Phascogale tapoatafa (F. A. Meyer, 1793)

Russell A. Mittermeier & Don E. Wilson, 2015, Dasyuridae, Handbook of the Mammals of the World – Volume 5 Monotremes and Marsupials, Barcelona: Lynx Edicions, pp. 232-348 : 325-327

46.

Common Brush-tailed Phascogale

Phascogale tapoatafa

French: Grand Phascogale / German: Grof 3e 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).

Figures

On following pages: 41. Short-furred Dasyure (Murexia longicaudata); 42. Broad-striped Dasyure (Murexia rothschildi); 43. Long-nosed Dasyure (Murexia naso); 44. Red-tailed Phascogale (Phascogale calura); 45. Northern Brush-tailed Phascogale (Phascogale pirata); 46. Common Brush-tailed Phascogale (Phascogale tapoatafa); 47. Giles's Planigale (Planigale giles); 48. Long-tailed Planigale (Planigale ingrami); 49. Common Planigale (Planigale maculata); 50. Papuan Planigale (Planigale novaeguineae); 51. Narrow-nosed Planigale (Planigale tenuirostris); 52. Wongai Ningaui (Ningauiride); 53. Pilbara Ningaui (Ningaui timealeyi); 54. Southern Ningaui (Ningaui yvonneae); 55. Kultarr (Antechinomys laniger).

Distribution. Australia, patchily distributed in SW & NW Western Australia, NE, E & SE Queensland, NE New South Wales, and Victoria. Presumed extinct in South Australia.