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Microchip-automated devices have the potential to provide individual free-living animals with safe nesting areas and act as a method of targeted food delivery, while excluding competitors and predators. Wildlife have been successfully trained to use such devices in captivity but never in the wild. Bringing animals into captivity may not always be feasible or appropriate due to the high cost, likely increased stress on the animals, and potential biosecurity risk. Therefore to demonstrate proof of concept that wildlife could be trained in situ to use commercially available microchip-automated devices, a brush-tailed phascogale in the wild was exposed to a microchip-automated door attached to a nest box. The phascogale was successfully trained within 15 days to use the microchip-automated door.

There are many limitations when using traditional laboratory species. Limits on variation, may result in limited outcomes, at both the species and individual level, due to different individuals/species having diverse physiological processes, or differing molecular and genetic mechanisms. By using a variety of model species, we will be able to develop creative solutions to biological problems and identify differences of which we were not previously aware. The laboratory mouse has been a suitable model species for various mammalian studies, however most are bred specifically for laboratory research with limited variability due to selective breeding. Marsupial models offer unique research opportunities compared to eutherian models. We believe that there should be an expansion in marsupial model species, and the introduction of the red-tailed phascogale (Phascogale calura), a dasyurid marsupial, should be one of them. Phascogales are easily managed in captivity, and there are now multiple studies involving their development, reproduction, nutrition, behavior and immune system, which can serve as a baseline for future studies. The addition of the phascogale as a model species will improve future mammalian studies by introducing variability and offer alternate solutions to biological problems, particularly in the areas of genetics, nutrition, immunology, the neuro-endocrine system, and ageing, due to their semelparous reproductive strategy and hence, subsequent predictive physiology. In this review, we provide information based on existing research on red-tailed phascogales to support their inclusion as a model species.

Hearing is essential for communication, to locate prey and to avoid predators. We addressed the paucity of information regarding hearing in Australian native mammals by specifically assessing the hearing range and sensitivity of the red-tailed phascogale (Phascogale calura), the kultarr (Antechinomys laniger) and the spinifex hopping-mouse (Notomys alexis). Auditory brainstem response (ABR) audiograms were used to estimate hearing thresholds within the range of 1–84 kHz, over a dynamic range of 0–80 dB sound pressure level (SPL). Phascogales had a hearing range of 1–40 kHz, kultarrs 1–35 kHz and hopping-mice 1–35 kHz, with a dynamic range of 17–59 dB SPL, 20–80 dB SPL and 30–73 dB SPL, respectively. Hearing for all species was most sensitive at 8 kHz. Age showed no influence on optimal hearing, but younger animals had more diverse optimal hearing frequencies. There was a relationship between males and their optimal hearing frequency, and greater interaural distances of individual males may be related to optimal hearing frequency. Because nocturnal animals use high-range hearing for prey or predator detection, our study suggests this may also be the case for the species examined in this study. Future studies should investigate their vocalizations and behaviour in their natural environments, and by exposing them to different auditory stimuli.

Dasyurus tafa White in É. Geoffroy Saint-Hilaire, 1803 Dasyurus tafa White in É. Geoffroy Saint-Hilaire, 1803b: 259. COMMON NAME. — Brush-tailed Phascogale. CURRENT NAME. — Phascogale tapoatafa (Meyer, 1793). See Jackson & Groves (2015: 64). COLLECTOR/ S. — Charles-Alexandre Lesueur, expedition Nicolas Baudin, contra de Beaufort (1966: 519) who mentioned both Péron and Lesueur as its collectors. COLLECTION LOCALITY. — Near Port Jackson (Sydney), New South Wales, Australia. COLLECTION DATE. — Between 3 July and 18 November 1802. SPECIMEN NUMBER/S. — No specimen present in MNHN collection. COMMENTS. — É. Geoffroy Saint-Hilaire (1804a: 360) noted that “I only know this species by the description and the figure by John White” (White 1790: 281). The illustration referred to above under Dasyurus penicillatus É. Geoffroy Saint-Hilaire, 1804a may also be attributable to this taxon. ILLUSTRATIONS. — No data., Published as part of Jackson, Stephen M., Jansen, Justin J. F. J., Baglione, Gabrielle & Callou, Cécile, 2021, Mammals collected and illustrated by the Baudin Expedition to Australia and Timor (1800 - 1804): A review of the current taxonomy of specimens in the Muséum national d'Histoire naturelle de Paris and the illustrations in the Muséum d'Histoire naturelle du Havre, pp. 387-548 in Zoosystema 43 (21) on pages 400-402, DOI: 10.5252/zoosystema2021v43a21, http://zenodo.org/record/5142972, {"references":["MEYER F. A. A. 1793. - Systematische-summarische Uebersicht der neuesten zoologischen Entdeckungen in Neuholland und Afrika. Nebst zwey andern zoologischen Abhandlungen. Dykischen Buchhandlung, Leipzig. https: // www. biodiversitylibrary. org / page / 27654933","JACKSON S. M. & GROVES S. M. 2015. - Taxonomy of Australian Mammals. CSIRO Publishing, Melbourne, 529 p.","BEAUFORT F. DE 1966. - Catalogue des types de mammiferes du Museum national d'Histoire naturelle. Paris. VI. Monotremata. - VII Marsupialia. Bulletin du Museum national d'Histoire naturelle. Section B. Botanique, Biologie et EcologieVegetables Phytochimie (2) 38: 509 - 553. https: // www. biodiversitylibrary. org / page / 55121975","WHITE J. 1790. - Journal of a Voyage to New South Wales; with sixtyfive plates of non descript animals, birds, lizards, serpents, curious cones of trees and other natural productions. J. Debrett, London. https: // www. biodiversitylibrary. org / page / 50707371"]}

A fifth of all species in the carnivorous‐insectivorous marsupial family Dasyuridae show obligate male semelparity (die‐off), where males die after their first mating season, but females may breed in more than one season. Life history strategies of many species in the genera Antechinus and Phascogale are well known, but the other dasyurid genus with apparently obligately semelparous males, Dasykaluta, is poorly known, and there have been no demographic studies of this monotypic genus in the wild. Both sexes in Antechinus and Phascogale species are highly polygamous. Comparative and experimental tests of insectivorous marsupials indicate that this polygamy is linked to the evolution of semelparity in more strongly seasonal breeders, through intense sperm competition. We assessed whether the kaluta (Dasykaluta rosamondae), a small dasyurid endemic to the broader Pilbara region of Western Australia, has obligate male semelparity, multiple paternity within litters, and positive testis allometry in the wild. We documented complete die‐off of males over two breeding seasons. Multiple paternity was confirmed in seven of eight litters, where either two or three fathers sired young. The seven young in the remaining litter appeared to be sired by one male. Intraspecific positive testis allometry was found in adult males. We conclude that the life history strategies of wild male and female kalutas are consistent with those of the two other genera of dasyurids with obligate male semelparity (Antechinus and Phascogale), and sperm competition is likely to be important in the mating system of kalutas. Our results show that male obligate semelparity has evolved twice in dasyurids.

Nest boxes are used to manage populations of tree-cavity dependent birds and mammals. Concerns have been raised that due to their poor insulative properties nest boxes may cause heat stress and occasionally death during summers of extreme maximum temperatures. Our study investigated whether this nest box heat stress hypothesis applies to two small cavity-dependent mammals (brush-tailed phascogales and sugar gliders). Focusing on days when ambient temperature reached ≥40 °C, we recorded: i) temperatures within occupied nest boxes, ii) temperatures within nearby unoccupied tree cavities, iii) the duration of temperatures of ≥40 °C within nest boxes, iv) whether direct mortality was observed, and v) the relative abundance of these species in nest boxes before and after a very hot summer. When ambient temperature reached ≥40 °C, nest boxes were equivalent to ambient or 1–2 °C cooler, whereas tree cavities were 3–7 °C cooler than ambient. Exposure in nest boxes to temperatures of ≥40 °C lasted an average of 2–8 h. We observed no mortality over 65 records of phascogales and 31 records of gliders in nest boxes on days when ambient reached ≥40 °C. No decline in abundance was recorded after a summer with 11 days of temperatures ≥40 °C, with each species subsequently occupying >40 nest boxes. Our observations suggest these species are tolerant of the high temperatures that occurred. Nonetheless, provision of nest boxes designed to minimise summer heating is recommended.

Constructing hollows or cavities in trees with chainsaws is an emerging approach to manage hollow-dependent species in hollow depleted landscapes. Small-scale experiments are required to refine this approach before implementing on a broad scale. We addressed two questions: i) are chainsaw hollows used by the regionally threatened brush-tailed phascogale (Phascogale tapoatafa) and the non-threatened sugar glider (Petaurus notatus), ii) do chainsaw hollows retain their integrity over time. We paired 45 chainsaw hollows with nest boxes of equivalent internal dimensions in southeastern Australia and inspected these on 20 occasions over 2.5 years. Camera traps revealed phascogales visited some cavities within hours of installation and monthly inspections revealed rapid uptake of these hollows by both target species. Overall, phascogales and sugar gliders used 32% and 84% of the chainsaw hollows respectively, and 21% and 82% of the nest boxes. We used multi-method occupancy to compare detection within the two types of cavities. Detection models that included cavity type had more support than those without. Detection of both species was substantially higher in the chainsaw hollows compared to the nest boxes. Over the 2.5-year monitoring period the faceplates of some chainsaw hollows showed signs of deformity. Callous regrowth over the faceplate was pronounced on some trees suggesting the need for periodic maintenance. Our study confirms the potential of chainsaw hollows to restore habitat for hollow-dependent mammals but highlights periodic maintenance is likely to be a feature of this approach as it is with nest boxes.

Expressed coding sequences for interleukin-6 (IL-6) and interleukin-6 receptor α (IL-6R) were examined in five marsupial species. Full length expressed coding sequences for IL-6 and IL-6R were identified and characterized in the gray short-tailed opossum (Monodelphis domestica). For IL-6, ∼225 bp fragments of the mRNA sequence were identified in the red-tailed phascogale (Phascogale calura), kultarr (Antechinomys laniger), and stripe-faced dunnart (Sminthopsis macroura), while ∼563 bp fragments of mRNA encoding IL-6R were identified in the red-tailed phascogale, kultarr, stripe-face dunnart and fat-tailed dunnart (Sminthopsis crassicaudata). Relative expression levels of IL-6 and IL-6R were examined in the heart, muscle, lung, liver, spleen and kidney of adult red-tailed phascogales, and IL-6 gene expression was found to be significantly higher in the lung and spleen than the other tissues examined, while the expression of IL-6R was significantly higher in the liver, lung and spleen. These results now serve as a reference point for examining the role and levels of IL-6 and IL-6R in the health and disease of these marsupial species. The pro-tumorigenic nature of IL-6 is of particular interest, and the identification of these IL-6 and IL-6R coding sequences provides a platform for further work to evaluate the potential role of IL-6 in marsupial cancers.

Marsupials are born immunologically premature, relying on cells and molecules in maternal milk for immune protection. Both immunoglobulin and complement proteins have been identified in marsupial milk, but the expression of specific complement proteins remains largely unexplored. We report partial cDNA sequences for two complement-activating proteins, C3, C1r, CFP and MASP2, in liver tissues from red-tailed phascogale (Phascogale calura). Conservation of functionally relevant motifs were identified in the translated cDNA sequences from phascogale C3, CFP and MASP2 and their eutherian homologues. Gene expression of representative molecules from each of the major complement pathways was also investigated in whole body tissues from 1 to 18 day old animals and liver tissues from 31-day to 14-month old animals. Average complement expression in whole bodies and liver tissues of C1r, CFP, MASP2 and C3 increased significantly in juveniles compared to pouch young, presumably due to the maturation of the young's own complement system. Comparing expression in liver tissues only, we found that the average CFP expression were higher in pouch young compared to juveniles, while results were still statistically similar to the average expression of all tissues for C1r, MASP2 and C3. The average complement expression then significantly decreased as the animals aged into adulthood.

The threatened northern brush-tailed phascogale (Phascogale pirata) is one of the most poorly known mammals in Australia. While the few available records indicate a decline in its distribution and abundance, it has not previously been subject to intensive targeted survey. Here, we trialled a specifically tailored methodology for detection of P. pirata, with the aim of informing ongoing survey and monitoring of this species. We deployed 50 motion-sensor cameras (spaced closely together in a grid 500×1000m) on Melville Island (Northern Territory, Australia), between June 2018 and May 2019. Cameras were baited and secured to trees ~3m above the ground on a bracket facing the trunk. We selected for large (>30cm diameter at breast height [DBH]) trunks of the dominant tree species (Eucalyptus miniata, E. tetrodonta and Corymbia nesophila). We detected P. pirata 16 times on eight cameras over the duration of the study, finding that detection was most likely on large (DBH >41.5cm) E. tetrodonta trees during the wet season. Our results indicate that survey effort for this species should be seasonally targeted and focussed on large trees. However, the efficacy of additional methods (nest boxes, Elliott traps) and a comparison between detections on arboreal versus ground-based cameras requires further investigation. We highlight the importance of conducting additional work on this species, as little is known about its ecology, population trends and threats, making it difficult to assess its conservation status. Without more targeted work, P. pirata is at risk of slipping into extinction unnoticed.

ContextFeral cats have benefitted from effective control of foxes in south-western Australia and, consequently, their impact on some threatened mammal species has increased. Control of feral cats in the region can be enhanced by use of the Eradicat® cat bait, but its impact on non-target animal populations requires investigation before widespread use. AimsThe aim of the present study was to determine through field trials whether consumption of Eradicat® baits by resident red-tailed phascogales, following a broadscale baiting operation to control feral cats, was sufficiently frequent to cause significant rates of mortality in wild populations of phascogales. MethodsNine radio-tagged red-tailed phascogales were monitored through an Eradicat® baiting event to determine their survival. Removal and consumption of toxic and non-toxic rhodamine B-labelled baits by a range of species were monitored with camera traps and by subsequent trapping of red-tailed phascogales and other mammals to sample whiskers for evidence of rhodamine uptake. Key resultsAlthough some phascogales showed interest in baits and sometimes moved them from the deposition site, all radio-tagged phascogales survived for at least 1 week after baiting, by which time very few or no baits remained. Examination of whiskers sampled from individuals exposed to rhodamine-labelled baits showed that consumption of non-toxic Eradicat® baits by phascogales was negligible; only one phascogale of 62 sampled showed any rhodamine banding. ConclusionsThe present study provided no evidence that red-tailed phascogales in the study region are at risk from an Eradicat® baiting episode in autumn. ImplicationsThe risk to red-tailed phascogale populations through the use of Eradicat® baiting to control cats in their habitat in the Great Southern region of Western Australia is likely to be low. Further research to elucidate any impact of repeated baiting on populations of this species at several locations is recommended.

Background The red-tailed phascogale is a ‘Near Threatened’ dasyurid marsupial. Males are semelparous and die off shortly after the breeding season in the wild due to a stress-related syndrome, which has many physiological and immunological repercussions. In captivity, males survive for more than 2 years but become infertile after their first breeding season. Meanwhile, females can breed for many years. This suggests that captive males develop similar endocrine changes as their wild counterparts and undergo accelerated aging. However, this remains to be confirmed. The health status and immune function of this species in captivity have also yet to be characterized. Results Through an integrative approach combining post-mortem examinations, blood biochemical and hematological analyses, we investigated the physiological and health status of captive phascogales before, during, and after the breeding season. Adult males showed only mild lesions compatible with an endocrine disorder. Both sexes globally maintained a good body condition throughout their lives, most likely due to a high quality diet. However, biochemistry changes potentially compatible with an early onset of renal or hepatic insufficiency were detected in older individuals. Masses and possible hypocalcemia were observed anecdotally in old females. With this increased knowledge of the physiological status of captive phascogales, interpretation of their immune profile at different age stages was then attempted. During the breeding season, males developed a stress leukogram characterized by a marked lymphopenia, further aggravated by a severe leukopenia after the breeding season. To determine whether these changes were limited to the peripheral blood or had more profound implications, histopathology of the spleen was performed opportunistically. Adult males showed white pulp atrophy, at various degrees. The atrophy was mainly lymphoid and more severe in 1.5-year-old males than in 3.5-year-old females. These results suggest that captive males undergo accelerated immunosenescence. Conclusions Functional studies are now needed to characterize the underlying mechanisms leading to immunosenescence in marsupials. Semelparous dasyurids present great potential for studying the effects of sex and stress on immunity in marsupials. Characterization of these immune-endocrine interactions may help refine veterinary treatment plans, husbandry protocols and conservation programs to maintain the health of captive and wild populations.

Cells expressing the surface markers CD3, CD4, CD79b, IgM, MHC class II, and ModoUG (nonclassical MHC class I) were detected in red-tailed phascogale tissues using immunohistochemistry, and the appearance and localization of cells observed here was consistent with previous observations in other marsupial species. CD3+ cells were first detected at one day postpartum (dpp) in the thymus, followed by ModoUG+ cells at 5-7 dpp in the thymus and lymph nodes. CD79b+ cells were first detected at 12-14 dpp in bone marrow, spleen, and lymph nodes. IgM+ cells were first detected at 12-14 dpp in thymus, bone marrow, spleen, and lymph nodes. MHC class II+ cells were first detected at 12-14 dpp in thymus, bone marrow, and lymph nodes. CD4+ cells were detected in adult thymus and spleen only. The presence of the mature immune cell populations and their localization to characteristic T and B cell zones in mature lymphoid tissues with normal histological structure indicates that red-tailed phascogales develop immunocompetence by the end of pouch life. Anat Rec, 302:1985-2002, 2019. © 2019 American Association for Anatomy.

Objective Marsupials suffer from an increasing number of stressors in this changing world. Functional studies are thus needed to broaden our understanding of the marsupial immune system. The red-tailed phascogale (Phascogale calura) is a small Australian marsupial previously used in descriptive immunological studies. Here, we aimed to develop functional assays by isolating and stimulating blood and spleen mononuclear cells in vitro. Results While peripheral blood mononuclear cell (PBMC) were relatively easy to isolate, only 105 mononuclear cells (> 90% purity and > 75% viability) could be recovered from the spleen, independently of the sex and age of the animal or the centrifugation time and speed tested. The pores of the mesh sieve used for tissue homogenization might have been too big to yield a single cell suspension. Nevertheless, in spite of the overall low number of cells recovered, PBMC and splenic mononuclear cells were successfully activated in preliminary trials with phytohemaglutinin. This activation state was evidenced by a change in shape and the presence of small cell aggregations in the mitogen-stimulated cultures. A non-radioactive colorimetric assay was also performed to confirm cell proliferation in these wells. This work highlights the importance of developing and reporting detailed methodological protocols in non-traditional research species.

Expressed coding sequences were identified for the T cell receptors TCRα, TCRβ, TCRγ, TCRδ, TCRμ, and co-receptors CD3e, CD4, and CD8α in the red-tailed phascogale (Phascogale calura). Expression of these genes was examined in the developing thymus and spleen of pouch young and juvenile individuals to detect any changes in expressionrelative to developmental stages. Transcripts of CD3e, CD4, TCRδ and TCRμ were first detected in the thorax at 1 day postpartum (dpp), and transcripts of all target genes were detected in the thymus from 3 dpp, and the spleen from 17 dpp. The relative expression of all target genes changed significantly over the course of pouch life and was associated with histological milestones of both the thymus and the spleen. Peak gene expression was observed early in pouch life in the thymus, with a decrease in expression associated with timing of thymic involution, while expression peaked later in the spleen and was associated with the initial differentiation of the red and white pulp and a population of T cells forming the periarterial lymphatic sheath (PALS). Active expression of these target genes within the thymus and spleen indicates that the young marsupials are actively developing immunocompetence during pouch life.

Red-tailed phascogale (Phascogale calura) pouch young at birth were relatively underdeveloped in comparison with their eutherian counterparts, and the lymphoid tissues of the immune system were found to be histologically immature. The phascogale thymus rapidly developed in the first few days of pouch life and was quickly populated with lymphocytes. By the end of pouch life, involution of the thymus was underway. The bone marrow started to develop in the early stage of pouch life, although adipocytes and megakaryocytes were not observed until slightly later. The liver was hematopoietic from birth and reached histological maturity toward the end of pouch life. The lymph nodes were difficult to detect macroscopically because of their small size, but were easily identified microscopically later in pouch life, particularly in the mesentery, and these lymph nodes exhibited germinal centers by the end of pouch life. The early spleen was predominately mesenchymal, but exhibited some erythropoiesis. Follicles with well-developed germinal centers were not observed until the latest stage of pouch life. Although intraepithelial lymphocytes were detected in the intestines early in pouch life, the discrete lymphoid aggregates and Peyer's patches characteristic of the gut-associated lymphoid tissue (GALT) were not detected until later in pouch life. This is the first report of histological development in phascogale pouch young, as well as the first report of the thymus, bone marrow, and lymph nodes in this dasyurid species at any age.

Torpor is an important physiological process used to reduce body temperature and thus energy needs in mammals and birds. Rewarming from torpor can be costly. In some small mammals basking is used to reduce the cost of rewarming from torpor. The use and duration of basking was measured in two native Australian marsupials, the red-tailed phascogale (Phascogale calura) and kultarr (Antechinomys laniger), using a heat rock. Animals were observed via video cameras to determine whether they used basking behaviours and whether this behaviour was affected by food availability. Both species became accustomed to the heat rock as time progressed. Absence of food increased basking duration and frequency in some individuals but not all animals. Generally kultarrs basked for longer periods than phascogales, and kultarrs would bask at all hours of the day while phascogales showed a strict nocturnal regime (1900-0700h). The results demonstrate that basking may be an important behavioural adaptation of the kultarr; while the phascogale requires further research.

Ixodes goliath n. sp. (Acari: Ixodidae), is described based on females collected from the eastern hyomys, Hyomys goliath (Milne-Edwards) (Rodentia: Muridae) from Papua New Guinea. Females of I. goliath n. sp. are similar to those of I. priscicollaris Schulze, 1932 but can be distinguished by the overall size, porose areas sculpture and shape of palpal segment I ventrally. For comparative purposes, the female of I. priscicollaris is redescribed. Studied females of I. priscicollaris were found on murid rodents and Phascogale sp. (Dasyuromorphia: Dasyuridae) from Indonesia (Papua Province) and Papua New Guinea.

Phascogale flavipes burrelli Le Souef, in Le Souef et al., 1926 Le Souef and Burrell, The Wild Animals of Australasia, p. 344, fig. 92. (October 1926). Common name. Brown Antechinus. Current name. Antechinus stuartii Macleay, 1841; following Jackson & Groves (2015). Holotype. M.2593, by original designation. Male, skull, study skin, Ebor, NSW, registered on 8 November 1915, presented by the Council of the Zoological Society of NSW. Collector and collection date not given. The original M Register entry is “ Phascogale flavipes Ebor, N.S. Wales Pres. Council Zoological Society”. Condition. Cranium and dentaries complete. Study skin has two bald patches on the ventral surface and is missing tail tip. Type locality. Localized to Ebor, NSW by Iredale & Troughton (1934), given as “ Highlands of Northern New South Wales ” in the original account. Comments. Although authorship attribution to Le Souef and Burrell is entrenched in the literature, in the original description it is clearly attributed to Le Souef alone, and is called “Burrell’s Phascogale ”. A single specimen is mentioned by registration number in the original account and we have not found any indication from the M Register that further specimens were involved. Iredale & Troughton (1934) note “ ex Le Souef MS”—presumably Le Souef had prepared a more detailed account in an unpublished manuscript. Troughton (1941, and subsequent editions) doubted the validity of this taxon and he stated that it was “ described as having a smaller body, comparatively larger feet, and slightly crested tail. These characters were possibly due to a period of captivity; so the race may be disregarded until wild specimens are found with similar characters.” This taxon was confused with A. flavipes but Wakefield & Warneke (1967) placed it with A. stuartii, sensu lato. Dickman et al. (1998), in describing A. agilis, placed burrelli with A. stuartii, sensu stricto. Skull measurements of the holotype of burrelli are given by Dickman et al. (1998) and Van Dyck & Crowther (2000)., Published as part of Parnaby, Harry E., Ingleby, Sandy & Divljan, Anja, 2017, Type Specimens of Non-fossil Mammals in the Australian Museum, Sydney, pp. 277-420 in Records of the Australian Museum 69 (5) on page 309, DOI: 10.3853/j.2201-4349.69.2017.1653, http://zenodo.org/record/5237800, {"references":["Le Souef, A. S., H. Burrell, and E. L. G. Troughton. 1926. The Wild Animals of Australasia: Embracing the mammals of New Guinea and the Nearer Pacific Islands. With a chapter on the bats of Australia and New Guinea by Ellis Le G. Troughton. Sydney: G. G. Harrap. 388 pp. + 105 pls","Macleay, W. S. 1841. Notice of a new genus of Mammalia discovered by J. Stuart, Esq., in New South Wales. Annals and Magazine of Natural History (series 1) 8 (51): 241 - 243, pl. vii.","Troughton, E. [L. G.] 1941. Furred Animals of Australia. First edition. Sydney: Angus & Robertson. xxvii + 374 pp.","Dickman, C. R., H. E. Parnaby, M. S. Crowther, and D. H. King. 1998. Antechinus agilis (Marsupialia: Dasyuridae), a new species from the A. stuartii complex in south-eastern Australia. Australian Journal of Zoology 46 (1): 1 - 26. https: // doi. org / 10.1071 / ZO 97036"]}

Phascogale macdonnellensis Spencer, 1895a Proc. R. Soc. Vic. 7: 222. (January 1895). Common name. Fat-tailed Pseudantechinus. Current name. Pseudantechinus macdonnellensis (Spencer, 1895a), following Jackson & Groves (2015). ? Syntype. M.1004,?male, adult, skull, skin mount, registered 8 September 1895, from “Central Australia ” = Northern Territory, presented by Baldwin Spencer, as a skin mount. Condition. Cranium is missing right zygomatic arch, right auditory capsule, occipital area and section of right lateral braincase wall and floor; hole in left frontal; both dentaries complete. Skin mount: faded, tail tip almost detached, left ear torn and tip missing, missing toes on the right manus, a fracture on the tail, sparse fur on the ventral surface and the right side. Considered to be adult, based on dentition and sutures. Sex not determined (skin mount has no signs of scrotum or pouch but, as an adult, it could be a male with scrotum removed). Type locality. Alice Springs district, Northern Territory, Australia. Topotypes. (2). M.1145, adult?female, skin mount, skull in situ; M.1146, adult male, skull, skin mount; both from Alice Springs, donated by Baldwin Spencer, registered 10 October 1896. Collector and collection date not given. There is no associated correspondence listed in the M Register for either specimen. Comments. Correspondence from Spencer dated 5 Sept 1895 (AM Archives AMS9, Letters Received, S:65/1895) clearly indicates that one male macdonnellensis was sent to the AM; it was registered as M.1004 but without further data. There was no indication that it was a type although Spencer used the term “type” in other correspondence. Spencer (1895a) did not indicate the number of specimens examined but more than one is implied by the statement that “the first specimen was found by Mounted Trooper South …”. As noted by Calaby (1996), Spencer (1895b), published in January, states that he had three specimens for his description, a male and two females. Dixon (1970) listed C7804 (a female) as a syntype, noting an attached label marked “type”, apparently Spencer’s original tag. She incorrectly cites Trooper South as the collector—he obtained a male specimen. Dixon states that the only other specimen listed in the MV register is a male; presumably she had reason to doubt the type status of this specimen. We are unable to determine whether either M.1004 or the MV male cited by Dixon (1970) is the syntype male. While it is possible that M.1004 (assuming that it is male) is the missing male syntype, it might be one of a series of specimens obtained after Spencer prepared the initial description. In his report on the Horn Expedition, Spencer (1896a) indicated that he had acquired 13 specimens by that time and M.1004 could have been one of these. Calaby (1996) provided insights into specimens sent to Spencer by collectors after the Horn Expedition. In a letter to Spencer dated 29 July 1895, F. J. Gillen, one of Spencer’s collectors, indicates that he was about to ship seven or eight specimens of this species to Spencer. The AM specimen could have been one of these, given that Spencer sent it to the AM in September 1895 and the fact that other shipments to Spencer were prior to November 1895 (Calaby, 1996). Mahoney & Ride (1988b) cited MV C7804 as the holotype, presumably because the ambiguity of Spencer’s original account could allow an interpretation that it was based on one specimen. We do not recognize the inference of a holotype by Mahoney & Ride (1988b) as constituting lectotype designation because, prior to their work, there was no agreement that the species had been based on a single specimen. To the contrary, Dixon (1970) had assumed the existence of a syntype series (see Article 74.6, the Code)., Published as part of Parnaby, Harry E., Ingleby, Sandy & Divljan, Anja, 2017, Type Specimens of Non-fossil Mammals in the Australian Museum, Sydney, pp. 277-420 in Records of the Australian Museum 69 (5) on page 310, DOI: 10.3853/j.2201-4349.69.2017.1653, http://zenodo.org/record/5237800, {"references":["Spencer, W. B. 1895 a. Preliminary notice of two new species of marsupials from central Australia. Proceedings of the Royal Society of Victoria (new series) 7: 222 - 224.","Calaby, J. H. 1996. Baldwin Spencer's post-Horn Expedition collectors in central Australia. In Exploring Central Australia: Society, the Environment and the 1894 Horn Expedition, ed. S. R. Morton and D. J. Mulvaney, pp. 188 - 208. Chipping Norton, NSW: Surrey Beatty.","Spencer, W. B. 1895 b. The Horn Expedition to Central Australia. Nature 51 (1314): 222 - 223. https: // doi. org / 10.1038 / 051222 b 0","Dixon, J. M. 1970. Catalogue of mammal types (Class Mammalia) in the National Museum of Victoria. Memoirs of the National Museum of Victoria 31: 105 - 114.","Spencer, W. B. 1896 a. Mammalia. In Report on the work of the Horn Scientific Expedition to Central Australia, Part 2. Zoology, ed. B. Spencer, pp. 1 - 52, pls 1 - 4. Melbourne: Melville, Mullen and Slade."]}

Microchip-automated feeders and doors allow individualised access to supplementary food and shelter during soft-release of wildlife. A wild-caught brush-tailed phascogale was used to test whether a wild animal could be trained to use microchip-automated devices. The phascogale was trained to use each device in less than a month.

We report the capture of individuals of two species of phascogale (Phascogale tapoatafa wambenger and P. calura) at the same locations within the same habitat and during the same survey event, something that has not been reported previously. This would suggest that these species may occur in sympatry where their distributions overlap. Further investigations are required to confirm the extent of sympatry as this could have implications for the conservation and management of both species.

The red-tailed phascogale is a small arboreal dasyurid marsupial that inhabits semi-arid to arid regions of Western Australia's wheat belt. Its body mass (34.7 g) is only ∼15% of that predicted based on its phylogenetic position among other dasyuromorphs; we interpret this as an adaptation to its scansorial and semi-arid/arid lifestyle. The standard physiology of this species at a thermoneutral ambient temperature of 30 °C conforms to that of other dasyurid marsupials; body temperature (34.7 ± 0.37 °C), basal metabolic rate (0.83 ± 0.076 mL O2 g−1 h−1), evaporative water loss (1.68 ± 0.218 mg H2O g−1 h−1) and wet thermal conductance (3.8 ± 0.26 J g−1 h−1 °C−1) all fall within the 95% predication limits for the respective allometric relationships for other dasyurid species. Thermolability confers an energy savings at low Ta and water savings at high Ta. Torpor, observed at low Ta, was found to be more beneficial for energy savings than for water economy. The red-tailed phascogale therefore has a physiology suitable for the challenges of arid environments without any obvious requirement for adaptations to its scansorial lifestyle, other than its considerably lower-than-expected body mass.

Lyn is a tyrosine kinase molecule required for modulation of signalling cascades in cell populations including B lymphocytes of the mammalian immune system. We have characterised the coding domain of the marsupial lyn gene of two macropod marsupials; the Tammar Wallaby (Macropus eugenii) and the Bridled Nailtail Wallaby (Onychogalea fraenata) and show the co-expression of two Lyn isoforms in cells and tissues of these and three other marsupials (Brush-tail Possum, Trichosurus vulpecula; American Grey Short-tailed Opossum, Monodelphis domestica and Red-tailed Phascogale, Phascogale calura). The predicted Lyn proteins (LynA and LynB) were highly conserved across vertebrate species, with amino acid identities of 94% with their human orthologues and conservation of key tyrosine kinase motifs that suggests that marsupial Lyn most likely functions in cell signalling. Comparison of our cDNA data to annotations for Lyn transcripts (available through the Ensembl Genome Browser) for the Tammar Wallaby confirm splice sites for a number of exons in the wallaby transcript that are missing from the current annotation. This is the first report of the expression of kinase signalling molecules that influence immunity in metatherian mammals and provides key information to support ongoing studies of immune regulation in marsupials.

Antimicrobial substances in serum include circulating complement proteins and acute phase proteins (APPs). We identified gene sequences for APPs, haptoglobin (Hp), C-reactive protein (CRP) and serum amyloid A (SAA) in marsupial genomes. Hp and SAA levels were measured in red-tailed phascogale ( Phascogale calura ) sera using commercially available assays. Hp levels were higher in males than females, while SAA levels suggest the phascogales used in this study were healthy. Serum was co-cultured with four bacterial species. Bacterial growth was inhibited after incubation at 37 °C, however effectiveness differed with bacteria and incubation time. The least amount of bacterial growth was noticed after introduction to K. pneumoniae , and most when introduced to P. aeruginosa . Despite marsupials not having mature immune tissues at birth, and unable to mount specific immune responses, this study suggests other immune strategies, such as APPs in serum likely aid marsupials in their defence against pathogens.

BackgroundReference genes serve an important role as an endogenous control/standard for data normalisation in gene expression studies. Although reference genes have recently been suggested for marsupials, independent analysis of reference genes on different immune tissues is yet to be tested. Therefore, an assessment of reference genes is needed for the selection of stable, expressed genes across different marsupial tissues.MethodsThe study was conducted on red-tailed phascogales (Phascogale calura) using five juvenile and five adult males. The stability of five reference genes (glyceraldehyde-3-phosphate dehydrogenase,GAPDH;β-actin,ACTB;18SrRNA,18S; 28SrRNA, 28S;and ribosomal protein L13A,RPL13A) was investigated using SYBR Green and analysed with the geNorm application available in qBasePLUSsoftware.ResultsGene stability for juvenile and adult tissue samples combined show thatGAPDHwas most stable in liver and lung tissue, and18Sin small intestine and spleen. While all reference genes were suitable for small intestine and spleen tissues, all reference genes except28Swere stable for lung and only18Sand28Swere stable for liver tissue. Separating the two age groups, we found that two different reference genes were considered stable in juveniles (ACTBandGAPDH) and adults (18Sand28S), andRPL13Awas not stable for juvenile small intestine tissue. Except for28S, all reference genes were stable in juvenile and adult lungs, and all five reference genes were stable in spleen tissue.DiscussionBased on expression stability,ACTBandGAPDHare suitable for all tissues when studying the expression of marsupials in two age groups, except for adult liver tissues. The expression stability between juvenile and adult liver tissue was most unstable, as the stable reference genes for juveniles and adults were different. Juvenile and adult lung, small intestine and spleen share similar stable reference genes, except for small intestine tissues where all reference genes were stable in adults butRPL13Awas not suitable in juveniles.

Ecological monitoring is widely used to measure change through time in ecosystems. The current extinction crisis has resulted in a wealth of monitoring programs focussed on tracking the status of threatened species, and the perceived importance of monitoring has seen it become the cornerstone of many biodiversity conservation programs. However, many monitoring programs fail to produce useful outcomes due to inherent flaws. Here we use a monitoring program from south-eastern Australia as a case study to illustrate the potential of such endeavours. The threatened carnivorous marsupial, the brush-tailed phascogale (Phascogale tapoatafa), has been monitored at various locations between 2000 and 2010. We present strong evidence for a decline in relative abundance during this period, and also describe relationships with environmental variables. These results provide insights likely to be valuable in guiding future management of the species. In the absence of the monitoring program, informed management would not be possible. While early detection of population declines is important, knowledge of the processes driving such declines is required for effective intervention. We argue that monitoring programs will be most effective as a tool for enhanced conservation management if they test specific hypotheses relating to changes in population trajectories. Greater emphasis should be placed on rigorous statistical analysis of monitoring datasets in order to capitalise on the resources devoted to monitoring activities. Many datasets are likely to exist for which careful analysis of results would have benefits for determining management directions.

Red-tailed phascogales Phascogales calura are near-threatened (Friend, 2008) arboreal Dasyurids. A breeding programme was established at Alice Springs Desert Park in 2001 to aid species recovery. Twenty-five captive-bred phascogales were released into a suitable habitat at the park in 2006. If shown to be successful, the initial release was to be expanded with the release of further captive-bred phascogales into a suitable habitat in the nearby National Park and into South Australia. In this study, a dietary analysis was conducted to determine the preferred diet of the translocated phascogales in the park environment. Scats were collected during July–October, 2006 and January–March, 2007 from nesting sites within the park. Faecal samples were weighed, soaked in hot water and particles were separated through sieves before examination under a microscope. Scat analysis methods identified that red-tailed phascogales were primarily insectivorous with 92.6% of all scats containing arthropods. They are also opportunistic predators within the park, consuming birds (51.6%), small mammals (33.3%) and on occasion reptiles, and plant material (27.4%). Seasonal comparison of data through SIMPER analyses showed there was significant variation (P=0.009) between spring and summer, due to a large portion of birds present in the diet in spring. The red-tailed phascogale is able to exploit a number of prey types and it is therefore likely that they would survive a ‘hard’ translocation into the wild provided the site chosen has adequate food supply.

The brush-tailed phascogale (Phascogale tapoatafa tapoatafa) is a marsupial threatened in Victoria by habitat fragmentation and modification. As part of the development of infrastructure in phascogale habitat, Melbourne Water was required to improve habitat for the phascogale. We aimed to develop and test a method for monitoring phascogales on Melbourne Water land, and to assess the effectiveness of habitat improvements. Trapping, nest boxes, and cameras were all successful in detecting phascogales; however, cameras had the highest detection per unit effort. We suggest that future monitoring should favour camera surveys with trapping potentially every 3–5 years for acquisition of physical data.

It is commonly accepted that dasyurids (Marsupialia: Dasyuridae) radiated in the late Miocene or early Pliocene in response to a drying trend in Australia's climate as evidenced from the high diversity of dasyurids from modern arid environments compared with Miocene rainforest assemblages. However, mid-Pleistocene dasyurid assemblages from cave deposits at Mt Etna, Queensland are more diverse than any previously known from rainforest habitats. New taxa will be described elsewhere, but include three new genera as well as new species of Dasyurus, Antechinus and Phascogale. Comparison of dasyurids from Mt Etna sites that are interpreted as rainforest palaeoenvironments with fossil and extant assemblages indicate that they are at least as diverse as those from modern arid environments. Thus Neogene diversification of dasyurids occurred in both arid and rainforest habitats, but only the former survived continuing aridification. Hence, aridification cannot be invoked for the diversification of all dasyurid lineages.

The red-tailed phascogale (Phascogale calura) is an endangered dasyurid from southwestern Western Australia that has been bred in captivity since 2001. It is an annual breeder that has a restricted breeding period, and males only participate in a single breeding season. We examined the reproductive biology of female red-tailed phascogales, assessing methods for detection of estrus, variation in the timing of breeding in captivity across facilities and years, and reproductive success. Body mass and pouch changes were useful indicators of estrus, whereas assays for fecal estradiol and progesterone did not allow for accurate prediction of ovulation. Females mated with multiple males and matings occurred over a period of at least 5 days, with females storing sperm in the lumen and crypts of the oviduct. Births tended to occur in July at Alice Spring Desert Park, with births at Adelaide Zoo occurring from early June to late August. The predictable births at Alice Springs are similar to those observed with Antechinus, but with flexibility remaining in the reproductive strategy of the species as observed at Adelaide. Of the 146 breeding females at Alice Springs, 127 females produced 846 pouch young, of which 68% were weaned. A female bias was observed in weaned young. Information gained from this study has been incorporated into the captive-breeding program for this species.

The quolls are the largest native predators remaining on mainland Australia. We describe the cross-species testing of all available microsatellite loci for representative across the entire distribution of quolls, with the additional characterization of five new microsatellite loci from the Chuditch. All primers produced clear and polymorphic amplification patterns containing between nine and 17 alleles and with high levels of diagnostic variability. These highly polymorphic primers make them useful additional tools in planning conservation strategies across related conspecifics, many of which are under threat.

Five species of dasyurid marsupials endemic to New Guinea and herein referred to as Murexia longicaudata, M. rothschildi, M. habbema, M. naso, and M. melanurus have a varied taxonomic history. Until 1984, the latter 3 species had been assigned to the otherwise strictly Australian genus Antechinus, but comparisons of phallic morphology showed that this arrangement was untenable. Immunological and DNA hybridization studies of dasyurids suggested that New Guinean “antechinuses” were part of a clade that included M. longicaudata and M. rothschildi, a finding that was confirmed by early DNA sequence comparisons. Comprehensive analyses of cytochrome b, 12S rRNA, and protamine P1 sequences from all recognized dasyurid species led to the recommendation that these 5 species be included in a single genus (Murexia) that, along with Phascogale and Antechinus, constitutes the dasyurid tribe Phascogalini. Recently, however, a morphocladistic study concluded that these species are neither closely related to one...

Genus Phascogale Temminck, 1824 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. Remarks. Iredale & Troughton (1934) were the first to recognise the combination Phascogale tapoatafa (Meyer, 1793)., Published as part of 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, pp. 1-73 in Zootaxa 4055 (1) on page 23, DOI: 10.11646/zootaxa.4055.1.1, http://zenodo.org/record/240153, {"references":["Temminck, C. J. (1824) Monographies de Mammalogie, ou description de quelques genres de Mammiferes, dont les especes ont ete observees dans les differens Musees de l'Europe, & c. Ouvrage accompagne de planches d'Osteologie, pouvant servir de suit et de complement aux notices sur les animaux vivans, publiees par M. le Baron G. Cuvier, dans ses recherches sur les ossemens fossils. Tome 1. G. Dufour et E. D'Ocagne, Paris, 72 pp.","Shaw, G. (1800) General Zoology or Systematic Natural History. Vol. 2. F. Kearsley, London, 502 pp. http: // dx. doi. org / 10.5962 / bhl. title. 53459","Meyer, F. A. A. (1793) Systematische-summarische Uebersicht der neuesten zoologischen Entdeckungen in Neuholland und Afrika. Nebst zwey andern zoologischen Abhandlungen. im Verlage der Dykischen Buchhandlung, Leipzip, 208 pp.","Gould, J. (1844) Exhibition and character of a number of animals, & c. transmitted from Australia by Mr. Gilbert. Proceedings of the Zoological Society of London, 12 (1844), 103 - 107.","Thomas, O. (1904) On a collection of mammals made by Mr. J. T. Tunney in Arnhem Land, Northern Territory of South Australia. In: Rothschild, W. Hartert, E. & Jordan, K. (Eds.), Novitates Zoologicae. Vol. XI. Zoological Museum, Tring, pp. 222 - 229.","Iredale, T. & Troughton, E. le G. (1934) A Checklist of the Mammals Recorded From Australia. Memoir VI. The Australian Museum, Sydney, 122 pp."]}

Phascogale tapoatafa wambenger Rhind & Aplin subsp. nov. 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. 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. 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). 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 222g for males and to 166g 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'. Phascogale t. wambenger has been the focus of a major field study in the jarrah forest (Rhind 1996, 1998, 2002, 2003, 2004; Rhind & Bradley 2002). Outside of this habitat, nothing is recorded of its natural history., Published as part of 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, pp. 1-73 in Zootaxa 4055 (1) on pages 35-37, DOI: 10.11646/zootaxa.4055.1.1, http://zenodo.org/record/240153, {"references":["Bindon, P. & Chadwick, R. (2002) A Nyoongar Wordlist from the South West of Western Australia. Western Australian Museum, Perth, 496 pp.","Rhind, S. G. & Bradley, J. S. (2002) The effect of drought on body size, growth and abundance of wild brush-tailed phascogales (Phascogale tapoatafa) in south-western Australia. Wildlife Research, 29, 235 - 245. http: // dx. doi. org / 10.1071 / WR 01014","Spencer, P. B. S., Rhind S. G. & Eldridge, M. D. B. (2001) Phylogeographic structure within Phascogale, (Marsupialia: Dasyuridae) based on partial cytochrome b sequence. Australian Journal of Zoology, 49, 369 - 377. http: // dx. doi. org / 10.1071 / ZO 00080","Rhind, S. G. (2004) Direct impacts of logging and forest management on the brush-tailed phascogale Phascogale tapoatafa and other arboreal marsupials in a jarrah forest of Western Australia. In: Lunney, D. (Ed.), Conservation of Australia's Forest Fauna. 2 nd Edition. Royal Zoological Society of NSW, Mosman, pp. 639 - 655.","Lundelius, E. L. & Turnbull, W. D. (1973) The mammalian fauna of Madura Cave, Western Australia; Part I. Fieldiana Geology, 31, 1 - 35.","Lundelius, E. L. & Turnbull, W. D. (1978) The mammalian fauna of Madura Cave, Western Australia; Part III. Fieldiana Geology, 38, 1 - 120.","Baynes, A. (1987) The original mammal fauna of the Nullarbor and southern peripheral regions: evidence from skeletal remains in superficial cave deposits. In: McKenzie, N. L. & Robinson, A. C. (Eds.), A Biological Survey of the Nullarbor Region, South and Western Australia in 1984. National Parks and Wildlife Service, Department of Environment and Planning, South Australia, Adelaide, pp. 139 - 145, 400 - 401.","Baynes, A. (2000) Original mammal faunas of the Carnarvon Basin, based on fossil material from the surfaces of small caves. Supplementary Table 2. In: McKenzie, N. L., Hall, N. J. & Muir, W. P. Non-volant mammals of the southern Carnarvon Basin, Western Australia. Records of the Western Australian Museum, 61 (Supplement), 479 - 510.","Baynes, A. & Jones, B. (1993) The mammals of Cape Range Peninsula, Western Australia. Records of the Western Australian Museum, 45 (Supplement Number), 207 - 225.","Rhind, S. G. (1996) Habitat tree requirements and the effects of removal during logging on the marsupial Brush-tailed Phascogale (Phascogale tapoatafa) in Western Australia. The Western Australian Naturalist, 21, 1 - 22. [note: errata Vol 22]"]}

Phascogale tapoatafa (Meyer, 1793) Phascogale tapoatafa featured as the 'Tapoa Tafa' (or 'Tapha') in White's 'Journal of a Voyage to New South Wales' (1790), an adaptation of the local Aboriginal name tapoatafa. This name was first used in the formal combination Viverra tapoatafa Meyer, 1793. The species group names penicillata Shaw, 1800, and tafa White, 1803, were proposed on the basis of the same original account and thus are objective junior synonyms of tapoatafa Meyer, 1793. Apart from pirata Thomas, 1904, there are no other available species level names for members of the P. tapoatafa group. Previous workers have recognized only one subspecies apart from the nominotypical form��� P. tapoatafa pirata Thomas 1904, applied to some or all of the various northern Australian populations (e.g. Iredale & Troughton 1934; Tate 1947; Troughton 1967; Cuttle 1983, Soderquist 1995 a). We recognize P. pirata as a very distinct species and distinguish three subspecies within P. tapoatafa, two of them newly described herein., Published as part of 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, pp. 1-73 in Zootaxa 4055 (1) on page 23, DOI: 10.11646/zootaxa.4055.1.1, http://zenodo.org/record/240153, {"references":["Meyer, F. A. A. (1793) Systematische-summarische Uebersicht der neuesten zoologischen Entdeckungen in Neuholland und Afrika. Nebst zwey andern zoologischen Abhandlungen. im Verlage der Dykischen Buchhandlung, Leipzip, 208 pp.","Shaw, G. (1800) General Zoology or Systematic Natural History. Vol. 2. F. Kearsley, London, 502 pp. http: // dx. doi. org / 10.5962 / bhl. title. 53459","White, J. (1803) in Geoffroy Saint-Hilaire, E. Note sur les especes du genre dasyure. Bulletin des sciences par la Societe Philomatique de Paris, 3 (81), 158 - 159. [misprinted as should be pages 258 - 259]","Thomas, O. (1904) On a collection of mammals made by Mr. J. T. Tunney in Arnhem Land, Northern Territory of South Australia. In: Rothschild, W. Hartert, E. & Jordan, K. (Eds.), Novitates Zoologicae. Vol. XI. Zoological Museum, Tring, pp. 222 - 229.","Iredale, T. & Troughton, E. le G. (1934) A Checklist of the Mammals Recorded From Australia. Memoir VI. The Australian Museum, Sydney, 122 pp.","Tate, G. H. H. (1947) Results of the Archbold Expeditions. No 56: On the anatomy and classification of the Dasyuridae (Marsupialia). Bulletin of the American Museum of Natural History, 88, 97 - 156.","Troughton, E. (1967) Furred Animals of Australia. 9 th Edition. Angus and Roberston, Sydney, 376 pp.","Cuttle, P. (1983) Brushtailed Phascogale. In: Strahan, R. (Ed.), The Australian Museum Complete Book of Australian Mammals. Angus and Robertson Publishers, Sydney, pp. 34 - 35.","Soderquist, T. R. (1995 a) Spatial organisation of the arboreal carnivorous marsupial Phascogale tapoatafa. Australian Journal of Zoology, 237, 385 - 398."]}

Phascogale pirata Thomas, 1904 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. 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������., Published as part of 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, pp. 1-73 in Zootaxa 4055 (1) on pages 43-48, DOI: 10.11646/zootaxa.4055.1.1, http://zenodo.org/record/240153, {"references":["Thomas, O. (1904) On a collection of mammals made by Mr. J. T. Tunney in Arnhem Land, Northern Territory of South Australia. In: Rothschild, W. Hartert, E. & Jordan, K. (Eds.), Novitates Zoologicae. Vol. XI. Zoological Museum, Tring, pp. 222 - 229.","Spencer, P. B. S., Rhind S. G. & Eldridge, M. D. B. (2001) Phylogeographic structure within Phascogale, (Marsupialia: Dasyuridae) based on partial cytochrome b sequence. Australian Journal of Zoology, 49, 369 - 377. http: // dx. doi. org / 10.1071 / ZO 00080","Woinarski, J. C. Z., Ward, S., Mahney, T., Bradley, J., Brennan, K., Ziembicki, M. & Fisher, A. (2011) The mammal fauna of the Sir Edward Pellew island group, Northern Territory, Australia: refuge and death-trap. Wildlife Research, 38, 307 - 322. http: // dx. doi. org / 10.1071 / WR 10184","MacDonald, J. D. (1969) Notes on the taxonomy of Neositta. Emu, 69, 169 - 174. http: // dx. doi. org / 10.1071 / MU 969169","Ford, J. (1987) Minor isolates and minor geographical barriers in avian speciation in continental Australia. Emu, 87, 90 - 102. http: // dx. doi. org / 10.1071 / MU 9870090","Schodde, R. & Mason, I. J. (1999) The Directory of Australian Birds. Passerines. CSIRO Publishing, Collingwood, 851 pp."]}

Phascogale tapoatafa kimberleyensis Aplin & Rhind subsp. nov. 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. Etymology. Referring to the geographic region of occurrence of this taxon. 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. 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 3.5mm) 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)., Published as part of 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, pp. 1-73 in Zootaxa 4055 (1) on pages 38-40, DOI: 10.11646/zootaxa.4055.1.1, http://zenodo.org/record/240153, {"references":["Troughton, E. (1967) Furred Animals of Australia. 9 th Edition. Angus and Roberston, Sydney, 376 pp.","Start, A. N., Burbidge, A. A., McDowell, M. C. & McKenzie, N. L. (2012) The status of non-volant mammals along a rainfall gradient in the south-west Kimberley, Western Australia. Australian Mammalogy, 34, 36 - 48. http: // dx. doi. org / 10.1071 / AM 10026"]}

Phascogale tapoatafa ���Cape York population��� This population remains poorly understood. Although we have examined a total of 15 specimens from three areas [Atherton Tableland (n = 1); Cooktown area (n = 5); and Lockhart River (n = 9); see Fig. 22 & Supplementary Table II for details], only two are represented by cleaned skulls and one of these (CM 12699) is shattered. Sparse museum records of this form are supplemented by sightings at various other localities in north Queensland. These suggest a geographic range extending at least from Iron Range in the north, to Elizabeth Creek, Mt Surprise, in the south. Unvouchered but otherwise reliable records of Phascogales are available from two localities on the margin of the Gulf of Carpentaria (Figs 18 & 22). Several individuals including a female with seven young were recorded by Donald Thomson in October 1929 at Edward River on the western side of Cape York (Dixon & Huxley 1985: 32��� 33); and two individuals were captured and released in 1988 on West Island in the Sir Edward Pellew group (Johnson & Kerle 1991). The latter population was referred to P. p i r at a by Woinarski et al. (2011), though without explicit justification. On biogeographic grounds, these Carpentarian populations might represent outliers of either P. tapoatafa or P. p i r a t a and they are here left unallocated to species. A recent survey of the Sir Edward Pellew Island Group failed to detect populations of Brush-tailed Phascogales on any island (Woinarski et al. 2011). External measurements taken from museum specimen records are summarised in Table 4. The adult male specimen from the Atherton Tableland (CM 12699) is smaller than similarly-aged individuals of the nominotypical subspecies from southeast Queensland but it closely resembles this form in most aspects of external morphology. The dorsal fur is lush and pure grey, with no indication of mid-dorsal darkening, and without the brown tinge of P. pirata. The venter has a narrow mid-ventral band of creamy white fur that continues forward onto the throat and chin. Hairs on the upper surface of the pes are white intermingled with grey, changing to pure white on the digits. Significant differences are observed in the plantar pads of the pes and in the patterning of the tail stem. In the pes, the outer metatarsal tubercle is very short (6.1 mm) and more widely separated from the fourth interdigital pad (gap of 3.7 mm). The tail stem is relatively long (95 mm) compared with the brush (135 mm) and is clothed above and below in dark hairs with cream tips. This contrasts with the condition in nominotypical P. tapoatafa where the upper surface and sides of the tail stem are clothed in white hairs with lack tips. The Cooktown sample includes several adult males (JM 14678 and JM 11205) that approach the size of adult males from southeast Queensland. In general, the external morphology also falls within the range of variation of the nominotypical race, but with several significant departures. One concerns the patterning of the tail brush, which in most individuals matches the condition described above for CM 12699. However, in one adult male (JM 14678) hairs on the upper surface of the tail stem are entirely black, contrasting with white on the flanks and black to russet frosted with white below; the overall effect is unusual and different from that observed in any other population. The fur on the upper surface of the pes in most individuals is a mixture of white and dark hairs, giving way to pure white on the toes. JM 14678 differs again in having the pes almost entirely white, with dark hairs restricted to around the ankle. The condition of the plantar pads also varies somewhat more than usual within a population. In JM 14678 and JM 14477 (an adult female) the outer metatarsal tubercle is very short (6.5 and 6.1 mm, respectively), and thus comparable to CM 12699 from the Atherton Tableland. In contrast, three other specimens in the Cooktown series have elongate outer metatarsal tubercles, measuring 8.7���10.3 mm. In all specimens the inner metatarsal tubercle is fused with the first interdigital pad; in all but JM 15411 (a subadult male) the point of union is marked by a clear kink in the pad. The adult female (JM 14477) has six small but clearly defined teats inside a poorly developed pouch. This contrasts with eight teats in nominotypical P. tapoatafa. The series from Lockhart River on the eastern side of Cape York closely resembles those from Cooktown in external morphology. The only fully adult specimen (DTC 304 from Bare Hill) is represented by a skull alone; this is illustrated in Fig. 23. This is relatively small compared with series from southeastern Queensland but slightly larger than CM 12699 from the Atherton Tableland. The morphology of the skull and teeth is very close to specimens from southeast Queensland and northern New South Wales (e.g. CM 24425 from Coraki; Fig. 16) except that the molars and auditory bullae are proportionally smaller. The northern Queensland specimens of P. tapoatafa thus conform in most regards (e.g. body colouration, cranial morphology) with the nominotypical form of P. tapoatafa but share some features with the Kimberley subspecies (e.g. smaller size, reduced teat number, tail stem patterning). However, these morphological comparisons are clouded somewhat by the variation observed within and between the series. We obtained a single cyt b sequence from within this geographic region (CM 12699 from Atherton Tableland; Table 1); attempts to obtain sequence from other individuals failed, probably due to their being fixed in formalin. The sequence from CM 12699 is about equidistant from P. t. kimberleyensis (4.4���4.9 %; n = 2) and P. t. tapoatafa from New South Wales and southeast Queensland (3.0��� 5.3 %; n = 4). Pending further information on the northern Queensland populations of P. tapoatafa, we elect to leave them unallocated at subspecies level. As noted earlier, a subfossil record of P. tapoatafa from Christmas Creek, inland of Townsville, raises the possibility that Brush-tailed Phascogales once occurred along the entire length of the Queensland coast, perhaps with some narrow discontinuities through the driest areas., Published as part of 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, pp. 1-73 in Zootaxa 4055 (1) on pages 41-42, DOI: 10.11646/zootaxa.4055.1.1, http://zenodo.org/record/240153, {"references":["Dixon, J. M. & Huxley, L. M. (1985) Donald Thomson's Mammals and Fishes of Northern Australia. Thomas Nelson, Melbourne, 210 pp.","Johnson, K. A. & Kerle, J. A. (Eds.) (1991) Flora and Vertebrate Fauna of the Sir Edward Pellew Group of Islands, Northern Territory. Report to the Australian Heritage Commission. Conservation Commission of the Northern Territory, Alice Springs,","Woinarski, J. C. Z., Ward, S., Mahney, T., Bradley, J., Brennan, K., Ziembicki, M. & Fisher, A. (2011) The mammal fauna of the Sir Edward Pellew island group, Northern Territory, Australia: refuge and death-trap. Wildlife Research, 38, 307 - 322. http: // dx. doi. org / 10.1071 / WR 10184"]}

Phascogale tapoatafa tapoatafa (Meyer, 1793) 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 311g for males and 212g 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. 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. Phascogale t. tapoatafa has been the focus of several field studies in Victoria (Cuttle 1978, 1982; Soderquist 1993 a, 1993 b, 1994 a, 1994 b, 1995 a; Soderquist & Ealey 1994; Soderquist & Lill 1995) but little is known of the species��� ecology in New South Wales or Queensland. The difference between these populations in the degree of sexual dimorphism suggests caution in assuming a uniformity of all aspects of ecology among populations of this widespread subspecies. Dixon & Huxley (1989) reported extracts concerning Victorian P. tapoatafa from the natural history notes of Donald F. Thomson, drawn primarily from sources on the Bellarine Peninsula where the species was so common in the 1940 s��� 1960 s that it invaded local residences and was regarded as a nuisance. There are no modern records of the species in this, Published as part of 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, pp. 1-73 in Zootaxa 4055 (1) on pages 23-35, DOI: 10.11646/zootaxa.4055.1.1, http://zenodo.org/record/240153, {"references":["Meyer, F. A. A. (1793) Systematische-summarische Uebersicht der neuesten zoologischen Entdeckungen in Neuholland und Afrika. Nebst zwey andern zoologischen Abhandlungen. im Verlage der Dykischen Buchhandlung, Leipzip, 208 pp.","Mahoney, J. A. & Ride, W. D. L. (1988) Dasyuridae. In: Walton, D. W. (Ed.), Zoological Catalogue of Australia. Vol. 5. Mammalia. Australian Government Printers, Canberra, pp. 14 - 33.","White, J. (1790) Journal of a Voyage to New South Wales. J. Debrett, London. p. 281.","Troughton, E. (1967) Furred Animals of Australia. 9 th Edition. Angus and Roberston, Sydney, 376 pp.","Thomas, O. (1888) Catalogue of the Marsupialia and Monotremata in the Collection of the British Museum (Natural History).","Soderquist, T. R. (1995 c) Ontogeny of sexual dimorphism in size among polytocous mammals: tests of two carnivorous marsupials. Journal of Mammalogy, 76, 376 - 390.","Cuttle, P. (1982) Life history strategy of the dasyurid marsupial Phascogale tapoatafa. In: Archer, M (Ed.), Carnivorous Marsupials. Royal Zoological Society of New South Wales. Surrey Beatty, NSW, pp. 13 - 22.","Soderquist, T. R. (1993 a) Maternal strategies of Phascogale tapoatafa (Marsupialia: Dasyuridae). I. Breeding seasonality and maternal investment. Australian Journal of Zoology, 41, 549 - 566.","Rhind, S. G. & Bradley, J. S. (2002) The effect of drought on body size, growth and abundance of wild brush-tailed phascogales (Phascogale tapoatafa) in south-western Australia. Wildlife Research, 29, 235 - 245. http: // dx. doi. org / 10.1071 / WR 01014","Wood Jones, F. (1923) The Mammals of South Australia. Part I. R. E. E. Rogers, Government Printer, Adelaide, pp. 98 - 101.","Wakefield, N. A. (1964) Mammal remains. Supplementary Table (pp. 494 - 498) to: Mulvaney, D. J. (1964). Archeological excavations at Fromm's Landing on the lower Murray River, South Australia. Proceedings of the Royal Society Victoria, 77, 479 - 516.","Baynes, A. & Johnson, K. A. (1996) The contributions of the Horn Expedition and cave deposits to knowledge of the original mammal fauna of central Australia. In: Morton, S. R. & Mulvaney, D. J. (Eds.), Exploring Central Australia: Society, the Environment and the 1894 Horn Expedition. Surrey Beatty & Sons Pty Ltd, Chipping Norton, pp. 168 - 186.","Cuttle, P. (1978) The Behaviour in Captivity of the Dasyurid Marsupial Phascogale tapoatafa (Meyer). M. Sc. Thesis, Monash University, Melbourne, 241 pp.","Soderquist, T. R. (1993 b) Maternal strategies of Phascogale tapoatafa (Marsupialia: Dasyuridae). II. Juvenile thermoregulation and maternal attendance. Australian Journal of Zoology, 41, 567 - 576.","Soderquist, T. R. (1994 a) Reproductive Strategies of Phascogale tapoatafa (Marsupialia: Dasyuridae). Ph. D. Thesis, Monash University, Melbourne. [unkown pagination]","Soderquist, T. R. (1994 b) Anti-predator behaviour of the brushtailed phascogale (Phascogale tapoatafa). Victorian Naturalist, 111, 22 - 24.","Soderquist, T. R. (1995 a) Spatial organisation of the arboreal carnivorous marsupial Phascogale tapoatafa. Australian Journal of Zoology, 237, 385 - 398.","Soderquist, T. R. & Ealey, L. (1994) Social interactions and mating strategies of a solitary carnivorous marsupial Phascogale tapoatafa, in the wild. Wildlife Research, 21, 527 - 542. http: // dx. doi. org / 10.1071 / WR 9940527","Soderquist, T. R. & Lill, A. (1995) Natal dispersal and philopatry in the carnivorous marsupial Phascogale tapoatafa (Dasyuridae). Ethology, 99, 297 - 312. http: // dx. doi. org / 10.1111 / j. 1439 - 0310.1995. tb 00904. x","Dixon, J. M. & Huxley, L. M. (1989) Mammals of Victoria from the collection of Donald F. Thomson. Victorian Naturalist, 106, 4 - 25."]}

46. Common Brush-tailed Phascogale Phascogale tapoatafa French: Grand Phascogale / German: Grof3e Pinselschwanzbeutelmaus / Spanish: Fascogalo de cola de cepillo meridional Other common names: Black-tailed Phascogale, Brush-tailed Phascogale, Common Wambenger, Tuan Taxonomy. Viverra tapoatafa F. A. Meyer, 1793, Sydney, New South Wales, Australia. There were originally two subspecies of P. tapoatafa: nominate tapoatafa from both eastern and south-western Australia and pirata from the Northern Territory and Kimberley (Western Australia). Fieldbased studies and morphological data on eastern and western populations showed that these allopatric populations differed in size, sexual dimorphism, and teat number. There are no clear external differences between eastern and south-western P. tapoatafa, but there are cranio-dental ones. A study in 2001 measured genetic (mtD-NA) differentiation among P. tapoatafa (from south-eastern and south-western Australia), P. calura, and P. pirata (then recognized as the subspecies P. t. pirata from Northern Territory). P. tapoatafa was clearly recovered as monophyletic, with south-eastern and south-western populations forming strongly supported, reciprocally monophyletic clades. P. pirata was monophyletic and poorly resolved assister to P. tapoatafa; P. calura was also monophyletic. Within P. tapoatafa, south-eastern and south-western populations were 8:7-13-9% divergent, south-eastern populations were 0-3-6% divergent, and south-western populations were 0-03-3-6% divergent. On average, 16% sequence divergence (range 12-21-9%) separated P. tapoatafa and P. calura, which supported clear morphological distinctions between the two species. Genetic divergence of P. tapoatafa and P. pirata averaged 10-9% (range 9-4-18-5%), which also corroborated their clear morphological distinction. Based on this combined research, three subspecies of P. tapoatafa were suggested (south-western Western Australia; Kimberley, northern Western Australia; and eastern Australia), but a conservative approach is followed here. Monotypic. Distribution. Australia, patchily distributed in SW & NW Western Australia, NE, E & SE Queensland, NE New South Wales, and Victoria. Presumed extinct in South Australia. Descriptive notes. Head-body 16-26.1 cm (males) and 14.8-22.3 cm (females), tail 17.5-23.4 cm (males) and 1622-6 cm (females); weight 175-311 g (males) and 106— 212 g (females); all measurements from Common Brush-tailed Phascogales in Victoria (phascogales in other areas are 20-30% smaller). Growth of the Common Brush-tailed Phascogale is often inhibited during environmentally poor years (drought), especially among males. The Common Brush-tailed Phascogale is sexually dimorphic for size. It is uniform grizzled gray above and cream-to-white below, and it has large, naked ears and a conspicuous black tail, with a brush of hairs up to 55 mm long. Habitat. Open forest with sparse groundcover, but also a broad variety of habitats from low woodlands and shrublands (mallee) to rainforest. Formerly, Common Brush-tailed Phascogales occurred throughout dry sclerophyll forests and woodlands of temperate and tropical Australia. Food and Feeding. Common Brush-tailed Phascogales are one of the most arboreal dasyurids, but they sometimes feed on the ground. Foraging success is greatest on mature trees, large logs, and dead standing trees. The Common Brush-tailed Phascogale is an agile hunter, spiraling up tree trunks, darting along or underneath major branches and leaping as far as 2 m between trees. Its hindfoot can be rotated an astonishing 180° at the ankle, enabling it to climb upward or downward with relative ease. Most foraging occurs on rough-barked trees where the going is easier than on smooth-barked trees. Hidden prey is captured by tearing away bark with their sturdy front incisors and using their dexterous hands to probe expertly into crevices. Common Brush-tailed Phascogales feed mainly on large insects and spiders, and occasionally small vertebrates and, given the opportunity, chickens. In south-western Western Australia, stomachs of eleven individuals and 45 feces obtained during summer and autumn were studied. The Common Brush-tailed Phascogale was a generalist; its diet consisted predominantly of invertebrates. Nevertheless,it is opportunistic, and scavenging of vertebrates may contribute to the diet. Nectar was also eaten, but its availability is probably patchy. Spider and beetle remains made up the bulk of most feces in both seasons. Ants and orthopterans also frequently appeared in fecal samples. In summer, cockroaches were often eaten, appearing in 13 of 20 fecal samples (65%). Earwigs represented a small proportion of the diet in summer but were a major component in autumn, featuring in 16 of 25 samples (62%). Other invertebrates in feces included orthopterans, ticks, dipterans, moth larvae, fleas, and psyllids (Hemiptera). Stomach analyses corroborated the importance of spiders, beetles, orthopterans, and earwigs in the diet. Another study conducted in Perup Nature Reserve, ¢.270 km south of Perth (Western Australia), investigated distribution of food resources available to Common Brushtailed Phascogales and compared prey biomass on bark vs. in the litter layer. Spiders dominated biomass of bark fauna, but cockroaches, beetles, and ants also were present. Comparatively, litter was dominated by orthopterans; invertebrate biomass was much greater in the litter layer than on tree trunks. Prey items in bark were more abundant in Melaleuca (Myrtaceae) swamps than in surrounding dry sclerophyll woodland. Breeding. Breeding pairs of Common Brush-tailed Phascogales typically mate during the day in a tree hollow. Breeding season extends over a three-week period between mid-May and early July, varying with locality and between years. Gestation is c¢.30 days. Litter size typically equals number of teats, which is eight throughout most of the distribution and 6-9 in south-western Australia. As pregnancy advances, exposed mammary area deepens and a fleshy rim develops, which completely encloses newborns. Despite rapid growth of young, they remain fully protected until more than five weeks old when they barely protrude from the pouch. At c¢.7 weeks old, young are deposited in a nest by their mother. Females return from foraging every few hours to warm and nurse their litters. Hairless and weighing only c.4 g, nestlings can lose heat rapidly on cold nights, becoming torpid within an hour of their mother’s departure. Nest attendance gradually decreases until young are c.20 weeks old; the mother then does not return until dawn, and weaning is complete. As in other dasyurids, mothers only rarely carry their young on their backs and only for short distances near nests. All male Common Brush-tailed Phascogales die after the breeding season. Unlike antechinuses, they show little physical decline during this period; indeed, they continue to forage and groom while roaming several kilometers each night in search of mates. Eventually, weakened by stress-induced illnesses such as gastric ulcers, males will regularly fall prey to owls, Red Fox (Vulpes vulpes), and domestic and feral cats. As lactation progresses, energy demands on mothers ensure they are less wary while foraging and more vulnerable to predators. Although females can live three years in the wild, most only survive to raise one litter. One study of three populations of Common Brush-tailed Phascogales in Victoria found that two-year-old females typically gave birth earlier than first-year females; births were spread, on average, over 15 days, and in some years, they occurred two weeks earlier than average. Females have eight teats, but litters of 1-6 young comprised 29% of 45 litters, with litter size averaging 6-6 young. Sex ratio of litters produced by second-year females was significantly male-biased (1:0-62), but for first-year females, it was close to parity. When juvenilesfirst relinquished their mother’s teats at c¢.48 days of age, they weighed c.4 g. Weight gain by captive juveniles was more rapid than in wild individuals prior to weaning, but skeletal growth rate and morphological development were similar. Researchers found that investment in lactation by the Common Brush-tailed Phascogale was far greater than expected for a dasyurid of its size (wild litters at weaning averaged 313% of maternal weight; captive litters average 370%). High mortality of wild adult female Brush-tailed Phascogales during lactation may reflect this energetic drain. In south-western Australia, late breeding by females in one year suggested that sperm storage may be extensive. During a drought, individuals were in poor condition, which may have influenced birth times and sex ratios in litters. Another study documented physical characteristics of selected nest trees and described spatial and temporal patterns of nest-tree use by radio-tracking individuals. Nine Common Brush-tailed Phascogales (seven females and two males) were radio-tracked in March—July in an area where most woodland habitat was confined to linear strips along roads and streams or small patches and scattered trees in cleared farmland. Female Common Brush-tailed Phascogales were monitored for 13-35 days over periods of 5-15 weeks, and two males were monitored for 2-9 days, respectively. A total of 185 nest-tree fixes was collected; all nests occupied by Common Brush-tailed Phascogales were in standing trees. Eighty-three nest trees were identified, with diameters at breast height of 25-171 cm. In this study, Common Brush-tailed Phascogales did not discriminate among canopy tree species as nest trees but exhibited significant selection for trees in the largestsize class. All individuals used multiple nest trees, with the seven females occupying an average of 11-4 nest trees from a mean number of 25 diurnal locales. Number of nest trees used by Common Brush-tailed Phascogales continued to increase throughout the study, suggesting that even more would be identified during a longer or more intensive study. Occupied nest trees were located throughout each individual’s home range, illustrating the importance of a continuous spatial distribution of suitable nest trees across the landscape. Nesting trees were also found in adjacent farmland, up to 225 m from roadside vegetation, demonstrating the value that scattered clumps and even single trees in farmland can have for conservation of the Common Brush-tailed Phascogale. Activity patterns. Although very little is known about specific activity patterns of Common Brush-tailed Phascogales, they are typically nocturnal and may spend much of a night foraging for nectar in a single heavily flowering eucalypt. Movements, Home range and Social organization. When alarmed, the Common Brush-tailed Phascogale repeatedly taps its hands on tree bark, which purportedly sends a signal to a predator that it has been spotted. While foraging, males rub their chest gland against trees, leaving scent that females keenly investigate. Males also pursue females, but a female can repel a pursuing male by as little as a vocal threat, even during height of breeding. Play among juveniles is limited to brief chases; juveniles explore and learn to hunt alone. Dispersal of mobile young Common Brush-tailed Phascogales occurs in mid-summer. Juvenile males move many kilometers, and females settle in nearby vacant territories or occasionally share their mother’s territory. The Common Brush-tailed Phascogale is solitary, and individuals rarely nest together after dispersal, except during nutritionally stressful years when poor body condition can encourage communal nesting with its associated thermal benefits. Home ranges of females typically are 20-40 ha but can be as small as 5 ha in high-quality habitat. Females will exclude unrelated females from their territory, so their distribution is typically sparse. Male home ranges are very large, often more than 100 ha, overlapping with those of females and other males; males expand their home ranges during the breeding season. An individual can use more than 40 nests in a single year, in a variety of places, including hollow trees, rotted stumps, house ceilings, and bird nests. Lactating females prefer large tree cavities with small, secure entrances; females build spherical nests of bark strips, feathers, and fur. A female may drag bark strips up to one-third of her body weight and construct a nest more than 30 times her own size. Nest hollows often contain an accumulation of pungent black feces that may serve as a territorial marker. Feral honey bees (Apis mellifera) and Sugar Gliders (Petaurus breviceps) sometimes take over nest cavities of Common Brush-tailed Phascogale, providing serious competition where suitable nursery hollows are rare. In one study, nine Common Brush-tailed Phascogales (seven females and two males) were radio-tracked in March—July to investigate spatial organization of a population in Victoria. In this area, only 3:6% of the original woodland vegetation remained after 150 years of agricultural clearing since settlement. Most wooded habitat is now confined to narrow linear strips along roads and streams. These remnants are on fertile soil, and because they have been spared intensive harvesting, density of large, old trees is more than ten times that found in nearby protected areas. Female Common Brush-tailed Phascogales were monitored for 13-38 days during 5-15 weeks. Sizes of home ranges of females were 2-3-8 ha and averaged 5 ha. This value was one-eighth of mean home range size typically recorded for Common Brush-tailed Phascogales in contiguous forest in Victoria. All individuals used multiple nest trees; nests were generally in trees with diameters at breast height greater than 80 cm. The dense population of Common Brush-tailed Phascogales in these remnants suggested that before agricultural clearing and timber harvesting, they were much more common in Victoria than at present. In another study of wild Common Brush-tailed Phascogales in Victoria, play ofjuveniles entailed brief, non-contact chases, which apparently provided social practice prior to the essentially solitary, post-dispersal life of adults. Interactions between adults very rarely included physical contact; most encounters (63%) comprised chases. Wild females readily deterred males from approaching too close by vocal threatening, even during peak breeding, so forced copulation (as seen in the related species of Antechinus) was unlikely. Scent presentation suggested that sternal marking by males was part of intersexual communication; this behavior may serve, along with chases, to familiarize females with future mates. Status and Conservation. Classified as Near Threatened on The IUCN Red List. The Common Brush-tailed Phascogale is in decline at a rate of less than 30% over ten years due to habitat clearing, degradation, and fragmentation and impacts of introduced predators; it is therefore close to qualifying as Vulnerable. Prior to European settlement, distribution of the Common Brush-tailed Phascogale encompassed almost the entire eastern and south-eastern coast of the Australian mainland, south-western Western Australia, and northern Western Australia (Kimberley). Now, distributions of the two populations of Common Brush-tailed Phascogales in Western Australian have retracted markedly. Following European settlement, much of the prime habitat of Common Brush-tailed Phascogales on fertile soil was cleared for agriculture. Remaining habitat in temperate areas is often degraded and fragmented, thus reducing densities of Common Brush-tailed Phascogales; over time, this isolates small populations and impedes genetic exchange. Alarmingly, recent distributional contractions have also occurred in relatively unfragmented habitats of the Kimberley, and numbers can be low in otherwise apparently suitable habitat. This makes the Common Brush-tailed Phascogale vulnerable throughoutits distribution, even though it is perhaps surprisingly not listed federally in Australia. Sparse densities (large home range), fluctuating reproductive success, and annual die-off of males make the Common Brush-tailed Phascogale susceptible to local extinction. Thus, long-term monitoring is required to detect further distributional contractions and more clearly define impacts of habitat fragmentation and degradation, logging, fire, and predators on population viability. The Common Brush-tailed Phascogale is apparently vulnerable to baiting for Red Fox. In one study,field trials were undertaken in box-ironbark woodland at Puckapunyal Military Area in central Victoria to assess bait uptake by Common Brush-tailed Phascogales during simulated baiting exercises for Red Fox (Vulpes vulpes). The marker rhodamine B was used in non-toxic baits to detect non-target bait consumption. Trials demonstrated that free-living Common Brush-tailed Phascogales were capable of accessing non-toxic fox baits buried under 10 cm of sand; rhodamine B markings were detected in six (15%) of 40 Common Brush-tailed Phascogales captured during the study. Reduced predation pressure by red fox can change behavior of Common Brushtailed Phascogales; reduction in Red Fox numbers following baiting may have prompted Common Brush-tailed Phascogales to spend relatively more time foraging on the ground and thus their potential exposure to fox baits was higher. In this study, Common Brush-tailed Phascogales spent up to 40% of their foraging time on the ground during summer, which was much more time spent on the ground than predicted in other research. Bibliography. Burbidge & McKenzie (1978), Covacevich et al. (1994), Cuttle (1982), Fairbridge et al. (2003), Menkhorst, Rhind & Ellis (2008), Meyer (1793), Millis et al. (1999), van der Ree, Bennett & Soderquist (2006), van der Ree, Soderquist & Bennett (2001), Rhind (2002, 2003, 2004), Rhind & Bradley (2002), Rhind et al. (2001), Scarff & Bradley (2006), Scarff et al. (1998), Soderquist (1993a, 1993b, 1995a), Soderquist & Ealey (1994), Soderquist & Lill (1995), Soderquist & Rhind (2008), Spenceret al. (2001), Traill & Coates (1993)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2015, Dasyuridae, pp. 232-348 in Handbook of the Mammals of the World – Volume 5 Monotremes and Marsupials, Barcelona :Lynx Edicions on pages 325-327, DOI: 10.5281/zenodo.6608102

44. Red-tailed Phascogale Phascogale calura French: Petit Phascogale / German: Kleine Pinselschwanzbeutelmaus / Spanish: Fascogalo de cola roja Other common names: Red-tailed \Wambenger Taxonomy. Phascogale calura Gould, 1844, Williams River, Military Station, Western Australia, Australia. The type specimen of P. calura was collected by J. Gilbert at the “Military Station on the Williams River,” 150 km south-east of Perth (Western Australia’s capital) in 1843, the early years of European settlement. J. Gould formally named the species a year later. At the time of capture, Gilbert commented “I was indebted to a domestic cat who had captured it in the night. The soldiers informed me that they had often met with it in the storeroom of the Station, but they could give me no other information respecting it, except that specimens with much larger or bushier tails were sometimes seen.” Subsequent surveys extended the species’ distribution to the junction of the Murray and Darling rivers in eastern Australia by the mid-19" century, to central Australia in the late-19"™ century, and to the Canning Stock Route in Western Australia in 1931. A live specimen was also apparently taken around Adelaide at some time before 1888. Nevertheless, for all localities other than south-western Western Australia, these were the first and last records for the species. In 2001, measurements of genetic differ entiation among P. calura, P. tapoatafa, and P. pirata (then recognized as the subspecies Pt. pirata) were estimated using mtDNA (cytochrome-b gene); 16% sequence divergence separated P. caluraand P. tapoatafa, which supported clear morphological distinctions between the two species. P. caluraand P. pirata averaged 14-6% genetic divergence, which is also corroborated by their clear morphological distinction. Monotypic. Distribution. SW Australia, now entirely confined to the Southern wheat belt region in Western Australia. Descriptive notes. Head-body 10-5-12:2 cm (males) and 9.3-10.5 cm (females), tail 13.4-14.5 cm (males) and 11.9-14.4 cm (females); weight 39-68 g (males) and 38-48 ¢ (females). The Red-tailed Phascogale is ashy gray above and cream to white below; there is a blackish patch in front of eye. Ears and upper proximal part oftail are distinctive reddish brown; distal half oftail has a brush of long black hairs. Habitat. A variety of habitats with greater canopy density and greater abundance of hollows than unoccupied sites. Red-tailed Phascogales are found in isolated patches of forest that receive 300-600 mm of rain/year. They prefer habitats that are generally denser and taller climax vegetation within wandoo (Eucalyptus wandoo, Myrtaceae) and rock sheoak (Allocasuarina huegeliana, Casuarinaceae); hollows in wandoo provide nesting sites. One study found that the Red-tailed Phascogale occurs mostly in woodland habitat with old-growth hollow-producing eucalypts, primarily wandoo or york gum (E. loxophleba). Records from the periphery of its current distribution appear to represent a broader variety of habitats, including shrublands, mosaics of woodland, and scrub-heath. Red-tailed Phascogales occupy areas of remnant vegetation of varying sizes from very small to very large, many on privately owned agricultural land. Large connected areas, such as riverine corridors and stands of upland remnant vegetation, appear important to long-term persistence of the Red-tailed Phascogale; however, sites isolated by increasing distance from other occupied sites tended to be unoccupied by the Red-tailed Phascogale. Food and Feeding. Like its congeners, the Red-tailed Phascogale is arboreal. It can leap up to 2 m, covering considerable distances while foraging in the forest canopy. Red-tailed Phascogales often feed on the ground and are opportunists, taking a wide variety of insects and spiders (preferring prey less than 1 cm in length); they will also eat small birds and mammals (particularly the House Mouse, Mus musculus). Prey satisfies water requirements. One study examined feces from Red-tailed Phascogales that had been translocated to the Alice Springs Desert Park; they were primarily insectivorous, with 92:6% of all feces containing arthropods. They also consumed birds (51:6% of feces), small mammals (33-:3%), plant material (27-4%), and occasionally reptiles; birds were a large proportion of the diet in spring. Breeding. Female Red-tailed Phascogales show an increase in weight prior to estrus, and mating occurs during a three-week period in July. Captive copulations haves lasted up to six hours. Individual females may be receptive to males for at least five days, and births may be tightly synchronized or spread over a couple of weeks. Females are able to store viable sperm for at least four days prior to ovulation; gestation is 28-30 days. Females ovulate c.16 eggs, with up to 13 young observed at birth, despite availability of only eight teats. At least two males can sire young within a single litter. Unlike males, which all die synchronously at the conclusion of breeding, female Red-tailed Phascogales are capable of surviving and reproducing in a second or even third season. At birth, young weigh c.15 mg and are c¢.5 mm long. Fur on young begins to appear at c.34 days, and young first detach from teats while in a nest at c¢.45 days. Weaning of young occurs at ¢.90 days when they weigh c.20 g. In the wild, males live for only 11-5 months (similar to male Antechinus, the sister genus). Captive males can survive for up to five years. A recent study reported successful captive breeding in four-year-old female Red-tailed Phascogale; females can survive until at least five years of age in captivity. These findings have implications for captive breeding programs, providing evidence that older females can be successfully bred. Males are reproductively senescent after their first breeding season. Prior to mating, male germ cells needed for production of sperm are depleted, and sperm production ceases. Sperm stored in the epididymis during the mating period thus sires the next generation. Activity patterns. Red-tailed Phascogales are mainly nocturnal, but they have been seen emerging during the day to investigate potential food sources. Movements, Home range and Social organization. Home ranges of male Red-tailed Phascogales increase about tenfold during the mating period. Home ranges of females remain at c.8 ha. Status and Conservation. Classified as Near Threatened on The IUCN Red List. Listed as endangered in Australia. There are probably fewer than 10,000 mature Red-tailed Phascogales in the wild, and the overall population may be decreasing. Rate of decrease is likely less than 10% over ten years, so it is close to qualifying as Vulnerable under criterion C of the IUCN. Populations of Red-tailed Phascogales apparently fluctuate with rainfall. There are no apparent current population declines, but there were large declines historically. Survival of Red-tailed Phascogales in Western Australia may be related to presence of Gastrolobium and Oxylobium, legume-like plants that produce monosodium fluoracetate (‘1080’), a chemical that can be lethal to introduced species. Native species have evolved a tolerance to the fluoroacetate; their bodies can contain enough of it to poison introduced carnivores, such as the Red Fox (Vulpes vulpes), that pose threats to their survival. Too frequent burning of forests also negatively affects survival of the Red-tailed Phascogale by destroying nesting hollows and dense, protective foliage. Suitable habitat for the Red-tailed Phascogale appears widespread, but it is under threat over the longer term. Increasing dryland salinity in lowland areas, loss of dense stands of rock sheoak in upland areas, and continuing loss of corridors of vegetation along roadsides due to road widening contribute to loss of habitat and habitat connectivity. The Red-tailed Phascogale now occupies less than 1% ofits former distribution. Today, it occurs in parts of the Avon Wheatbelt, Jarrah Forest, Mallee, and Esperance Plains bioregions. Red-tailed Phascogales persist only in areas that have been extensively cleared for agriculture and where remaining bush land is highly fragmented. It does not appear to extend into unfragmented habitat in the Jarrah Forest to the west or Mallee region to the east. Predation by domestic and feral cats and red fox also reduce numbers of Red-tailed Phascogale. Taken together, these factors, particularly fragmentation and lack of suitable nesting hollows, suggest that the long-term persistence of Red-tailed Phascogales in areas beyond the wandoo belt is far from assured. One study on a translocated population found that Red-tailed Phascogales are able to exploit various prey types and therefore are likely candidates to survive a “hard” (without acclimation prior to release) translocation into the wild, provided the site chosen had adequate food supply. Bibliography. Bradley (1997), Bradley et al. (2008), Foster & Taggart (2008), Foster et al. (2006), Friend (2008b), Gould (1844, 1863), Kitchener (1981), Short & Hide (2012), Short et al. (2011), Spencer et al. (2001), Stannard, Borthwick et al. (2013), Stannard, Caton & Old (2010)., Published as part of Russell A. Mittermeier & Don E. Wilson, 2015, Dasyuridae, pp. 232-348 in Handbook of the Mammals of the World – Volume 5 Monotremes and Marsupials, Barcelona :Lynx Edicions on pages 324-325, DOI: 10.5281/zenodo.6608102