DOI: 10.18195/issn.0313-122x.78(1).2010.285-298 Records of the Western Australian Museum, Supplement 78: 285–298 (2010).

The original fauna of the biogeographic region of north-western

Alexander Baynes1 and Matthew C. McDowell2

1Research Associate, Department of Earth and Planetary Sciences, Western Australian Museum, Locked Bag 49, Welshpool DC, 6986, Australia. Email: [email protected]

2Mammal Section, South Australian Museum, North Tce, Adelaide 5000; and School of Biological Science, Flinders University of South Australia, PO Box 2100, Adelaide, South Australia 5001, Australia.

Abstract – The hills and ranges of the Pilbara contain large numbers of caves, but only a miniscule proportion of these holds mammal remains that can be used to reconstruct the original (i.e. pre-European) fauna. During two fi eld seasons in 1985 and 2004, only 12 sites with bones were discovered. Material from three other sites lodged in the collections of the Western Australian Museum was also used. Thirty-six native mammal species plus the introduced House Mouse were identifi ed from cave surface remains. Compared with the fauna known from live-caught specimen records and the results of the Pilbara Biodiversity Survey, rodent species are comprehensively represented among the remains, dasyurid and are moderately well represented, macropodoids are poorly represented, and bats (except two cave-roosting species) hardly represented at all. Results of this study indicate that the Central Rock-rat (Zyzomys pedunculatus), and probably the Golden-backed Tree-rat (Mesembriomys macrurus), originally occurred throughout the Pilbara ranges. Chuditch (Dasyurus geoffroii) is recorded from the Pilbara for the fi rst time, suggesting that it may have occurred in the lowlands, while Northern (D. hallucatus) occupied the ranges. New records of Melomys burtoni and Pseudomys fi eldi represent substantial extensions of geographic range. Overall, these results confi rm the Pilbara as the western outpost of several northern mammal species, but show that the original fauna is less distinctive than previously thought, with a higher proportion of species widespread in the arid zone.

INTRODUCTION specimens representing 11 species (one, Taphozous hilli Kitchener, 1980a, not recognised as distinct at The original (i.e. immediately pre-European) non- the time) have been found (Appendix 1). The main volant mammal fauna of the Pilbara biogeographic collecting methods used were fi rearms and shelter region is incompletely known. The holotype destruction. This approach produced 34 of Mesembriomys macrurus (Peters, 1876) was specimens, 18 bats, four dasyurids and no rodents. collected from a mainland creek on the southern One of the most interesting specimens is the side of Mermaid Strait in the western Dampier , M3432, which Archer (1976) considered Archipelago (Mahoney and Richardson 1988), but to possibly represent an undescribed species th the inland Pilbara was not visited by 19 century within the . Far more mammal specimens collectors of (Glauert 1950; Ride 1968). resulted from collecting work by E.H.M. (‘Tim’) The mammal collection of the Western Australian Ealey and colleagues on Abydos, Eginbah, Mount Museum contains a small number of specimens Edgar, White Springs and Woodstock Stations received during the 1930s and 1940s from Pilbara in the north-eastern Pilbara between 1953 and pastoral stations, but the mammal fauna remained 1962 (Archer 1975). The collections of the Western relatively unknown until the 1950s. The Western Australian Museum contain at least 148 specimens Australian Museum sent an expedition to the representing 15 species of non-volant mammals Hamersley Range in July and August 1958. Ride collected by them (including Pseudantechinus roryi (1959) reported that the expedition obtained, Cooper, Aplin and Adams, 2000, not recognised as apart from many insects, birds, reptiles and fi sh, distinct at the time). The Ealey collection provided about 50 specimens of 13 species of mammals. most of the type series of Dasykaluta rosamondae The specimens from the expedition are diffi cult to (Ride, 1964), the fi rst specimen of timealeyi trace in the Museum mammal collection because Archer, 1975, named after Ealey, and the first they were not registered in a single group, but 56 specimens of Pseudomys chapmani Kitchener, 1980b 286 A. Baynes, M.C. McDowell

(Appendix 1). This work was followed by further mammals, accumulated by predators that used the collecting in the Pilbara, in particular by A.M. caves as shelters. In general the most important Douglas, who concentrated on bats, and W.H. of these are Barn Owls, Tyto alba. Many mammal Butler. These collections contained few specimens species have suffered drastic range reductions of mammal species that fall within the critical (some to ) since the arrival of Europeans weight range (mean adult weight of 35 g to 5500 g) in Australia, and fossil material has been used to identifi ed by Burbidge and McKenzie (1989), and in reconstruct their past distributions and the original particular no large rodents. faunas of regions (e.g. Lundelius 1957; Kendrick In subsequent decades, biological surveys of and Porter 1974; Baynes 1990; Baynes and Baird proposed resource developments and national 1992; Baynes and Jones 1993; McDowell and Medlin parks (e.g. Dunlop and Sawle 1983), and a survey 2010). The vertebrate palaeontological collections by the Western Australian Museum of the Abydos- of the Western Australian Museum contain many Woodstock Reserve (How et al. 1991; How and samples of such remains collected throughout Cooper 2002), have resulted in many mammal the State during the 1960s and 1970s. Only one of specimens being added to the collections of these, however, is from the Pilbara: registration the Western Australian Museum. These have numbers 76.6.21–76.6.25, collected by W.H. Butler in included some additions to the general Pilbara 1976 at Marandoo near Mt Bruce. Although small, non-volant mammal fauna, but not of species of this sample includes the fi rst remains identifi ed the critical weight range (Gibson and McKenzie from the Pilbara of four medium to large rodent 2009). However, faunas from mammal remains species—Notomys longicaudatus, Pseudomys desertor, from the caves in the karst limestone of Cape Range Pseudomys nanus and Rattus tunneyi. to the west of the Pilbara (Kendrick and Porter Two periods of dedicated fi eld work to attempt 1974; Baynes and Jones 1993), suggested that north- to fi nd more such material in the Pilbara were western Australian non-volant mammal faunas carried out by A.B. and Thomas A. Smith between once included several more species. 25 July and 9 August 1985 and by the authors and T.A. Smith from 13 to 27 August 2004, as part of MATERIALS AND METHODS the Pilbara Biodiversity Survey (McKenzie et al. 2009). Very large numbers of rock shelters and rock Not only deep limestone caves but also small overhangs were searched each time but very little caves and holes in cliffs and ‘breakaway’ mesas was found. In 1985, only one large sample was can contain remains of vertebrates, particularly discovered, at Coppin Gap in the east Pilbara. Small

Figure 1 Location of the sites in which mammal remains have been found in the Pilbara biogeographic region. The boundaries of the region are shown in bold, and the four (named) sub-regions in stipple. For abbreviations of site names see Results section. Original mammal fauna 287 samples were found on Yarrie and Pilga Stations in airport, from surface to depth of 1 ft (30 cm) in pot- the east Pilbara, near Wittenoom in the Hamersley hole in honeycombed limestone ridge intersecting Range, and near Urandy Bore and Pannawonica mud flats, 20˚25′S 118˚06′E, E. Atkins (Utah in the west Pilbara. A medium-sized sample was Construction Co.), August 1965. Western Australian found at Shay Gap, just over the eastern boundary Museum vertebrate palaeontological collection, of the Pilbara bioregion. In 2004, one large sample catalogue numbers 65.10.206–65.10.211. was found in a breakaway near Herbert Well on Ord Ranges (OR). Small hole in an outcrop at Station in the west Pilbara, and two about 20˚15′S 119˚05′E, A. Baynes and T.A. Smith, 1 small samples in a valley in the northern face of August 1985. the Hamersley Range. Further substantial samples Island Hill (IH). Small caves in Island Hill, Pilga were also collected from a large sinkhole in the Station, 21˚19′S 119˚20′E, PIL1, A. Baynes and T.A. Ripon Hills, east Pilbara, previously discovered Smith, 28 July 1985. Field numbers ABRS 108/1, and sampled by P.G. Kendrick and J. Angus in 108/2. November 1997. Coppin Gap (CG). Overhang in rocks at top In the laboratory, identifiable bones (mainly of scree slope on eastern side of Coppin Gap, jaws) were sorted from the samples, and identifi ed 20˚53′11″S 120˚07′03″E, PIL1, A. Baynes, 29 July 1985. by comparison with material in the vertebrate Field number ABRS 109. palaeontological and modern mammal collections of the Western Australian and South Australian Shay Gap (SG). Cave in sandstone escarpment Museums. on west side of townsite, 20˚29′55″S 120˚09′44″E, GSD1, A. Baynes and T.A. Smith, 31 July 1985. Field number ABRS 111. RESULTS (YS). Cave in hill at 20˚35′14″S Sites 120˚15′21″E, PIL1, A. Baynes and T.A. Smith, 30 July Site details are given in west to east order. 1985. Field number ABRS 110. The positions of the sites within the Pilbara East Pilbara sinkhole (EP-1). Deep vertical biogeographic region are shown in Figure 1. sinkhole in dolomite at 21˚50′S 120˚30′E, bone ‘Herbert Well’ (HW). Tubular hole in a breakaway material collected by R. Darren Brooks, 17 July 1994, cliff face north of Herbert Well on Yarraloola held by Western Australian Museum (BES 4389). Station, at 21˚39′22″S 115˚54′07″E, WGS 84, 91 m, A. Ripon Hills sinkhole (RHS). Large sinkhole Baynes, M.C. McDowell and T.A. Smith, 18 August in Ripon Hills at 21˚13′27″S 120˚41′40″E, WGS 84, 2004. 396 m. Huge overhangs on northern (Nth) and ‘Urandy Bore’ (UB). Breakaway caves south of southern (Sth) sides of central rock pile. Material Urandy Bore at 22˚25′S 116˚18′E, A. Baynes and T.A. previously collected from the northern side by Smith, 5 August 1985. Peter Kendrick (PK) in November 1997, and held at ‘Pannawonica’ (P). Cave in north side of mesa the Western Australian Museum. Further samples 10 km ESE of Pannawonica, MR 394 035, 21˚40′S were collected from both northern (B2, T1) and 116˚25′E, A. Baynes and T.A. Smith, 4 August 1985. southern (T2) ‘caves’ and the cave fl oor surface (S) by A. Baynes, M.C. McDowell and T.A. Smith, 21–23 ‘Mt Bruce’ (MB). Floor of breakaway cave 6 km August 2004. SSE of Mt Bruce, on the former Marandoo iron ore ′ ′ prospect, 22˚40 S 118˚10 E, W.H. Butler, 26 April Fauna 1976. Western Australian Museum vertebrate palaeontological collection, catalogue numbers The mammal species identifi ed from the remains 76.6.21–76.6.25. from the various Pilbara sites are listed in Table 1. Most species are represented by well-preserved ‘Wittenoom’ (W). Hole in low cliff 2 km ESE of material, particularly in the large samples, giving ′ ′ Wittenoom townsite, MR 388 387, 22˚15 S 118˚20 E, rise to confi dent identifi cations. Dasyurus geoffroii, PIL3, A. Baynes and T.A. Smith, 26 July 1985. Field however, is recorded only from the Wittenoom site, number ABRS 107. in which the mammal remains consist of small Hamersley Range site 1 (HR1). Small cave in a broken bone fragments and isolated teeth. The D. valley in the northern face of the Hamersley Range, geoffroii specimen consists of only a small fragment 22˚18′26″S 118˚28′32″E, WGS 84, 517 m, T.A. Smith and of left dentary with the alveoli for M4, but this is A. Baynes, 15 August 2004. suffi cient for it to be confi dently distinguished from Hamersley Range site 2 (HR2). Very small cave Isoodon on shape and the smaller D. hallucatus on in a valley in the northern face of the Hamersley size. Two sites contain other interesting specimens Range, 22˚18′23″S 118o28′31″E, WGS 84, 467 m, M.C. that cannot be identifi ed to species. The material McDowell, A. Baynes and T.A. Smith, 15 August 2004. from the Wittenoom site also includes a lower Port Hedland (PH). Limestone ridge north of the permanent premolar cap of a small macropodid. 288 A. Baynes, M.C. McDowell

It is closest in form to that of Lagorchestes hirsutus, include a and medium to large rodent but the match is not exact and it has therefore not species now lost from the region. The variation been identifi ed beyond Macropodidae indet. The in representation of species, particularly small specimens recorded as Macropodidae indet. from dasyurids, between the subsamples from the the Herbert Well and Hamersley Range 1 sites Ripon Hills sinkhole suggests that even the large (Table 1) consist of fragments of long bones of large aggregate sample from that site may not be large of indeterminable species and represent enough to contain all the species in the original other taxa. The Port Hedland site material includes local fauna. a small dasyurid right dentary (Western Australian Museum vertebrate palaeontological collection DISCUSSION number 65.10.207) with the fi rst three lower molars in place; unfortunately these are very worn. The How representative are the fossil samples? jaw is similar in size to those of Sminthopsis ooldea and S. youngsoni, but the molars appear broader, Three factors bias samples of mammal remains and there are only alveoli for two premolars, in caves compared to the communities from which whereas species of Sminthopsis all have three. they are drawn: the sample size, the accumulating This specimen is therefore not identifi able beyond agent and the timing of accumulation. The indet. The Dasyuridae indet. from site nature of the statistical relationship between B2 in the Ripon Hills sinkhole (Table 1) consist sample size and number of species detected (e.g. of edentulous jaws of other taxa, not identifi able May 1975) dictates that no practical sample will beyond family level. contain a completely comprehensive local mammal fauna, even without the taphonomic fi lter of the The modern mammal fauna of the Pilbara (i.e. accumulating agent or agents. The most important that based upon live-caught ) is listed in accumulating agent in most caves in arid Australia Table 2, and compared with the fauna obtained is the Barn Owl, Tyto alba. Obviously, there is an from cave remains from the Pilbara, and the upper weight limit to the mammal species that it original mammal faunas of Cape Range peninsula can catch and eat, at about the size of an adult rat; and southern mainland Carnarvon Basin. There although it can prey upon juveniles of bandicoots are slight differences between the faunas listed in and rat-kangaroos (Higgins 1999). A further bias is Table 2 and those in their sources, including Gibson introduced by the nomadic behaviour of Barn Owls and McKenzie (2009), whose faunal list (their Table (Morton and Martin 1979). This is an adaptation to 1) includes information provided by our study. the population variations of native rodents. While Species that cannot be distinguised beyond generic the populations of insectivorous marsupials of the level as cave remains, such as Dasycercus spp., are arid zone remain at relatively constant, but low, listed as sp. indet. in Table 2. Lagorchestes hirsutus levels (e.g. Reid et al. 1993; Heywood and Pavey is listed in the fauna by Gibson and McKenzie 2002), those of the local rodents tend to fl uctuate (2009). This record is based upon a preliminary through several fold (e.g. Finlayson 1939; Newsome identifi cation of the premolar from the Wittenoom and Corbett 1975), with peaks following major cave site, that, as noted above, is now not considered rain events (e.g. How et al. 1991; Reid et al. 1993; to be specifi cally identifi able. Leporillus apicalis is Dickman et al. 1999; McDowell and Medlin 2009). listed in the fauna by Gibson and McKenzie (2009). Barn Owls move to take advantage of these booms The source of this record is unclear, as the northern in rodent numbers (Schodde and Mason 1980), limits of the species’ range lie in the Carnarvon and therefore contribute most bones to deposits and Gascoyne regions. Leporillus conditor was not when rodent numbers are locally high. Both the recorded from any of the Cape Range cave samples high relative abundance of the living rodents, and studied by Baynes and Jones (1993), but remains probably the owls’ search images for these species, of the species have subsequently been found in a bias the resulting accumulations of remains toward cave in foothills on the eastern side of the Range (A. disproportionately high rodent and low dasyurid Baynes, unpublished observation). Pseudomys fi eldi relative abundances (e.g. Morton and Martin 1979; has been identifi ed in remains from the Ripon Hills Baynes and Baird 1992; McDowell and Medlin sinkhole site since Gibson and McKenzie (2009) was 2009). This effect biases representation of species completed. among remains in cave deposits in favour of Comparison of mammal faunas from modern rodents. and cave records (Tables 1 and 2) shows two general patterns. Most bat species are poorly represented The original non-volant mammal fauna of the in the cave material, although remains of the cave- Pilbara roosting genera Taphozous and Macroderma were As noted above, the results (Table 1) suggest found; and there is a consistency between the that the original non-volant mammal fauna was deposits in the non-volant mammal faunas, which fairly constant throughout the Pilbara. For reasons Original mammal fauna 289 Nth Sth PK B2 T1 T2 S orded; (X) = record based on quills rather (X) = record orded; ...... X X .. .. X ...... X ...... X ...... X ...... X ...... X ...... X ...... X .. .. cf...... X X ...... X ...... cf...... cf...... X ...... cf...... X .. X ...... X .. X ...... X .. X .. .. (X).. (X) .. .. (X) ...... (X) X ...... X...... X X...... X....X.. X X.. ..XX ..X XX X...... X....X...... X .. .. X XX...... X .. .. XXXX X.. X .. .. X ..X X .. ..X X ...... X...... XX...... XX...... X .. ..X X ....XXXXX....XXX ...... X .. XX...... X..X...... XX....XXX .. X .. X X X X X .. X .. .. X X .. .. X .. X .. X X ...... X X ?? .. .. X .. X ...... X X ...... X ...... X X X X X ...... X X X X X .. X .. .. X X .. X X X .. X X .. .. X X .. X .. X .. X .. X .. .. X X .. .. X X .. .. X .. X X .. X X .. X X X X X X ...... X X X X X .. .. X .. X ...... X...... XX ..XX...... X ...... X X ...... X ...... X X .. X X ...... X X .. cf. X ...... f...... cf...... cf...... X ...... X X X X .. X X X .. sp. indet. X ...... X ...... X X .. P. ingrami than bone material; cf. = less certain record; ?? = possible record; .. = not recorded. ?? = possible record; than bone material; cf. = less certain record; eldi p ne...... X.. ..X...... indet.X...... XX.... sp. sp. indet...... X ...... X X .. X X .. sp. indet. X X ...... X .. .. X .. X ...... sp. cf. sp. given in ‘Results’. X = rec are Site abbreviations sites in the Pilbara region. from recorded are Mammal species whose remains arpddeidtX...X...... X..X...... indet.X Pseudantechinus roryi Pseudantechinus Sminthopsis macroura Dasyuridae indet. auratus Isoodon rothschildiPetrogale Macropodidae Taphozous .. gigas Macroderma .. Leggadina .. macrurus Mesembriomys .. .. delicatulus Pseudomys ..Pseudomys hermansburgensis .. X ...... X ...... Planigale maculata Planigale Sminthopsis longicaudata Sminthopsis youngsoni Trichosurus vulpecula Macropus robustus Vespertilionidae indet.Mormopterus beccarii Melomys burtoni ..Mus musculus Notomys alexis longicaudatus Notomys .. chapmani Pseudomys ..Pseudomys desertor Pseudomys fi ..Pseudomys nanus Rattus tunneyi XZyzomys argurus pedunculatus Zyzomys Muridae .. indet...... X ...... X X .. X .. .. X ...... Site HW UB P MB W HR1 HR2 PH OR IH CG SG YS EP-1 RHS Antechinomys laniger Antechinomys hallucatus Dasyurus Species aculeatus Tachyglossus Dasycercus rosamondae Dasykaluta geoffroii Dasyurus timealeyiNingaui calura Planigale Table 1 Table 290 A. Baynes, M.C. McDowell

Table 2 The original native mammal fauna of the Pilbara, identifi ed from remains from caves (data from this study) and modern native mammal fauna (data from W.A. Museum specimen records; Gibson and McKenzie 2009; McKenzie and Bullen 2009), compared with the total (original + modern) native mammal faunas recorded from the mainland southern Carnarvon Basin (i.e. excluding the Shark Bay peninsulas and islands) (data from Baynes 2000; McKenzie et al. 2000; McKenzie and Muir 2000) and Cape Range (data from Baynes and Jones 1993). X = recorded; ? = less certain record; .. = not recorded. * = locally extinct on the Pilbara main- land; ** = totally extinct. # discussed in Results section.

Species Carnarvon Basin Cape Range Original Pilbara Modern Pilbara

Tachyglossus aculeatus XX XX

Antechinomys laniger XX X.. Dasycercus sp. indet. XX XX Dasykaluta rosamondae ?X XX Dasyurus geoffroii* XX X.. Dasyurus hallucatus .. X X X Ningaui timealeyi .. X X X Phascogale calura* XX X.. Planigale sp. cf. P. ingrami .. .. X X Planigale maculata .. X X X Pseudantechinus roryi .. .. X X Pseudantechinus woolleyae ...... X Pseudantechinus sp. indet. .. X X .. Sminthopsis dolichura X...... Sminthopsis hirtipes X...... Sminthopsis longicaudata .. X X X Sminthopsis macroura XX XX Sminthopsis ooldea .. ? .. X Sminthopsis youngsoni XX XX

Chaeropus ecaudatus** X...... Isoodon auratus* XX XX bougainville* X X .. .. lagotis XX ..X

Trichosurus vulpecula .. X X X

Bettongia lesueur* X X .. .. Lagorchestes conspicillatus ...... X Macropus robustus XX XX Macropus rufus .. X .. X Petrogale lateralis .. X .. X Petrogale rothschildi .. .. X X

Pteropus alecto X....X Pteropus scapulatus XX ..X Saccolaimus fl aviventris X....X Taphozous georgianus XX ?X Species Carnarvon Basin Cape Range Original Pilbara Modern Pilbara Original mammal fauna 291

Table 2 (continued)

Taphozous hilli ...... X Macroderma gigas .. .. X X Rhinonicteris aurantius ...... X Chalinolobus gouldii XX ..X Chalinolobus morio ...... X Nyctophilus arnhemensis ...... X Nyctophilus bifax daedalus ...... X Nyctophilus geoffroyi geoffroyi X....X Nyctophilus geoffroyi pallescens ...... X Scotorepens balstoni X...... Scotorepens greyii X....X Vespadelus fi nlaysoni XX ..X Chaerephon jobensis X....X Mormopterus beccarii X..?X Mormopterus loriae ...... X Mormopterus planiceps X...... Tadarida australis XX ..X

Hydromys chrysogaster ...... X Leggadina lakedownensis ?? ?X Leporillus apicalis** X X .. ..# Leporillus conditor* XX#.... Melomys burtoni* .. .. X .. Mesembriomys macrurus* .. X X .. Notomys alexis XX XX Notomys amplus** X X .. .. Notomys longicaudatus** XX X.. Pseudomys albocinereus X...... Pseudomys chapmani .. X X X Pseudomys delicatulus .. .. X X Pseudomys desertor XX XX Pseudomys fi eldi* XX X#.. Pseudomys hermannsburgensis XX XX Pseudomys nanus* XX X.. Rattus tunneyi* XX X.. Zyzomys argurus .. .. X X Zyzomys pedunculatus* XX X..

Canis lupus .. X .. .. 292 A. Baynes, M.C. McDowell discussed in the section above, absence of a pedunculatus occurred throughout the Pilbara species from a particular sample is not necessarily ranges, something that was previously unknown. meaningful. But where samples are large, the Notomys longicaudatus, too, was very widespread remaining differences between the faunas are and is represented in many of the samples. mainly explicable in terms of the presence or Pseudomys fi eldi was not previously known to have absence of suitable habitat for the species around originally occurred in the Pilbara, and the Ripon the sites. Thus, for example, remains of Leggadina Hills sinkhole record represents a substantial are present in quite large numbers in the sample extension of range, east from the Cape Range and from the Herbert Well site because there are north from the Gibson Desert. extensive plains (its preferred habitat) around The most surprising record is that of Dasyurus the breakaway that contains that site, but are geoffroii from the Wittenoom site. It has been present in only very small numbers in one of thought (e.g. the distribution map in Serena and the samples from the Ripon Hills sinkhole site, Soderquist 1995) that the species did not occur in which is surrounded by rocky hills. Moreover, the Pilbara, and that only D. hallucatus was present. most of the Ripon Hills samples are larger than The actual situation seems to have been more that from Herbert Well. Relative abundances of complex. Although the record of D. geoffroii is based other rodent species similarly refl ect the quantity upon a very fragmentary specimen, it appears to of habitat around sites. For example, although be a reliable identifi cation. The Wittenoom site is the rock-rat Zyzomys pedunculatus is recorded in the northern foothills of the Hamersley Range, from Herbert Well, it is present in only very low overlooking the valley. There is numbers, whereas its remains are quite abundant likely to have been ecological separation between in the Ripon Hills sinkhole, and also in the caves of the two species of Dasyurus, probably primarily by Cape Range (Baynes and Jones 1993). This pattern habitat, with D. hallucatus occurring in the rocky is consistent with the results obtained by Gibson habitats of the ranges and D. geoffroii probably and McKenzie (2009), whose analyses showed that restricted to the valley and plain habitats in the substrate was the most important predictor of region. The specimen in the Wittenoom site is occurrence at a particular site for the small ground probably from an individual caught in the valley mammal species still extant in the Pilbara. and transferred into the site by its predator, most Among the most interesting records (Table 1) likely a Dingo on the basis of the highly fragmented are those of Mesembriomys macrurus from both nature of most of the mammal bones in the site. the Herbert Well site and Ripon Hills sinkhole, Occurrence of only D. hallucatus in the Ripon suggesting that the species originally had a much Hills site, both as remains and as a living wider distribution in the Pilbara than previously observed by M.C. McDowell in August 2004, is indicated by the one modern specimen from the consistent with this proposed pattern because that coast near Roebourne and fairly scarce remains site is entirely surrounded by rocky hills. Although in Cape Range cave deposits (Kendrick and the two Hamersley Range sites, from one of which Porter 1974; Baynes and Jones 1993). Although D. hallucatus is recorded (see Table 1), are in a an arboreal species, extant populations in the similar situation to the Wittenoom cave, they are north-west Kimberley have been recorded from a considerably higher and further into the Range, and wide variety of habitats, ranging in density from neither appears suitable as a Dingo lair. rainforest to woodlands over hummock grassland Melomys burtoni is probably recorded from just and rugged sandstone screes (McKenzie and Kerle the Port Hedland site because the species originally 1995). The animals spend quite a lot time on the had a very restricted distribution in the Pilbara, ground, easily crossing open vegetation formations. occurring only on the coast, possibly in or around Like many Australian rodents, M. macrurus is the mangrove formations. The final record of omnivorous, feeding on fl owers, fruits and insects particular interest is that of Phascogale calura in (particularly termites), and other vegetable foods the Herbert Well site deposit. The signifi cance of (McKenzie and Kerle 1995); it is even reported to this is more diffi cult to assess because its apparent have a liking for marine molluscs (Watts and Aslin absence from the Ripon Hills sinkhole could be due 1981). In the Pilbara, fi g trees (Ficus platypoda) may to the fact that the small dasyurid material in that have provided several habitat requirements of M. deposit is poorly preserved, generally fragmented macrurus, including fruit and arboreal nesting sites. and very diffi cult to identify. In this context it is The site in the breakaway range near Herbert Well unfortunate that no further material was obtained is situated in a deep gully containing several large during the survey from the EP-1 sinkhole, because fi g trees. Fig trees also occur in the Cape Range the bones from that site are beautifully preserved. (Keighery and Gibson 1993), sometimes marking One species that appears to be under-represented the entrances to caves. in the cave deposits is Trichosurus vulpecula, which The results (Table 1) also suggest that Zyzomys is recorded only from the Herbert Well site. Original mammal fauna 293

Trichosurus vulpecula often uses caves as shelters, ranges in the arid zone, are probably present and its remains are common in the Cape Range in the Pilbara and Cape Range but absent from caves (Baynes and Jones 1993). It is particularly the southern Carnarvon Basin because the latter surprising that no remains of it have been found lacks suitable ranges. Conversely, remains of the in the Ripon Hills sinkhole. It is possible that most sand habitat specialists Perameles bougainville and of the Pilbara caves are not deep or cool enough Notomys amplus are both very rare in Cape Range to provide shelter of adequate quality for this deposits (Baynes and Jones 1993) and not recorded species. Although the Ripon Hills sinkhole may from the Pilbara, even though both originally provide a suitable shelter for the species, it may occurred in the (Western be surrounded by such inhospitable terrain that Australian Museum record M1489 for Perameles; A. Trichosurus was not able to reach it. Baynes unpublished observation for N. amplus), as well as the southern Carnarvon Basin. Biogeography Finally, there is a group of mammal species In Table 2 the Pilbara mammal fauna is compared restricted to north-western Australia: Dasykaluta with those of the mainland southern Carnarvon rosamondae, Ningaui timealeyi, Petrogale rothschildi Basin and the Cape Range (in the northern and Pseudomys chapmani. Of these, only Petrogale Carnarvon Basin) to the west. Generally, the faunas rothschildi is confi ned to the Pilbara. When their are quite similar, but from south-west to north-east systematics are resolved, it is likely that the two there is a loss of southern species and addition of or species of Planigale recorded from the Pilbara will replacement by northern species. be added to this group. The original ranges of three mammal species widespread in the drier inland of south-western ACKNOWLEDGEMENTS Australia—Sminthopsis dolichura, Mormopterus planiceps and Pseudomys albocinereus—extended The 1985 fi eld work was fi nanced by a grant to as far north as the southern mainland Carnarvon A. Baynes from the Australian Biological Resources Basin, but no further. Sminthopsis hirtipes of the Study. We thank CALM/DEC (through the auspices southern arid zone sandy habitats drops out of Norm McKenzie) for funding for fi eld work in in favour of the more northerly distributed S. 2004, and for contracts to collect the specimens youngsoni. ecaudatus originally reached in 2004 and write them up in 2007. Tom Smith its northern range limit in the southern Carnarvon provided skilled assistance during the fi eld work. Basin, and both species of Leporillus originally We are very grateful to Tom Smith and Sara Peet reached their northern range limits on the Cape for sorting material from Herbert Well, Hamersley Range peninsula. Range and Ripon Hills. Brent Johnson is thanked for assistance with specimen preparation. Gavin Northern forms recorded from the Pilbara, and in Prideaux provided advice on identification of some cases also the Cape Range, include Dasyurus macropodoid remains. We thanks our referees Anne hallucatus, Lagorchestes conspicillatus, Rhinonicteris Kerle and Jim Mead, and editor Alex George, for aurantius, Nyctophilus arnhemensis, Nyctophilus bifax, constructive suggestions that improved the paper. Mormopterus loriae, Melomys burtoni, Mesembriomys macrurus, Pseudomys delicatulus and Zyzomys argurus. Of these, Mormopterus loriae and Melomys REFERENCES burtoni have or had coastal distributions because Archer, M. (1975). 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APPENDIX 1. EARLY PILBARA MAMMAL M3566 Tambrey (Station) 23 Jul 1958 RECORDS FROM THE WESTERN M4958 Millstream (Station) 1958 AUSTRALIAN MUSEUM M4959 Millstream (Station) 1958 M4960 Millstream (Station) 1958 Hamersley Expedition mammals M4961 Millstream (Station) 1958 Dasyurus hallucatus M4962 Millstream (Station) 1958 M4518 Hamersley Range area 1958 29 records

Planigale sp. Macropus rufus M3432 Tambrey 3 Aug 1958 (Archer 1976, p. 357) M3507 Tambrey Station 4 Aug 1958 M3508 Station Jul 1958 Pseudantechinus ‘macdonnellensis’ M3528 Marillana Station Jul 1958 M3430 Mount Herbert 27 Jul 1958 M4956 Kangiangi Station 21 Jul 1958 M3445 30 Jul 1958 M4957 1958 Macropus robustus 5 records

M3503 Tambrey Station 31 Jul 1958 Pteropus alecto M3504 Coolawanyah Station 26 Jul 1958 M5125 Millstream Station 29 Jul 1958 M3505 Millstream Station 19 Jul 1958 M5126 Millstream Station 22 Jul 1958 M3506 Tambrey Station 31 Jul 1958 M5127 Millstream Station 22 Jul 1958 M3509 Millstream Station 16 Jul 1958 M5128 Millstream Station 26 Jul 1958 M3513 Coolawanyah Station 26 Jul 1958 M5129 Millstream Station 27 Jul 1958 M3514 Tambrey Station 29 Jul 1958 M5130 Millstream Station 22 Jul 1958 M3515 [Coolawanyah Station] 26 Jul 1958 M5131 Millstream Station 22 Jul 1958 M3516 Kangiangi Station 24 Jul 1958 M5132 Millstream Station 27 Jul 1958 M3517 Millstream Station 19 Jul 1958 8 records M3518 Millstream 19 Jul 1958 M3519 Millstream 19 Jul 1958 Taphozous georgianus M3523 Millstream 1958 M3431 Millstream Station 22 Jul 1958 M3526 Coolawanyah Station 4 Aug 1958 M3448 Millstream Station 22 Jul 1958 M3527 Millstream Station 20 Jul 1958 M4749 Tambrey Station 30 Jul 1958 M3529 Coolawanyah Station 29 Jul 1958 M4750 Tambrey Station 30 Jul 1958 M3530 Millstream Station 20 Jul 1958 4 records M3559 Marillana Station Jul 1958 Taphozous hilli M3560 Marillana Station Jul 1958 M4747 Tambrey Station 31 Jul 1958 M3561 Marillana Station Jul 1958 M3562 Marillana Station Jul 1958 Macroderma gigas M3564 Marillana Station Jul 1958 M3415 Tambrey Station 1 Aug 1958 M3565 Marillana Station Jul 1958 M4413 Tambrey Station 4 Aug 1958 296 A. Baynes, M.C. McDowell

Chalinolobus gouldii M10646 Woodstock Station M6310 Tambrey Station 3 Aug 1958 M10647 Woodstock Station Vespadelus fi nlaysoni M10648 Woodstock Station M10649 Woodstock Station M4751 Tambrey Station 30 Jul 1958 M10650 Woodstock Station M4752 Tambrey Station 3 Aug 1958 28 records Tim Ealey and co-worker specimens Ningaui timealeyi Tachglossus aculeatus M5076 Abydos Station Nov 1962 M4932 Pullcunah Hill 24 Oct 1958 (R.M. Sadlier) Pseudantechinus ‘macdonnellensis’ Dasycercus sp. indet. M3659 Woodstock Station 25 Feb 1958 M4268 Abydos Station Apr 1958 M5279 Woodstock Station 6 Feb 1958 M5509 Woodstock Station 25 Jan 1958 Dasykaluta rosamondae M5510 Woodstock Station 10 Feb 1958 M3416 Abydos Station 16 Aug 1956 M5769 Woodstock Station 1959 M3421 Woodstock Station M6056 Woodstock Station 4 Apr 1958 M5507 Woodstock Station Oct 1957 6 records M5508 White Springs Station Jul 1954 M5512 Woodstock Station Jun 1956 Pseudantechinus roryi M5513 Woodstock Station 3 Apr 1957 M4291 Eginbah Station Aug 1959 M5514 Pullcunah Hill (near) 4 Apr 1958 M5280 Woodstock Station 6 Feb 1958 M6054 Woodstock Station Apr 1957 M5511 Woodstock Station 11 Feb 1958 M6055 Abydos Station 8 Nov 1956 Macrotis lagotis 9 records M4196 Woodstock Station pick up, donated Sep 1960 Dasyurus hallucatus Lagorchestes conspicillatus M3469 Woodstock Station M3496 accessed 1959 M3501 Woodstock Station M3500 Woodstock Station accessed 1959 M3502 Woodstock Station M3571 Woodstock Station Macropus robustus M3651 Woodstock Station M4696 Woodstock Station 6 Nov 1959 M3652 Woodstock Station M4697 Woodstock Station Nov 1959 M3653 Woodstock Station M5840 Woodstock Station 2 Aug 1954 M3654 Woodstock Station M6095 Mount Edgar Station 1958 M3655 Woodstock Station M6096 Mount Edgar Station 1958 M3656 Woodstock Station 28 Sep 1960 M6097 Mount Edgar Station 1958 M3657 Woodstock Station M6098 Mount Edgar Station 1958 M3989 Woodstock Station 1958 M6099 Mount Edgar Station 1958 M3993 Woodstock Station 23 Apr 1958 M6100 Mount Edgar Station 1958 M6140 Woodstock Station 1958 M6101 Mount Edgar Station 1958 M6141 Woodstock Station 10 Apr 1958 M6102 Mount Edgar Station 1958 M6142 Woodstock or Mount Edgar Station M6103 Mount Edgar Station 1958 M10529 Woodstock Station M6104 Mount Edgar Station 1958 M10607 Woodstock Station M6105 Mount Edgar Station 1958 M10608 Woodstock Station M6106 Mount Edgar Station 1958 M10609 Woodstock Station M6107 Mount Edgar Station 1958 M10610 Woodstock Station M6108 Mount Edgar Station 1958 M10611 Woodstock Station M6109 Mount Edgar Station 1958 M10645 Woodstock Station M6110 Mount Edgar Station 1958 Original mammal fauna 297

M6111 Mount Edgar Station 1958 M4287 Woodstock Station 4 Dec 1957 M6112 Mount Edgar Station 1958 M4440 Woodstock Station M6113 Mount Edgar Station 1958 M4443 Woodstock Station Feb 1958 M6114 Mount Edgar Station 1958 M6220 Woodstock Station 5 Oct 1957 M6115 Mount Edgar Station 1958 11 records M6116 Mount Edgar Station 1958 Pseudomys chapmani M6117 Mount Edgar Station 1958 M5767 Mount Edgar Station 8 Oct 1957 M6118 Mount Edgar Station 1958 M5864 White Springs Station 18 Sep 1956 M6119 Mount Edgar Station 1958 M5865 White Springs Station 18 Sep 1956 M6120 Mount Edgar Station 1958 M5866 White Springs Station 18 Sep 1956 M6121 Mount Edgar Station 1958 M5867 White Springs Station 18 Sep 1956 M6122 Mount Edgar Station 1958 M5868 White Springs Station 18 Sep 1956 M6123 Mount Edgar Station 1958 M5870 White Springs Station 18 Sep 1956 M6124 Mount Edgar Station 1958 7 records M6125 Mount Edgar Station 1958 M6126 Mount Edgar Station 1958 Pseudomys hermannsburgensis M6127 Mount Edgar Station 1958 M3452 Woodstock Station accessed 1959 M6128 Mount Edgar Station 1958 Zyzomys argurus M6129 Mount Edgar Station 1958 M3296 Woodstock Station Dec 1957 M6130 Mount Edgar Station 1958 M5733 Woodstock Station 24 Jan 1958 M6131 Mount Edgar Station 1958 M5739 Woodstock Station 12 Dec 1957 M6132 Mount Edgar Station 1958 M5740 Woodstock Station 16 Dec 1957 M6133 Mount Edgar Station 1958 M5741 Woodstock Station 28 Jan 1958 M6134 Mount Edgar Station 1958 M5749 Woodstock Station 1959 M6135 Mount Edgar Station 1958 M5750 Woodstock Station 1959 M6136 Mount Edgar Station 1958 M5751 Woodstock Station 1959 M6137 Mount Edgar Station 1958 M5752 Woodstock Station 1959 M6138 Mount Edgar Station 1958 M5753 Woodstock Station 1959 M6139 Mount Edgar Station 1958 M5754 Woodstock Station 1959 M6171 Mount Edgar Station 1958 M5755 Woodstock Station 1959 M6172 Mount Edgar Station 1958 M5756 Woodstock Station 1959 M6173/1 Mount Edgar Station 3 Mar 1960 M5757 Woodstock Station 1959 M6173/2 Mount Edgar Station 3 Mar 1960 M5758 Woodstock Station 1959 M6246 Woodstock or Mount Edgar Station 1958 M5759 Woodstock Station 16 Dec 1957 53 records M5760 Woodstock Station 1957 Macropus rufus M5761 Woodstock Station 1957 M3510 Woodstock Station 2 Oct 1955 M5762 Woodstock Station 21 Jan 1958 M3511 accessed 1959 M5763 Woodstock Station 6 Feb 1958 M6170 Mount Edgar or Woodstock Station 1 May 1955 M5764 Woodstock Station 17 Dec 1957 21 records Petrogale rothschildi M3324 Woodstock Station Other records M3325 Woodstock Station Sminthopsis youngsoni M3499 Woodstock Station M32738 Woodstock Station 2 Mar 1990 M3650 Woodstock Station M3881 Pullcunah Hill 18 Sep 1957 Trichosurus vulpecula M4259 Pullcunah Hill 21 Nov 1959 M2767 Red Hill Station 22o08’S 116o04’E 1948 M4274 Pullcunah Hill 3 Dec 1957 M19550 Bonney Downs 22o10’S 119o56’E 1981