t W

este

rnA

ustr

alia

Dev

il’s L

air,

orS

outh

wes

t of

Witc

hclif

fe,

Sm

all

cave

in

aeol

ian

Dep

osits

con

side

red

(Lun

deliu

sS

urfic

ial

laye

rs d

escr

ibed

as

Dor

tch

and

Nan

nup

Cav

e of

sout

h w

est

Wes

tern

calc

aren

ite. T

rave

rtin

e flo

or19

60)

to b

e a

resu

lt of

Sar

coph

ilus,

rece

nt a

s 32

0 ye

ars

BP.

Oth

erM

erril

ees

(197

1)Lu

ndel

ius

(196

0;A

ustr

alia

over

red

cla

yey

sand

she

nce

the

nam

e D

evil’

s La

ir.de

posi

ts e

xten

ding

to

35,1

6019

66)

and

thou

ght

to b

e w

ashe

d in

.D

ortc

h an

d M

erril

ees

(197

1)ye

ars

BP

Bal

me

et a

l. (1

978)

.D

ortc

h an

dC

ook

(196

0).

*D

epos

its d

escr

ibed

fro

m t

wo

desc

ribed

as

arte

fact

s an

d fo

odLu

ndel

ius

(196

0) d

ated

the

Mer

rilee

s (1

973)

leve

ls:

imm

edia

tely

bel

owre

mai

ns p

resu

mab

ly le

ft as

a r

esul

t of

uppe

r le

vel

8,50

0 ±

160

year

str

aver

tine

floor

; an

d at

Abo

rigin

al u

se o

f th

e ca

ve.

Som

eB

P, w

ith d

epos

its a

t 4

feet

as

Lund

eliu

s (1

966)

dept

h of

4 f

eet.

rem

ains

may

hav

e be

en l

eft

by12

,175

± 2

75 y

ears

BP

Sar

coph

ilus.

Bay

nes

et a

l. (1

975)

Dor

tch

and

Mer

rilee

s (1

973)

Bal

me

et a

l.R

epre

sent

ed c

onsi

sten

tlyco

nsid

ered

upp

er l

ayer

s m

ay h

ave

liste

d up

perm

ost

stra

tigra

phic

(197

8)th

roug

hout

dep

osit

(all

ages

)be

en a

ccum

ulat

ed b

y hu

man

s an

dun

it to

be

less

tha

n 12

,000

belie

ved

Sar

coph

ilus

was

res

pons

ible

year

s ol

d, w

ith o

ther

maj

orB

ayne

s et

al.

Dep

osits

con

side

red

to b

e pa

rtfo

r su

bseq

uent

ly ‘w

orki

ng o

ver’

stra

tigra

phic

uni

ts a

s 19

,000

–(1

975)

of t

he m

oder

n fa

una

(occ

urrin

gth

e de

posi

ts.

25,0

00 y

ears

old

and

the

in t

he a

rea

imm

edia

tely

pre

botto

m d

epos

it m

ore

than

Eur

opea

n ar

rival

)31

,000

yea

rs o

ld.

Sub

sequ

ent

datin

g te

chni

ques

sup

port

edLu

ndel

ius

(196

0)B

oth

leve

lsth

ese

date

s fo

r up

per

laye

rs a

ndin

dica

ted

low

er l

evel

s m

ay b

e in

Coo

k (1

960)

exce

ss o

f 40

,000

yea

rs o

ld(T

urne

y et

al.

2001

).

Page 32: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

44 P.J. de Torres et al.

Str

ong’

s C

ave,

or

7 m

iles

sout

h of

As

for

Mam

mot

h C

ave

has

aC

entr

al p

ortio

n of

tal

us s

lope

Coo

k (1

963)

Teet

h on

lyS

tron

gs’ C

ave

ofM

amm

oth

Cav

eco

mpl

ex d

evel

opm

ent

has

mor

e re

cent

, yo

ung

depo

sits

,C

ook

(196

3)hi

stor

y. M

ain

depo

sits

in

but

exca

vatio

ns f

rom

str

eam

bed

Mer

rilee

s (1

968)

entr

ance

cha

mbe

r. H

asco

nsid

ered

old

er,

but

youn

ger

talu

s sl

ope

unde

rmin

ed b

yth

an M

amm

oth

Cav

e. S

ome

ofst

ream

act

ion

& c

entr

alM

amm

oth

Cav

e de

posi

ts n

otpo

rtio

n of

tal

us c

olla

psin

g.pr

esen

t in

Str

ong’

s C

ave

also

Mat

eria

l w

ashe

d an

dim

ply

Str

ong’

s C

ave

depo

sits

tran

spor

ted

by s

trea

m a

ndm

ay b

e yo

unge

rth

en m

ixed

with

sed

imen

tary

depo

sits

Two

smal

lN

ear

Turn

er B

rook

,S

urfa

ce/s

ub-s

urfa

ce c

ave

Dep

osits

inc

lude

d un

wor

n43

0 ±

160

BP

bas

ed o

n ra

dio

Arc

her

and

Pre

sent

in

Cav

e 3

only

un-n

amed

cav

esA

ugus

tade

posi

ts w

ithin

8cm

of

Sar

coph

ilus

teet

h w

ith p

oorly

for

med

carb

on d

ated

hai

r fr

om C

ave

1.B

ayne

s (1

972)

(Cav

e 1

and

surf

ace

(Cav

e 1)

and

13c

mro

ots.

Int

erpr

eted

as

juve

nile

ani

mal

s.C

ave

3 m

ay b

e ol

der.

Nei

ther

Cav

e 3)

of s

urfa

ce (

Cav

e 3)

Thi

s, c

ombi

ned

with

kno

wn

cave

had

dep

osits

of

the

geog

raph

ic r

ange

of

med

ium

siz

ein

trod

uced

mam

mal

s (h

ouse

owl

spec

ies

and

the

inac

cess

ibili

ty o

fm

ouse

, bl

ack

rat

or r

abbi

t),

all o

fca

ves,

sug

gest

s de

posi

ts a

res

ult

ofw

hich

may

hav

e ar

rived

pos

tow

l pr

edat

ion.

accu

mul

atio

n of

dep

osits

Brid

e’s

Cav

eM

arga

ret

Riv

er a

rea,

not

desc

ribed

not

give

nG

laue

rt (

1948

)so

uth

wes

t W

este

rnA

ustr

alia

Sku

ll C

ave

Cap

e Le

euw

in r

egio

n of

Com

pris

ed o

f a

larg

e m

ain

May

hav

e fu

nctio

ned

as a

pit

trap

Upp

er l

ayer

(21

–28c

m)

aged

at

Por

ter

(197

9)P

rese

nt a

t de

pths

to

170c

mso

uth

wes

t W

este

rnch

ambe

r w

ith p

artia

llyw

ith a

dditi

onal

dep

osits

res

ultin

g2,

900

± 80

yea

rs B

PA

ustr

alia

colla

psed

roo

f an

d a

smal

ler

from

use

by

owls

Inte

rmed

iate

lay

er (

100–

115c

m)

cham

ber

lead

ing

from

7,87

5 ±

100

year

s B

P. D

epth

of

wes

tern

par

t of

mai

n19

0cm

con

side

red

late

cham

ber.

Bot

h ch

ambe

rsP

leis

toce

nepa

rtia

lly f

illed

with

san

dyse

dim

ents

, po

ssib

ly w

ashe

d in

Sco

tt R

iver

Coa

stal

dun

es s

outh

of

Exp

osed

fos

sil

soils

and

not

give

n, p

rese

nce

of f

ox a

ndB

utle

r (1

969)

the

Sco

tt R

iver

surf

ace

depo

sits

fro

mra

bbit

rem

ains

impl

y de

posi

t is

mob

ile-p

artly

sta

bilis

edm

ix o

f fo

ssil

and

mod

ern

anim

alsa

nd d

unes

rem

ains

Sco

tt R

iver

Bet

wee

n Le

dge

Poi

ntB

ones

col

lect

ed f

rom

colle

cted

in

perio

d 12

Mar

ch 1

976

not

date

d, p

resu

mab

ly s

ub f

ossi

lK

abay

and

Sta

rtP

resu

mab

ly s

ub f

ossi

l or

rec

ent

Are

a *

and

Bla

ck P

oint

sand

dun

esto

22

Mar

ch 1

976

or r

ecen

t. P

rese

nce

of r

abbi

t(1

976)

rem

ains

with

oth

er s

peci

men

sim

plie

s th

ese

are

rece

nt (

part

of

the

mod

ern

faun

a)

Tabl

e 1

(co

nt.)

Site

nam

eLo

catio

nD

escr

iptio

nC

omm

ent

Age

Sou

rce

Com

men

t on

S

eton

ix r

ecor

d

Page 33: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka 45

War

ren

Riv

er /

Des

crib

ed a

s bo

unde

dS

peci

men

s co

llect

ed f

rom

colle

cted

in

perio

d 27

to

31 M

ayno

t da

ted,

pre

sum

ably

sub

Kab

ay a

nd S

tart

Pre

sum

ably

sub

fos

sil

or r

ecen

tD

onne

lly R

iver

by t

he W

arre

n an

dsa

nd d

unes

19

76

foss

il or

rec

ent

(197

6)a

rea

Don

nelly

riv

ers,

the

sout

h co

ast

and

Vas

seH

ighw

ay.

Bon

e m

ater

ial

from

Yea

geru

p S

and

Du

ne

s

Nor

man

’sS

and

blow

out

sout

h of

desc

ribed

onl

y as

bon

e(s)

colle

cted

in

perio

d 15

to

20no

t da

ted,

pre

sum

ably

sub

Kab

ay a

nd S

tart

Pre

sum

ably

sub

fos

sil

or r

ecen

tB

each

/P

lant

agen

et l

ocat

ion

colle

cted

Aug

ust

1976

foss

il or

rec

ent

(197

6)P

lant

agen

et43

48 a

nd 1

km

eas

t of

Nor

man

’s B

each

Wal

pole

Des

crib

ed a

s co

asta

lS

peci

es l

iste

d as

obt

aine

dco

llect

ed i

n pe

riod

15 t

o 20

not

date

d, h

owev

er t

heK

abay

and

Sta

rtP

resu

mab

ly s

ub f

ossi

l or

rec

ent

sand

blo

w o

ut,

in s

and

dune

sA

ugus

t 19

76pr

esen

ce o

f ra

bbit

and

fox

(197

6)W

alpo

le a

rea

rem

ains

with

oth

er s

peci

men

sco

llect

ed i

mpl

ies

thes

e ar

ere

cent

dep

osits

(pa

rt o

f th

em

oder

n fa

una)

Two

Peo

ples

Bay

Eas

t of

Alb

any

Spe

cies

lis

ted

as o

btai

ned

colle

cted

in

perio

d 23

Apr

il 19

76 t

o 5

not

date

d, p

resu

mab

ly s

ubK

abay

and

Sta

rtP

resu

mab

ly s

ub f

ossi

l or

rec

ent

Nat

ure

Res

erve

in s

and

dune

sM

ay 1

976

and

15 t

o 20

Aug

ust

1976

foss

il or

rec

ent.

Pre

senc

e of

(197

6)ra

bbit

rem

ains

with

oth

ersp

ecim

ens

impl

ies

thes

e ar

ere

cent

(pa

rt o

f th

e m

oder

nfa

una)

Pla

ntag

enet

San

d du

nes,

sou

th o

fde

scrib

ed o

nly

as b

one(

s)co

llect

ed i

n pe

riod

15 t

o 20

not

date

d, p

resu

mab

ly s

ubK

abay

and

Sta

rtP

resu

mab

ly s

ub f

ossi

l or

rec

ent

Pla

ntag

enet

Loc

atio

nco

llect

edA

ugus

t 19

76fo

ssil

or r

ecen

t(1

976)

41

30

Will

iam

Bay

Spe

cies

lis

ted

as m

ater

ial

colle

cted

in

perio

d 4

to 1

0no

t da

ted,

pre

sum

ably

sub

Kab

ay a

nd S

tart

Pre

sum

ably

sub

fos

sil

or r

ecen

tN

atio

nal

Par

kpi

cked

up

in s

and

dune

sF

ebru

ary

1976

foss

il or

rec

ent

(197

6)

Boa

t Har

bour

San

d bl

owou

t at

Boa

tde

scrib

ed o

nly

as b

one(

s)co

llect

ed i

n pe

riod

15 t

o 20

not

date

d, p

resu

mab

ly s

ubK

abay

and

Sta

rtP

resu

mab

ly s

ub f

ossi

l or

rec

ent

Har

bour

eas

t of

colle

cted

Aug

ust

1976

foss

il or

rec

ent

(197

6)R

eser

ve 7

723

Page 34: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

46P.J. de Torres et al.

Table 2

Australian Museum records of the quokka, Setonix brachyurus. Additional records by Masters lacking location records are not shown.

Year of Location Collector Form of Comments Australian Museumcollection specimens Registration number

Undated Rottnest Island brain in Formalin M 18429

Undated Rottnest Island G. P. Whitley-Staff skull and mandibles Shown as G. P. Whitney-Staff in Australian Museum database S 1936

Undated Rottnest Island G. P. Whitley-Staff skull only Shown as G. P. Whitney-Staff in Australian Museum database S 1937

Undated Rottnest Island G. P. Whitley-Staff skull, odd mandible Shown as G. P. Whitney-Staff in Australian Museum database S 1938

Undated Rottnest Island Taronga Park Trust skin M 7994

Undated Rottnest Island Taronga Park Zoo skin skull M 8177

1866 King George’s Sound Masters mount Year of collection nominally shown here as 1866, as Masters is known to have P 1048collected in King George’s Sound area from Jan to April 1866 (Abbott 1999; Glauert1950) and Sept 1868 to April 1869 (Glauert 1950), however this record not listed byKrefft (1867; 1869). No co-ordinates listed, co-ordinates for Albany used, howeverMasters known to have collected as far north as the Pallinup River (Salt River)(Glauert 1950) and the Stirling Range (Abbott 1999)

1866 King George’s Sound Masters mount See comment for record P 1048 P 1049

1866 King George’s Sound Masters spirit See comment for record P 1048 P 1060

1866 King George’s Sound Masters spirit See comment for record P 1048 P 1061

1866 King George’s Sound Masters spirit See comment for record P 1048 P 1062

1920 Nornalup near Denmark A. S. Le Souef skin skull Year of collection not listed. Nominally listed here 1920 as Le Souef known to have M 4212collected in Porongurup area in 1920 (Abbott 1999). Co-ordinates map to SouthernOcean, directly south of Nornalup Inlet and Walpole, corrected to map near Nornalup

1920 Nornalup, near Denmark A. S. Le Souef skull only No co-ordinates with this record, corrected co-ordinates for Le Souef record (M 4212) S 1799used. Other comments for record M 4212 apply.

1920 Nornalup, near Denmark A. S. Le Souef skull only No co-ordinates with this record, corrected co-ordinates for Le Souef record (M 4212) S 1800used. Other comments for record M 4212 apply.

1921 King River, 10 miles E. Le G. Troughton skin skull Shown as LE G. Troughton in Australian Museum database M 3083from Albany J. Wright

1921 King River, 10 miles E. Le G. Troughton skin skull Shown as LE G. Troughton in Australian Museum database M 3084from Albany J. Wright

1921 King River, 10 miles E. Le G. Troughton Shown as LE G. Troughton in Australian Museum database M 3085from Albany J. Wright

Page 35: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka 47

19

21

Kin

g R

iver

, 10

mile

sE

. Le

G.

Trou

ghto

nsk

in s

kull

Sho

wn

as L

E G

. Tr

ough

ton

in A

ustr

alia

n M

useu

m d

atab

ase

M 3

086

from

Alb

any

J. W

right

19

21

Prin

cess

Roy

al B

ay,

E.

Le G

. Tr

ough

ton

skin

sku

llS

how

n as

LE

G.

Trou

ghto

n in

Aus

tral

ian

Mus

eum

dat

abas

e. C

o-or

dina

tes

M 3

087

Alb

an

yJ.

Wrig

ht(3

5 de

g 3

min

Sou

th a

nd 1

17 d

eg 5

3min

Eas

t) m

ap t

o w

ithin

Prin

cess

Roy

alH

arbo

ur,

corr

ecte

d to

map

to

map

on

Prin

cess

Roy

al H

arbo

ur f

ores

hore

nea

rA

lban

y to

wns

ite

19

35

Rot

tnes

t Is

land

S.

Larn

ach

spiri

tM

182

14

19

35

Rot

tnes

t Is

land

S.

Larn

ach

spiri

tM

182

15

19

35

Rot

tnes

t Is

land

S.

Larn

ach

spiri

tM

182

16

19

35

Rot

tnes

t Is

land

S.

Larn

ach

spiri

tM

182

17

19

35

Rot

tnes

t Is

land

S.

Larn

ach

spiri

tM

182

18

19

35

Rot

tnes

t Is

land

S.

Larn

ach

spiri

tM

182

19

19

61

Rot

tnes

t Is

land

Taro

nga

Par

ksk

in s

kull

M 9

056

19

87

edge

of

Lake

Bag

dad,

Gav

in G

atte

nby

ske

leto

nM

186

26R

ottn

est

Isla

nd

19

97

Rot

tnes

t Is

land

E.

Dov

eysk

ull

M 3

3146

Page 36: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

48P.J. de Torres et al.

Table 3

Published records of occurrence of the quokka, Setonix brachyurus. Records are listed chronologically.

Year of Location or Source Commentrecord Site name

1658 Rottnest Island Samuel Volckersen (Volckertzoon), Considered the first record of quokka sighting by a European and only the second record of a marsupial from Australiacited by (Alexander 1914) by a EuropeanGlauert (1950)

1696 Rottnest Island Willem de Vlamingh, cited by Mistook the quokka on Rottnest island for ‘ … A kind of rat as big as a common cat … ‘(Alexander 1914) Glauert (1950)

1801 Rottnest Island M. Freycinet,, cited by M. Freycinet, observed a quadruped ‘which the old Dutch navigators actually mistook for a rat’Alexander (1916)

1826 King George’s Sound Records of Quoy and Gaimard Reported the new species, the specimen described as recently dead and ... ‘probably from disease’as cited by Alexander (1916)

1829 King George’s Sound Scott Nind, cited by (Alexander Scott Nind, medical officer at the King George’s Sound penal settlement recorded the presence of the quokka1916) and Glauert (1950)

1829 Rottnest Island Dr T.B. Wilson, cited by Dr T.B. Wilson, visitor to the Swan River Settlement, when visiting Rottnest Island recorded ‘the dogs caught two wallabi’Alexander (1918)

1830 King George Sound Walton (1988) The type specimen of Quoy and Gaimard

1837 Swan River, Perth George Gray, cited by Glauert (1950) reported Gray’s list of species from the Swan River. The list included the quokka Kitchener et al. (1978)

Glauert (1950) noted Gray’s reference to ‘Swan River’ may have included the Darling Range & the York & Avon River valleys. Nominallymapped as Piesse Brook

1842 Coastal areas Gilbert, cited by Gould (1863) Gilbert found the quokka abundant in all the swampy tracts which ‘skirt nearly the whole of Western Australia at a shortdistance from the sea’. Presumably this was referring to the south west of WA only. Gilbert was known to have visited WAand collected from this area in 1839–1840 and 1842–1843 Abbott (1999). Date nominally listed here as 1842 andlocations nominally shown as Albany and Margaret River area

1905 Bald Island Shortridge (1909) see note 1, below

1905 Rottnest Island Shortridge (1909) see note 1, below

1905 Margaret River, Burnside Shortridge (1909) see note 1, below

1905 Busselton, Yallingup Shortridge (1909) see note 1, below

1905 Albany, King River Shortridge (1909) see note 1, below

1905 Albany, King River Thomas (1906) Descriptions of collections made by Shortridge. Listed as collected during end of 1904 and during 1905. Nominally listedhere as 1905 and mapped as same location as Shortridge (1909) for King River

Page 37: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka 49

19

05

Alb

any,

Big

Gro

ve,

Tho

mas

(19

06)

Des

crip

tions

of

colle

ctio

ns m

ade

by S

hort

ridge

. Li

sted

as

colle

cted

dur

ing

end

of 1

904

and

durin

g 19

05.

Nom

inal

ly l

iste

dK

ing

Geo

rge’

s S

ound

here

as

1905

and

nom

inal

ly m

appe

d as

the

bet

wee

n A

lban

y an

d K

ing

Riv

er

19

07

Mar

gare

t R

iver

are

aW

. H

. Loa

ring,

cite

d by

Quo

kkas

rec

orde

d as

ver

y nu

mer

ous

1907

–191

0. L

ocat

ion

nom

inal

ly m

appe

d as

cre

eklin

e in

for

este

d ar

ea,

imm

edia

tely

Whi

te (

1952

)w

est

of M

arga

ret

Riv

er

19

12

Irw

in I

nlet

– M

t F

rank

land

Dia

ries

of S

.W.

Jack

son,

Jack

son’

s di

arie

s fo

r 19

12–1

913

reco

rd ‘c

augh

t a

wal

laby

in

trap

’ and

sle

epin

g un

der

swor

d gr

ass

here

(D

eep

Riv

er)

are

aci

ted

by A

bbot

t (1

998)

inte

rpre

ted

by A

bbot

t as

ref

errin

g to

quo

kkas

. N

omin

ally

sho

wn

as 1

912

and

nom

inal

ly m

appe

d as

Dee

p R

iver

.

19

19

Low

er B

lack

woo

d V

alle

yP

erry

(19

71)

Rec

orde

d th

e qu

okka

as

num

erou

s an

d fr

eque

ntly

see

n in

the

Low

er B

lack

woo

d va

lley.

App

roxi

mat

e lo

catio

n on

ly

19

20

Mar

gare

t R

iver

Hoy

, ci

ted

by S

hort

and

From

col

lect

ions

by

Cha

rles

Hoy

, 19

19–1

922.

Hoy

col

lect

ed 9

spe

cim

ens

from

the

Mar

gare

t R

iver

are

a in

192

0. W

ith t

heC

alab

y (2

001)

exce

ptio

n of

Tric

hosu

rus

vulp

ecul

a, h

e de

scri

bed

none

of

the

colle

cted

mam

mal

s as

ple

ntifu

l. T

he q

uokk

a w

as d

escr

ibed

as s

eldo

m s

een

(due

to

noct

urna

l ha

bits

). L

ocat

ion

nom

inal

ly m

appe

d as

im

med

iate

ly n

orth

wes

t of

Mar

gare

t R

iver

19

20

Un-

nam

ed c

ave

near

Roe

(19

71)

Roe

(19

71)

reco

rded

thi

s lo

catio

n as

a f

ossi

l/sub

foss

il re

cord

and

not

ed l

ong

term

res

iden

ts w

ere

aw

are

quok

kas

(alo

ngG

ingi

nw

ith b

oodi

es,

Bet

tong

ia l

esue

ur,

and

bilb

ies,

Mac

rotis

lag

otis

) w

ere

pres

ent

in t

he d

istr

ict

until

arr

ival

of

the

fox

19

20

Bic

kley

, D

arlin

g S

carp

,W

. H

. Loa

ring,

cite

d by

Not

ed a

s pa

rtic

ular

ly p

lent

iful

in t

he e

arly

192

0s w

ith e

xten

sive

run

way

s in

thi

ck s

crub

bor

derin

g st

ream

sno

w c

onsi

dere

d a

subu

rbW

hite

(19

52)

of P

erth

19

20

Dar

ling

Ran

ge,

Pie

sse

W.H

. Loa

ring

inT

he q

uokk

a de

scrib

ed t

o ha

ve ‘v

anis

hed

from

its

gul

ly h

aunt

s …

30

year

s ag

o’. T

his

reco

rd i

s no

min

ally

sho

wn

asB

rook

- B

ickl

eyS

erve

nty

et a

l. (1

954)

pres

ent

here

30

year

s ea

rlier

(19

20)

and

loca

tion

map

ped

as P

iess

e B

rook

(P

iess

e G

ully

)

19

22

Yarl

oop,

Log

ue B

rook

,W

hite

(19

52)

Rec

orde

d as

num

erou

s in

low

tan

gled

scr

ub w

hen

auth

or w

as a

sch

oolb

oy a

nd s

till

plen

tiful

unt

il 19

26 w

hen

he l

eft

Dar

ling

Ran

ge b

etw

een

scho

ol.

App

roxi

mat

e lo

catio

n on

ly,

nom

inal

ly m

appe

d as

upp

er c

atch

men

t of

Log

ue B

rook

and

dat

e no

min

ally

sho

wn

1922

Per

th a

nd H

arve

y

19

29

Bu

sse

lton

Whi

te (

1952

)R

ecor

ded

as ‘

in g

reat

num

bers

’, lo

catio

n ap

prox

imat

e

19

29

Cap

e Le

euw

in –

Cap

eW

hite

(19

52)

Rec

orde

d as

‘in

gre

at n

umbe

rs’,

loca

tion

appr

oxim

ate

Nat

ural

iste

19

20

s/H

elen

a R

iver

, ne

arP

erry

(19

73)

Rec

orde

d as

abu

ndan

t in

den

se l

ow c

over

alo

ng c

reek

s an

d riv

er c

ours

es f

rom

val

ley

of H

elen

a R

iver

eas

twar

ds.

19

30

sM

unda

ring

Loca

tion

iden

tifie

d as

Gre

ysto

nes

pine

pla

ntat

ion

whi

ch w

as p

lant

ed b

y D

ick

Per

ry i

n th

e 19

20s/

1930

s an

d re

cord

assu

med

to

be f

rom

the

192

0s t

o 19

30s

(Ian

Abb

ott,

pers

. co

m.

to P

JdeT

).

19

30

Nor

thcl

iffe

area

Dau

bney

et

al.

(und

ated

)D

iscu

ssio

ns b

etw

een

one

of u

s (P

JdeT

) an

d lo

cal

land

hold

ers

plus

rep

orts

in

Dau

bney

et

al.

(und

ated

) in

dica

te t

hequ

okka

was

con

side

rabl

y m

ore

abun

dant

in

the

Nor

thcl

iffe

area

prio

r to

the

arr

ival

of

the

fox.

Gla

dys

Buc

king

ham

, sc

hool

teac

her

in N

orth

cliff

e, r

ecal

led

of t

he 1

930s

‘eve

ry s

wam

p w

as a

maz

e of

tra

cks

and

tunn

els

of w

alla

bies

, qu

okka

s,ba

ndic

oots

…’

(Dau

bney

et

al.

unda

ted)

. Lo

catio

n ap

prox

imat

e an

d m

appe

d as

im

med

iate

ly e

ast

of N

orth

cliff

e to

wns

ite,

date

her

e no

min

ally

lis

ted

as 1

930

19

31

Vas

se E

stua

ry,

near

Whi

te (

1952

)Q

uokk

as ‘i

n nu

mbe

rs’ i

n lo

w s

crub

bet

wee

n co

asta

l du

nes,

loc

atio

n ap

prox

imat

eB

uss

elto

n

19

32

Pin

e pl

anta

tions

,S

tew

art

(193

6)R

ecor

ded

as r

espo

nsib

le f

or d

amag

e to

pin

e pl

anta

tions

with

in k

arri

fore

st a

reas

and

rec

orde

d gr

azin

g up

to

2 km

s fr

omP

em

be

rto

nsw

amps

. Lo

catio

n m

appe

d he

re n

omin

ally

as

pine

pla

ntat

ions

, im

med

iate

ly e

ast

of P

embe

rton

19

32

Rot

tnes

t Is

land

Sto

rr (

1963

)S

torr

cite

d H

.T.P

ears

e, a

s th

e fir

st r

efer

ence

(19

32)

to t

he l

arge

num

ber

of q

uokk

as o

n R

ottn

est

Isla

nd b

eing

reg

arde

das

a p

est

Page 38: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

50 P.J. de Torres et al.

19

33

Com

men

ts o

n ge

nera

lG

laue

rt (

1933

)P

ublis

hed

to i

ndic

ate

the

pres

ent

rang

e (in

193

3) o

f m

arsu

pial

fau

na i

n W

este

rn A

ustr

alia

and

as

an u

pdat

e to

the

dist

ribu

tion

dist

ribut

ions

des

crib

ed b

y S

hort

ridge

(19

09).

Des

crib

ed d

istr

ibut

ion

of t

he q

uokk

a as

ext

endi

ng f

rom

Moo

re R

iver

to

the

sout

h co

ast

and

incl

usiv

e of

Rot

tnes

t Is

land

and

Bal

d Is

land

. N

o re

fere

nce

to t

he p

rese

nce

on t

he i

slan

ds o

ff E

sper

ance

as s

ugge

sted

by

Sho

rtrid

ge.

Des

crib

ed a

s ab

unda

nt i

n su

itabl

e sw

ampy

loc

aliti

es.

Nom

inal

ly m

appe

d he

re a

s A

lban

yan

d D

wel

lingu

p, H

olyo

ake

19

33

Bic

kley

, D

arlin

g S

carp

,W

. H

. Loa

ring,

cite

d by

Quo

kkas

stil

l pr

esen

t in

gul

lies

in 1

933–

34no

w c

onsi

dere

d a

subu

rbW

hite

(19

52)

of P

erth

19

33

Mar

gare

t R

iver

are

aW

. H

. Loa

ring,

cite

d by

Quo

kkas

rep

orte

d as

far

les

s pl

entif

ul t

han

prev

ious

ly r

ecor

ded

in 1

907–

10.

Loca

tion

nom

inal

ly m

appe

d as

for

190

7–W

hite

(19

52)

1910

and

as

cree

klin

e in

for

este

d ar

ea,

imm

edia

tely

wes

t of

Mar

gare

t R

iver

19

33

Can

al R

ocks

, ne

ar Y

allin

gup

Whi

te (

1952

)R

ecor

ded

as p

artic

ular

ly n

umer

ous

in 1

933,

stil

l pr

esen

t 4

year

s la

ter,

les

s no

ticea

ble

with

in a

few

yea

rs

19

54

Man

jimup

, P

erup

are

aA

.D.

Jone

s in

Ser

vent

y et

al.

Rec

ogni

sed

the

area

bou

nded

by

the

Tone

and

Per

up r

iver

s, n

orth

war

d to

lat

itude

34

degr

ees

12 m

inut

es S

outh

was

(195

4)ric

h in

mar

supi

als.

Dat

e no

min

ally

lis

ted

here

as

1954

and

loc

atio

n m

appe

d as

with

in t

he P

erup

For

est,

and

betw

een

Tone

& P

erup

riv

ers,

nor

thea

st o

f M

anjim

up

19

54

Man

jimup

, M

oral

lup

A.D

. Jo

nes

in S

erve

nty

et a

l.R

ecor

d re

port

ed t

o A

.D.

Jone

s (in

Ser

vent

y et

al.

1954

) as

a s

ight

ing

10 m

iles

nort

h of

Mor

dallu

p. P

resu

mab

ly t

his

is a

(195

4)re

fere

nce

to M

oral

lup,

whi

ch p

lace

s th

is r

ecor

d ap

prox

imat

ely

23km

nor

thea

st o

f M

anjim

up.

Dat

e no

min

ally

lis

ted

here

as

1954

19

54

Com

men

ts o

n ge

nera

lL.

Gla

uert

in

Ser

vent

y et

al.

Des

crib

ed a

s pl

entif

ul o

n R

ottn

est

Isla

nd a

nd ‘s

ome

isla

nds

off

the

sout

h co

ast’

and

in s

wam

py c

ount

ry i

n th

e lo

wer

dist

ribu

tion

(195

4)so

uth

wes

t. N

omin

ally

map

ped

here

as

Alb

any

and

the

Bro

ke I

nlet

(D

’Ent

reca

stea

ux)

area

s, t

he l

atte

r as

map

ped

byC

hris

tens

en e

t al

. (1

985)

19

54

Com

men

ts o

n ge

nera

lL.

Gla

uert

in

Ser

vent

y et

al.

Des

crib

ed a

s no

lon

ger

pres

ent

in t

he v

alle

ys o

f th

e D

arlin

g R

ange

. P

resu

mab

ly t

he ‘i

slan

ds’ r

efer

ence

is

refe

rrin

g to

Bal

ddi

strib

utio

n (c

ont)

fro

m L

.(1

954)

Isla

nd,

but

this

may

als

o be

a r

efer

ence

to

the

isla

nds

of E

sper

ance

whi

ch w

ere

othe

rwis

e re

ferr

ed t

o on

ly b

yG

laue

rt i

n S

even

tyS

hort

ridge

(19

09).

et a

l. (1

954)

19

54

Yarl

oop,

Dar

ling

Ran

geS

erve

nty

et a

l. (1

954)

Ser

vent

y no

ted

quok

kas

seen

occ

asio

nally

, w

ith p

lent

y of

sig

ns i

n al

l sw

amps

. N

omin

ally

rec

orde

d as

195

4. E

xact

betw

een

Per

th a

nd H

arve

ylo

catio

n un

clea

r, n

omin

ally

map

ped

as L

ogue

Bro

ok a

rea

and

the

sam

e lo

catio

n as

Whi

te (

1952

). S

erve

nty’

s re

port

appe

ars

to c

ontr

adic

t th

e re

port

s of

Gla

uert

and

Loa

ring,

bot

h al

so i

n S

erve

nty

et a

l. (1

954)

, w

ho b

elie

ved

quok

kas

had

vani

shed

fro

m g

ully

hau

nts

of t

he D

arlin

g R

ange

30

year

s pr

evio

usly

. H

owev

er,

the

area

ref

erre

d to

by

Ser

vent

y et

al.

(195

4) i

s 10

0km

Sou

th o

f th

e se

ctio

n of

Dar

ling

Ran

ge L

oarin

g an

d G

laue

rt m

ay h

ave

been

ref

errin

g to

.

19

54

Man

jimup

, ne

arS

erve

nty

et a

l. (1

954)

Rec

orde

d as

firs

t re

cord

for

10

year

s. N

omin

ally

rec

orde

d as

195

4, e

xact

loc

atio

n un

clea

r. P

ossi

bly

refe

rrin

g to

the

‘B

roke

Bro

ke I

nlet

tur

noff

Inle

t tu

rn-o

ff’ f

rom

the

Sou

th W

este

rn H

ighw

ay (

Man

jimup

-Wal

pole

Hig

hway

), m

appe

d at

thi

s lo

catio

n

19

54

Tool

brun

up,

Stir

ling

Sha

rman

(19

54)

Exa

min

ed q

uokk

a ch

rom

osom

e nu

mbe

r an

d ur

ogen

ital

syst

em a

nd c

ompa

red

with

rel

ated

spe

cies

. N

oted

quo

kkas

Ra

ng

es

form

erly

wid

ely

dist

ribut

ed,

now

com

mon

onl

y on

Rot

tnes

t an

d B

ald

isla

nds.

Not

ed a

sku

ll co

llect

ed f

rom

Stir

ling

Ran

ges.

Loca

tion

nom

inal

ly s

how

n as

Blu

ff K

noll,

Stir

ling

Ran

ge.

Tabl

e 3

(con

t.)

Year

of

Loca

tion

orS

ourc

eC

omm

ent

reco

rdS

ite n

ame

Page 39: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka 51

19

55

Wal

pole

J.A

. R

ate,

cite

d by

Cap

ture

of

an i

mm

atur

e qu

okka

. A

ppro

xim

ate

loca

tion

only

, no

min

ally

map

ped

as w

ithin

Wal

pole

-Nor

nalu

p N

atio

nal

Par

kB

arke

r et

al.

(195

7)

19

56

Rot

tnes

t Is

land

Dun

net

(196

3)E

xam

ined

quo

kka

sub

popu

latio

ns a

t B

agda

d an

d S

erpe

ntin

e so

aks

at e

aste

rn/c

entr

al a

rea

of R

ottn

est

Isla

nd i

n th

epe

riod

1954

–195

8. N

omin

ally

lis

ted

here

as

1956

, se

e al

so D

unne

t (1

962)

. E

stim

ates

of

popu

latio

n si

ze w

ere

varia

ble

and

diffe

red

betw

een

and

with

in s

ampl

ing

perio

ds.

Dun

net

caut

ione

d th

ese

estim

ates

wer

e al

so s

ubje

ct t

o bi

as

19

56

Alb

any

Hig

hway

, ne

arR

. A

itken

, ci

ted

by B

arke

rTw

o ro

adki

ll re

cord

s. L

ocat

ion

appr

oxim

ate

only

and

ide

ntifi

ed f

rom

map

in

Sad

leir

(195

9).

See

196

6 re

cord

by

Ken

tTr

avel

lers

’ Arm

set

al.

(195

7)W

illia

ms

(Tab

le 4

) an

d 19

96 r

ecor

d by

de

Tore

s, D

illon

, To

mki

nson

and

Bue

hrig

(Ta

ble

6)(s

outh

east

of

Per

th)

19

56

Rot

tnes

t Is

land

Shi

eld

(195

9)45

0 qu

okka

s sa

mpl

ed i

n 19

56–5

7 as

par

t of

stu

dy a

sses

sing

whe

ther

quo

kka

popu

latio

n w

as l

imite

d by

env

ironm

enta

l/ph

ysio

logi

cal

fact

ors.

No

estim

ate

of t

he s

ize

of t

he R

ottn

est

popu

latio

n

19

56

Rot

tnes

t Is

land

War

ing

(195

6)N

oted

quo

kkas

kno

wn

from

tw

o of

f-sh

ore

isla

nds

only

(R

ottn

est

and

Bal

d) a

nd c

onsi

dere

d th

e on

ly m

ainl

and

popu

latio

nto

be

‘a r

emna

nt p

opul

atio

n in

Kar

ri fo

rest

’ in

the

sout

h-w

est.

Ref

erre

d to

the

Rot

tnes

t po

pula

tion

as c

onsi

stin

g of

‘per

haps

5,0

00 i

ndiv

idua

ls’.

19

57

Byf

ord,

Man

jeda

l B

rook

Bar

ker

et a

l. (1

957)

Thr

ee q

uokk

as t

rapp

ed i

n th

e pe

riod

8 M

ay t

o 6

June

195

7. T

rapp

ing

surv

ey i

nitia

ted

in r

espo

nse

to a

new

spap

er a

rtic

leon

the

sou

thea

st e

dge

ofw

hich

sug

gest

ed q

uokk

as h

ad b

ecom

e ex

tinct

on

the

mai

nlan

d.th

e P

erth

met

ropo

litan

are

a

19

58

Rot

tnes

t Is

land

Her

rick

(196

1)E

xam

ined

adr

enal

fun

ctio

n of

the

quo

kka

and

incl

uded

ani

mal

s co

llect

ed f

rom

Rot

tnes

t Is

land

(S

umm

er 1

958)

and

obse

rvat

ions

fro

m t

wo

site

s on

Rot

tnes

t

19

58

Man

jeda

l B

rook

, no

rth

Sto

rr (

1964

b)S

tudy

of

quok

ka h

abita

t, no

est

imat

es o

f po

pula

tion

size

. P

resu

mab

ly t

he s

ite(s

) us

ed b

y S

adle

ir (1

959)

and

of J

arra

hdal

eB

arke

r et

al.

(195

7)

19

58

Rot

tnes

t Is

land

Sto

rr (

1964

a)N

utrit

iona

l st

udy,

511

quo

kkas

cau

ght

over

uns

peci

fied

num

ber

of m

onth

ly v

isits

in

1958

. N

o es

timat

e of

pop

ulat

ion

size

19

59

Rot

tnes

t Is

land

Hol

swor

th (

1967

)E

xam

ined

hom

e ra

nge

and

terr

itory

use

in

the

perio

d 19

54 t

o 19

64.

Nom

inal

ly s

how

n he

re a

s 19

59.

No

estim

ate

ofpo

pula

tion

size

, ho

wev

er s

how

ed v

aria

tion

in t

he n

umbe

r of

ind

ivid

ual

anim

als

caug

ht f

rom

38

in 1

960

to 8

13 i

n 19

63

19

59

Bal

d Is

land

Sto

rr (

1965

)R

ecor

ded

quok

kas

as a

bund

ant,

occu

rrin

g fr

om s

ea l

evel

to

the

peak

, bu

t de

nsity

var

ied

grea

tly

19

59

Way

chin

icup

Sto

rr (

1965

)N

one

trap

ped

or s

nare

d, a

bund

ant

evid

ence

in

‘sw

ampy

val

leys

dra

inin

g in

to t

he l

ower

Way

chin

icup

’. E

xact

loc

atio

n no

tgi

ven,

map

ped

as a

ppro

xim

ate

loca

tion

only

.

19

63

Rot

tnes

t Is

land

Pac

ker

(196

5)O

bser

vatio

nal

stud

y in

196

3, n

o es

timat

e of

pop

ulat

ion

size

19

67

Rot

tnes

t Is

land

Nic

holls

(19

71)

Cau

ght

and

mon

itore

d 27

quo

kkas

fro

m W

est

End

of

Rot

tnes

t Is

land

to

dete

rmin

e ho

me

rang

e an

d m

ovem

ents

,19

67–1

968.

No

estim

ate

of p

opul

atio

n/su

b-po

pula

tion

size

19

69

Rot

tnes

t Is

land

Kitc

hene

r (1

973)

Rec

orde

d an

ave

rage

of

53 r

esid

ent

quok

kas

at t

he B

arke

rs S

wam

p st

udy

site

, w

ith a

vera

ge d

ensi

ty o

f 1

quok

ka p

er0.

06ha

and

not

ed t

his

was

con

side

rabl

y gr

eate

r th

an t

he a

vera

ge o

f 1

quok

ka p

er 0

.4 –

1.2

ha r

ecor

ded

by M

ain

and

Yada

v (1

971)

. T

he l

ocal

ised

hig

h de

nsity

at

Bar

kers

Sw

amp

was

con

side

red

to r

efle

ct t

he l

ocal

ised

hig

h qu

ality

site

and

the

high

den

sity

was

mai

ntai

ned

by ‘

recr

uitm

ent

of c

onsi

dera

ble

num

ber

of q

uokk

as i

nto

the

loca

l po

pula

tion’

19

70

Rot

tnes

t Is

land

Kitc

hene

r (1

972)

Rep

orte

d on

obs

erve

d be

havi

ours

, in

tera

ctio

ns a

nd d

omin

ance

of

sub

popu

latio

n at

Bar

kers

Sw

amp.

Sub

pop

ulat

ion

estim

ated

by

dire

ct c

ount

dur

ing

noct

urna

l ob

serv

atio

ns.

How

ever

, su

bpop

ulat

ion

estim

ate

not

repo

rted

Page 40: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

52 P.J. de Torres et al.

19

70

Dar

ling

Ran

ge,

clos

eR

ide

(197

0)R

ecor

ded

as r

are

on t

he m

ainl

and,

kno

wn

only

fro

m a

few

sw

ampy

val

leys

in

the

Dar

ling

Ran

ge.

Nom

inal

ly s

how

n he

reto

Per

thas

Dw

ellin

gup

(Hol

yoak

e) w

here

kno

wn

to o

ccur

in

the

1970

s

19

71

Com

men

ts o

n ge

nera

lC

alab

y (1

971)

Cal

aby

note

d th

e qu

okka

was

kno

wn

from

a f

ew s

wam

py a

reas

on

the

mai

nlan

d an

d w

as s

till

com

mon

on

Rot

tnes

t an

ddi

stri

butio

nB

ald

isla

nds.

Nom

inal

ly m

appe

d as

Dw

ellin

gup,

Hol

yoak

e

19

71

Rot

tnes

t Is

land

Mai

n an

d Ya

dav

(197

1)R

evie

wed

the

con

serv

atio

n in

form

atio

n fo

r m

acro

pods

fro

m B

arro

w I

slan

d an

d co

mpa

red

this

to

othe

r is

land

s. G

ave

anes

timat

e of

quo

kka

popu

latio

n de

nsity

for

Rot

tnes

t Is

land

as

rang

ing

from

1 q

uokk

a pe

r 1.

2ha

to 1

per

0.4

ha.

How

ever

,no

dat

a su

pplie

d to

sho

w h

ow t

hese

val

ues

wer

e de

rived

and

den

sity

may

hav

e be

en d

eter

min

ed o

n ba

sis

of t

he e

ntire

isla

nd b

eing

occ

upie

d. D

ate

nom

inal

ly s

how

n he

re a

s 19

71

19

71

Bal

d Is

land

Mai

n an

d Ya

dav

(197

1)G

ave

an e

stim

ate

of q

uokk

a po

pula

tion

size

at

Bal

d Is

land

as

rang

ing

from

600

–190

0 qu

okka

s (if

ent

ire i

slan

d w

asoc

cupi

ed).

Qua

lifie

d, i

n re

cogn

ition

not

all

of t

he i

slan

d w

as o

ccup

ied,

to

give

an

uppe

r es

timat

e of

600

. D

ate

nom

inal

lylis

ted

here

as

1971

19

71

Dw

ellin

gup

Sch

mid

t an

d M

ason

(19

73)

Rec

orde

d a

suite

of

mam

mal

spe

cies

in

fore

st n

ear

Dw

ellin

gup

in 1

971

and

conc

lude

d po

pula

tions

wer

e co

ncen

trat

ed i

nsw

amps

and

oth

er a

reas

of

dens

e ve

geta

tion.

Quo

kkas

inc

lude

d in

lis

t of

spe

cies

rec

orde

d, n

o si

te d

etai

ls,

nom

inal

lym

appe

d he

re a

s H

olyo

ake

19

72

Dw

ellin

gup,

Wre

ns R

oad

Chr

iste

nsen

and

Kim

ber

(197

5)P

opul

atio

n tr

appe

d (1

972–

74)

befo

re a

nd a

fter

a pa

rtia

l bu

rn.

Trap

suc

cess

rat

e in

crea

sed

afte

r th

e bu

rn.

Indi

ces

toS

wam

p (e

ast)

abun

danc

e im

ply

mod

erat

e si

ze p

opul

atio

n, h

owev

er n

o es

timat

e of

pop

ulat

ion

size

. D

ate

assu

med

to

be 1

972,

see

Chr

iste

nsen

(un

date

d) i

n ta

ble

of u

npub

lishe

d re

cord

s

19

72

Alb

any

Hig

hway

,C

rabb

(19

73)

Roa

dkill

, ne

ar t

he ‘2

7 m

ile p

eg o

n A

lban

y H

ighw

ay’

sout

heas

t of

Per

th

19

74

Dw

ellin

gup,

Dun

cans

Chr

iste

nsen

and

Kim

ber

(197

5)P

opul

atio

n tr

appe

d 19

72–7

4. T

his

site

unb

urnt

, cf

Wre

ns R

oad.

Tra

p su

cces

s co

mp

arab

le t

o po

st b

urn

capt

ure

succ

ess

Roa

d S

wam

pat

Wre

ns R

oad.

Ind

ices

to

abun

danc

e im

ply

a m

oder

ate

size

pop

ulat

ion,

no

estim

ate

of p

opul

atio

n si

ze.

Dat

e no

min

ally

show

n as

197

4

19

75

Kar

ri, n

orth

wes

t of

Chr

iste

nsen

et

al.

(198

5)Li

sted

and

map

ped

by C

hris

tens

en e

t al

. (1

985)

as

Sur

vey

Are

a L,

Kar

ri. S

ee n

ote

2, b

elow

Nor

thcl

iffe

19

75

Yea

ga

rup,

be

twe

en

th

eC

hris

tens

en e

t al

. (1

985)

List

ed a

nd m

appe

d by

Chr

iste

nsen

et

al.

(198

5) a

s S

urve

y A

rea

A,

Yeag

arup

. S

ee n

ote

2, b

elow

Don

nelly

and

War

ren

river

s

19

75

Mitc

hell,

nor

th n

orth

wes

tC

hris

tens

en e

t al

. (1

985)

List

ed a

nd m

appe

d by

Chr

iste

nsen

et

al.

(198

5) a

s S

urve

y A

rea

I, M

itche

ll. S

ee n

ote

2, b

elow

of W

alpo

le

19

75

Soh

o, n

orth

east

of

Wal

pole

Chr

iste

nsen

et

al.

(198

5)Li

sted

and

map

ped

by C

hris

tens

en e

t al

. (1

985)

as

Sur

vey

Are

a H

, S

oho.

See

not

e 2,

bel

ow

19

75

Bor

anup

, so

uth

ofC

hris

tens

en e

t al

. (1

985)

List

ed a

nd m

appe

d by

Chr

iste

nsen

et

al.

(198

5) a

s S

urve

y A

rea

D,

Bor

anup

. S

ee n

ote

2, b

elow

Mar

gare

t R

iver

Tabl

e 3

(con

t.)

Year

of

Loca

tion

orS

ourc

eC

omm

ent

reco

rdS

ite n

ame

Page 41: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka 53

19

75

Gia

nts,

bet

wee

n N

orna

lup

Chr

iste

nsen

et

al.

(198

5)Li

sted

and

map

ped

by C

hris

tens

en e

t al

. (1

985)

as

Sur

vey

Are

a O

, G

iant

s. S

ee n

ote

2, b

elow

Inle

t an

d Ir

win

Inl

et,

east

of

Nor

nalu

p

19

75

Dom

baku

p, w

est

Chr

iste

nsen

et

al.

(198

5)O

ne o

f tw

o si

tes

liste

d an

d m

appe

d by

Chr

iste

nsen

et

al.

(198

5) a

s S

urve

y A

rea

C,

Dom

baku

p. S

ee n

ote

2, b

elow

nort

hwes

t of

Nor

thcl

iffe

19

75

Woo

lbal

es,

sout

heas

t of

Chr

iste

nsen

et

al.

(198

5)Li

sted

and

map

ped

by C

hris

tens

en e

t al

. (1

985)

as

Sur

vey

Are

a B

, W

oolb

ales

. S

ee n

ote

2, b

elow

Bro

ke I

nlet

19

75

Dom

baku

p, w

est

of B

roke

Chr

iste

nsen

et

al.

(198

5)O

ne o

f tw

o si

tes

liste

d an

d m

appe

d by

Chr

iste

nsen

et

al.

(198

5) a

s S

urve

y A

rea

C,

Dom

baku

p. S

ee n

ote

2, b

elow

Inle

t (D

’Ent

reca

stea

uxN

atio

nal

Par

k)

19

75

Sun

klan

ds,

sout

h-C

hris

tens

en e

t al

. (1

985)

List

ed a

nd m

appe

d by

Chr

iste

nsen

et

al.

(198

5) a

s S

urve

y A

rea

E,

Sun

klan

ds.

See

not

e 2,

bel

owso

uthe

ast

of B

usse

lton

19

76

Nan

nup,

Mow

en/S

toat

sH

art

et a

l. (1

986)

Trap

ped

as p

art

of c

ompa

rativ

e st

udy

of S

alm

onel

la i

nfec

tion.

Im

plie

d to

be

at l

ow d

ensi

tyR

oad

Cro

ssin

g

19

76

Mud

dy L

ake,

sou

th o

fH

art

et a

l. (1

986)

Trap

ped

as p

art

of c

ompa

rativ

e st

udy

of S

alm

onel

la i

nfec

tion.

Im

plie

d to

be

at l

ow d

ensi

tyB

un

bu

ry

19

76

Dw

ellin

gup,

Hol

yoak

eH

art

et a

l. (1

986)

Rep

orte

d hi

gh t

rap

yiel

d (1

0 an

imal

s fr

om 1

00 t

rap

nigh

ts)

usin

g el

abor

ate

trap

s co

mpr

ised

of

fenc

e lin

es a

nden

clos

ures

, w

here

mul

tiple

tra

p ni

ghts

ass

umed

for

eac

h tr

ap

19

76

Byf

ord

Har

t et

al.

(198

6)Tr

appe

d as

par

t of

com

para

tive

stud

y of

Sal

mon

ella

inf

ectio

n. I

mpl

ied

to b

e at

low

den

sity

. P

resu

mab

ly s

ame

site

as

trap

ped

by B

arke

r et

al.

(195

7) a

nd S

adle

ir (1

959)

19

77

Dw

ellin

gup,

Hol

yoak

eH

art

et a

l. (1

986)

Rep

orte

d hi

gh t

rap

yiel

d (2

7 an

imal

s fr

om 2

04 t

rap

nigh

ts)

usin

g el

abor

ate

trap

s co

mpr

ised

of

fenc

e lin

es a

nden

clos

ures

, w

here

mul

tiple

tra

p ni

ghts

ass

umed

for

eac

h tr

ap

19

80

Bal

d is

land

Har

t et

al.

(198

6)14

quo

kkas

cau

ght

from

a s

ingl

e on

e-ni

ght

trip

. N

o es

timat

e of

abu

ndan

ce r

epor

ted,

how

ever

pop

ulat

ion

leve

l re

port

edto

flu

ctua

te a

s a

resu

lt of

sum

mer

sta

rvat

ion

19

83

Dar

ling

Sca

rpD

ell

(198

3)T

his

publ

icat

ion

prod

uced

to

upda

te t

he W

este

rn A

ustr

alia

n M

useu

m m

amm

al r

ecor

ds w

ith r

ecen

t ob

serv

atio

ns a

ndre

view

the

lite

ratu

re.

The

quo

kka

was

lis

ted

as o

ne o

f se

vera

l sp

ecie

s to

hav

e de

clin

ed.

Nom

inal

ly m

appe

d he

re a

sD

wel

lingu

p, U

rbra

e F

ores

t B

lock

19

83

Gre

en R

ange

, 60

kmK

irke

(198

3)Q

uokk

as r

epor

ted

as f

leei

ng f

rom

bur

ning

bus

hlan

d. S

kele

ton

subs

eque

ntly

col

lect

ed.

The

onl

y re

cord

at

this

loc

atio

nno

rthe

ast

of A

lban

ypr

ior

to t

his

was

fro

m P

earc

e (1

969)

, se

e 19

69 e

ntry

in

Tabl

e 4,

Unp

ublis

hed

Rec

ords

.

19

85

Way

chin

icup

Inl

et,

east

Har

t et

al.

(198

6)Tr

appe

d as

par

t of

com

para

tive

stud

y of

Sal

mon

ella

inf

ectio

n. P

opul

atio

n im

plie

d to

be

at l

ow d

ensi

tyof

Alb

any

19

85

Pt

D’E

ntre

cast

eaux

How

et

al.

(198

7)S

urve

y co

nduc

ted

in t

he p

erio

d M

arch

198

5, O

ctob

er t

o N

ovem

ber

1985

and

Jan

uary

to

Feb

ruar

y 19

86.

Nom

inal

lysh

own

here

as

1985

. Q

uokk

a re

port

ed f

rom

onl

y 1

of 1

0 lo

catio

ns s

urve

yed.

Thi

s re

cord

of

pres

ence

im

plie

d by

How

et

al.

(198

7) t

o be

que

stio

nabl

e. L

ocat

ion

appr

oxim

ate

only

Page 42: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

54 P.J. de Torres et al.

19

85

Pem

bert

on a

nd D

wel

lingu

pM

ead

et a

l. (1

985)

Ani

mal

s co

llect

ed f

rom

mai

nlan

d (D

wel

lingu

p an

d P

embe

rton

) an

d is

land

s fo

r la

bora

tory

ass

essm

ent

of t

oler

ance

to

1080

. S

ampl

e si

ze n

ot s

peci

fied

and

no e

stim

ate

of p

opul

atio

n si

ze a

t co

llect

ion

poin

t. D

ate

nom

inal

ly s

how

n he

re a

s19

85.

Loca

tion

map

ped

nom

inal

ly a

s st

ate

fore

st,

imm

edia

tely

eas

t of

Pem

bert

on a

nd D

wel

lingu

p (H

olyo

ake)

19

85

Bal

d Is

land

Mea

d et

al.

(198

5)A

nim

als

colle

cted

for

lab

orat

ory

asse

ssm

ent

of t

oler

ance

to

1080

. S

ampl

e si

ze n

ot s

peci

fied

and

no e

stim

ate

ofpo

pula

tion

size

at

colle

ctio

n po

int.

Dat

e no

min

ally

sho

wn

here

as

1985

19

85

Rot

tnes

t Is

land

Mea

d et

al.

(198

5)A

nim

als

colle

cted

for

lab

orat

ory

asse

ssm

ent

of t

oler

ance

to

1080

. S

ampl

e si

ze n

ot s

peci

fied

and

no e

stim

ate

ofpo

pula

tion

size

at

colle

ctio

n po

int.

Dat

e no

min

ally

sho

wn

here

as

1985

20

00

Col

lie,

Vic

tor

Roa

d,H

ayw

ard

et a

l.(20

03)

Trap

ped

over

the

per

iod

1998

–200

0. P

opul

atio

n es

timat

ed t

o be

9 +

/- 1

(Jo

lly-S

eber

mar

k-re

capt

ure

deriv

ed e

stim

ates

)H

amilt

on F

ores

t B

lock

20

00

Col

lie/H

arve

y, H

adfie

ldH

ayw

ard

et a

l.(20

03)

Trap

ped

over

the

per

iod

1998

–200

0. P

opul

atio

n es

timat

ed t

o be

29

+/-

5 (

Jolly

-Seb

er m

ark-

reca

ptur

e de

rived

est

imat

es)

For

est

Blo

ck

20

00

Dw

ellin

gup,

Kes

ners

Hay

war

d et

al.(

2003

)Tr

appe

d ov

er t

he p

erio

d 19

98–2

000.

Pop

ulat

ion

estim

ated

to

be 3

6 +

/- 6

(Jo

lly-S

eber

mar

k-re

capt

ure

deriv

ed e

stim

ates

)S

wam

p, T

urne

r F

ores

tB

lock

20

00

Jarr

ahda

le,

Cha

ndle

r R

oad

Hay

war

d et

al.(

2003

)Tr

appe

d ov

er t

he p

erio

d 19

98–2

000.

Pop

ulat

ion

estim

ated

to

be 1

0 (J

olly

-Seb

er m

ark-

reca

ptur

e de

rived

est

imat

es)

Site

, C

hand

ler

For

est

Blo

ck

20

00

Jarr

ahda

le,

Ros

ella

Roa

d,H

ayw

ard

et a

l.(20

03)

Trap

ped

over

the

per

iod

1998

–200

0. N

o po

pula

tion

estim

ate,

onl

y on

e an

imal

tra

pped

Gor

don

For

est

Blo

ck

Not

es:

1.S

hort

ridge

col

lect

ed 3

8 sp

ecim

ens

and

desc

ribed

the

quo

kka

as p

lent

iful

amon

g co

asta

l th

icke

ts a

nd s

wam

ps o

f th

e so

uth

wes

t, no

t ex

tend

ing

inla

nd.

The

dis

trib

utio

n w

as r

ecor

ded

as e

xten

ding

from

the

Moo

re R

iver

in

the

nort

h, a

nd m

appe

d to

inc

lude

the

sou

th c

oast

as

far

east

as

Esp

eran

ce. T

he e

aste

rn e

xten

t ap

pear

s to

be

base

d of

rep

orts

fro

m T

win

Pea

k &

oth

er i

slan

ds o

ffE

sper

ance

. T

his

is t

he o

nly

repo

rt o

f qu

okka

occ

urre

nce

from

the

isl

ands

off

Esp

eran

ce a

nd n

o co

llect

ions

hav

e be

en d

ocum

ente

d fr

om t

hese

isl

ands

. S

hort

ridge

’s r

ecor

d is

ass

umed

to

bere

ferr

ing

to t

amm

ars,

see

tex

t.2.

Chr

iste

nsen

et

al.

(198

5) f

rom

sur

veys

197

0–19

82 n

oted

the

quo

kka

as w

ides

prea

d an

d lo

cally

com

mon

. E

stim

ates

of

popu

latio

n si

ze w

ere

not

prov

ided

and

loc

atio

ns a

re a

ppro

xim

ate

and

rem

appe

d fr

om m

ap o

n p4

of

Chr

iste

nsen

et

al.

(198

5).

Dat

e no

min

ally

sho

wn

as 1

975

Tabl

e 3

(con

t.)

Year

of

Loca

tion

orS

ourc

eC

omm

ent

reco

rdS

ite n

ame

Page 43: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka

55

Table 4

Unpublished records of occurrence of the quokka, Setonix brachyurus. Records are listed chronologically.

Year of Location or Source Commentrecord Site name

1940 Northcliffe, Bashford Road L. Wilson (personal communication Quokkas observed in large numbers in 1940s and earlier and known to feed in paddocks away from vegetatedarea, 10km east to PJdeT) creeklines. Reported as ‘last seen in number when fleeing from fire’ in the 1940s. Location approximate and shownof Northcliffe nominally as 1940

1958 Rottnest Island Sadleir (1959) Conducted a trapping program at the Byford/Manjedal Swamp site on the mainland and Rottnest Island to examinephysiological differences between the two populations

1958 Byford, Manjedal Sadleir (1959) Trapped at Manjedal Swamp in April – May and September – October 1958. Six quokkas only caught, all in theSwamp/Brook April – May period

1962 Rottnest Island Holsworth (1964) Trapped in period May 1961 to May 1963 and incorporated data from 1954 to 1961 and from December 1963 to April1964. Nominally shown here as 1962. Population estimates were derived for 4 areas within the West End only. Eacharea was comprised of several group territories. Population estimates were described by Holsworth as stable for theperiod 1955 to 1963, however data indicates large variations in abundance.

1965 Travellers’ Arms, Albany Kent Williams (pers. comm. Quokkas successfully trapped in 1965 at swamp on the opposite side of Albany highway from the Travellers’ Arms Hotel.Highway, south-east of the to PJdeT) Subsequent trapping in the same year, post fire was unable to confirm continued presence. See record for 1996Perth metropolitan area (Table 6) when presence at this location was re-confirmed

1969 Two Peoples Bay Nature Bannister (1970) Reported carcass found in 1969 by N. Robinson, in thick scrub at bottom of a shallow gully, near CSIRO hut. OldReserve, near CSIRO hut evidence and no indication of recent activity when surveyed by Bannister (1970). Location mapped here is approximate

1969 Rottnest Island Kitchener (1970) Estimated a mean monthly population size from a study area comprised of 48 quadrats, each 12m x 12m at BarkersSwamp, near the centre/eastern end of the Island. Estimates derived from data collected in the period December 1967to December 1969. Listed here as 1969. Estimates were from direct observations and included permanent andtemporary residents, recruits and transients. Mean monthly estimate was approximately 45 adults and 13 juveniles withinthe Barkers Swamp study site.

1969 Green Range, 50 miles Pearce (1969) Local landholder recorded the presence of small kangaroos. Subsequent investigation by WA Department of Fisheries(30km) northeast of Albany Fauna Warden detected spoor consistent with quokka. Drawing of spoor provided in Departmental file – we have

presumed this to be a quokka. Presence recorded here in 1983, see 1983 entry in Table 3.

1970 Two Peoples Bay Nature Bannister (1970) Unable to confirm by trapping, presence inferred by runs and scats detected in thickly vegetated gullies, near weatherReserve, near weather station above CSIRO hut. Location mapped here is approximatestation above CSIRO hut

1970 Two Peoples Bay Nature Bannister (1970) Unable to confirm by trapping, presence inferred by runs and scats detected in thickly vegetated gullies, bottom ofReserve, south-west of valley, south-west of Pt GardnerPt Gardner

1970 Two Peoples Bay Nature Bannister (1970) Unable to confirm by trapping, presence inferred by runs and scats detected in thickly vegetated gullies, thick swampyReserve, Moates Lagoon areas at west end of Moates Lagoon

Page 44: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

56 P.J. de Torres et al.

19

72

Dw

ellin

gup,

Wre

ns R

oad

Chr

iste

nsen

(un

date

d)Tr

appi

ng p

rogr

am b

efor

e (A

ugus

t to

Nov

embe

r 19

72)

and

afte

r (1

2 to

21

Dec

embe

r 19

72)

a lo

w i

nten

sity

bur

n of

60%

of

Sw

amp

(wes

t),

Urb

rae

quok

ka h

abita

t w

ithin

the

sw

amp.

No

estim

ates

of

dens

ity,

how

ever

a t

otal

of

22 i

ndiv

idua

ls t

rapp

ed.

For

est

Blo

ck

19

72

Dw

ellin

gup,

Wre

ns R

oad

Chr

iste

nsen

(un

date

d)Tr

appi

ng p

rogr

am b

efor

e (A

ugus

t to

Nov

embe

r 19

72)

and

afte

r (1

2 to

21

Dec

embe

r 19

72)

a lo

w i

nten

sity

bur

n of

60%

of

Sw

amp

(eas

t),

Whi

tequ

okka

hab

itat

with

in t

he s

wam

p. N

o es

timat

es o

f de

nsity

, ho

wev

er a

tot

al o

f 22

ind

ivid

uals

tra

pped

.F

ores

t B

lock

19

75

Den

bark

er,

5km

eas

t of

Kab

ay a

nd S

tart

(19

76),

150

cage

tra

p ni

ghts

and

180

sna

re t

rap

nigh

ts o

ver

9 ni

ghts

(28

–30

July

197

5 an

d 11

–20

Aug

ust

1975

). S

ix q

uokk

asD

enba

rker

Roa

d br

idge

note

1,

belo

wtr

appe

d or

sna

red

over

Mitc

hell

Riv

er

19

75

Mou

nt M

anyp

eaks

, N

orth

Kab

ay a

nd S

tart

(19

76),

130

cage

tra

p ni

ghts

ove

r th

ree

nigh

ts p

lus

60 s

nare

tra

p ni

ghts

ove

r th

ree

nigh

ts (

5–7

July

197

5).

One

quo

kka

only

of M

ount

Man

ypea

ksno

te 1

, be

low

seen

, ru

nway

s an

d sc

ats

foun

d

19

75

Two

Peo

ples

Bay

,K

abay

and

Sta

rt (

1976

),18

0 ca

ge t

rap

nigh

ts o

ver

4 ni

ghts

(1–

8 Ju

ly 1

975)

. N

o qu

okka

cap

ture

s, b

ut p

rese

nce

infe

rred

by

pres

ence

of

scat

sM

t G

ardn

erno

te 1

, be

low

and

runw

ays

in g

ullie

s an

d in

den

se h

eath

bor

derin

g gu

llies

19

75

Nan

nup,

Mow

en R

oad

Kab

ay a

nd S

tart

(19

76),

Sur

veye

d Ju

ne 1

975

with

50

cage

tra

p ni

ghts

and

10

snar

e tr

ap n

ight

s ov

er o

ne n

ight

(in

per

iod

6–17

Jun

e 19

75).

cree

k cr

ossi

ng,

east

of

itsno

te 1

, be

low

Thr

ee q

uokk

as t

rapp

ed,

scat

s an

d ru

nway

s pr

esen

tju

nctio

n w

ith S

toat

s R

oad

19

76

Bal

d Is

land

Kab

ay a

nd S

tart

(19

76),

360

cage

tra

p ni

ghts

ove

r th

ree

nigh

ts (

29 M

ay –

3 J

une

1976

). O

ne q

uokk

a tr

appe

d, l

arge

num

ber

seen

and

a n

umbe

rno

te 1

, be

low

of s

kulls

col

lect

ed

19

76

Two

Peo

ples

Bay

,K

abay

and

Sta

rt (

1976

),36

0 ca

ge t

rap

nigh

ts o

ver

4–9

nigh

ts a

t a

rang

e of

loc

atio

ns (

13 A

pril

– 2

May

197

6). T

wo

juve

nile

quo

kkas

tra

pped

at

‘Rob

inso

n’s

Gul

ly’

note

1,

belo

w‘R

obin

son’

s G

ully

’. A

ppro

xim

ate

loca

tion

only

map

ped

here

19

76

Bun

bury

, M

uddy

Lak

e,K

abay

and

Sta

rt (

1976

),15

0 ca

ge t

rap

nigh

ts o

ver

thre

e ni

ghts

Jun

e 19

75 (

10–1

2 Ju

ne 1

975)

. Q

uokk

a sk

elet

al m

ater

ial

reco

vere

d, r

unw

ays

and

11km

Sou

th o

f B

unbu

ryno

te 1

, be

low

scat

s co

mm

on.

No

live

anim

als

trap

ped

in J

une

1975

, bu

t w

ere

seen

sub

sequ

ently

(4

Feb

197

6) b

y th

ese

inve

stig

ator

s.C

o-or

dina

tes

corr

ecte

d to

plo

t to

loc

atio

n of

Mud

dy L

ake

19

77

Dw

ellin

gup,

Hol

yoak

eH

art

(197

7)Tr

appe

d 10

ind

ivid

ual

quok

kas

over

the

per

iod

6–11

Nov

embe

r 19

76 a

nd 2

7 ov

er t

he p

erio

d 17

–24

Mar

ch 1

977

19

79

Jan

da

kot

Aus

tin (

1979

)E

stim

ated

pop

ulat

ion

size

of

a tr

ansl

ocat

ed,

fenc

ed p

opul

atio

n, w

here

393

quo

kkas

ex

Rot

tnes

t Is

land

had

bee

nre

leas

ed i

n th

e pe

riod

197

2–19

78.

Pop

ulat

ion

estim

ated

to

be a

ppro

xim

atel

y 84

ani

mal

s in

197

9. T

rans

loca

ted

popu

latio

n su

bseq

uent

ly s

how

n to

hav

e fa

iled

to p

ersi

st

19

82

Jarr

ahda

le (

Mun

dilu

pD

. G

iles

(per

sona

l co

mm

unic

atio

nR

ecor

d of

roa

dkill

(s)

repo

rted

fro

m t

he e

arly

198

0s (

nom

inal

ly 1

982)

by

loca

l la

ndho

lder

. R

epor

ted

to D

oug

Gile

s,F

ores

t B

lock

)to

PJd

eT)

Dep

artm

ent

of C

onse

rvat

ion

and

Land

Man

agem

ent,

Jarr

ahda

le

19

86

Stir

ling

Ran

ge N

atio

nal

A.N

. S

tart

, no

te 2

bel

owC

onfir

med

rec

ords

of

pres

ence

(si

ghtin

gs)

by A

llan

Ros

e in

the

per

iod

from

198

6 to

199

1. N

omin

ally

lis

ted

and

map

ped

Par

k –

Blu

ff K

noll

here

as

1986

, se

e al

so r

ecor

d fo

r 19

91(1

98

6–

19

91

)

Tabl

e 4

(con

t.)

Year

of

Loca

tion

orS

ourc

eC

omm

ent

reco

rdS

ite n

ame

Page 45: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka 57

19

88

Wal

pole

-Nor

nalu

p N

atio

nal

K.

Gill

en,

note

2 b

elow

Rec

ords

of

scat

s an

d ru

nway

s (e

ast

of M

t H

opki

ns,

Kar

ri H

ill G

ully

eas

t of

Cry

stal

Lak

e an

d so

uth

of D

eep

Riv

er)

over

the

Par

k, N

uyts

Wild

erne

sspe

riod

June

198

6 to

Jun

e 19

88.

Nom

inal

ly s

how

n he

re a

s 19

88 a

nd m

appe

d at

eas

t of

Cry

stal

Lak

e

19

88

Mou

nt M

anyp

eaks

are

aK

. G

illen

, no

te 2

, be

low

Sev

eral

rec

ords

of

quok

ka s

cats

and

run

way

s no

ted

in p

erio

d S

epte

mbe

r 19

87 t

o S

epte

mbe

r 19

89.

Map

ped

loca

tion

isap

prox

imat

e on

ly a

nd l

iste

d as

198

8

19

89

Mou

nt F

rank

land

Nat

iona

lA

.N.

Sta

rt,

note

2 b

elow

Inci

dent

al s

ight

ing

by A

ndre

w M

orto

n. S

ight

ing

of o

ne a

nim

al o

nly

Par

k, M

iddl

e R

oad

19

89

Two

Peo

ples

Bay

Nat

ure

K.

Gill

en,

note

2 b

elow

Mul

tiple

rec

ords

of

quok

ka s

cats

and

run

way

s no

ted

in p

erio

d Ju

ne 1

985

to J

uly

1991

. T

hree

con

firm

ed s

ight

ings

in

July

Res

erve

, M

t G

ardn

er a

rea

1988

(Lo

wer

Rob

inso

n’s

Gul

ly/C

offin

Gul

ly),

Jul

y 19

89 (

Cof

fin G

ully

) an

d Ju

ly 1

990

(Rob

inso

n’s

Gul

ly/C

offin

Gul

ly)

19

89

Bal

d Is

land

K.

Gill

en,

note

2 b

elow

Rec

ord

of s

cats

and

sub

sequ

ent

quok

ka s

ight

ing

on r

ocks

nea

r w

este

rn s

hore

19

91

Stir

ling

Ran

ge N

atio

nal

Sm

ith,

R.

Car

cass

rec

orde

d by

Bob

Sm

ith (

CA

LM M

anjim

up),

via

Lac

hie

McC

aw (

CA

LM M

anjim

up).

Quo

kka

thou

ght

to b

eP

ark,

Suc

cess

Rid

ge T

rack

disp

lace

d as

a r

esul

t of

fire

19

91

Na

nn

up

, L

ew

an

aA

.N.

Sta

rt,

note

2 b

elow

Car

cass

of

juve

nile

fox

-kill

ed q

uokk

a co

llect

ed i

n pi

ne p

lant

atio

n. A

ppro

xim

ate

map

ping

of

loca

tion

only

. N

o de

tails

of

Pla

nta

tion

sour

ce p

opul

atio

n

19

91

Stir

ling

Ran

ge N

atio

nal

A.N

. S

tart

, no

te 2

bel

owC

onfir

med

rec

ords

of

pres

ence

(si

ghtin

gs)

by A

llan

Ros

e in

the

per

iod

from

198

6 to

199

1. N

omin

ally

lis

ted

here

as

1991

,P

ark

- B

luff

Kno

llse

e al

so 1

986

reco

rd a

bove

. A

rea

burn

t in

Apr

il 19

91 a

nd o

nly

one

reco

rd a

t th

is l

ocat

ion

sinc

e (s

ee 1

991

reco

rd b

elow

).(1

98

6–

19

91

)P

rese

nce

othe

rwis

e su

gges

ted

by s

cats

, se

e 19

95 r

ecor

d of

Bar

rett

(199

6).

19

91

Stir

ling

Ran

ge N

atio

nal

A.N

. S

tart

, no

te 2

bel

owC

arca

ss (

bone

s on

ly)

colle

cted

by

Alla

n R

ose.

Pre

sum

ed k

illed

as

a re

sult

of f

ireP

ark

- B

luff

Kno

ll (1

991)

19

91

Jarr

ahda

le,

Ros

ella

A.N

. S

tart

, no

te 2

bel

owP

rese

nce

repo

rted

fro

m s

cat

only

. S

ubse

quen

tly t

rapp

ed t

o co

nfir

m p

rese

nce

in 1

995

(pre

viou

sly

unpu

blis

hed

resu

lts,

Roa

d, G

ordo

n F

ores

t B

lock

see

Tabl

e 9)

and

to

estim

ate

popu

latio

n si

ze 1

998–

2000

(H

ayw

ard

et a

l. 20

03)

19

91

Jarr

ahda

le,

Kar

net

A.N

. S

tart

, no

te 2

bel

owS

ight

ed b

y G

rant

Pro

nk w

hen

flush

ed f

rom

veg

etat

ion

as e

scap

ing

bush

fire

For

est

Blo

ck

19

92

Wal

pole

, R

oe F

ores

t B

lock

A.N

. S

tart

, no

te 2

bel

owK

now

n fo

r on

e ro

adki

ll ca

rcas

s re

cord

ed b

y P

hil

Dur

ell

19

94

Mt

Man

ypea

ksK

. G

illen

, no

te 2

bel

owR

epor

ted

as d

ead

sub-

adul

t m

ale

19

94

Two

Peo

ples

Bay

(Sin

clai

r 19

99)

Non

e ca

ught

dur

ing

Sin

clai

r’s s

urve

y pe

riod.

Sub

sequ

ently

cau

ght

(by

WA

Dep

t of

Con

serv

atio

n an

d La

nd M

anag

emen

tN

atur

e R

eser

vest

aff)

fro

m D

ecem

ber

1994

onw

ards

fro

m s

ever

al d

iffer

ent

loca

tions

. Lo

catio

n co

-ord

inat

es h

ere

are

ther

efor

eap

prox

imat

e

19

95

Stir

ling

Ran

ge N

atio

nal

Bar

rett

(199

6)P

rese

nce

indi

cate

d by

det

ectio

n of

hai

r (h

air

tube

). F

resh

sca

ts a

nd r

unw

ays

also

obs

erve

d by

Sin

clai

r (1

999)

.P

ark

- E

llen

Pea

k

19

95

Stir

ling

Ran

ge N

atio

nal

Bar

rett

(199

6)P

rese

nce

indi

cate

d by

det

ectio

n of

sca

tsP

ark

- To

olbr

unup

Pea

k

19

95

Stir

ling

Ran

ge N

atio

nal

Bar

rett

(199

6)P

rese

nce

indi

cate

d by

det

ectio

n of

sca

ts f

rom

sou

ther

n sl

opes

and

cas

cade

are

aP

ark

- B

luff

Kno

ll

Page 46: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

58 P.J. de Torres et al.

19

95

Mou

nt M

anyp

eaks

Bar

rett

(199

6)P

rese

nce

indi

cate

d by

det

ectio

n of

hai

r (h

air

tube

)N

atur

e R

eser

ve

19

95

Stir

ling

Ran

ge N

atio

nal

Bar

rett

(199

6)P

rese

nce

indi

cate

d by

det

ectio

n of

sca

tsP

ark

- H

ume

Pea

k

19

96

Pem

bert

on,

Poo

le F

ores

tG

. Li

ddel

ow,

note

3 b

elow

App

roxi

mat

e da

te.

Abu

ndan

ce r

ated

as

1 (lo

w)

on s

cale

of

1–3

for

exte

nt o

f vi

sibl

e ac

tivity

(sc

ats

with

in r

unw

ays,

ext

ent

Blo

ck (

seco

nd s

ite)

of r

unw

ays

and

spoo

r)

19

96

Pem

bert

on,

Tin

kers

Bro

ok,

G.

Lidd

elow

, no

te 3

bel

owA

ppro

xim

ate

date

. A

bund

ance

rat

ed a

s 1

(low

) on

sca

le o

f 1–

3 fo

r ex

tent

of

visi

ble

activ

ity (

scat

s w

ithin

run

way

s, e

xten

tS

utto

n F

ores

t B

lock

of r

unw

ays

and

spoo

r)

19

96

Pem

bert

on,

Poo

leG

. Li

ddel

ow,

note

3 b

elow

App

roxi

mat

e da

te.

Abu

ndan

ce r

ated

as

1 (lo

w)

on s

cale

of

1–3

for

exte

nt o

f vi

sibl

e ac

tivity

(sc

ats

with

in r

unw

ays,

ext

ent

For

est

Blo

ckof

run

way

s an

d sp

oor)

19

96

Jarr

ahda

le,

Cha

ndle

rD

. G

iles

(per

sona

l co

mm

unic

atio

nK

now

n fr

om o

ne s

ight

ed i

ndiv

idua

l. P

resu

mab

le c

ontig

uous

and

/or

from

Cha

ndle

r R

oad

popu

latio

nR

oad

Orc

hard

to P

JdeT

)

19

96

Stir

ling

Ran

ge N

atio

nal

Sin

clai

r (1

999)

Not

ed a

s a

sing

le d

ead

anim

al f

ound

on

Che

ster

Pas

s R

oad

in 1

996.

Dat

e pr

ovid

ed a

nd l

ocat

ion

give

n to

Mar

k R

oddy

,P

ark,

Che

ster

Pas

s R

oad

Par

k R

ange

r, as

opp

osite

Tol

l P

eak

car

park

, C

hest

er P

ass

Roa

d

19

97

Jarr

ahda

le,

Gor

don

P. B

att

Gile

s (p

erso

nal

Iden

tifie

d fr

om e

xten

sive

run

way

s, m

ay b

e co

ntin

uos

with

the

Ros

ella

Roa

d po

pula

tion

For

est

Blo

ckco

mm

unic

atio

n to

PJd

eT)

19

98

Two

Peo

ples

Bay

Nat

ure

Frie

nd a

nd B

utle

r (n

omin

ally

199

8)In

cide

ntal

, no

n-ta

rget

cap

ture

of

quok

kas

at 3

site

s at

Tw

o P

eopl

es B

ay N

atur

e R

eser

ve w

hen

mon

itorin

g/su

rvey

ing

for

Res

erve

, M

t G

ardn

er a

rea

Gilb

ert’s

pot

oroo

. S

ites

wer

e E

ast

Fire

brea

k, L

ower

Fire

brea

k &

Wes

t 6.

Map

ped

here

as

1 lo

catio

n, n

ear

Mt

Gar

dner

19

99

Two

Peo

ples

Bay

Nat

ure

Frie

nd a

nd B

utle

r (n

omin

ally

199

9)In

cide

ntal

, no

n-ta

rget

cap

ture

of

quok

kas

at 4

site

s at

Tw

o P

eopl

es B

ay N

atur

e R

eser

ve w

hen

mon

itorin

g/su

rvey

ing

for

Res

erve

, M

t G

ardn

er a

rea

Gilb

ert’s

pot

oroo

. S

ites

wer

e E

ast

Fire

brea

k, L

ower

Fire

brea

k, N

orth

Fire

brea

k &

Hak

ea.

Map

ped

here

as

1 lo

catio

n, n

ear

Mt

Gar

dner

20

01

Per

th w

ater

sup

ply

J. L

iddi

ngto

n (p

erso

nal

One

ind

ivid

ual

quok

ka t

rapp

ed a

s a

non-

targ

et i

ncid

enta

l ca

ptur

e w

hen

trap

ping

for

pig

s. F

irst

repo

rted

cap

ture

catc

hmen

t, C

hurc

hman

com

mun

icat

ion

to P

JdeT

)fr

om t

his

site

For

est

Blo

ck

20

01

Per

th w

ater

sup

ply

J. L

iddi

ngto

n (p

erso

nal

Quo

kka

caug

ht a

s a

non-

targ

et i

ncid

enta

l ca

ptur

e w

hen

trap

ping

for

pig

s. S

econ

d ca

ptur

e fr

om t

his

catc

hmen

t, w

ater

com

mun

icat

ion

to P

JdeT

)se

ctio

n of

cat

chm

ent

catc

hmen

t ex

clus

ion

with

inC

hurc

hman

For

est

Blo

ck

20

02

Per

th w

ater

sup

ply

J. L

iddi

ngto

n (p

erso

nal

Quo

kka

caug

ht a

s a

non-

targ

et i

ncid

enta

l ca

ptur

e w

hen

trap

ping

for

pig

s. T

hird

cap

ture

fro

m t

his

sect

ion

of c

atch

men

tca

tchm

ent,

Chu

rchm

anco

mm

unic

atio

n to

PJd

eT)

For

est

Blo

ck (

seco

nd s

ite)

Tabl

e 4

(con

t.)

Year

of

Loca

tion

orS

ourc

eC

omm

ent

reco

rdS

ite n

ame

Page 47: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka 59

20

02

Per

th w

ater

sup

ply

J. L

iddi

ngto

n (p

erso

nal

Con

firm

ed r

oadk

ill.

Fou

rth

reco

rd w

ithin

12–

15 m

onth

s fr

om t

his

sect

ion

of c

atch

men

tca

tchm

ent,

Chu

rchm

anco

mm

unic

atio

n to

PJd

eT)

For

est

Blo

ck (

third

site

)

20

02

Nor

thcl

iffe,

Nai

rn F

ores

tM

. S

heeh

an (

pers

onal

Roa

dsid

e si

ghtin

g of

liv

e qu

okka

s an

d se

vera

l co

nfirm

ed r

oadk

ill r

ecor

ds.

Roa

dkill

s ap

pear

to

be a

ssoc

iate

d w

ithB

lock

com

mun

icat

ion

to P

JdeT

)di

stur

banc

e re

sulti

ng f

rom

roa

ding

act

ivity

, pr

ior

to t

imbe

r ha

rves

ting

20

02

Dw

ellin

gup,

Urb

rae

R.

Sta

ines

(pe

rson

alT

hree

quo

kkas

tra

pped

and

one

car

cass

fou

nd i

n pe

riod

21

Febr

uary

to

11 M

arch

200

2. T

rapp

ed q

uokk

as w

ere

non-

For

est

Blo

ckco

mm

unic

atio

n to

PJd

eT)

targ

et i

ncid

enta

l ca

ptur

es w

hen

trap

ping

for

pig

s. W

ithin

3.5

km o

f su

b ca

tchm

ent

whe

re C

hris

tens

en (

unda

ted)

tra

pped

in 1

972

20

02

Dw

ellin

gup,

Kya

bram

R.

Sta

ines

(pe

rson

alW

ithin

Alc

oa’s

Will

owda

le m

ine

site

. Tra

pped

as

a no

n-ta

rget

inc

iden

tal

capt

ure

whe

n tr

appi

ng f

or p

igs.

Kno

wn

at t

his

site

Bro

ok,

Par

k F

ores

t B

lock

com

mun

icat

ion

to P

JdeT

)fr

om o

ne c

aptu

re o

nly

20

02

Qua

rrum

Nat

ure

Res

erve

,G

. F

reeb

ury

(per

sona

lQ

uokk

a pr

esen

ce c

onfid

ently

inf

erre

d by

det

ectio

n of

sca

ts i

n ch

arac

teris

tic q

uokk

a r

unw

ays

east

of

Wal

pole

com

mun

icat

ion

to P

JdeT

)

20

02

Pea

k F

ores

t B

lock

,G

. F

reeb

ury

(per

sona

lQ

uokk

a pr

esen

ce c

onfid

ently

inf

erre

d by

det

ectio

n of

sca

ts i

n ch

arac

teris

tic q

uokk

a ru

nway

sN

orth

of

Wal

pole

com

mun

icat

ion

to P

JdeT

)

20

02

Mud

dy L

ake,

sou

th(D

ell

& H

yder

-Grif

fiths

200

2)T

he a

utho

rs r

epor

ted

‘road

kill

spe

cim

ens

had

been

col

lect

ed r

ecen

tly’ a

nd ‘c

arca

sses

had

bee

n br

ough

t …

in

… u

p un

tilof

Bun

bury

abou

t tw

o ye

ars

ago’

. A

sku

ll co

llect

ed f

orm

the

site

on

25 S

epte

mbe

r 20

02 h

as b

een

conf

irmed

as

a qu

okka

(W

AM

reco

rd M

5413

2),

how

ever

the

sku

ll is

not

age

d (N

orah

Coo

per,

per

s. c

omm

. to

PJd

eT)

and

quok

ka p

rese

nce

is o

ther

wis

eon

ly i

nfer

red

from

pre

senc

e of

run

way

s

20

03

Dw

ellin

gup

(Ban

ksia

dale

M.

Bur

t (p

erso

nal

com

mun

icat

ion

Roa

dkill

. C

arca

ss r

epor

ted

to M

erv

Bur

t, D

wel

lingu

p. C

arca

ss c

onfir

med

at

swam

p si

te,

not

colle

cted

For

est

Blo

ck)

- w

ithin

to P

JdeT

)A

lcoa

’s H

untly

Min

e S

item

inin

g en

velo

pe

20

03

Del

Par

k R

oad,

Dw

ellin

gup

M.

Bur

t (p

erso

nal

com

mun

icat

ion

Roa

dkill

. C

arca

ss r

epor

ted

and

colle

cted

by

Mer

v B

urt,

Dw

ellin

gup.

Dec

ompo

sed

carc

ass

colle

cted

(Tur

ner/

Mar

rinu

p F

ores

tto

PJd

eT)

Blo

ck)

20

03

Big

hill

Bro

ok (

Nai

rnM

. S

heeh

an (

pers

onal

Roa

dkill

, W

heat

ley

Coa

st R

oad,

300

m N

orth

cliff

e si

de o

f B

ighi

ll B

rook

For

est

Blo

ck)

com

mun

icat

ion

to P

JdeT

)

20

03

Nan

nup

(Bor

onia

G.

Voi

gt (

pers

onal

com

mun

icat

ion

Cha

ract

eris

tic q

uokk

a ru

nway

s id

entif

ied

and

conf

irmed

as

quok

ka b

y G

. Li

ddel

ow,

Dep

artm

ent

of C

onse

rvat

ion

and

For

est

Blo

ck)

to P

JdeT

)La

nd M

anag

emen

t. Id

entif

ied

prio

r to

pro

pose

d si

lvic

ultu

ral

burn

20

03

Nan

nup

(Nel

son

For

est

M.

Max

wel

l (pe

rson

alD

epar

tmen

t of

Con

serv

atio

n an

d La

nd M

anag

emen

t B

lack

woo

d D

istr

ict

reco

rd n

o 70

, no

ctur

nal

sigh

ting

Blo

ck)

com

mun

icat

ion

to P

JdeT

)

20

03

Nan

nup

(Bea

ton

For

est

M.

Max

wel

l (pe

rson

alD

epar

tmen

t of

Con

serv

atio

n an

d La

nd M

anag

emen

t B

lack

woo

d D

istr

ict

reco

rd n

o 74

. D

ead,

pre

sum

ably

a r

oadk

illB

lock

)co

mm

unic

atio

n to

PJd

eT)

20

03

Nan

nup

(Gre

gory

For

est

M.

Max

wel

l (pe

rson

alD

epar

tmen

t of

Con

serv

atio

n an

d La

nd M

anag

emen

t B

lack

woo

d D

istr

ict

reco

rd n

o 76

. D

ead,

pre

sum

ably

a r

oadk

illB

lock

)co

mm

unic

atio

n to

PJd

eT)

Page 48: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

60 P.J. de Torres et al.

20

03

Mt

Chu

dalu

p, W

indy

M.

She

ehan

(pe

rson

alTw

o qu

okka

s, n

ight

sig

htin

g, e

aste

rn s

ide

of r

oad

near

‘w

ater

poin

t’ at

Mt

Chu

dalu

p m

onad

nock

Har

bour

Roa

dco

mm

unic

atio

n to

PJd

eT)

20

03

Kar

ri G

ully

, D

alga

rup

K.

Red

man

(pe

rson

alT

hree

sep

arat

e ro

ad k

ill r

ecor

ds o

n B

rock

man

Hig

hway

, be

twee

n N

annu

p an

d B

ridge

tow

n, a

ll w

ithin

the

firs

t ha

lf of

200

3F

ores

t, B

rock

man

hig

hway

,co

mm

unic

atio

n to

PJd

eT)

betw

een

Nan

nup

and

Bri

dg

eto

wn

20

03

Sal

mon

Bea

ch R

oad,

off

M.

She

ehan

(pe

rson

alF

our

sepa

rate

sig

htin

gs o

ver

thre

e w

eeks

on

road

side

in

coas

tal

heat

h/st

unte

d pe

pper

min

t, ne

ar p

ump

shed

Win

dy H

arbo

ur R

oad

com

mun

icat

ion

to P

JdeT

)

20

03

Jarr

ahda

le,

Fro

llet

D.

Gile

s (p

erso

nal

com

mun

icat

ion

Fou

r qu

okka

s si

ghte

d fle

eing

a b

urn

Pla

ntat

ion

(bet

wee

nto

PJd

eT)

Mun

dlim

up a

nd C

obia

cfo

rest

blo

cks)

20

03

Nor

thcl

iffe,

Whe

atle

yP.

Sar

giso

n (p

erso

nal

One

sub

-adu

lt m

ale

road

kill,

200

–300

m s

outh

of

Orc

hid

Roa

d, o

n ea

ster

n si

de o

f W

heat

ley

Coa

st R

oad.

Car

cass

Coa

st R

oad

com

mun

icat

ion

to P

JdeT

)re

tain

ed f

or c

olle

ctio

n by

Dep

artm

ent

of C

onse

rvat

ion

and

Land

Man

agem

ent

staf

f

20

03

Na

nn

up

/Bri

dg

eto

wn

,K

. R

edm

an (

pers

onal

One

adu

lt m

ale

road

kill

. H

alf

way

bet

wee

n N

annu

p an

d B

ridge

tow

n –

400m

W o

f Ja

rrah

Par

kB

rock

man

Hig

hway

com

mun

icat

ion

to P

JdeT

)

20

03

Gre

en R

ange

J.A

. F

riend

(pe

rson

alH

air

sam

ple

colle

cted

whe

n su

rvey

ing

(hai

r tu

bes)

for

Gilb

ert’s

Pot

oroo

.co

mm

unic

atio

n to

PJd

eT)

Not

es:

1K

abay

and

Sta

rt (

1976

) S

ough

t in

form

atio

n fr

om t

he p

ublic

, fr

om f

ield

sta

ff fr

om r

elev

ant

gove

rnm

ent

agen

cies

and

car

ried

out

biol

ogic

al s

urve

y to

det

ect

the

pres

ence

of

Gilb

ert’s

pot

oroo

,P

otor

ous

trid

acty

lus

gilb

ertii

, (n

ow P

. gi

lber

tii)

and

the

broa

d fa

ced

poto

roo,

P.

plat

yops

, in

the

per

iod

Apr

il 19

75 t

o O

ctob

er 1

976.

Quo

kka

pres

ence

was

als

o re

cord

ed a

t nu

mer

ous

site

sin

vest

iga

ted

.2.

Unp

ublis

hed

data

base

rec

ords

of

A.N

. S

tart

, W

este

rn A

ustr

alia

n D

epar

tmen

t of

Con

serv

atio

n an

d La

nd M

anag

emen

t.3.

Unp

ublis

hed

reco

rds

from

inc

iden

tal

sigh

tings

and

opp

ortu

nist

ic t

rapp

ing

by G

. Li

ddel

ow,

Wes

tern

Aus

tral

ian

Dep

artm

ent

of C

onse

rvat

ion

and

Land

Man

agem

ent

Tabl

e 4

(con

t.)

Year

of

Loca

tion

orS

ourc

eC

omm

ent

reco

rdS

ite n

ame

Page 49: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka 61

Table 5

Records of occurrence of the quokka, Setonix brachyurus, extracted from unpublished Department of Conservationand Land Management databases (CALM unpublished; Gilfillan unpublished). Original source for most records wasDepartmental operational district staff. Records are predominantly from incidental and opportunistic sightings. Recordsare listed chronologically.

Date of Location Commentrecord

1987 Pemberton, Brockman National Park, near Pemberton-Northcliffe Road carcass – 3 animals

1992 William Bay National Park Information forwarded by Greg Freebury, WADepartment of Conservation and LandManagement. Date is estimate. Diurnal sighting of1 individual

1993 Pemberton, Brockman National Park roadkill

1994 Manjimup, Lewin Forest Block, Davidson Road / Pine Creek gully system No other details

1994 Manjimup, Lewin Forest Block. Davidson Road / Pine Creek gully system

1994 Manjimup, 7 Day Road / Court Rd No other details

1994 Manjimup

1995 Manjimup, Lewin Forest Block, Junction Davidson & Easter Rd No other details

1995 Manjimup, Andrew Forest Block No other details

1995 Manjimup, Lewin Forest Block, junction of Davidson and Easter roads No other details

1995 Manjimup, Lewin Forest Block, junction of Davidson and Easter roads

1995 Manjimup, Lewin Forest Block, junction Davidson and Easter roads

1995 Manjimup, Andrew Forest Block

1996 Walpole-Nornalup National Park, Cemetery Road Reserve 31362

1996 Walpole-Nornalup National Park, Monastery Road, 1.5 km fromSouth Coast Highway

1996 Walpole-Nornalup National Park, South Coast Highway, 0.5 km westof Gully Road

1996 Walpole-Nornalup National Park, near Spike Road Reserve 31362

1996 Walpole-Nornalup National Park, Frankland River, nearMonastery Landing

1996 Walpole-Nornalup National Park, Cemetery Road Reserve 31362

1996 Walpole-Nornalup National Park, Allen Road, 0.5 km from Hilltop Road

1996 Walpole-Nornalup National Park, Pool Road

1996 Walpole-Nornalup National Park, eastern boundary of Reserve 31362

1996 Walpole, Valley of the Giants, Valley of the Giants Road

1996 Walpole, Loc. 10190, 1 km northwest of junction of Jones Road andHilltop Road

1996 Walpole, Valley of the Giants, Bohall Road, 2.5 km from South CoastHighway

1996 Walpole, Valley of the Giants, I km along track running southeast ofPedro firebreak

Page 50: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

62 P.J. de Torres et al.

1996 Walpole, Valley of the Giants, northern end of Twin Creek Road

1996 Walpole, Valley of the Giants, Court Road, 1 km from South Coast Hwy

1996 Walpole, Valley of the Giants, South Coast Highway, 250 m west ofConspicuous Beach Road.

1996 Walpole, 1.5 km northwest of junction of Jones Road and Hilltop Road

1996 Walpole, Valley of the Giants, Valley of the Giants Road

1996 Walpole-Nornalup National Park, Monastery Road, 0.5 km west ofZig Zag Road

1996 Walpole, Valley of the Giants, Valley of the Giants Road

1996 Walpole, Valley of the Giants, Rate Road 3.3 km north ofSouth Coast Hwy

1996 Pemberton, Brockman NP, adjacent to farmland Diurnal sighting of 2 quokka disturbed by andescaping from machine (dozer) disturbance

1996 Walpole, Valley of the Giants, South Coast Hwy, 1.1 km east ofConspicuous Beach Road

1996 Frankland, Ford Road 3 km from Talbot Road

1996 Walpole Inlet, near Collier Creek, eastern edge

1996 Nornalup, Nornalup Bridge, South Coast Highway.

1996 Manjimup, Solai Forest Block, Solai Rd HC6395 No other details

1996 Walpole, Valley of the Giants, South Coast Highway, 1km southof 28 Mile Road

1996 Manjimup, Yardup Forest Block, 1km North along Edwards Roadfrom Perup Road Database records indicate two different sightings,

same day. The source is identified for one sightingonly and is possibly a duplicate record. The Yarduprecord(s), combined with the 1954 record by A.D.Jones in Serventy et al. (1954) represent the onlyquokka records from the Perup forest.

1996 Manjimup, Lewin Forest Block, Easter & Eastwin roads No other details

1996 Manjimup, Nelson Forest Block, Bibbulmun Track approx 500m south No other detailsof Willow Springs

1996 Walpole, Valley of the Giants, near South Coast Highway, 2 km westof Nut Road

1996 Manjimup, Solai Forest Block, Solai Road

1996 Manjimup, Yardup Forest Block, 1km north along Edwards Road from No other detailsPerup Road

1996 Roe, first creek system west of Claude Road, on Roe Road.(near Mt. Roe)

1996 D’Entrecasteaux National Park, Mandalay Beach Road, near beach

1996 Walpole-Nornalup National Park, Tinglewood Road, 1.5 km from SouthCoast Highway

1996 Manjimup, Nelson Forest Block, Bibbulmun Track approx 500m south ofWillow Springs

1996 Walpole, Keystone Rd. 1.5 km north of South Coast Highway

Table 5 (cont.)

Date of Location Commentrecord

Page 51: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka 63

1996 Manjimup, Easter Forest Block, Easter and Eastwin roads

1996 Walpole-Nornalup National Park, Shelleys Beach

1996 Walpole-Nornalup National Park, South Coast Highway, 0.75 km westof Tinglewood Road

1996 Walpole-Nornalup National Park, walking track between Sealers Coveand Circus Beach

1996 Walpole-Nornalup National Park, gully north-northwest of Circus Beach

1996 Walpole-Nornalup National Park, 1 km northwest of Circus Beach

1996 Walpole, Rest Point Road, 1 km from South Coast Highway

1996 Denbarker, Denmark- Mt. Barker Road, near Mitchell River Bridge

1997 Manjimup, Netic Forest Block, Pool Road

1997 Walpole, Approximately 200m east of the junction of Tree Top Walk RoadkillRoad and Valley of the Giants Road on Valley of the Giants Road

1997 Pemberton, D’Entrecasteaux National Park, near Landslide Road in diurnal sighting of 1 adult quokkaaerial burn area DC16

1997 Walpole, Valley of the Giants Road approximately 200m west of the Roadkill, carcass identifiedTree Top Walk turn-off

1997 Manjimup, Graphite Forest Block, Austin Road

1997 Manjimup, Andrew Forest Block, Top Road

1997 Manjimup, Netic Forest Block, Netic quokka exclusion (WHAT IS THIS)

1997 Manjimup, Netic Forest Block, Pool Road

1997 Manjimup, Netic Forest Block, junction of Sexton and Parky Roads

1997 Manjimup, Netic Forest Block, Sexton Road

1997 Manjimup, Netic Forest Block, Kanny Road

1997 Manjimup, Andrew Forest Block, Dalberg Road waterpoint – 2.5 kmwest of Austin Road

1997 Pemberton, D’Entrecasteaux National Park Roadkill on Salmon Beach Road 1km from WindyHarbour. Definite signs of quokka activity in 4yearold heath north of road

1997 Manjimup, Mack Forest Block, Well Road No other details

1997 Manjimup, Netic Forest Block, Kanny / Parky Roads

1997 Pemberton, Crowea Forest Block, McAlpine Road waterpoint Presence inferred by signs (scats, runways)

1997 Walpole, Walpole-Nornalup National Park, walk trail from town to Diurnal sighting of one individual, crossingCoalmine Beach walk trail

1997 Manjimup, Solai Forest Block, Monk / Solai Roads

1997 Pemberton, Crowea Forest Block, thick ti-tree between coupes Presence inferred by signs (scats, runways)on Crowea Road

1997 Pemberton, Crowea Forest Block Diurnal sighting

1997 Pemberton, Crowea Forest Block, track crossing Orchid Rd,1.1km from Crowea Road

1997 Pemberton, Crowea Forest Block, south-west corner 1984 regen Presence inferred by signs (scats, runways)on south side of road

Date of Location Commentrecord

Page 52: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

64 P.J. de Torres et al.

1997 Pemberton, Crowea Forest Block, McAlpine Road, water point Presence inferred by signs (scats, runways)near Hawkins Road and Dan Road

1997 Pemberton, Crowea Forest Block, Rowney Road north of Crowea Rd Presence inferred by signs (scats, runways)

1997 Pemberton, Crowea Forest Block, Karri/Marri gully 1.9km fromWheatley Coast Road

1997 Pemberton, Crowea Forest Block, track crossing Cederman Road400m from Wheatley Coast Road. Karri/Marri regrowth

1997 Walpole, Wye Forest Block Presence inferred by signs (spoor)

1997 Manjimup, Mack Forest Block, Mt Mack Road / Tom Road intersection

1997 Manjimup, Netic Forest Block, Pool Road No other details

1997 Walpole, Rocky Forest Block No detail, presumably a sighting

1997 Walpole, Rocky Forest Block Old (not fresh) evidence in unburnt gully

1997 Walpole, Sharpe Forest Block Presumably determined by evidence of activity, atcreek in Sharpe Block (North of Sharpe 6), flowsinto the Deep River

1997 Manjimup, Solai Forest Block, Monk / Solai Roads No other details

1997 Manjimup, Andrew Forest Block, Top Road No other details

1997 Manjimup, Graphite Forest Block, Austin Road No other details

1997 Manjimup, Graphite Forest Block, Cow Brook – Davidson Road No other details

1997 Manjimup, Gordon Forest Block, Mobil Road No other details

1997 Manjimup, Andrew Forest Block, Dalberg Road waterpoint – 2.5 km No other details

west of Austin Road

1997 Manjimup, Mack Forest Block, McNab Well No other details

1997 Manjimup, Netic Forest Block, Penny Road No other details

1997 Manjimup, Netic Forest Block, Kanny Road No other details

1997 Manjimup, Netic Forest Block, Kanny / Parky Roads No other details

1997 Manjimup, Netic Forest Block, Junction of Sexton and Parky Roads No other details

1997 Manjimup, Netic Forest Block, Pool Road No other details

1997 Manjimup, Netic Forest Block No other details

1997 Manjimup, Mack Forest Block, Mt Mack Road / Tom Road intersection No other details

1997 Walpole, Rocky Forest Block Recorded as gully crossing road, presumablypresence inferred by evidence of activity

1997 Manjimup, Mack Forest Block, McNab Well

1997 Manjimup, Gordon Forest Block, Mobil Road

1997 Manjimup, Netic Forest Block, Penny Road

1997 Manjimup, Graphite Forest Block, Cow Brook – Davidson Road

1997 Manjimup, Netic Forest Block, Sexton Road No other details

Table 5 (cont.)

Date of Location Commentrecord

Page 53: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka 65

1997 Walpole, Ford Road approximately 2.0km south west from the corner diurnal sighting of 1 quokka crossing roadof Ford and Collis roads

1997 Walpole, approximately 200m west of the Conspicuous Beach Roadkillturn-off on the South Coast Highway (northern verge)

1997 Manjimup, Mack Forest Block, Well Road

1998 Walpole, Walpole-Nornalup National Park, Valley of the diurnal sighting of 1 quokka crossing carparkGiants old carpark

1998 Walpole, Walpole-Nornalup National Park, Valley of the Giants Road, night sighting of adult quokka crossing roadapproximately 500m south of Howe Road

1998 Walpole, 100m west on Beardmore and Thomson roads diurnal sighting of 1 quokka crossing road

1998 Walpole, approximately 400m west of the Jack Rate Lookout on nocturnal sighting of 1 quokka at roadsidethe South West Highway

1999 Walpole, South Coast Highway approximately 3.5km east of Walpole Roadkill 1 adult

1999 Walpole, Jones Road, 1km south of Clarke Road Nocturnal sighting of 3 quokkas, at least 1 adult.Runways present in vegetation

1999 Manjimup, Beavis Forest Block, Beavis 8 diurnal sighting of 2 quokkas

1999 Manjimup, Lewin Forest Block, Pine Creek Road, 1.3 km from CarcassDavidson Road

1999 Walpole, Walpole-Nornalup National Park, 100m west of Nornalup Bridge Roadkill, carcass identified

1999 Manjimup, 220 metres east of Boundary Road on Willow Spring Road identified from carcass

1999 Manjimup, Andrew Forest Block diurnal sighting of 2 quokkas

1999 Manjimup, near intersection Penny Road and Willow Spring Road diurnal sighting of 1 quokka

1999 Stirling Range National Park, Pyungoorup Peak Carcass forwarded to Western Australian Museum

1999 Two Peoples Bay Nature Reserve, on access track to research quarters Nocturnal sighting of 1 quokka

1999 Walpole, South West Highway, approximately 400m west of the Roadkill 1 adultJack Rate Lookout and approximately 700m east of Tinglewood Road

2000 Waychinicup National Park carcass

2000 Two Peoples Bay Nature Reserve, Little Beach Road, start of slashed Nocturnal sighting of 1 quokkafirebreak to south of road

2000 Two Peoples Bay Nature Reserve, approximately 2–300 m from Nocturnal sighting of 2 quokkasstart of Little Beach Road

2000 Waychinicup National Park, Waychinicup Road, just north carcassof creek crossing

2000 Manjimup, Nelsons Location 9466 Jones Rd, Yanmah / Glenoran area day sighting of 3 individuals, feeding onhousehold scraps

2000 Waychinicup National Park, campsite Nocturnal sighting of 2 quokkas

2000 Two Peoples Bay Nature Reserve, Little Beach Road, start of slashed carcassfirebreak to south of road

2000 Manjimup, Beavis Forest Block

2000 Manjimup, Donnelly Mill Road nocturnal sighting of 1 quokka

2000 Manjimup

2000 Manjimup, 100m South of Palings Road and Coronation Road Carcass

Date of Location Commentrecord

Page 54: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

66 P.J. de Torres et al.

2000 Manjimup, 1km east along Telephone Road from Coronation Road Carcass

2000 Manjimup one individual sighted running from road to scrub

2001 Walpole, 1/2 way between Deep River and Crystal Springs on carcassSouth Western Highway

2001 Walpole, Crystal Springs, Coalmine Beach turnoff on South Coast CarcassHighway, approximately 3km east of Walpole

2001 Stirling Range National Park, Mt. James Road capture of 1 quokka during Western Shieldmonitoring

2001 Dwellingup, Del Park Road, south of Alcoa minesite Roadkill 1 adult

Table 5 (cont.)

Date of Location Commentrecord

Page 55: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka

67

Table 6

Previously unpublished records of occurrence of the quokka, Setonix brachyurus, held by the authors. Records are from opportunistic sightings and trapping programsimplemented to determine presence. Records are listed chronologically.

Year of Location or Source Commentrecord Site name

1971 Dwellingup, Lewis (Wild Pig) Swamp Dillon (1993) see note 1, below Reported on the 1971 survey of the wetter western creek systems in vicinity of Dwellingup. A total of 52quokkas caught at 6 sites. Population density estimates of 1 animal per 2 ha in swamps unburnt for 5–10years. Locations nominally mapped as Lewis and Holyoake swamps

1971 Dwellingup, Holyoake Dillon (1993) see note 1, below As above

1988 Dwellingup, Alcoa’s Huntly Mine Site Dillon (1993) see note 1, below Four of 5 sites inspected in 1988 showed evidence of quokka activity. Nominally mapped as Lewisenvelope Swamp, within mine site envelope. No estimates of population size

1992 Dwellingup, Holyoake Dillon (1993) see note 1, below 15 of 30 swamps surveyed in vicinity of Dwellingup in 1992 showed evidence of quokka activity. Three ofthese were then trapped unsuccessfully (fence funnel trap), subsequently one quokka only was trapped(wire cage trap) at the Holyoake site only.

1992 Collie/Mornington, Hadfield R. Brazell, note 2 below Population presumed to be contiguous with population trapped and population size reported byForest Block Hayward et al. (2003)

1992 Collie/Mornington, Gervasse R. Brazell, note 2 below This site known to support a relatively large population. Population estimate reported in this study (seeForest Block Table 7)

1992 Collie/Mornington, Hamilton R. Brazell, note 2 below 3 quokkas trapped when site trapped to determine presence in 1992. Population presumably contiguousForest Block with the population trapped by Hayward et al. (2003)

1992 Collie/Mornington, Victor Road Site, R. Brazell, note 2 below This site known to support a relatively large population. Population presumably contiguous with theHamilton Forest Block population trapped by Hayward et al. (2003)

1992 Collie/Mornington, Hadfield R. Brazell, note 2 below This site known to support a quokka population – population estimate reported by Hayward et al (2002).Forest Block Other incidental quokka reportings and opportunistic trapping from Hadfield Forest Block presumed to be

contiguous with this population

1995 Pemberton, Chudalup Forest Block Dillon (1996), note 3 below Presence indicated, abundance rated at 1 on scale of 0 (absence) to 3 (relatively abundant). Abundanceconsidered comparable with survey in 1978–79

1995 Manjimup, Andrew Forest Block Dillon (1996), note 3 below Presence indicated, abundance rated at 2 on scale of 0 (absence) to 3 (relatively abundant). Unknown if(second site) abundance has changed since survey in 1978–79

1995 Manjimup, Lindsay Forest Block Dillon (1996), note 3 below Presence indicated. Abundance rated at 1 on scale of 0 (absence) to 3 (relatively abundant) andconsidered to be reduced from previous survey in 1978–79. Logging operations close to swamp.

1995 Nannup, Mack Forest Block Dillon (1996), note 3 below Presumably contiguous with the Andrew Forest Block (third site) record. Presence indicated, abundancerated at 1 on scale of 0 (absence) to 3 (relatively abundant). Abundance considered comparable withsurvey in 1992–93

Page 56: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

68 P.J. de Torres et al.

19

95

Wal

pole

, W

eld

For

est

Blo

ckD

illon

(19

96),

not

e 3

belo

wP

rese

nce

indi

cate

d, a

bund

ance

rat

ed a

t 1

on s

cale

of

0 (a

bsen

ce)

to 3

(re

lativ

ely

abun

dant

). A

bund

ance

cons

ider

ed c

ompa

rabl

e w

ith s

urve

y in

197

8–79

19

95

Wal

pole

-Nor

nalu

p N

atio

nal

Par

kD

illon

(19

96),

not

e 3

belo

wP

rese

nce

indi

cate

d, a

bund

ance

rat

ed a

t 2

on s

cale

of

0 (a

bsen

ce)

to 3

(re

lativ

ely

abun

dant

). A

bund

ance

(Hill

top

Roa

d or

for

mer

lyco

nsid

ered

com

para

ble

with

sur

vey

in 1

978–

79.

Was

bur

nt i

n w

ildfir

e in

198

7, q

uokk

as s

een

disp

ersi

ng a

tH

illto

p F

ores

t B

lock

)tim

e of

fire

19

95

Wal

pole

, C

ross

ing

For

est

Blo

ckD

illon

(19

96),

not

e 3

belo

wP

rese

nce

indi

cate

d, a

bund

ance

rat

ed a

t 1

on s

cale

of

0 (a

bsen

ce)

to 3

(re

lativ

ely

abun

dant

). A

bund

ance

cons

ider

ed c

ompa

rabl

e w

ith s

urve

y in

197

8–79

19

95

Wal

pole

, G

iant

s F

ores

t B

lock

Dill

on (

1996

), n

ote

3 be

low

Pre

senc

e in

dica

ted,

abu

ndan

ce r

ated

at

1 on

sca

le o

f 0

(abs

ence

) to

3 (

rela

tivel

y ab

unda

nt).

Abu

ndan

ceco

nsid

ered

com

para

ble

with

sur

vey

in 1

978–

79

19

95

Nan

nup,

Gre

gory

For

est

Blo

ckD

illon

(19

96),

not

e 3

belo

wP

rese

nce

indi

cate

d, a

bund

ance

rat

ed a

t 1

on s

cale

of

0 (a

bsen

ce)

to 3

(re

lativ

ely

abun

dant

). A

bund

ance

cons

ider

ed c

ompa

rabl

e w

ith s

urve

y in

199

2–93

. S

ite i

s lo

ng u

nbur

nt

19

95

Nan

nup,

Bea

ton

For

est

Blo

ckD

illon

(19

96),

not

e 3

belo

wP

rese

nce

indi

cate

d, a

bund

ance

rat

ed a

t 1

on s

cale

of

0 (a

bsen

ce)

to 3

(re

lativ

ely

abun

dant

). A

bund

ance

cons

ider

ed c

ompa

rabl

e w

ith s

urve

y in

199

2–93

19

95

Nan

nup,

Bea

ton

For

est

Blo

ckD

illon

(19

96),

not

e 3

belo

wP

rese

nce

indi

cate

d, a

bund

ance

rat

ed a

t 1

on s

cale

of

0 (a

bsen

ce)

to 3

(re

lativ

ely

abun

dant

). A

bund

ance

(thi

rd s

ite)

cons

ider

ed c

ompa

rabl

e w

ith s

urve

y in

199

2–93

19

95

Man

jimup

, A

ndre

w F

ores

t B

lock

Dill

on (

1996

), n

ote

3 be

low

Pre

sum

ably

con

tiguo

us w

ith t

he M

ack

For

est

Blo

ck r

ecor

d. P

rese

nce

indi

cate

d. A

bund

ance

rat

ed a

t 2

on(t

hird

site

)sc

ale

of 0

(ab

senc

e) t

o 3

(rel

ativ

ely

abun

dant

) an

d co

nsid

ered

to

be c

ompa

rabl

e w

ith p

revi

ous

surv

ey i

n19

78–7

9. S

ite d

istu

rbed

by

cons

truc

tion

of n

ewly

dug

wat

er p

oint

.

19

95

Nan

nup,

Hel

ms

For

est

Blo

ckD

illon

(19

96),

not

e 3

belo

wP

rese

nce

indi

cate

d, a

bund

ance

rat

ed a

t 1

on s

cale

of

0 (a

bsen

ce)

to 3

(re

lativ

ely

abun

dant

). A

bund

ance

cons

ider

ed r

educ

ed f

rom

sur

vey

in 1

975.

sw

amp

patc

h bu

rnt

in 1

994

(1 y

ear

prio

r to

thi

s as

sess

men

t)

19

95

Nan

nup,

Bea

ton

For

est

Blo

ckD

illon

(19

96),

not

e 3

belo

wP

rese

nce

indi

cate

d, a

bund

ance

rat

ed a

t 1

on s

cale

of

0 (a

bsen

ce)

to 3

(re

lativ

ely

abun

dant

). U

nabl

e to

(sec

ond

site

)de

term

ine

if ch

ange

in

abun

danc

e fr

om p

revi

ous

surv

ey

19

95

Nan

nup,

Bea

ton

For

est

Blo

ckD

illon

(19

96),

not

e 3

belo

wP

rese

nce

indi

cate

d, a

bund

ance

rat

ed a

t 1

on s

cale

of

0 (a

bsen

ce)

to 3

(re

lativ

ely

abun

dant

). U

nabl

e to

(fift

h si

te)

dete

rmin

e if

abun

danc

e ha

s ch

ange

d si

nce

prev

ious

sur

vey

19

95

Man

jimup

, A

ndre

w F

ores

t B

lock

Dill

on (

1996

), n

ote

3 be

low

Pre

senc

e in

dica

ted,

abu

ndan

ce r

ated

at

2 on

sca

le o

f 0

(abs

ence

) to

3 (

rela

tivel

y ab

unda

nt).

Unk

now

n if

abun

danc

e ha

s ch

ange

d si

nce

surv

ey i

n 19

78–7

9

19

95

Nan

nup,

Bea

ton

For

est

Blo

ckD

illon

(19

96),

not

e 3

belo

wP

rese

nce

indi

cate

d, a

bund

ance

rat

ed a

t 2

on s

cale

of

0 (a

bsen

ce)

to 3

(re

lativ

ely

abun

dant

). U

nabl

e to

(fou

rth

site

)de

term

ine

if ab

unda

nce

chan

ged

from

pre

viou

s su

rvey

19

95

Nan

nup,

Gre

gory

For

est

Blo

ckD

illon

(19

96),

not

e 3

belo

wP

rese

nce

indi

cate

d, a

bund

ance

rat

ed a

t 1

on s

cale

of

0 (a

bsen

ce)

to 3

(re

lativ

ely

abun

dant

). u

nabl

e to

(sec

ond

site

)de

term

ine

if ab

unda

nce

has

chan

ged

sinc

e la

st s

urve

y. r

unw

ays

appe

ar u

nuse

d, o

ld s

cat

only

det

ecte

d

Tabl

e 6

(con

t.)

Year

of

Loca

tion

orS

ourc

eC

omm

ent

reco

rdS

ite n

ame

Page 57: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka 69

19

95

Dw

ellin

gup,

Kes

ners

Sw

amp,

P. d

e To

res,

M.

Dill

on,

A. T

omki

nson

Kno

wn

to s

uppo

rt a

pop

ulat

ion

in t

he 1

970s

, po

pula

tion

pres

ence

con

firm

ed t

hrou

gh t

rapp

ing

inM

arrin

up F

ores

t B

lock

and

R.

Bue

hrig

, no

te 4

bel

owF

ebru

ary

to M

arch

199

5. P

opul

atio

n si

ze e

stim

ated

by

Hay

war

d (2

003)

19

95

Dw

ellin

gup,

Lew

is (

Wild

Pig

) S

wam

p,P.

de

Tore

s, M

. D

illon

, A

. Tom

kins

onS

how

ed s

igns

of

fres

h ac

tivity

in

1995

, pr

esen

ce u

nabl

e to

be

conf

irmed

by

trap

ping

. P

rese

nce

also

Wils

on F

ores

t B

lock

and

R.

Bue

hrig

, no

te 4

bel

owun

able

to

be c

onfir

med

in

2000

by

Hay

war

d et

al.

(200

3) a

nd t

his

loca

tion

may

no

long

er s

uppo

rts

ap

op

ula

tion

19

95

Dw

ellin

gup,

Hol

yoak

e F

ores

t B

lock

P. d

e To

res,

M.

Dill

on,

A. T

omki

nson

Sho

wed

evi

denc

e of

act

ivity

in

1995

, ho

wev

er,

by 2

000

cons

ider

ed t

o no

lon

ger

supp

ort

a po

pula

tion

and

R.

Bue

hrig

, no

te 4

bel

ow

19

95

Jarr

ahda

le,

Ros

ella

Roa

d,P.

de

Tore

s, M

. D

illon

, A

. Tom

kins

onTr

appe

d to

con

firm

pre

senc

e. S

even

cap

ture

eve

nts

of 4

quo

kkas

, A

pril

1995

Gor

don

For

est

Blo

ckan

d R

. B

uehr

ig,

note

4 b

elow

19

96

Man

jimup

, S

torr

y F

ores

t B

lock

Dill

on (

1996

), n

ote

3 be

low

Pre

senc

e in

dica

ted,

abu

ndan

ce r

ated

at

1 on

sca

le o

f 0

(abs

ence

) to

3 (

rela

tivel

y ab

unda

nt).

Una

ble

tode

term

ine

if ab

unda

nce

has

chan

ged

sinc

e la

st s

urve

y

19

96

Nan

nup,

Net

ic F

ores

t B

lock

Dill

on (

1996

), n

ote

3 be

low

Pre

senc

e in

dica

ted,

abu

ndan

ce r

ated

at

2 on

sca

le o

f 0

(abs

ence

) to

3 (

rela

tivel

y ab

unda

nt).

Una

ble

tode

term

ine

if ab

unda

nce

has

chan

ged

sinc

e pr

evio

us s

urve

y. f

resh

tra

cks

and

activ

ity b

eyon

d sw

amp

area

19

96

Nan

nup,

Nel

son

For

est

Blo

ckD

illon

(19

96),

not

e 3

belo

wP

rese

nce

indi

cate

d, a

bund

ance

rat

ed a

t 1

on s

cale

of

0 (a

bsen

ce)

to 3

(re

lativ

ely

abun

dant

). U

nabl

e to

dete

rmin

e if

abun

danc

e ha

s ch

ange

d si

nce

prev

ious

sur

vey.

app

ears

6–1

0 ye

ars

sinc

e la

st (

patc

hy)

burn

19

96

Jarr

ahda

le,

Cha

ndle

r R

oad,

Cha

ndle

rP.

de

Tore

s, M

. D

illon

, A

. Tom

kins

onK

now

n to

sup

port

a p

opul

atio

n -p

opul

atio

n es

timat

e re

port

ed b

y H

ayw

ard

et a

l. (2

003)

For

est

Blo

ckan

d R

. B

uehr

ig,

note

4 b

elow

19

96

Jarr

ahda

le,

Alb

any

Hig

hway

Site

.P.

de

Tore

s, M

. D

illon

, A

. Tom

kins

onP

opul

atio

n re

port

ed f

rom

a r

oadk

ill i

n 19

96 a

nd s

ubse

quen

t fo

llow

up

trap

ping

(in

the

sam

e ye

ar)

(Tra

velle

rs’ A

rms

site

).an

d R

. B

uehr

ig,

note

4 b

elow

conf

irm

ed p

rese

nce.

Thi

s si

te p

resu

med

to

be t

he T

rave

llers

’ Arm

s si

te r

efer

red

to i

n 19

56 b

y R

. A

itken

,ci

ted

by B

arke

r et

al.

(195

7) (

Tabl

e 3)

and

tra

pped

by

Ken

t W

illia

ms

in 1

966

(Tab

le 4

)

19

96

Col

lie/

Mor

ning

ton

Roa

d,R

. B

raze

ll, n

ote

2 be

low

Kno

wn

from

1 r

oadk

ill,

sour

ce p

opul

atio

n no

t co

nfirm

edH

amilt

on F

ores

t B

lock

20

00

Col

lie,

Dav

is F

ores

t B

lock

M.

Hay

war

d, n

ote

5 be

low

Pre

senc

e in

ferr

ed b

y de

tect

ion

of s

cats

with

in r

unw

ays

20

00

Dw

ellin

gup,

Mar

rinup

For

est

Blo

ckM

. H

ayw

ard,

not

e 5

belo

wP

rese

nce

infe

rred

by

dete

ctio

n of

sca

ts w

ithin

run

way

s

20

00

Dw

ellin

gup,

Fed

eral

For

est

Blo

ckM

. H

ayw

ard,

not

e 5

belo

wP

rese

nce

infe

rred

by

dete

ctio

n of

sca

ts w

ithin

run

way

s

20

00

Pem

bert

on,

Dic

kson

Roa

dM

. H

ayw

ard,

not

e 5

belo

wP

rese

nce

infe

rred

by

dete

ctio

n of

sca

ts w

ithin

run

way

s

20

00

Dw

ellin

gup,

Fin

lay

Bro

ok,

Wils

onM

. H

ayw

ard,

not

e 5

belo

wP

rese

nce

infe

rred

by

dete

ctio

n of

sca

ts w

ithin

run

way

s. S

poor

sub

sequ

ently

obs

erve

d by

For

est

Blo

ckde

Tor

es a

nd D

illon

20

00

Dw

ellin

gup,

Clin

ton

For

est

Blo

ckM

. H

ayw

ard,

not

e 5

belo

wP

rese

nce

infe

rred

by

dete

ctio

n of

sca

ts w

ithin

run

way

s.

20

00

Dw

ellin

gup,

Alc

oa C

onve

yor

Bel

tM

. H

ayw

ard,

not

e 5

belo

wP

rese

nce

infe

rred

by

dete

ctio

n of

sca

ts w

ithin

run

way

s.

20

00

Dw

ellin

gup,

Sou

th D

anda

lup

Dam

M.

Hay

war

d, n

ote

5 be

low

Pre

senc

e in

ferr

ed b

y de

tect

ion

of s

cats

with

in r

unw

ays.

Als

o kn

own

prev

ious

ly f

rom

inc

iden

tal

sigh

tings

.P

opul

atio

n m

ay b

e co

ntig

uous

with

pop

ulat

ion

at K

esne

rs S

wam

p an

d th

e C

onve

yor

Bel

t S

ite

Page 58: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

70 P.J. de Torres et al.

20

00

Dw

ellin

gup,

Kea

ts F

ores

t B

lock

M.

Hay

war

d, n

ote

5 be

low

Pre

senc

e in

ferr

ed b

y de

tect

ion

of s

cats

with

in r

unw

ays.

20

00

Dw

ellin

gup,

Tur

ner

For

est

Blo

ckP.

de

Tore

s, n

ote

4 be

low

Inci

dent

al s

ight

ing

in M

ay 2

000,

sub

sequ

ent

addi

tiona

l in

cide

ntal

sig

htin

g re

port

ed i

n O

ctob

er 2

000

and

road

kill

(car

cass

col

lect

ed b

y M

. M

axw

ell

and

C. T

reth

owan

) in

Dec

embe

r 20

00.

Als

o su

bseq

uent

lyco

nsid

ered

by

Hay

war

d to

sup

port

a q

uokk

a po

pula

tion

Not

es:

1.D

illon

(19

93)

repo

rted

on

a 19

71 s

urve

y of

cre

ek s

yste

ms

in t

he w

ette

r w

este

rn c

reek

sys

tem

s in

vic

inity

of

Dw

ellin

gup,

a 1

988

surv

ey w

ithin

Alc

oa’s

Hun

tly m

ine

site

env

elop

e ne

ar D

wel

lingu

p an

d19

92 b

road

scal

e su

rvey

to

dete

rmin

e pr

esen

ce i

n sw

amps

nea

r D

wel

lingu

p2.

Unp

ublis

hed

reco

rds

from

inc

iden

tal

sigh

tings

and

opp

ortu

nist

ic t

rapp

ing

by R

.I. B

raze

ll, W

este

rn A

ustr

alia

n D

epar

tmen

t of

Con

serv

atio

n an

d La

nd M

anag

emen

t.3.

Dill

on (

1996

) re

port

on

surv

ey o

f si

tes

surv

eyed

fro

m 1

995

–199

6, w

here

pre

senc

e w

as c

onfir

med

thr

ough

tra

ppin

g, o

r in

dica

ted

by s

cats

obs

erve

d w

ithin

run

way

s an

d ot

her

fres

h si

gns

of a

ctiv

ity–

scat

s al

one

not

cons

ider

ed s

uffic

ient

to

conf

irm p

rese

nce.

4.U

npub

lishe

d re

cord

s fr

om i

ncid

enta

l si

ghtin

gs,

trap

ping

pro

gram

s in

itiat

ed t

o de

term

ine

pres

ence

and

opp

ortu

nist

ic t

rapp

ing

by P

. J.

de

Tore

s, M

. J.

Dill

on,

A.

Tom

kins

on a

nd R

. B

uehr

ig,

Wes

tern

Aus

tral

ian

Dep

artm

ent

of C

onse

rvat

ion

and

Land

Man

agem

ent.

5.R

esul

ts f

rom

unp

ublis

hed

field

sur

vey

cond

ucte

d by

M.

W.

Hay

war

d be

twee

n S

epte

mbe

r 19

98 a

nd S

epte

mbe

r 20

00 w

here

a r

ange

of

pote

ntia

l lo

catio

ns w

ere

exam

ined

for

evi

denc

e of

quo

kka

pres

ence

. W

ith t

he e

xcep

tion

of t

he S

outh

Dan

dalu

p D

am s

ite,

the

site

s lis

ted

wer

e pr

evio

us n

ot k

now

n to

sup

port

quo

kkas

.

Tabl

e 6

(con

t.)

Year

of

Loca

tion

orS

ourc

eC

omm

ent

reco

rdS

ite n

ame

Page 59: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka 71

Table 7

Previously unpublished trapping records for the quokka, Setonix brachyurus, from sites in the northern jarrah forest ofsouth-west Western Australia

Site name Y ear(s) Purpose of trapping Method of analysis Resulttrapped program

Gervasse Forest 1992 to Long-term monitoring Abundance estimates derived from the Population estimate ofBlock, near Collie 2000 of abundance and ‘deaths and no immigration’ Jolly-Seber 49.21 ±7.85 individuals

survivorship model Survivorship estimates derivedfrom the Program MARK Cormack-Jolly-Seber model

Rosella Road, April 1995 Opportunistic trapping to Listing of total number of captured 7 captures of 4 quokkasnear Jarrahdale determine presence only individuals

Subsequently trapped byHayward et al. (2003) toderive populationestimates

Albany Highway, 1996 Opportunistic trapping to Listing of total number of captured 1 capture of an adult malesoutheast of the determine presence only individualsPerth metropolitanarea. This location ispresumed to be theTravelers’ Arms sitereferred to by Barkeret al. (1957) (Table 3)and Williams in 1966(Table 4)

Holyoake, August Opportunistic trapping to N/A No captures, no evidenceDwellingup 1995 determine presence only of recent presence. Old

runways and no freshscats

Lewis (Wild Pig) August Opportunistic trapping to N/A No captures, someSwamp, Dwellingup 1995 determine presence only evidence of recent activity

in runways

Kesners Swamp, February to Opportunistic trapping to Listing of total number of captured 22 captures of 10 quokkasDwellingup June 1995 determine presence only individuals

Subsequently trapped byHayward et al. (2003) toderive population estimates

Page 60: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

72 P.J. de Torres et al.

Table 8

Location records for the quokka, Setonix brachyurus, excluded from maps used to infer distribution and excluded fromestimates of extent of occurrence. Records listed and/or the location described were either unable to be confirmed, orwere from captive colonies/sanctuaries

Location Year of Source of Commentrecord record

North Twin 1905 Shortridge (1909) Shortridge included ‘Twin Peak’ island as a known site for the quokka. UnclearPeak Island if this was a reference to North Twin Peak Island or South Twin Peak Island.

Quokkas have not been recorded from either island by any other author andthe record is considered to be a tammar wallaby, see comments in text

South Twin 1905 Shortridge (1909) Comments for the North Twin Peak Island, above apply.Peak Island Western Australian Museum Identification confirmed as a quokka in 1982 by D.J. Kitchener (WA Museum)Point Culver records. WAM registration (Norah Cooper pers. comm. to PJdeT). The record is outside the known

number 024346 historic range and the range as determined by the fossil record and istherefore considered spurious. No additional location details available and nodate of collection or any other details. Accuracy of location thereforeconsidered insufficient to include within inferred distribution.

Coorow 1949 Western Australian Museum Validity of record questioned by Nora Cooper (WA Museum). Remains wererecords. WAM registration collected from a cave in 1949, subsequently destroyed and unable to benumber 002781 verified (Norah Cooper pers. comm. to PJdeT).

Breaksea 1975 Western Australian Museum See comments in text.Island records. WAM registration

number 002781

Department of 1958 Western Australian Museum Specimen presumed to be from the captive colony held at the University ofZoology, and records. WAM registration Western Australia, ex Rottnest IslandUniversity of 1959 numbers 003365 andWestern 003618Australia

Karakamia 1998 Smitz (1998) Four quokkas originally sourced from mainland locations and released withinSanctuary, a predator proof fenced sanctuary. Three quokkas released in 1996, two ofChidlow, east which were originally sourced from the Gervasse population, near Collie andof Perth held by the ‘Big Swamp Wildlife Park’, Bunbury. The third was a captive bred

young of the first two. The fourth animal sourced from the Rosella Road siteand released in 1997. One death has been recorded and at least 1 younghas been produced, population size unknown at May 2002

Harry Waring 1979 Austin (1979) Population sourced from Rottnest Island and released at the fenced HarryMarsupial Waring Marsupial Reserve. The colony failed to persist.Reserve,Jandakot

Muddy Lake, 2002 Dell and Hyder-Griffiths Unconfirmed record – See listing in Table 4south of (2002)Bunbury

Water 2004 Jim Lane (pers. comm. Unconfirmed recordCorporation to PJdeT)reserve,Dunsborough

Page 61: Review of the distribution, causes for the decline and ... · Marsupialia) (Quoy & Gaimard 1830) is a small to medium sized macropodid marsupial, endemic to the mainland of south-west

Distribution of the quokka 73

Table 10

Locations previously known to support populations of the quokka, Setonix brachyurus, re-surveyed in the period1995–1996 by Dillon (1996) and found to show no evidence of presence or where presence was unable to beconfirmed.

Location Year of last Commentknown record

Netic Forest Block, 1978–79 0 to 2 years since last burnt. Quokkas subsequently reported from other locations within thisManjimup and neighbouring forest block (see Table 5)

Shannon, Walpole 1978–79 0 to 2 years since last burnt, swamp vegetation completely burnt. Quokkas subsequentlyreported from other locations within Shannon River area (see Table 5)

Giants Forest Block, 1978–79 Very heavy litter layer and 11 years or more since last burn. Previously known from a roadkillWalpole only at this site. Quokkas subsequently reported from other locations within this forest block

(see Tables 5 and 6)

Walpole-Nornalup 1978–79 Very heavy litter layer and 11 years or more since last burn. No recent evidence of activity.National Park (Hilltop Quokkas subsequently reported from other locations within this vicinity and may be at highRoad area), Walpole density (see text and Tables 5 and 6)

Sheepwash Forest 1978–79 6 to 10 years since last burn. No sign of recent activityBlock, Denmark

Farmland area, 1976 No evidence of activity, habitat now water logged. Location determined on basis of incorrectBunbury geographic co-ordinates for Muddy Lake listed by Kabay and Start (1976)

Urbrae Forest Block, 1988 11 years or more since last burn. Quokkas subsequently reported from other locations withinDwellingup this forest block (see Table 6)

Holyoake, 1992 3 to 5 years since last burn. Subsequently trapped more intensively (August 1995) (Table 7)Dwellingup with no evidence of quokka activity or presence

Inglehope Forest 1972–73 6 to 10 years since last burn. No sign of activity when surveyed by Dillon (1996) nor whenBlock, Dwellingup surveyed by Hayward (2002) in 1999–2000.

Table 9

Known To Be Alive (KTBA) estimates for the quokka,Setonix brachyurus, at the Gervasse Forest Block site, forthe period 1992 to 2002.

Month and Year KTBA estimateof trapping

June/July 1992 First trapping session, no KTBA estimateavailable

March 1993 14

June/July 1993 20

January 1994 22

August 1994 20

April 1995 19

January 1996 15

January 1997 12

February 1998 12

March 1999 11

February 2000 Last trapping session, no KTBA estimateavailable