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The white-plumed honeyeater (Ptilotula penicillata, formerly Lichenostomus penicillatus ) is a small passerine bird endemic to Australia. The name “penicillata” derives from the latin word “penicillis” meaning brush tip, referring to the white plume across the side of the neck. White-plumed honeyeaters are common around water and are often seen in backyards and suburbs with vegetation cover. The species is probably of Miocene origin, with a fossil fragment found in Riversleigh, Qld appearing very similar to the white-plumed honeyeater leg bone.

Description
Plumages do not differ between sexes. The distinguishing characteristic is a conspicuous white plume across the neck from the throat to the edge of the nape. The top of head and neck are olive, with a yellow eye-ring surrounding a black-brown to olive-brown eye. Cheeks and ear coverts are yellow-olive, with the rear coverts tipped black, creating a short black stripe along the anterior edge of the neck plume. The chin and throat are dull yellowish-olive.

Upperparts are uniform grey-olive and the uppertail coverts have a yellowish tinge. Upperwing coverts are mostly brown with olive to yellow-olive tips or edges, creating a somewhat scalloped appearance. Remiges are dark brown with yellowish edges to secondaries, forming a yellow-olive panel when the wing is folded. Uppertail is olive-brown with yellow-olive outer edges. Underbody is mainly light brown-grey, with pale yellow streaks in the centre of the breast, pale yellow on the upper belly, flanks and undertail coverts, and cream on the lower belly. Underwing coverts are off-white with brown-grey remiges. Undertail is brownish grey.

Feet and legs are pinkish to purplish grey. During breeding season, the bill and gape are black; however outside of breeding the bill obtains an orange-yellow or orange-brown base (only on lower mandible in some individuals) with a yellow gape.

Four subspecies are recognised: penicillatus, leilavalensis, carteri and calconi. P.p.leilavalensis is paler and yellower, carteri is strongly tinged yellow, and calconi yellower. These subspecies intrograde where ranges overlap. In all four races, males are slightly larger than females. P.p.penicillatus is the largest, with males averaging 20.5g and females 18.1g. P.p.leilavalensis and P.p.carteri are similar in size, where males are around 18g and females 16g. P.p.calconi is the smallest, with males around 17.5g and females around 16g. Wing lengths are much larger in penicillatus, and bill lengths differ only slightly with carteri somewhat longer than the other three subspecies.

Juveniles are similar to adults, but with duller colours on the top of head and neck and brighter colours on the eye ring and ear coverts. The plume is shorter and less distinct, often lacking the black border of the ear coverts. Underbody is slightly more brown. The bill is pink or yellowish with a brown tip in young individuals, changing to black with yellow-orange or pinkish base; while the gape is yellow and noticeably puffy looking. In newly fledged birds the feet and legs are a paler pinkish-brown and appear swollen compared to the adult.

Distribution
The species is widely distributed throughout south-eastern Australia (excluding Tasmania), up towards central Australia with patches occurring in central and western Queensland, Northern Territory and Western Australia. They are absent from desert areas but can persist in arid regions where trees and especially standing water are present.

Nominate race P.p.penicillatus occurs throughout south-eastern Australia to the SA gulf and throughout the Murray-Darling basin. P.p.leilavalensis is found from Lake Eyre south to Flinders Ranges in SA, east to Barrier Ranges in western NSW, west to the edges of the Gibson and Great Sandy Deserts, and north to Qld. P.p.carteri occurs in the Pilbara region of WA from Geraldton to the Fortescue Ranges, and east to the Western Deserts; while P.p.calconi is known only from the southern Kimberly region.

The species has undergone range expansion in the last century. It was formerly linked closely to the distribution of River red gum Eucalyptus camaldulensis and a few other riverine species; but has since expanded into coastal areas to become resident in Sydney and Newcastle where they were previously uncommon or seasonal visitors. Banding studies indicate the species is primarily sedentary with 99.8% of recaptures occurring less than 10km from the banding site. Small local movements occur, perhaps in response to environmental conditions.

Habitat
Oligotypic, mainly found in open sclerophyllous woodland, often near water sources such as wetlands, swamps, creeks and dams. They are found extensively in River red gum woodlands but also riparian woodlands dominated by Eucalyptus, Melaleuca or Casuarina. They may also be found in nearby river flats or open forest and woodland.

Diet
The white-plumed honeyeater diet consists mainly of nectar, insects and their products (E.G. Honeydew and lerp), manna, fruit and very occasionally seeds. They may also peck at berries to feed on juice.

Where sufficient standing water occurs, P.penicillata can be observed drinking at least twice per day. In xeric habitats, additional moisture is obtained from food. A study in the Pilbara region of WA found white-plumed honeyeaters must eat around 100 berries to obtain 1mL of water. Needing 5-10mL per day, white-plumed honeyeaters were able to feed on 500-1000 berries in a five hour period to obtain both caloric requirements and sufficient water intake.

Foraging behaviour
White-plumed honeyeaters mainly feed by gleaning leaf surfaces. The tongue contains brush like filaments consisting of about 60 bristles which is capable of mopping up nectar. Arthropod prey is usually taken by gleaning, but some prey is taken on the wing.

In urban areas, they are typically canopy foragers but foraging heights can vary depending on the influence of other resident species. For example, in the presence of aggressive competitors such as fuscous honeyeaters, white-plumed honeyeaters avoid conflict by modifying their foraging behaviour. In one location where ranges of the two species overlapped, white-plumed honeyeaters fed at lower heights in the overlap zone than they did in the riparian zone where fuscous honeyeaters were absent.

Information on diet comes from scats, stomach contents and foraging observations. Stomach contents are heavily biased toward strong tissues from plants and arthropods. In 1980 it was found that manna, honeydew and lerp were extremely important food resources. Other research indicated that insects were only a small part of the diet, when they had previously thought to have been major components. Insects are sometimes taken on the wing, but probably only provide a protein and mineral source as a surplus of energy is available in nectar, lerp and manna; and more energy is spent hawking prey than can be redeemed from the prey.

Social Behaviour
This is a gregarious species, often found in groups of 12 or more individuals during the non breeding season which dissipates during breeding. P.penicillata is considered colonial, with observations of colonies throughout the range. There is some suggestion of cooperative breeding, but if occurring it is uncommon. Communal breeding is likely, with conspecifics grouping to attack predators near nests. However, individual territories within communities are maintained, with territorial song frequency increasing during the breeding season.

Experiments with captive populations indicate hierarchies form based on plumage colour, gape flange characters and voice, but no data from wild populations exists. Observations of aggressive interactions between conspecifics and other species such as Yellow-faced honeyeater (Caligavis chrysops), Willie Wagtail (Rhipidura leucophrys), Red wattlebird (Anthochaera carunculata) and smaller species such as pardalotes (Pardalotus spp.), Mistletoebird (Dicaeum hirundinaceum) and small honeyeaters have been made. There are some observations of attacks on Fuscous honeyeater but the two species tend to avoid each other where occurring sympatrically. In some regions smaller birds are absent due to the aggressive nature of white-plumed honeyeaters and other species of similar size, resulting in the exclusion of these poorer competitors.

Foraging groups may use a “chip-chip” contact call, or a song which is repeated by nearby individuals. Corroborees of up to 12 or more individuals sitting together on a branch have been observed which engage in extensive calling, followed by rapid dispersal.

Reproduction
Females are relatively more ready to breed from late winter through to summer, however males maintain enlarged testes throughout the year. Breeding occurs throughout the range with records in all months, and clutches typically produced 2-3 times per year. Breeding events usually coincide with outbreaks of herbivorous insects.

Nests are small and cup-shaped, 5-6cm diameter and similar depth. These are skilfully woven from grasses, bound with spider’s silk, animal hair or wool; and can include feathers and plant material. The female appears to be responsible for construction but both sexes maintain and make repairs. Nests are usually located among the foliage in crowns of trees or shrubs, and only rarely seen in mistletoe. Sometimes nests are constructed in forks or on branches, and rarely in dead foliage. There is some evidence of site fidelity, and nest trees may be shared with active nests maintained by several other species including wagtails, magpie-lark, woodswallows and other honeyeater species.

Males undertake a song flight display, with a slight climbing and undulating flight above tree tops while singing. At the song’s completion, the male dives quickly into a nearby tree. This is performed throughout the day during breeding season, but less commonly in the early morning. Playback of this song does not induce territorial responses, suggesting the display is sexual in function. Other songs heard commonly after the breeding period begins may also have courtship functions.

The clutch is usually 2-3 eggs, varying from 1-4 overall. The second egg is typically laid within 24 hours of the first. Eggs are approximately 20 x 15cm, weighing 2.45g; oval, smooth, finely grained and may be slightly glossy. Colour varies from white to yellowish, light red or pink. Markings may vary from uniform reddish freckles to purplish or dark reddish-brown spots. Marks are sometimes in greater frequency at the large end. Eggs laid late in season tend to be lighter, and those from inland populations tend to be white with fewer markings.

Details about incubation duties are unknown, although it is thought to be performed only by the female. Eggs are incubated for 10-14 days before hatching. Both sexes feed offspring with females brooding, leaving only when the male arrives to feed chicks. Both parents remove faecal sacs and egg shell fragments. Parents undertake distraction displays when predators approach nests, mainly in the final days before nestlings fledge. Fledging occurs 12-13 days after hatching and juveniles are fed for two or more weeks after leaving nest.

White-plumed honeyeaters are parasitised by pallid and fan-tailed cuckoos, as well as Horsfield’s and shining bronze-cuckoos.

Systematics and Taxonomy
First described by John Gould in 1837, the specimen he examined came from “the interior of New South Wales”. He placed the species in Meliphaga, where it would remain for almost 150 years. In 1975, Schodde split Meliphaga and placed the species in Lichenostomus. With the introduction of molecular studies, it was apparent that the white-plumed honeyeater belonged in a clade within Lichenostomus. This had first been suggested by Mathews in 1915, who placed it with the Yellow-tinted honeyeater in a genus he named Ptilotula; but this was rejected by the Royal Australian Ornithological Union. The Ptilotula clade was finally promoted to genus after new molecular evidence was published in 2011.

Conservation Status
WPHE is listed as Least Concern.

Threats
Exotic vertebrate predators such as cats and dogs are a direct threat to white-plumed honeyeater populations. Loss of trees from watercourses will impact a population, however they are able to adapt to suburban environments, especially where native gardens are present.

Climate effects
WPHE body size has been observed to change over recent decades, with indivudals becoming smaller in extended drought, but an overall trend to larger body size since the 1960s. This is correlated with increasing temperatures in the regions surveyed, with body size increasing by 0.064% per year.