taxonID	type	description	language	source
03A087E6FFE38328D788FD6CD759FB43.taxon	materials_examined	Type species. Didelphis penicillata Shaw, 1800 (in the combination Dasyurus penicillata) [= Phascogale tapoatafa (F. Meyer, 1793)] by subsequent monotypy. Specific content. P. tapoatafa (Meyer, 1793), P. calura Gould, 1844, and P. pirata Thomas, 1904. Generic rediagnosis. Distinguished from all other genera of Phascogalinae by distal portion of tail with discrete brush of long black hairs; inner metatarsal tubercle united with first interdigital pad; outer metatarsal tubercle elongate; pinna with well-developed posterior notch and curled supratragus; bold facial patterning comprised of dark mid-rostral stripe, coronal patch and loreal stripe, and pale preauricular tuft, orbital crescents, cheek patch and supralabial bar; fur on upper chest and throat pure white to hair bases; ventral fur colour paler than dorsum and flanks; enlarged auditory bullae; well-developed maxillonasolabialis fossa extending onto anterior root of zygomatic arch; I 1 enlarged and procumbent; I 2 moderately to greatly enlarged; upper and lower canines enlarged.	en	Have, Ten (2015): Taxonomic revision of Phascogale tapoatafa (Meyer, 1793) (Dasyuridae; Marsupialia), including descriptions of two new subspecies and confirmation of P. pirata Thomas, 1904 as a ‘ Top End’ endemic. Zootaxa 4055 (1): 1-73, DOI: 10.11646/zootaxa.4055.1.1
03A087E6FFE38328D788FD6CD759FB43.taxon	discussion	Remarks. Iredale & Troughton (1934) were the first to recognise the combination Phascogale tapoatafa (Meyer, 1793).	en	Have, Ten (2015): Taxonomic revision of Phascogale tapoatafa (Meyer, 1793) (Dasyuridae; Marsupialia), including descriptions of two new subspecies and confirmation of P. pirata Thomas, 1904 as a ‘ Top End’ endemic. Zootaxa 4055 (1): 1-73, DOI: 10.11646/zootaxa.4055.1.1
03A087E6FFE3831CD788F8D1D4DDFE2C.taxon	materials_examined	Holotype: As explained by Mahoney & Ride (1988: 25), “ the holotype is the specimen described and figured under the name Tapoa Tafa by Anon. [Hunter, J.] in White, J. (1790). Journal of a Voyage to New South Wales with sixty-five plates of non descript animals, birds, lizards, serpents, curious cones of trees and other natural productions. London: J. Debrett xvii 335 pp. (281 – 284 pl 58); preparation 3758 of Owen, R. (1841). Descriptive and Illustrated Catalogue of the Physiological Series of Comparative Anatomy Contained in the Museum of the Royal College of Surgeons in London. Products of generation. London: R. & J. E. Taylor Vol. 5 iv + xxxvii 284 pp. (209) is part of the holotype specimen) ”. Although Troughton (e. g. 1967: 31) notes that “ the specimen figured was preserved in the Museum of the Royal College of Surgeons ”, neither Thomas (1888) nor Mahoney & Ride were able to locate an appropriate candidate, either in this collection or in the Natural History Museum, London. However, the original plate in White (1790) is unambiguously diagnostic (provided locality data are also taken into account) and neotype designation is unwarranted. Geographic Region HV (mm) TV (mm) Wgt (g) Geographic Region HV (mm) TV (mm) Wgt (g) Southwest Western Australia Mean 179.95 199 190 s. d. 19.61 16.16 n 11 11 1 Max 210 230 Min 153 182 Victoria Mean 193.25 206.25 155.75 s. d. 20.30 12.34 41.44 n 4 4 4 Max 213 222 187 Min 165 195 96 New South Wales / Southern Queensland Mean 186 189.5 183 s. d. 15.56 23.33 n 2 2 1 Max 197 206 Min 175 173 ' Top End', Northern Territory Mean 174.5 185.5 125.75 s. d. 11.90 5.20 17.67 n 4 4 4 Max 188 192 150 Min 160 180 108 Kimberley, Western Australia Mean 145 144 s. d. n 1 1 Max Min Revised description. This account is primarily based on specimens from the central to north coastal region of New South Wales, including one topotypical specimen (AM 932) collected in 1894 at Kingswood near Sydney. An adult male specimen from Coraki in northern New South Wales (CM 20990) is illustrated in Fig. 4. Regional variation is noted after the main account. External measurements taken from museum specimen records are summarised in Table 4. Adults in a Victorian population attained body weights of 311 g for males and 212 g for females (Soderquist 1995 c). The dorsal fur is relatively dense and lush, the contour hairs measuring 12 – 13 mm in the centre of the back. The overall colour of the back and flanks is grey with a silvery frosting, this effect imparted by narrow white tipping on grey-based contour hairs. Abundant black-tipped guard hairs are distributed evenly over the body and project as much as 9 mm through the fur. The mid-dorsal zone is distinctly darker than the flanks owing to a more intense basal pigmentation. Ventral fur colouration varies from grey with an extensive cream wash to predominantly cream (e. g., CM 3740). In most specimens, the central strip at least consists of pure cream fur, merging laterally into grey-based fur with a progressively shorter cream tipping. In all specimens, the pale ventral colouration extends anteriorly to the chin, posteriorly to surround the cloaca, and laterally to clothe the inner surface of the fore- and hind-limbs. The scrotum is heavily pigmented and densely clothed in cream fur. A pigmented gular gland is evident on some adult specimens of both sexes but the location of this gland is not marked by any obvious external discolouration of the fur. In some specimens the fur in this region has a slight russet tinge basally. The ventral fur in juveniles tends to be duskier, with a diffuse pale cream wash. Table 5 a: Males Geographic Region SL ZB OBW BS FMW C 1 - M 4 M 1 - 4 SouthwestW. A. Mean 46.34 27.49 18.61 5.67 6.80 17.60 10.28 s. d. 1.10 1.01 0.43 0.42 0.18 0.51 0.42 n 10 10 10 10 9 10 10 Range 44.57 – 47.79 26.17 – 28.81 17.97 – 19.09 4.94 – 6.32 6.40 – 7.02 16.89 – 18.69 9.70 – 11.23 SouthAustralia Mean 50.47 29.98 18.92 6.13 6.85 18.83 10.94 s. d. 0.86 0.66 0.58 0.87 0.31 0.67 0.54 n 3 3 3 3 3 3 3 Range 49.56 – 51.28 29.42 – 30.70 18.25 – 19.30 5.42 – 7.10 6.50 – 7.05 18.23 – 19.55 10.54 – 11.56 Victoria Mean 47.29 27.75 18.76 6.01 6.86 18.33 10.71 s. d. 2.09 2.12 0.46 0.63 0.53 0.55 0.25 n 8 7.00 7 7 7 8 8 Range 45.24 – 51.39 24.73 – 30.61 18.20 – 19.45 5.22 – 7.00 6.43 – 7.98 17.77 – 19.49 10.29 – 11.17. S. W. / Mean 47.55 28.45 18.95 5.76 6.41 17.78 10.34 Queensland s. d. 2.15 1.65 0.86 0.39 0.20 0.44 0.23 n 15 17 18 18 17 17 18 Range 42.60 – 51.73 24.02 – 30.00 17.31 – 19.95 4.95 – 6.35 6.04 – 6.74 16.78 – 18.59 9.91 – 10.70 NorthQueensland Mean 47.50 29.33 18.61 6.51 6.39 17.25 9.82 s. d. n 1 1 1 1 1 1 1 Range TopEnd', N. T. Mean 46.55 27.37 17.21 5.31 5.84 17.13 9.46 s. d. 0.98 1.84 0.25 0.38 0.04 0.31 0.31 n 3 4 4 4 3 4 5 Range 45.91 – 47.67 25.58 – 29.94 16.97 – 17.50 5.02 – 5.87 5.81 – 5.89 16.83 – 17.55 9.07 – 9.92 Kimberley, W. A. Mean 44.16 25.28 17.75 5.01 6.01 16.79 9.91 s. d. 0.85 1.35 1.09 0.33 0.24 0.15 0.26 n 2 2 2 2 2 2 3 Range 43.56 – 44.76 24.32 – 26.23 16.98 – 18.52 4.77 – 5.24 5.84 – 6.18 16.68 – 16.89 9.74 – 10.21 TOTAL 42 44 45 45 42 45 48 …… continued on the next page TABLE 5 a. (Continued) TABLE 5 b. Females Range 42.99 – 46.84 24.98 – 28.17 18.23 – 18.78 4.92 – 5.83 6.57 – 6.92 16.86 – 17.79 9.99 – 10.72 SouthAustralia Mean 47.33 26.96 19.00 5.37 6.95 18.66 11.01 s. d. 1.14 1.22 0.76 0.22 0.08 N 1 2 2 2 1 2 2 Range 26.15 – 27.76 18.14 – 19.86 4.83 – 5.90 18.5 – 18.81 10.95 – 11.07 Victoria Mean 46.64 27.20 18.42 5.70 6.65 18.10 10.69 s. d. 1.16 1.19 0.74 0.48 0.39 0.34 0.35 N 6 6 6 6 6 6 7 Range 45.06 – 47.75 25.14 – 18.11 17.51 – 19.42 5.22 – 6.37 6.06 – 7.23 17.61 – 18.52 10.19 – 11.07. S. W. / Sth Mean 44.29 26.24 18.52 5.35 6.51 16.75 10.24 Queensland s. d. 1.07 0.45 0.32 0.44 0.22 1.41 0.30 N 8 9 9 9 9 9 9 Range 42.05 – 45.5 25.63 – 27.10 18.0 – 18.97 4.90 – 6.32 6.10 – 6.80 16.39 – 18.20 9.79 – 10.70 TopEnd', N. T. Mean 41.29 23.80 16.14 4.80 5.98 16.08 9.28 s. d. 1.10 0.85 0.71 0.40 0.26 0.34 0.17 n 6 6 6 6 6 6 6 Range 40.09 – 42.66 22.43 – 24.67 15.05 – 16.98 4.28 – 5.24 5.68 – 6.46 15.49 – 16.38 9.04 – 9.50 Kimberley, W. A. Mean 41.51 16.66 5.42 6.70 16.58 10.19 s. d. n 1 1 1 1 1 1 Range TOTAL 31 32 33 33 31 33 35 ……. continued on the next page TABLE 5 b. (Continued) Range 13.78 – 15.46 7.69 – 8.90 16.12 – 18.81 7.94 – 8.94 4.75 – 6.03 2.80 – 3.10 1.76 – 2.02 SouthAustralia Mean 15.39 8.39 17.85 8.32 5.25 s. d. 0.40 0.09 0.19 0.55 N 1 2 2 2 2 Range 8.11 – 8.67 17.70 – 17.91 8.18 – 8.45 4.86 – 5.64 Victoria Mean 15.16 8.81 18.00 8.61 5.41 2.72 1.84 s. d. 0.66 0.55 0.62 0.17 0.11 0.11 0.03 N 6 6 6 6 7 2 2 Range 14.30 – 15.81 7.79 – 9.30 16.82 – 18.63 8.44 – 8.80 4.93 – 5.84 2.62 – 2.80 1.82 – 1.86 S. W. / Sth Mean 14.53 8.34 16.91 8.07 5.37 2.64 1.92 Queensland s. d. 0.28 0.18 0.36 0.44 0.37 0.13 0.05 N 9 9 9 9 6 9 9 Range 13.95 –– 14.91 8.03 – 8.54 16.31 – 17.50 7.42 – 9.00 4.81 – 5.78 2.50 – 2.86 1.82 – 2.00 TopEnd', N. T. Mean 12.64 7.38 14.57 6.82 5.18 2.23 1.54 s. d. 0.43 0.54 0.50 0.17 0.32 0.12 0.05 n 6 6 6 6 6 4 4 Range 11.82 – 12.97 6.71 – 8.08 16.31 – 17.50 6.61 – 7.05 4.68 – 5.60 2.12 – 2.40 1.50 – 1.58 Kimberley, W. A. Mean 13.10 7.60 7.08 6.06 2.40 1.60 s. d. n 1 1 1 1 1 1 Range TOTAL 32 33 32 33 32 21 21 Geographic region SL ZW OBW BS FMW C 1 - M 4 M 1 - 4 Southwest W. A. F 2 8.42 5.24 NS NS NS NS NS d. f. 1 1 P 0.010 0.035 South Australia / Victoria F 2 NS NS NS NS NS NS NS d. f. P. S. W. / Southeast Queensland F 2 15.99 15.23 NS 6.13 NS 7.93 NS d. f. 1 1 1 1 P 0.001 0.001 0.020 0.010 Top End', Northern Territory F 2 48.79 NS 8.08 NS NS 24.17 NS d. f. 1 1 1 P 0.000 0.022 0.001 Continued. Geographic region PALW ARW PRW IOW RD LCL LCW Southwest W. A. F 2 NS NS 6.36 NS 7.78 NS NS d. f. 1 1 P 0.022 0.012 South Australia / Victoria F 2 NS NS NS NS NS 12.01 10.73 d. f. 1 1.00 P 0.041 0.047. S. W. / Southeast Queensland F 2 17.25 7.34 12.09 NS NS 68.43 33.19 d. f. 1 1 1 1 1 P 0.000 0.013 0.002 0.000 0.000 Top End', Northern Territory F 2 NS 15.74 10.69 NS NS 25.49 31.11 d. f. 1 1 1 1 P 0.004 0.011 0.004 0.003 The fur on the head is pale grey on the sides of the face and dark grey with diffuse frosting on the crown. Additional facial patterning includes a well-developed, dark russet mid-rostral stripe, a broad and indistinct, dark grey loreal stripe and small, black postorbital spot, distinct off-white infra- and supra-orbital crescents, and a variably expressed cream-tipped preauricular tuft. Preauricular patches are indistinct in around 30 % of specimens and seem to be less conspicuous in juveniles than in adults. In most specimens, the cream fur of the chin and throat extends around the corner of the mouth and merges into a supralabial bar, while the cheek patch is cream peppered with black. The rostral vibrissal pads are densely clothed in short grey fur and the basal papillae are unpigmented and not visible. All facial vibrissae are thick and black, sometimes with gingery tipping. The pinnae are moderately elongate and broad, weakly to moderately pigmented and sparsely furred. The anterior border of each pinna supports a narrow band of short black hairs. The basal portion of the tail is thickly furred above and on the sides, and supports numerous projecting guard hairs, continuing the colour and texture of the rump (Figs 4 and 6). The tail narrows rapidly to form a distinct ‘ stem’, which is clothed above and below by long, adpressed hairs. Dorsally and around the sides of the tail, these hairs are white with short black tips, while those on the ventral surface are black with short cream tips grading to dark russet at the base of the brush. The tail ‘ brush’ is lush and measures up to 165 mm; it consists of jet black hairs except for a short section of dark russet hairs ventrally, at the base of the brush. The basal portion of the tail, from cloaca to base of brush, measures to 90 mm. The upper surface of the pes supports a mixture of white, dark grey and russet hairs, giving it an overall grizzled appearance. This merges into white fur on the sides of the foot and around the heel. The digits are densely clothed in white hairs. The plantar surface of the pes is granular and supports a total of five conspicuously striated pads. The inner metatarsal and first interdigital pads are fused, usually with no obvious line of demarcation; their combined length is 13.1 mm. The outer metatarsal pad is moderately long; on AM 932 it measures 7.4 mm and is separated by a gap of 3.9 mm from the fourth interdigital pad. The second to fourth interdigital pads are elongate and of subequal length. Both feet on one specimen (CM 1316) show a clear line of demarcation between the conjoined inner metatarsal and first interdigital pads. Females in all populations usually have eight teats in the pouch (e. g. Cuttle 1982; Soderquist 1993 a); occasional individuals have seven teats. Cranial and mandibular anatomy of a fully adult male (CM 24425) and subadult male (CM 1316) are illustrated in Figure 16. Cranio-mandibular and dental measurements for each sex are provided in Table 5. The New South Wales to southeast Queensland populations of P. t. tapoatafa show clear male-biased sexual dimorphism in cranio-dental measurements (Table 6 & Fig. 9). For all dimensions bar one (FMW), mean values are higher in males than females and for nine dimensions the contrast is statistically significant. This latter subset includes measures of overall cranial size (SL, ZW, PALW, PRW) and of the lower canine (C 1 L; C 1 W), as well as measures that are sensitive to canine enlargement (C 1 - M 4 L, ARW). The combined South Australian and Victorian sample shows a less pronounced male-biased dimorphism in which statistical significance is attained only for measures of the lower canine. In this population, the contrast in PRW also approaches significance. These observations on the Victorian population are consistent with Soderquist’s (1995 c) finding that the marked sexual dimorphism in body weight among Victorian P. tapoatafa is due primarily to somatic energy stores that are laid down in preparation for the short breeding season when foraging effort is curtailed. In the Victorian population studied by Soderquist the ratio of average adult male to female body weights is 1.62 (Soderquist unpublished data, reported by Rhind & Bradley 2002). Geographic variation. Possible clinal variation within P. t. tapoatafa is observed in various cranial and dental dimensions but with somewhat different expression in each sex. Among males there is a very slight and gradual decrease in cranial length (Fig. 17 a) from South Australia to Queensland, coupled with a more pronounced shortening of the upper molar row that also shows a more abrupt break between southern and northeastern populations (Fig. 17 b). Females show a more pronounced decrease in both cranial and dental dimensions (Figs 17 d – e). Both sexes show an abrupt and significant contrast between southern and northeastern populations in the breadth of the foramen magnum (Fig 17 c, f); this variable is little affected by overall growth of the cranium. The different pattern of geographic variation between the sexes of P. t. tapoatafa is largely a product of the variable expression of sexual dimorphism between the southern and southeastern Australian populations. Essentially, males of P. t. tapoatafa vary only slightly in cranial size throughout their range, while the more pronounced clinal variation in female cranial size, from large in the south to smaller in the north, creates a widening contrast between the two sexes. Overlaid on this pattern is a more pronounced geographic variation in molar size, with southern animals of both sexes having larger teeth on average than those from New South Wales and Queensland. On present evidence, this variation appears to be manifested in different ways between the sexes — step-like in males vs clinal in females — but larger samples from critical areas are needed to be certain of this.	en	Have, Ten (2015): Taxonomic revision of Phascogale tapoatafa (Meyer, 1793) (Dasyuridae; Marsupialia), including descriptions of two new subspecies and confirmation of P. pirata Thomas, 1904 as a ‘ Top End’ endemic. Zootaxa 4055 (1): 1-73, DOI: 10.11646/zootaxa.4055.1.1
03A087E6FFE3831CD788F8D1D4DDFE2C.taxon	distribution	Distribution and conservation status. The historical distribution of P. t tapoatafa corresponded to the more or less continuous tract of forested country that extended from the southeastern corner of South Australia, extending across most of Victoria (probably excluding only the northwestern mallee region and country above the treeline) and then running north along the coastal strip and mountainous hinterland of New South Wales and Queensland, north at least to Mackay (Fig. 18). A recently collected sub-fossil specimen from Ukerbarley station, 7 km NW of Coonabarabran (Fig. 18 & Table 9), indicates that populations of this taxon once occurred on the western slopes of the Great Dividing Range in New South Wales. Whether any of these survived into historical times is not known. At the western end of its range, a population of P. t. tapoatafa was recorded historically in the Mt Lofty Ranges of South Australia (Fig. 18). Although Wood Jones (1923) understood the range of this species (as P. penicillata) to extend to the Murray River, sub-fossil collections from several archaeological sites in this area have failed to confirm its former presence (Wakefield 1964). Morphologically, the few available specimens from the Mt Lofty population appear to be indistinguishable from Victorian specimens. However, in the absence of genetic data, it is not possible to decide whether the Mt Lofty population was genuinely isolated or not. Caution in this regard is further indicated by a subfossil record of P. tapoatafa from a cave deposit of mid-Holocene age on Kangaroo Island (Fig. 18 & Table 9), indicating a wider distributional and habitat range than might be inferred from the few historical records alone. As reported in a later section, populations of Phascogale tapoatafa are also present in northern Queensland. These resemble the nominate race in most morphological features but show a moderate level of genetic differentiation (albeit based on a single individual from the northern population). Today these populations appear to be separated from those in the south by a gap of more than 400 km of significantly drier habitat. However, a subfossil record of P. tapoatafa from Christmas Creek, inland of Townsville (Table 9) signals the possibility of recent continuity between these populations. The northern Queensland population is herein left unallocated at subspecific level. Although Victoria, New South Wales and Queensland still host populations of P. t. tapoatafa, its geographic range is now extremely fragmented and many regional populations are in decline. Indeed, the only region in which the taxon might be said to be secure is southeastern Queensland. Phascogale tapoatafa is probably extinct in South Australia, with the last vouchered record dating to 1969. Wood Jones (1923: 101 – 102) remarked on the former abundance of the species in South Australia and noted that it was known to “ the South Australian Murray River natives’ as “ pundi ”. He also speculated on the possible existence of populations of this taxon “ over a wide area in the Centre ”. It is possible that these reports referred instead to P. calura, which Wood Jones tentatively included in the South Australian fauna on the basis of an early record from ‘ Adelaide’ but otherwise knew little about. In fact, Spencer and Gillen had obtained several specimens of P. c a l u r a from Central Australia prior to 1901 (Baynes & Johnson 1996), although no further specimens were obtained from this area.	en	Have, Ten (2015): Taxonomic revision of Phascogale tapoatafa (Meyer, 1793) (Dasyuridae; Marsupialia), including descriptions of two new subspecies and confirmation of P. pirata Thomas, 1904 as a ‘ Top End’ endemic. Zootaxa 4055 (1): 1-73, DOI: 10.11646/zootaxa.4055.1.1
03A087E6FFD7831AD788FE22D328F872.taxon	materials_examined	Holotype: WAM 7674. Adult male. Quindalup, Western Australia, 33 0 40 S’ 115 0 00 ’ E. Collected by Mr R. C. Hislop, 17 th March 1967. Puppet skin and skull. Paratype: WAM 5006. Adult female. Balingup Brook, Western Australia, 33 0 41 ’ S 116 0 05 ’ E. Collected by Ms J. Kyd, 17 th March 1962.	en	Have, Ten (2015): Taxonomic revision of Phascogale tapoatafa (Meyer, 1793) (Dasyuridae; Marsupialia), including descriptions of two new subspecies and confirmation of P. pirata Thomas, 1904 as a ‘ Top End’ endemic. Zootaxa 4055 (1): 1-73, DOI: 10.11646/zootaxa.4055.1.1
03A087E6FFD7831AD788FE22D328F872.taxon	etymology	Etymology. The word ‘ wambenger’ is in common local use in southwestern Western Australia for the Brushtailed Phascogale. The name is listed by Dixon (1990; cited by Bindon & Chadwick 2002) as an ‘ Australian Aboriginal Word in English’, and identified as ‘ possibly Nyungar’ in origin. Bindon & Chadwick (2002) gave the transliteration wam-bing-ga, following the usage of Isaacs (n. d.) in a work evidently compiled prior to 1949. Isaacs’ source is not specified and there appear to be no earlier published records of the word. Bindon & Chadwick (2002) report two other Nyoongar words for ' squirrel' — kumining and comingcoming — either of which might be applicable to the Phascogale which is the most ‘ squirrel-like’ of the native mammals of the region.	en	Have, Ten (2015): Taxonomic revision of Phascogale tapoatafa (Meyer, 1793) (Dasyuridae; Marsupialia), including descriptions of two new subspecies and confirmation of P. pirata Thomas, 1904 as a ‘ Top End’ endemic. Zootaxa 4055 (1): 1-73, DOI: 10.11646/zootaxa.4055.1.1
03A087E6FFD7831AD788FE22D328F872.taxon	diagnosis	Diagnosis. A moderately large-bodied race of P. tapoatafa that differs from the typical race of southeastern Australia in the following ways: externally, with a narrower zone of cream-tipped fur on venter, cream preauricular patch rarely developed, generally longer tail brush, slightly larger pinnae, less densely furred muzzle, narrower zone of contrasting pale fur on inside of fore- and hind-limbs, variable number (6 – 8) and commonly fewer teats; cranially, very similar to nominotypical P. tapoatafa but with less prominent sexual dimorphism in canine size (reflecting relative canine reduction in males and enlargement in females) and corresponding relatively narrower rostrum (measured across canines); genetically, with 3.7 – 6.7 % sequence divergence for cyt b from other populations. Differs from P. t. kimberleyensis in the following ways: externally, with lusher fur and generally darker dorsal colouration, grey-based ventral fur, much narrower zone of contrasting pale fur on inside of fore- and hind-limbs, preauricular patch rarely developed, relatively longer tail with more extensive furring of the base and longer ‘ brush’, much larger pinnae, inner metatarsal pad smoothly united with first interdigital pad; cranially, larger in all dimensions but with proportionally broader foramen magnum, relatively shallower rostrum and proportionally smaller molars; genetically, with 5.7 – 7.8 % sequence divergence for cyt b from other populations. Differences from P. p i r at a are listed under the account of that taxon (see below).	en	Have, Ten (2015): Taxonomic revision of Phascogale tapoatafa (Meyer, 1793) (Dasyuridae; Marsupialia), including descriptions of two new subspecies and confirmation of P. pirata Thomas, 1904 as a ‘ Top End’ endemic. Zootaxa 4055 (1): 1-73, DOI: 10.11646/zootaxa.4055.1.1
03A087E6FFD7831AD788FE22D328F872.taxon	description	Description of holotype. External measurements, as taken from label: HB 200, TL 221, Pes 29, Ear 29. Pes measured on puppet skin c. 42 mm. The dorsal fur is relatively dense and lush, the contour hairs measuring 11 – 12 mm in the centre of the back. The overall colour of the back and flanks is grey with a silvery frosting, this effect imparted by a combination of narrow white tipping on grey-based contour hairs. Abundant black-tipped guard hairs are distributed evenly over the body and project as much as 7 mm through the fur. The mid-dorsal zone is distinctly darker than the flanks owing to a more intense basal pigmentation. The abdomen has a narrow, off-white central zone made up of pale cream hairs with short grey bases. This zone expands anteriorly to cover the pectoral region, and posteriorly, to surround the scrotum. The boundary between flank and ventral colouration is diffuse. The fore- and hind-limbs are grey externally but with a narrow zone of contrasting off-white fur extending along the inner surface. The scrotum is pigmented and thinly clothed in off-white fur. A narrow, elongate gular gland is evident as a zone of pigmented skin measuring 4 mm by 22 mm of the lower throat. The location of this gland is not marked by any obvious external discolouration of the fur, but hairs in this region have a russet tinge basally. The fur on the head is essentially pale grey on the sides of the face and dark grey with diffuse frosting on the crown. Additional facial patterning includes a well-developed, dark russet mid-rostral stripe, a broad and indistinct, dark grey loreal stripe and small, black postorbital spot, and distinct off-white orbital crescents. The vibrissal pads are thinly clothed in short grey fur. The mystacial vibrissal papillae are darkly pigmented and all facial vibrissae are thick and black to the tips. The pinnae are extremely large, weakly pigmented and sparsely furred internally except along their anterior and posterior borders, which carry a narrow strip of short black hairs. The external surface of the pinna is clothed in short dark hairs. The basal portion of the tail is thickly furred on all sides, and supports numerous projecting guard hairs, continuing the colour and texture of the rump (Fig. 6). This narrows gradually and gives way to a short tail ‘ stem’ that is clothed above and below by long, adpressed hairs; these are white with short black tips on the upper surface and sides of the stem, but are rich russet on the under-surface, blending forward into the base of the brush. The tail ‘ brush’ is lush and measures up to 170 mm; it consists of jet black hairs. The basal portion of the tail (cloaca to the base of the brush) measures 85 mm. The upper surface of the pes supports a mixture of white, dark grey and russet hairs, giving it an overall grizzled appearance. This merges into white fur on the sides of the foot and around the heel. The digits are densely clothed in white hairs. The plantar surface of the pes is granular and supports a total of five conspicuously striated pads. The inner metatarsal and first interdigital pads are fused with no obvious line of demarcation; their combined length is 13.1 mm. The outer metatarsal pad measures 7.4 mm and is separated by a gap of 3.9 mm from the fourth interdigital pad. The second to fourth interdigital pads are elongate and of subequal length. The cranium of WAM 7674 is illustrated in Figure 19. Measurements of the holotype and summary statistics for available specimens of both sexes are provided in Table 5. Male-biased sexual dimorphism is present in SL, ZW and PRW (Table 6). The lower canines of males are only slightly larger on average than those of females (Table 5; Fig. 13). Comparison with the nomino-typical race indicates that this is a consequence of a lesser degree of canine enlargement in males combined with a greater degree of enlargement in females. Other notable features of the cranium are the relatively great width of the foramen magnum in both sexes (a feature shared with South Australian and Victorian P. t. tapoatafa) and the relatively large auditory bullae of females (also matched by females of the New South Wales and Queensland population of P. t. tapoatafa) (Table 5). Observed variation. Adults attain body weights to 222 g for males and to 166 g for females (Rhind 2002). External measurements taken from museum specimen records are summarised in Table 4. The ratio of average body weight for adult males and females, measured in each of two years, was 1.24 and 1.44 (Rhind & Bradley 2002). Females have six to eight teats in the pouch, with the following frequencies (6 teats: 46 %; 7 teats: 18 %; 8 teats: 39 %; n = 82; data from Rhind et al. 2001: 362; also Rhind 2002). Fur inside the pouch of adult females is gingery (e. g., WAM 16027). Ventral colouration varies somewhat among the material examined; some individuals (e. g., CM 6810, WAM 16027) have a broad off-white to cream zone on the abdomen and extensive cream fur on the inside of the limbs, while others have less extensive pale wash on the chest and throat (e. g., WAM 2839, WAM 5006). One specimen (WAM 888) has poorly-developed cream-tipped preauricular patches; another (WAM 2839) lacks any darkening of the mid-dorsal fur. The outer metatarsal pad is variable in length; the longest measured (WAM 16027) is 9.1 mm, separated by 2.2 mm from the fourth interdigital pad. The pinnae on one specimen (WAM 2839) are more heavily pigmented. Taxonomic remarks. Rhind et al. (2001) identified some of the diagnostic cranial differences between southwestern and southeastern P. tapoatafa and recommended that they be recognised as distinct taxa. The same view was expressed by Spencer et al. (2001), based on the combined genetic and morphological evidence; however, their estimate of genetic divergence for cyt b was inflated, owing to a local sequence misalignment at position 322 (Spencer et al. 2001: Fig. 1). Former and present distribution. Phascogale t. wambenger was recorded historically from a wide region of southwestern Western Australia, from Lake Hinds in the north, to Kalgan in the southeast (Fig. 20). This broad distribution implies that the species once occupied a variety of different habitat types ranging from closed forest to open woodland. Surviving populations are largely restricted to forests dominated by jarrah (Eucalyptus marginata) (Rhind 2004). The Lake Hinds record and one from Nukarni represent significant inland outliers among the historical records. However, sub-fossil records indicate an even broader range in late prehistoric or early historic times, extending to the vicinity of Geraldton in the north and to the southern edge of the Nullarbor Plain in the east (Lundelius & Turnbull 1973, 1978; Baynes 1987; see Fig. 20 & Table 9). These records raise the possibility that P. t. wambenger formerly occupied much of the West Australian ‘ wheatbelt’ and was thus broadly sympatric with P. calura. Phascogale tapoatafa is not represented in more northerly sub-fossil localities in the Carnarvon Basin (Baynes 2000) or on Cape Range Peninsula (Baynes & Jones 1993). On a more general note, these widely distributed historical and sub-fossil records also point to an early and catastrophic historical decline of this species across much of its range, perhaps leaving only the most resilient local populations to persist, albeit tenuously, through to the commencement of systematic faunal survey and collection during the early decades of the 20 th century. In this regard, it is interesting to note Leake’s (1962: 51) statement that this species was often seen in the Kellerberrin district in the wheatbelt of Western Australia but, ' like other native animals, it disappeared in the nineties of the last [19 th] century'.	en	Have, Ten (2015): Taxonomic revision of Phascogale tapoatafa (Meyer, 1793) (Dasyuridae; Marsupialia), including descriptions of two new subspecies and confirmation of P. pirata Thomas, 1904 as a ‘ Top End’ endemic. Zootaxa 4055 (1): 1-73, DOI: 10.11646/zootaxa.4055.1.1
03A087E6FFD08319D788FBEAD74EFAE4.taxon	materials_examined	Holotype: WAM 16028. Adult male. Pago Mission, Napier Bay, near Broome, Western Australia, 14 0 8 ’ S 126 0 43 ’ E. Collected by Gerald F. Hill, 19 th February 1910. Puppet skin and skull. Paratype: WAM 16029. Subadult male. Locality and collector as for holotype. Collected on 20 th October 1909. Puppet skin and skull.	en	Have, Ten (2015): Taxonomic revision of Phascogale tapoatafa (Meyer, 1793) (Dasyuridae; Marsupialia), including descriptions of two new subspecies and confirmation of P. pirata Thomas, 1904 as a ‘ Top End’ endemic. Zootaxa 4055 (1): 1-73, DOI: 10.11646/zootaxa.4055.1.1
03A087E6FFD08319D788FBEAD74EFAE4.taxon	etymology	Etymology. Referring to the geographic region of occurrence of this taxon.	en	Have, Ten (2015): Taxonomic revision of Phascogale tapoatafa (Meyer, 1793) (Dasyuridae; Marsupialia), including descriptions of two new subspecies and confirmation of P. pirata Thomas, 1904 as a ‘ Top End’ endemic. Zootaxa 4055 (1): 1-73, DOI: 10.11646/zootaxa.4055.1.1
03A087E6FFD08319D788FBEAD74EFAE4.taxon	diagnosis	Diagnosis. A small-bodied race of P. tapoatafa that differs from the typical race of southeastern Australia in the following ways: externally, with shorter, crisper and generally paler dorsal fur, more extensive cream fur on venter, relatively shorter tail, more boldly patterned face with contrasting cream muzzle and cheek, inner metatarsal pad contacting but clearly distinct from first interdigital pad, probable lower number of teats; cranially, very similar to the other subspecies of P. tapoatafa but smaller in all cranial dimensions for both sexes and with proportionally narrower foramen magnum, relatively deeper rostrum and proportionally larger molars; genetically, with 3.0 – 5.8 % sequence divergence for cyt b from other populations. Additional differences from P. t. wambenger and P. pirat a are listed under the accounts of those taxa.	en	Have, Ten (2015): Taxonomic revision of Phascogale tapoatafa (Meyer, 1793) (Dasyuridae; Marsupialia), including descriptions of two new subspecies and confirmation of P. pirata Thomas, 1904 as a ‘ Top End’ endemic. Zootaxa 4055 (1): 1-73, DOI: 10.11646/zootaxa.4055.1.1
03A087E6FFD08319D788FBEAD74EFAE4.taxon	description	Description of holotype. External measurements, taken from label: Total length 350, Ear 30; as measured on puppet skin: HB c. 200, TL c. 205, Pes c. 35. The dorsal fur is relatively short and crisp, the contour hairs measuring 5 – 6 mm in the centre of the back. Overall colour of the back and flanks is pale grey with a silvery frosting. Abundant and evenly distributed blacktipped guard hairs project up to 7 mm beyond the contour hairs. The ventral fur is a uniform cream from the vent to the chin. A diffuse, yellowish patch on the throat overlies a pigmented gular gland that measures 25 mm long by 6 mm wide. The boundary between flank and ventral colouration is relatively sharp. The fore- and hind-limbs are grey externally and cream internally, with a sharp demarcation, especially on the hindlimb. The scrotum is lightly pigmented and thinly clothed in cream fur. The face is boldly patterned with a russet mid-rostral stripe that merges posteriorly into a dark grey patch on the crown, a cream preauricular patch, a dark loreal stripe, a dark postorbital spot, and distinct, black-rimmed orbital crescents. The cream fur of the chin and throat merges with a cream supralabial bar that extends partway along the upper lip, while the cheek patch is cream peppered with black. The vibrissal pads are thinly clothed in short cream fur, peppered with black. The mystacial vibrissae are thick and black, some with short gingery tipping, but the papillae are unpigmented. The pinnae are moderately large and thinly furred except along the anterior margin which supports a fringe of short black hairs. The tail supports a black ‘ brush’ measuring 135 mm. The basal portion of the tail is only moderately furred, producing the effect of an elongate ‘ stem’ (c. 67 mm). The dorsal fur of the stem is grey near the base, blending distally into off-white. The under-surface is grey blending into russet at the base of the brush. The upper surface of the pes is grey with a silvery frosting, changing abruptly to white on the sides and on the digits. The plantar surface of the pes is granular and supports six conspicuously striated, raised pads. The inner metatarsal and first interdigital pads are in contact but with an obvious line of demarcation; their combined length is 12.6 mm. The outer metatarsal pad measures 6.7 mm and is separated by a gap of 2.7 mm from the fourth interdigital pad. The second to fourth interdigital pads are elongate and of nearly equal length. The cranium of WAM 16028 is illustrated in Figure 21. Measurements are provided in Table 5. Too few specimens are available to demonstrate sexual dimorphism. However, the two adult males differ from the one adult female in ways that suggest male-biased dimorphism of the kind noted in the nomino-typical population — the male skulls are broader and more robust for their length, and have canines enlarged relative to near equal-sized cheekteeth. Observed variation. External measurements taken from museum specimen records are summarised in Table 4. The paratype, a subadult male, has softer, lusher fur of a more even grey colour. The tail ‘ stem’ is proportionally shorter (35 mm vs 116 mm for brush). WAM 1309, an adult female from Kunmunya Hill, has a well developed pouch with sparse, pale gingery hairs but the number of teats cannot be determined. The dorsal fur of this specimen is overall more gingery than the holotype or paratype but this is probably due to fading. The ventral fur is cream to the roots along a mid-ventral zone, from the chin to around the pouch. This is flanked on each side by a band of grey-based and cream tipped fur that extends onto the inside of the fore- and hind-limbs. The preauricular patch is more distinct in this specimen. The tail ‘ stem’ is proportionally shorter again, measuring 35 mm vs 125 mm for brush). WAM 53763, a damaged roadkill specimen, is an adult female with a damaged pouch; the number of teats cannot be determined. The pes measures 34 mm and the tail 207 mm with a basal stem of 55 mm. The outer metatarsal tubercle in this specimen is very short (4.7 mm; gap to fourth interdigital pad 3.5 mm) and the second interdigital pad is slightly longer than the third and fourth. The combined inner metatarsal tubercle and first interdigital pad are smoothly united on the left pes but distinguished by a clear kink on the left pes. Rhind et al. (2001) reported a female specimen from Mitchell Plateau (WAM 27208; as KIM 5) with six teats; it is not known whether this represents the typical teat number. Taxonomic remarks. This population has attracted little previous attention, although Troughton (1967: 32) remarked that “ the Northern Australian race (Phascogale tapoatafa pirata) … is also found in the Kimberley division, Western Australia ”. Rhind et al. (2001) also allied this population with P. t. pirata based on the small body size, slight brownish tinge to the dorsal fur and extensive cream venters. A total of five museum specimens are known from the entire Kimberley region (see Supplementary Table II for details). Former and present distribution. The few records of this taxon are scattered throughout the Kimberley region, from Yampi Sound in the west to near Kalumburu (Pago Mission) in the east (Fig. 22). There are recent records from Kimbolton Station near Derby (2003; WAM 53763) and from the King Leopold Range (2001; sighting by Mr Murray Ellis, pers. comm.). Undated subfossil occurrences in the southwest Kimberley are reported by Start et al. (2012).	en	Have, Ten (2015): Taxonomic revision of Phascogale tapoatafa (Meyer, 1793) (Dasyuridae; Marsupialia), including descriptions of two new subspecies and confirmation of P. pirata Thomas, 1904 as a ‘ Top End’ endemic. Zootaxa 4055 (1): 1-73, DOI: 10.11646/zootaxa.4055.1.1
03A087E6FFDF8311D788F8D8D754FABF.taxon	materials_examined	Holotype: BMNH 4.1.3.100. South Alligator River. Collected by J. T. Tunney, 24 th May 1903. Puppet skin and skull. Revised diagnosis. A relatively small-bodied Phascogale that differs from all races of P. tapoatafa in the following ways: externally, in having a distinctly brownish wash to fur of the dorsum and flanks, pure white fur on the upper surface of the pes, a more elongate outer metatarsal pad, a similarly elongate but narrower pinna, a pure white cheek patch, and a lack of differentiation in fur colour between the upper and lower surface of the tail ‘ stem’; dentally, in having a less enlarged and procumbent I 1, a less pronounced size gradient along I 2 - 4, a more compact upper premolar series with overlap between each tooth, a relatively more reduced P 1, a more elongate preparacrista on M 1, smaller protocones and metacones on M 4, smaller entoconids and less prominent buccal cingulids on M 1 - 3, a more anteriorly positioned paraconid on M 1, a more reduced taloned on M 4, a less enlarged I 1 and a less reduced I 3; and cranially, in being generally narrower across the braincase and zygomatic arches and in having an anteriorly less extensive lacrimal, a less prominent lacrimal preorbital crest, typically paired lacrimal foramina, a less prominent maxillonasolabialis fossa on the zygomatic arch, less forwardly rotated orbits, a shorter but deeper alisphenoid auditory bulla, and a posteriorly less inflated rostral tympanic process of the petrosal. Other proportional differences include more pronounced post-orbital constriction, a narrower foramen magnum and relatively smaller molars. It further differs from all southern and eastern races of P. tapoatafa in having shorter, crisper dorsal fur and extensive all-white ventral fur. It further differs from P. t. kimberleyensis in having the mid-dorsal fur darker than the flanks, at least some white-tipped guard hairs, whiter ventral fur, a more elongate and less heavily furred pinna, grey fur on the vibrissal pads, a more heavily pigmented scrotum, less elongate claws on the manus and a relatively longer tail (Table 4). It differs from P. cal ura in numerous features including much larger adult body size, essentially grey rather than brown dorsal colouration, a more elongate tail with more prominent brush, pure white fur on the upper surface of the pes, and absence of a contrasting postauricular patch. Genetically, P. p i r a t a differs from the various races of P. tapoatafa at 7.3 – 16.3 % sequence divergence for cyt b (10.8 – 16.3 % for P. t. tapoatafa; 7.3 – 10.5 % for P. t. wambenger; 11.7 – 14.4 % for P. t. kimberleyensis); and it differs from P. c al ur a at 10.3 – 11.4 % for the same gene.	en	Have, Ten (2015): Taxonomic revision of Phascogale tapoatafa (Meyer, 1793) (Dasyuridae; Marsupialia), including descriptions of two new subspecies and confirmation of P. pirata Thomas, 1904 as a ‘ Top End’ endemic. Zootaxa 4055 (1): 1-73, DOI: 10.11646/zootaxa.4055.1.1
03A087E6FFDF8311D788F8D8D754FABF.taxon	description	Description. External measurements taken from museum specimen records are summarised in Table 4. The dorsal fur is relatively short and crisp, the contour hairs measuring 6 – 7 mm in the centre of the back. The overall colour of the back and flanks is dull grey but with a slight brownish tinge, overlain by a fine peppering of white and black. This effect is produced by a combination of cream tipping on grey-based underfur, white tipping on grey based contour hairs, and an abundance of weakly emergent guard hairs that vary in colour from dark russet on the upper back and flanks to black on the rump. A small proportion of the guard hairs end in short white tips. The middorsal zone is distinctly darker than the flanks, especially on the upper back and extending forward onto the crown. The ventral fur is pure pale cream to the bases ranges along a central zone extending from the chin to the cloaca. On the central part of the body this merges laterally into a zone of fur that is pale grey-based but with progressively shorter cream tipping, giving an overall off-white appearance. The inner surface of the fore-limbs is clothed in short cream fur, while that on the hind-limbs is lusher and pale grey-based with cream tipping. The scrotum of adult males (e. g., CM 7225, CM 7729) is heavily pigmented and densely clothed in cream fur. The pouch of adult females (e. g., CM 8808, CM 11724) is thinly furred with pale russet hairs and contains six elongate teats. A pigmented gular gland is present in both sexes but is much larger and more conspicuous on adult male specimens where it measures 15 – 20 mm in length and 7 – 8 mm in width. The location of this gland is not marked by any obvious external discolouration of the fur in either sex. The head is boldly patterned. The coronal patch is dark grey and continuous with an indistinct to moderately well-defined mid-rostral stripe. This is flanked on each side by a band of pale grey fur flecked with black that extends onto the vibrissal pads. A distinct, dark grey loreal stripe runs from behind the mystacial vibrissae to the anterior corner of the eye. The eye itself is narrowly rimmed with lines of intense black hairs that merge posteriorly into a well-marked black to russet-brown postorbital spot. Distinct, all white orbital crescents are also present. The pale cream fur of the chin and throat extends around the corner of the mouth to merge with a supralabial bar and associated cheek patch rising to the level of the genal vibrissae. A narrow band of off-white fur, flecked with russet, separates the pale cream cheek patch from the white infraorbital crescent. The vibrissae are moderately thick and black, occasionally with short gingery tipping, and the vibrissal papillae are pigmented and visible through the fur. The pinna is elongate but moderately narrow, with a weakly developed posterior lobe. The external surface is very thinly furred, while the inner surface supports patches of white hairs anteriorly and posteriorly. The anterior margin of the pinna is emarginated by a narrow line of short black hairs. The basal portion of the tail is thickly furred above and on the sides, continuing the colour and texture of the rump, while the ventral surface is more sparsely clothed in long, adpressed pale cream hairs (Fig. 6). The tail narrows rapidly to form an elongate ‘ stem’ (to 85 mm) that is clothed above and below by elongate black-based but cream-tipped hairs, giving an overall grizzled appearance. The tail ‘ brush’ is relatively unexpanded and measures up to 170 mm; it consists of jet black hairs except for a short section of dark russet hairs ventrally, at the base of the brush. The upper surface of the pes is densely clothed in white hairs that extend around the heel and forward onto the digits. The plantar surface of the pes is granular and supports a total of five conspicuously striated pads. The inner metatarsal and first interdigital pads are fused with no obvious line of demarcation; their combined length is 11.7 mm. The outer metatarsal pad measures 7.5 mm and is separated by a gap of no more than 2 mm from the fourth interdigital pad. The second to fourth interdigital pads are elongate; the second is approximately 25 % longer than the third or fourth. The cranium of the holotype (BM 4.1.3.100, an adult male) is illustrated in Figure 24 alongside an adult male from the vicinity of el Sherana, South Alligator River area (CM 7225). Significant features were noted in an earlier section. Cranio-dental measurements for each sex are provided in Table 5. The small sample of P. p i r a t a shows clear male-biased sexual dimorphism in cranio-dental measurements (Tables 5 & 6). For all dimensions bar one (FMW), mean values are higher in males than females and for seven dimensions (SL, OBW, C-M 4, ARW, PRW, LCL, LCW) the contrast is statistically significant. Several other contrasts approach statistical significance, including key measures of overall cranial size (e. g. ZW). Sexual dimorphism in canine size is particularly striking in this species. Individual and geographic variation. The venter in some individuals (e. g., CM 7031, CM 8808) is entirely pale cream without the flanking grey-based zone. An adult male from Yirrkali (AM 7269), preserved as a flat skin, has an unusually narrow and short tail brush. We suspect that the animal was in poor health or perhaps recovering from being singed, an interpretation prompted by the observation that some parts of the tail support longer hairs that project through the brush, and the fact that fur on the lower back is very sparse. The skull of this specimen is shattered. Two specimens from Melville Island (CM 9983, CM 9989; both lacking skulls) differ from the mainland sample in having a browner tinge to the dorsal fur, considerably shorter tails with less densely furred brushes, and shorter pinnae. The ventral fur in the adult male (CM 9989) is shorter and less dense than in any mainland individual; the juvenile (CM 9983) has more luxuriant ventral fur but with the grey-based zone extending almost to the midline on the abdomen. This population may warrant subspecific distinction from typical pirata on the adjacent mainland; however, this action is deferred pending more detailed assessment of this population. Taxonomic remarks. Thomas’ (1904) description, based on two individuals of each sex, drew attention to all of the key diagnostic differences between pirata and tapoatafa. Nevertheless, he concluded that pirata ‘ is in general appearance exceedingly like the common Brush-tailed Phascogale of temperate Australia’. Spencer et al. (2001) reported the genetic distinction of P. p i r a t a from eastern and southwestern populations of P. tapoatafa but gave overinflated values due to sequence misalignment. The previous morphometric comparisons of ‘ pirata’ by Rhind et al. (2001) were compromised by the a priori pooling of the Top End, Kimberley and Cape York Phascogale populations on the assumption that these represented a single taxon. Recognition of P. pirata as a full species within the Phascogale tapoatafa group is justified by the numerous points of morphological distinction that distinguish P. p i r a t a from all other phascogales, including other northern populations formerly included within the composite subspecies ‘ P. t. pirata ’, and by the high level of mitochondrial sequence divergence between this taxon and all populations of P. tapoatafa (individual contrasts range 9.4 – 12.3 % sequence divergence for for cyt b). The differences in dental morphology between P. pi r a t a and P. tapoatafa are suggestive of significant ecological contrasts that remain to be elucidated by field observations. The patchwork geographic distribution of the two species also highlights their separate evolutionary histories since the range of P. pi r at a is effectively embedded within the more widely distributed and regionally differentiated populations of P. tapoatafa. Geographic distribution. The historical mainland range of P. pirata probably extended across much of the ‘ Top End’, with records extending from Litchfield in the Daly River catchment to Yirrkala on Gove Peninsula (Fig. 22). Most records come from the western part of this range, probably reflecting the historical focus of exploration and collecting. The only confirmed island population of the species is from Melville Island. A record from 1988 of a Phascogale on West Island in the Sir Edward Pellew Group is unvouchered; while this was referred to P. pirata by Woinarski et al. (2011), as noted above, it might also represent an outlier of the north Queensland population of P. tapoatafa. Phascogale pirata has declined across most of its former range and now appears to be everywhere scarce. The most recent sightings come from Kakadu and Litchfield National Parks and from near Batchelor. A recent survey of the Sir Edward Pellew Island Group failed to detect populations of phascogales on any island (Woinarski et al. 2011). The geographic relationship between populations of P. pirata and P. tapoatafa in the east is unclear owing to uncertainty over the specific identity of the populations on the Sir Edward Pellew islands and at Edward River on the western side of Cape York. However, some kind of range disjunction is more or less guaranteed by the presence of extensive tracts of essentially treeless habitats that occupy the country at the head of the Gulf of Carpenteria, the so-called ‘ Carpenteria Gap’ (MacDonald 1969; Ford 1987; Schodde & Mason 1999). On the western side, a narrower gap of around 300 km separates records of P. p i r a t a from the nearest known P. t. kimberleyensis. This gap corresponds to a stretch of essentially treeless black soil country in the Victoria and Keep River basins, which most likely represents a real barrier to dispersal by Brush-tailed Phascogales. The occurrence of P. tapoatafa populations both to the east and to the west of P. p i r a t a raises the possibility that populations of P. tapoatafa may still occur in the Northern Territory. The specific identity of all populations in this region should be reassessed in view of the newly identified specific distinction between P. p i r a t a and P. tapoatafa. The label attached to the holotype of P. p i r a t a contains the annotation “ caught in hollow tree, Aboriginal name ‘ Woomboo’ ”.	en	Have, Ten (2015): Taxonomic revision of Phascogale tapoatafa (Meyer, 1793) (Dasyuridae; Marsupialia), including descriptions of two new subspecies and confirmation of P. pirata Thomas, 1904 as a ‘ Top End’ endemic. Zootaxa 4055 (1): 1-73, DOI: 10.11646/zootaxa.4055.1.1
