Quail, Buttonquail and Plains-wanderer in Australia and New Zealand

By Joseph M Forshaw and Frank Knight

Although not closely related, quail, buttonquail and the Plains-wanderer have much in common. Quail, Buttonquail and Plains-wanderer in Australia and New Zealand examines 14 species of these small, secretive ground-dwelling birds, including Old World and New World quail, the endangered Buff-breasted Buttonquail, the elusive Plains-wanderer and the extinct New Zealand Quail.

Joseph Forshaw presents a comprehensive review of recent studies for these often hard to observe birds. Detailed species descriptions include key features, habitat, status, diet and breeding, along with information on eggs, calls and distribution. Each species is fully illustrated with exquisite colour identification plates by renowned wildlife artist Frank Knight. This is an essential reference for anyone fascinated by these elusive birds.

Certificate of Commendation, The Royal Zoological Society of NSW 2023 Whitley Awards: Highly Commended

• Language: English
• Publisher: CSIRO PUBLISHING
• Release date: May 1, 2023
• ISBN: 9781486312610

Joseph M Forshaw

Joseph M. Forshaw AM is a Research Associate in the Ornithology Section at the Australian Museum and the author of a number of award-winning natural history books. In 2015, he was made a Member of the Order of Australia for services to ornithology and wildlife conservation. His previous books include Vanished and Vanishing Parrots (CSIRO Publishing, 2017) and Pigeons and Doves in Australia (CSIRO Publishing, 2015).

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INTRODUCTION

It is similarities in lifestyle, not taxonomic relationships, that link together the species covered in this book. Apart from the two introduced New World quail, all are exclusively terrestrial, and their cryptic plumage coloration, together with shy, elusive habits, developed to afford protection, have been responsible for longstanding neglect by field researchers. Fortunately, this is changing and some species, especially those that are threatened, are being studied intensely. Also, unexpected taxonomic findings have resulted from molecular analyses, so resolving relationship uncertainties that had persisted for a very long time. Taxonomically the species may be divided into two major groups that are quite distantly related in that they are in different orders, and then each group may in turn be divided again into two groups that are in different families.

The so-called gallinaceous birds belonging to Galliformes are a worldwide assemblage of approximately 250 predominantly ground-frequenting species, among which are some of the most familiar species, especially the Gallus junglefowls, which give their name to the Order, and the Meleagris turkeys, from which are derived our domesticated chickens and turkeys. Included also are species, particularly pheasants, grouse and quail, commonly referred to as game birds because they are important quarry for hunters and often are reared commercially on farms specifically for release on hunting estates or reserves.

Cracraft (2013) points out that within Galliformes the identity of major groups and their overall interrelationships are moderately well known, having received extensive attention from investigators using morphological data in which there has been large taxon sampling. Although with more restricted taxon sampling, molecular studies have, in general, achieved results congruent with most morphological analyses at higher taxonomic levels. Cracraft recognises five families within Galliformes:

Order Galliformes

Families

Megapodidae (megapodes)

Cracidae (guans, currasows)

Numididae (guineafowl)

Odontophoridae (New World quail)

Phasianidae (pheasants, partridges, Old World quail)

This listing also adopts a phyletic sequencing which implies a set of hierarchical relationships, with each family being a sister-group to all subsequent families, so identifying Megapodidae as being the sister-group of all four families below it, Cracidae being the sister-group of the three families below it, and so on. New World quail in Odontophoridae and Old World quail in Phasianidae occur in Australia and New Zealand.

Apart from two aberrant African galliform species – Stone Partridge Ptilopachus petrosus and Nahan’s Francolin P. nahani, which are differentiated in a separate subfamily, all members of Odontphoridae are grouped together in the other subfamily – Odontphorinae and, being confined to the Americas, are collectively identified as New World quail. Probably the best known of the New World quail – the California Quail Callipepla californica has been introduced successfully as a game bird to many countries, including New Zealand and Australia, though in Australia it survives only on Norfolk Island and on King Island, in Bass Strait. Introduction to New Zealand of Northern Bobwhite Colinus virginianus, another popular target species for hunters, appears to have been unsuccessful, and occasional sightings of wild birds usually are attributed to escaped cagebirds or birds deliberately released from game bird farms.

Distributed worldwide, though with strongest representation in the Old World, especially in South-east Asia and Africa, and including some 155 species, Phasianidae is by far the largest family of gallinaceous birds. Many of the included species of pheasants, peafowl, junglefowls, partridges and Old World quail are well known as popular aviary birds or as game birds for hunters, and some species, particularly the Common Pheasant Phasianus colchicus and Chukar Partridge Alectoris chukar have been introduced to many countries. There are small feral populations of Common Pheasants in Australia, and in New Zealand the well-established feral population is regularly augmented by deliberate releases from game bird farms (Menkhorst et al. 2017; Heather and Robertson 2015). Uncertainties about interrelationships within the family have been reflected in differing taxonomic arrangements, and Cracraft points out that the substantial diversity eventually will need finer taxonomic recognition. Acceptance of two subfamilies is adopted quite commonly, but not so the composition of each family. The two subfamilies recognised by McGowan (1994) are Perdicinae (partridges and allies, including Old World quail) and Phasianinae (pheasants, junglefowl, peafowl). Cracraft recognises Rollulinae (Roulroul and allied partridges) and Phasianinae (all remaining species), and del Hoyo and Collar (2014) adopt a similar arrangement.

Very different from New World quail, both morphologically and behaviourally, Old World quail are represented in Australia by three native species, though only the Stubble Quail Coturnix pectoralis is endemic. The extinct New Zealand Quail Coturnix novaezelandiae was very similar to the Stubble Quail, and at times the two have been differentiated as separate species or as subspecies of C. novaezelandiae, but molecular analyses show that they are indeed separate species that diverged approximately five million years ago (Seabrook-Davison et al. 2009). The Brown Quail Synoicus ypsilophorus occurs also in New Guinea and the Sunda Islands, Indonesia, and has been introduced successfully to New Zealand and Fiji. The King Quail Synoicus chinensis is widely distributed from the Indian subcontinent and southern China through South-east Asia and the Philippines to the Indonesian Archipelago and New Guinea.

Although quail-like in appearance and habits, buttonquail and the highly distinctive Plains-wanderer Pedionomus torquatus are not related to Old World quail, but are included with shorebirds and their relatives in Charadriiformes, a cosmopolitan, diverse order that includes more than 300 species, which are waterbirds or birds that clearly are derived from waterbirds. Bock and McEvey (1969) pointed out that although the Plains-wanderer had been known to science for more than 100 years, its morphology had been little studied and its exact affinity with other birds remained disputed. The only anatomical study was a short contribution by Gadow (1891), who concluded that it should be placed with the buttonquail in a suborder of Gruiformes, and that arrangement was widely accepted, including by Peters (1934), who placed the buttonquail and the Plains-wanderer in separate families within the Suborder Turnices in the Order Gruiformes. Uncertainty was raised again when results of DNA-DNA hybridisation studies indicated that buttonquails seem to have no close living relatives (Sibley et al. 1988). One arrangement adopted to reflect that uncertainty was to treat buttonquail in their own order – Turniciformes incertae sedis (in Marchant and Higgins 1993).

In their osteological study, Bock and McEvey concluded that the skeleton of the Plains-wanderer was quite different from skeletons of the buttonquail, but it was not possible to determine the level of taxonomic distinctiveness so the general practice of placing the Plains-wanderer in a separate family should be maintained. No comment was made on the inclusion of both families in Gruiformes, so presumably that was not addressed. In another osteological examination, Olson and Steadman (1981) found that the Plains-wanderer is a typical member of Charadriiformes, but without significant characters in common with buttonquail, and it shares most similarities with the Neotropical seedsnipes of the family Thinocoridae, tending to bridge the gap between the aberrant seedsnipes and the more typical Charadriiformes, but being sufficiently distinct to merit its own family. Placement within Charadriiformes was subsequently supported by DNA-DNA hybridisation studies (Sibley and Ahlquist 1990).

Cracraft (2013) points out that relationships within Charadriiformes have received substantial study in recent years and, although morphological data are substantially incongruent, their interrelationships now are well resolved by molecular data. Three suborders – Charadrii, Scolopaci and Lari, can be recognised, with the first being the sister of the other two:

Order Charadriiformes

Suborder Charadrii (thick-knees, oystercatchers, stilts

 and avocets, plovers and lapwings)

Suborder Scolopaci

Parvorder Jacanida

Superfamily Thincoroidea

 Family Pedionomidae (Plains-wanderer)

 Family Thinocoridae (seedsnipes)

Superfamily Jacanoidea

 Family Rostratulidae (painted-snipes)

 Family Jacanidae (jacanas)

Parvorder Scolopacida

Superfamily Scolopacoidea

 Family Scolopacidae (waders, snipe, woodcocks, phalaropes)

Suborder Lari

Parvorder Turnicida

 Family Turnicidae (buttonquail)

Parvorder Larida

Superfamily Glareoloidea

 Family Dromadidae (Crab-Plover)

 Family Glareolidae (coursers and pratincoles)

Superfamily Alcoidea

 Family Alcidae (auks)

 Family Stercorariidae (skuas)

Superfamily Laroidea

 Family Laridae (gulls and terns)

Commenting further, Cracraft points out that finding buttonquail in the family Turnicidae to be the sister group of other families in Lari was one of the more surprising recent systematic discoveries and, now well supported by independent datasets, this finding has resolved the longstanding uncertainties about their relationships. Accepted also is the relationship between the Plains-wanderer and the Neotropical seedsnipes, though in separate families.

STATUS AND CONSERVATION

Introduction to New Zealand of two species of New World quail met with quite different outcomes. Multiple introductions of California Quail as game birds were carried out mostly by acclimatisation societies between 1860 and 1945. Williams (1952) attributed successful establishment of the species to these societies, but noted that a lack of adequate records in their annual reports hindered any attempt to document subsequent spread of the populations. Importation of Northern Bobwhite, also as game birds, is better documented, though there are slight differences in the two published accounts. Gurr (1953) noted that the first introduction was in 1898, when the Wellington Acclimatisation Society imported 400 birds. These and another 750 birds imported by the same society in 1899 were distributed throughout the country from Auckland to Southland. Westerskov (1957) does not mention an importation in 1899, and reported that five shipments arriving between 1899 and 1902 were supplied by Charles Payne, an American dealer in game birds resident in Wichita, Kansas. He noted also that the imported birds were liberated in many localities over a vast range from lat. 36°S in Northland to lat. 47°S on Stewart Island.

Despite the widespread presence of introduced mammalian predators, California Quail have thrived in New Zealand and, apart from Southland and West Coast, including the Southern Alps, in South Island and the Chatham Islands, they are abundant in favoured habitats. Comprehensive ecological studies were undertaken in the late 1940s by Gordon Williams and, although conceding that numbers had declined since their introductions, he stressed that the role of predation, especially following the introduction of mustelids, has been overestimated, and quality of habitat is more important in regulating numbers than is predation, hunting or disease (Williams 1952).

All available evidence indicates that New Zealand was the source of California Quail introduced to Lord Howe Island, Norfolk Island and Australia. Hindwood (1940) reported information provided by Mr WS Thompson, then one of the oldest residents on Lord Howe Island that these quail were introduced to the island by Captain Armstrong, who was the then resident magistrate. They increased in numbers and fairly large flocks were observed until ‘the coming of rats in 1918’, and then they were soon exterminated. Conversely, on nearby Norfolk Island, their introduction in 1895 or 1896 was successful, and Hindwood mentions the claim that this introduction was undertaken by a person connected with the Melanesian Mission. During frequent visits to Norfolk Island during the 1970s and 1980s, I found these quail to be one of the most conspicuous birds, being plentiful in all open areas. I suggest that an ecological study on Norfolk Island, where rats and feral cats are present, would be a worthwhile project for researchers telling us why on this island, as in New Zealand but not on Lord Howe Island, predation apparently has had less adverse impact on the population.

In his summary of introductions of California Quail to Australia, Long (1981) reports that birds were brought from New Zealand to Victoria and Tasmania between 1863 and 1874, and it was suspected that more than 260 birds may have been liberated in Victoria in 1873–1874. They thrived briefly and breeding was recorded, but all soon disappeared, and introduction attempts elsewhere in south-eastern Australia, occurring as late as 1944, were unsuccessful. Only on King Island in Bass Strait, Tasmania, where they were introduced to replace declining Brown Quail Synoicus ypsilophorus as a game species, did California Quail become established and, although common in the past, they now are encountered only in small, scattered parties and predominantly in the north of the island.

In marked contrast to widespread establishment of California Quail in New Zealand, introduction of Northern Bobwhite was not successful, and in most districts they disappeared within about 10 years. Gurr (1953) noted that after this time there were no records until 1923, when a number were caught at a coastal locality on the North Island. Up to 1949 there were reports of flocks of 10 to 20 birds being found in the Hawkes Bay district, North Island, where they were shot as game birds and appeared to be ‘holding their own’ (McKenzie 1953). Heather and Robertson (2015) list local occurrences in the 1950s as the most recent records, and it seems that later records are more likely to be birds deliberately released from captivity rather than remnant wild populations originating from the initial introductions.

As with most birds, it has been loss and degradation of habitat that has impacted adversely on populations of quail, buttonquail and the Plains-wanderer in Australia, especially in coastal and near coastal regions where urbanisation and agricultural developments have been concentrated. Williams and Morgan point out that following European settlement so extensive was clearing of native grasslands, the favoured habitat for many species, that in some regions less than 2 per cent of the original area now remains (in Antos and Schultz 2020). Strongly elusive habits of King Quail Synoicus chinensis hinders assessment of their status, and they may be more common than indicated by records, but there has been severe loss of their favoured moist habitats, particularly along the east coast where there is expanding urbanisation and widespread drainage of wetlands. Stubble Quail Coturnix pectoralis and, to a lesser extent, Brown Quail Synoicus ypsilophorus, have successfully colonised croplands, taking advantage of the seasonal abundance of fallen grain, together with farmlands and pastures, and both are invariably listed as being widespread and common. However, I am not aware of any comprehensive assessment of their status since the 1960s, when the Stubble Quail was the subject of long-term research.

It is not surprising that early European colonists in both Australia and New Zealand would view quail that they encountered primarily as game birds because of the close resemblance to the birds widely known in Europe as prized quarry for hunters. Consequently, we find in early published accounts, even from such eminent authors as John Gould and Walter Buller, much emphasis on their desirability as ‘table birds’, and frequent mention is made of the ease with which ‘many brace can be taken with dogs in a days hunting’. In New Zealand, where the New Zealand Quail Coturnix novaezelandiae had to contend with widespread loss of habitat as native grasslands were cultivated or subjected to livestock grazing, the introduction of mammalian predators, particularly mustelids, coupled with hunting undoubtedly contributed to their eventual extermination. In Australia, more specifically in the south-east, quail hunting was still widespread and intense in the early 1900s, and the poorly known buttonquail and Plains-wanderer Pedionomus torquatus were not excluded as quarry. Campbell (1901) referred to controversy in Victoria about timing of the ‘shooting season’, when claims were being made that Stubble Quail Coturnix pectoralis were still nesting when the season opened on 1 March, and of the Painted Buttonquail Turnix varius he wrote:

‘Old Bushman’ says that although you may occasionally kill an odd one during the winter in the Mordialloc district of Victoria, the majority of them come in September and leave in March.

The Painted Quail runs much upon the ground, and when on the wing has a wavering flight; it is therefore somewhat difficult to shoot.

North (1913) referred to this same conflict between sportsmen and ‘bird lovers’ in New South Wales when quoting from the Evening News, Sydney, 9 August 1904:

The wholesale destruction of Quail which is at present taking place in the north-western portion of the State, has awakened the susceptibilities of even the Quail shooters. According to one of these sportsmen, who has just returned from Narrabri with a bag of birds which he modestly estimated as not exceeding two hundred, the close season ought to be at once proclaimed. The birds, he states are not merely nesting, but are actually laying, and the havoc that is being wrought in their midst by the gun must seriously affect the preservation of this desirable game. At the present time shooting parties in the Moree district not infrequently return with bags running well into four figures, and the close season does not begin until the 15th September. Conscientious, as distinct from reckless sportsmen, are at one in urging in the present instance the immediate proclamation of the close season.

It was not until the 1960s that a comprehensive field study of a hunted species was undertaken. In 1966, the Fauna Protection Panel, later replaced by the National Parks and Wildlife Service, of New South Wales, where at the time quail were not game species, considered the reintroduction of open seasons, and to provide data to assist in management of populations of Stubble Quail, the most desired quarry species, long-term research was undertaken by the then Division of Wildlife Research in the Commonwealth Scientific and Industrial Research Organisation (CSIRO). That research program provided much of what we know about Stubble Quail (Frith et al. 1977).

Personally, I dislike the killing of native wildlife, whether it be quail or waterfowl, for sport, largely from a welfare aspect and because of the incidental killing of non-target species. Hunting quail does pose some risk to buttonquail and the endangered Plains-wanderer. However, from a management point of view I do not oppose it provided adequate safeguards are in place to eliminate, or at least minimise the risk to non-target species and, more importantly, that the necessary monitoring of recruitment levels is in place to ensure that harvesting is sustainable. Moloney and Hampton (2020) report that in Victoria the annual average harvest of Stubble Quail by hunters for the years 2009–2019 was 172 562 birds. I was surprised to learn that so many birds are killed each year in that state, and it raises the need for an independent comprehensive survey of Stubble Quail and Brown Quail, the two species categorised as game birds in some states. It is some 60 years since the ecological study of Stubble Quail in south-eastern Australia was undertaken by the CSIRO Division of Wildlife Research, so a new study would be warranted, with particular emphasis on the sustainability of present levels of harvesting by hunters. Effective non-lethal survey methods now are available, and would provide much needed data on distribution, regional status and breeding of targeted species. There is sufficient evidence to show that, as a secondary pressure, hunting contributed to extirpation of the New Zealand Quail, and it has been implicated in the dramatic decline of Stubble Quail in Tasmania, so we cannot be complacent about its potential impact on targeted species in states where licenced hunting is permitted.

Buttonquail are small to midsized birds very closely resembling unrelated Coturnix and Synoicus quail, so in Australia where buttonquail are far more prevalent than elsewhere, the two groups can be confused. Of the 17 species, seven occur in Australia, with all but one being endemic. Absence of a hind toe and reverse sexual dimorphism are characteristic external features, the latter being indicative of their normally polyandrous breeding behaviour. Widely distributed from the eastern Indonesian Archipelago and New Guinea to the eastern Solomon Islands and northern and eastern Australia, the Red-backed Buttonquail Turnix maculosus favours the same coastal and near coastal moist habitats as frequented by the King Quail Synoicus chinensis in Australia, and the two species can be found together. Also like King Quail, they are small, elusive and reluctant to take flight, so are easily overlooked and may be more common than indicated by records, but the same widespread urbanisation and drainage of wetlands undoubtedly has impacted adversely on populations, especially in the south-east.

Of the endemic buttonquail, most concern is held for the Buff-breasted Buttonquail Turnix olivii, which arguably is the least known of Australian birds. It is known only from the type specimen collected at an imprecise locality in the vicinity of Cooktown, north Queensland, in 1894 and seven specimens, together with clutches of eggs, collected farther north, in the Coen district, in 1922–1924, and there have been no subsequent confirmed records. In recent years, sightings have been reported from various localities in far north Queensland from eastern Cape York Peninsula south to the Atherton Tableland and the Charters Towers district, but all are without photographic confirmation. Furthermore, no road-killed specimens have been found and prolonged intensive searching in localities where sightings have been reported have located only Painted Buttonquail T. varius, so raising the very strong likelihood of misidentification (see Webster 2022). In conclusion, I can say only that, at the time of writing, status of the Buff-breasted Buttonquail remains unknown, and we have no irrefutable evidence of its continued occurrence.

Differing from other species in being rainforest specialists, Black-breasted Buttonquail T. melanogaster are endangered, and this has attracted research funding, so field studies being undertaken in south-eastern Queensland are providing much new information on status, habits and nesting behaviour. In the National Recovery Plan for the Black-breasted Buttonquail published in 2009, habitat loss and degradation are identified as the principal threat to existing populations, with predation by feral animals, notably wild dogs, cats and foxes, as a likely additional threat. There is some evidence, mostly unconfirmed, that these buttonquail formerly were much more widely distributed, but widespread land clearance for agriculture and urban development has resulted in severe fragmentation of their habitat with sporadic dispersal of surviving populations being confined almost entirely to south-eastern Queensland.

Variously treated as a separate species or, as I have done, as a subspecies of the Painted Buttonquail Turnix varius, the extinct New Caledonian Buttonquail T. v. novaecaledoniae is known only from the type specimen collected before 1889 and subfossil remains collected from surface deposits at a number of cave sites. Restricted to the Houtman Abrolhos Islands, Western Australia, T. v. scintillans is listed as endangered under Western Australian legislation, and seems already to have disappeared from North Island because of heavy grazing and trampling of vegetation by introduced Tammar Wallabies Macropus eugenii. Conversely, these buttonquail are locally common in their extensive range on the mainland, where they are closely associated with dry open forests or woodlands with a sparse, shrubby understorey and a groundcover of native grasses with a dense leaf litter, and recent surveys have resulted in an expansion of their known range in north Queensland and south-eastern Western Australia.

Probably because they range across northern Australia and are encountered less frequently by field observers, Chestnut-backed Buttonquail Turnix castanotus remain rather poorly known, but appear to be locally common in savanna woodland with a sparse understorey and much grass cover, typically occurring on flat or undulating country with a rocky or gravelly substrate. Again, recent surveys have resulted in an eastward expansion of their known range into far north-western Queensland, and have drawn attention to overlooked possible earlier records in the same region.

The superficially similar Little Buttonquail Turnix velox and the Red-chested Buttonquail T. pyrrhothorax are the most widely distributed species, and can be difficult to differentiate in the field. Both undertake significant movements in response to rainfall, especially Little Buttonquail which are highly nomadic and can appear in great numbers throughout the arid interior when unusually good rainfall results in luxuriant growth of tall grasses. At such times, when they become very numerous, some birds may come to coastal districts. Also in some districts, their movements may become more complex, and they may be considered residents or regular seasonal migrants. In the wheatbelt of south-western Western Australia, Little Buttonquail are listed as both migratory and nomadic, but in these agricultural districts an increased ease of observing birds when flushed during harvesting operations can result in a majority of records being made when harvesting takes place. I suspect that such changes in the ease of observation could be responsible for local reports of migration or nomadism. Movements undertaken by Red-chested Buttonquail appear to be more irruptive rather than nomadic. Field studies undertaken in the Mudgee district, central-eastern New South Wales, strongly suggest that occasional significant increases in numbers are the response of resident populations to favourable conditions rather than a result of long-distance movements.

A highly specialised endemic species with a strong dependence on native grasslands, the critically endangered Plains-wanderer has experienced an extraordinarily high loss and degradation of habitat, bringing it to the brink of extinction. Small populations survive in a few scattered expanses of preferred habitat, of which only two or three could be considered as strongholds, but even in these districts the decline continues. Fortunately, it has been, and continues to be well studied, and a National Recovery Plan has been adopted, with a National Recovery Team set up to oversee implementation of the Plan. Success has been achieved with a captive-breeding program, the objective being to release captive-bred birds to augment wild populations, and having a captive population also provides excellent opportunities to observe and document habits and behavioural characteristics of these normally, shy, elusive birds. Like buttonquail, the Plains-wanderer shows reverse sexual dimorphism, which again reflects a normally polyandrous breeding behaviour. Recruitment is influenced very much by climatic conditions, and breeding may cease altogether during prolonged periods of drought or at times of excessive rainfall.

Order GALLIFORMES

Included among the so-called gallinaceous birds belonging to this worldwide assemblage of predominantly ground-frequenting birds are some of the most familiar species, especially the Gallus junglefowls, which give their name to the Order, and the Meleagris turkeys, from which are derived our domesticated chickens and turkeys. Only species in two families – Odontophoridae and Phasianidae, are treated in this book.

Family ODONTOPHORIDAE

Apart from two aberrant African galliform species – Stone Partridge Ptilopachus petrosus and Nahan’s Francolin P. nahani, all members of this family are confined to the Americas. Because it was suggested that African Ptiopachus diverged some 374 million years ago in the Oligocene it has been differentiated in the subfamily Ptilopachinae, leaving all remaining species in Odontophorinae (Bowie et al. 2013).

Subfamily ODONTOPHORINAE

Belonging to this exclusively New World subfamily, the New World quail range naturally from northern United States south to north-eastern Argentina, with one familiar species – the California Quail Callipepla californica, having been introduced successfully at both northern and southern extremities in southern Canada and central Chile. It also is the species successfully introduced to our region.

New World quail bear little semblance to the more familiar Old World quail, being significantly larger, more robust birds with a short neck, rather long stout legs and strong feet capable of perching. In all but the three long-tailed tree-quails, the tail is short and squarish. Coronal ornamentation is present in some species, and varies from short crests to elongated ‘tear-drop’ feathers. Predominantly featuring brown, grey and white, the plumage coloration often is spectacularly mottled or scaled, offering cryptic camouflage among groundcover vegetation. Sexual dichromatism is pronounced in most species, females either lacking strong markings on the head and breast or having a more subdued overall pattern. They frequent a wide range of habitats from tropical rainforests to arid scrublands, with strongest representation being in the forests of Central and northern South America. Although they nest and mostly feed on the ground, they regularly roost at night or rest during the day among branches of trees and shrubs, at times at considerable height. Typically they lay large clutches of eggs, which hatch synchronously and, within a day of hatching, chicks leave the nest to be cared for by both parents.

Genus Callipepla Wagler

Callipepla Wagler, Isis von Oken, 1832, col. 277. Type, by monotypy, Callipepla strenua Wagler = Ortyx squamatus Vigors.

Plumage patterns of crested species belonging to this genus feature well-defined facial markings, strongly streaked flanks and prominently scaled or coloured underparts. Crests vary from short, erectile feathers to elongated ‘tear-drop’ plumes. Females and juveniles are duller than males and the crests are less prominent. The natural range is from north-western United States to central Mexico, and one species has been introduced successfully to our region.

CALIFORNIA QUAIL

Callipepla californica (Shaw)

Tetrao californicus Shaw, Nat. Misc., 9, 1798, pl. 345 and text. (California = Monterey.)

INTRODUCED

DESCRIPTION Length 25 cm.

ADULT MALE (ANWC B45958) Lores to forecrown white strongly streaked black; centre of crown black with black forward curving plumes; occiput to nape rufous-brown; white line from above eyes to sides of nape; feathers of hindneck and sides of neck white margined black to produce boldly mottled neck-collar; mantle grey, feathers finely margined black to produce vermiculated appearance; back to upper tail-coverts grey, feathers indistinctly margined greyish-white; primaries and primary-coverts dark brown; secondaries dark olive-brown tipped pale buff and on outer margins vermiculated dark brown; secondary-coverts olive-brown mottled dark and pale brown along margins; lesser and median coverts olive-brown faintly vermiculated and tipped pale brown; tertials olive-brown broadly margined cream-buff on inner webs to form bold stripes along folded wing; base of bill to behind eye to chin, throat and foreneck black bordered by U-shaped white band; breast bluish-grey; abdomen white, centrally suffused orange-buff and rufous-brown with all feathers margined black to give strongly scaled appearance; flanks and lower abdomen greyish-brown, feathers centrally streaked white; under tail-coverts cream-white, feathers centrally streaked brown; underwing-coverts grey; bill black; iris dark brown; legs grey-brown.

3 specimens (Norfolk and King Island, in ANWC and QVM collections): wing 109 mm, 111 mm, 111 mm; tail 76 mm, 79 mm, 81 mm; bill 11 mm, 11 mm, 13 mm; tarsus 24 mm, 27 mm, 27 mm; weight 174 g, 174 g, 175 g.

New Zealand (Williams 1952): 38 males: wing 103–117 (110.5) mm, exp. cul. 11.0–14.7 (13.4) mm, tarsus 31.0–40.0 (34.9) mm, weight 146–211 (183.8) g.

ADULT FEMALE (ANWC B45957) Overall duller than male and lacking black and white facial pattern; lores and forehead grey indistinctly streaked darker grey; crown to nape dull brown with much shorter black plumes; indistinct off-white line from behind eyes to sides of nape; hindneck and sides of neck pale greyish-white, feathers narrowly margined pale grey to produce indistinctly mottled neck-collar; mantle dark grey, feathers finely margined grey-black to produce indistinctly vermiculated appearance; back to upper tail-coverts grey without greyish-white margins to feathers; base of bill to cheeks, chin and throat off-white with dark grey margins to feathers giving streaked appearance; no orange-buff or rufous-brown suffusion on abdomen; bill dark grey-brown; iris mid-brown.

3 specimens (Norfolk and King Islands, in ANWC and QVM collections): wing 104 mm, 110 mm, 111 mm; tail 75 mm, 77 mm, 80 mm; bill 10 mm, 11 mm, 12 mm; tarsus 24 mm, 26 mm, 26 mm; weight 168 g, 182 g, 196 g.

New Zealand (Williams 1952): 32 females: wing 103–115 (109.0) mm, exp. cul. 11.0–14.2 (12.9) mm, tarsus 31.1–38.0 (34.8) mm, weight 150–209 (173.5) g.

JUVENILES (QVM 1968/2/115) Like adult female, but with much shorter, more brownish coronal plumes; crown and neck light brown streaked darker; no mottled neck-collar; scapulars brown, blotched darker and strongly streaked white; flight feathers, except outermost primaries, dark brownish-grey, on inner webs mottled pale brown and tipped white.

ANWC B45953 adult ♂

Browns Road, King Island, Tasmania, 1 December 1993.

ANWC B45957 adult ♀

Tin Mine Road, King Island, Tasmania, 1 December 1993.

QVM 1968/2/115 juvenile unsexed

Forestry Reserve, King Island, Tasmania, 7 March 1968.

downy chick (photographs)

DOWNY CHICKS (PHOTOGRAPHS) Head rich cinnamon-yellow with dark brown patch behind eye and chocolate-brown patch on crown; from hindcrown to mantle broad black band bordered white; upperparts buff-brown with broad black dorsal band continuing down from mantle and two narrower lateral black stripes, all bordered white; underparts buff-white with black blotches on thighs; bill pink; iris darkest brown; legs pink–brown.

DISTRIBUTION Feral populations established in New Zealand, on Norfolk Island, and on King Island in Bass Strait, Tasmania. Native to western North America, from south-western Oregon, north-western United States, south to Baja California, north-western Mexico.

SUBSPECIES

1. C. c. californica (Shaw)

The nominate subspecies occurs naturally in western North America, from south-eastern Oregon south through California, United States, to northern Baja California, north-western Mexico. Introduced to New Zealand.

2. C. c. brunnescens (Ridgway 1884)

Lophortyx californicus brunnescens Ridgway, Proc. Biol. Soc. Washington, 2, 1884, p. 94. (Santa Barbara, California, error = San Francisco.)

ADULTS Like californica, but with darker plumage coloration, upperparts being deep olive-brown; tertials broadly margined deeper ochraceous on inner margins; darker shaft-streaking to feathers on flanks.

Occurs naturally along the coastal plain in western United States, from south-western Oregon to central California. Introduced to New Zealand, Norfolk Island, and King Island, Tasmania.

It may no longer be possible to subspecifically identify feral populations, even though Heather and Robertson (2015) nominate brunnescens as the subspecies introduced to New Zealand, and consequently to Norfolk and King Islands. In his New Zealand Birds, Oliver (1955) titled his species text as ‘Californian Quail Lophortyx californica brunnescens’ but noted that both californica and brunnescens had been identified from New Zealand. Williams (1952) reported that 20 specimens were forwarded to the Museum of Vertebrate Zoology, Berkeley, California, for identification, and the response was that, rather than suggesting the browner brunnescens, the birds matched ‘variants only from the drier sections of the range of C. c. californica, such as the interior of Sacramento Valley and the mountain ranges of southern California near Los Angeles’. If introduced birds were of both subspecies, present populations could be expected to be quite variable, exhibiting characteristics of californica, brunnescens and intermediates, and that seems to be true (see Marchant and Higgins 1993).

GENERAL NOTES Williams (1952) noted that it is not possible to determine when or from where California Quail were imported into New Zealand during the years 1860 to 1870, but within the country the first liberation of birds was of two pairs by a Mr W Hay at Papakura, near Auckland, in 1862. Because they were introduced as game birds, most releases were organised by acclimatisation societies, and these commenced in 1865 when birds were released by the Nelson Acclimatisation Society. Williams listed a number of releases made at various localities in both North and South Islands between 1865 and 1945 and acknowledged that the acclimatisation societies were responsible for successful establishment of the species in New Zealand, though a lack of adequate records in their annual reports hinders any attempt to document subsequent spread of the populations. In his listing of early releases, Long (1981) includes numbers of birds liberated, and some were of quite high numbers, notably releases of 113 birds in 1867 and 42 birds in 1868 by the Auckland Society and of a queried 520 birds in 1871 by the Canterbury Society. Williams also noted that birds were introduced to the Chatham Islands at a date not determined, but almost certainly before 1900.

There is no evidence to suggest that New Zealand was not the source of California Quail introduced to Lord Howe Island, Norfolk Island and King Island, in Australia (see Williams 1952). Hindwood (1940) was informed by Mr W.S. Thompson, then one of the oldest residents on Lord Howe Island, that Captain Armstrong, who was the resident magistrate during 1879–1884, introduced these quail to the island in 1880. They increased in numbers and fairly large flocks were observed until ‘the coming of rats in 1918’, and then they were soon exterminated. Conversely, their introduction to Norfolk Island in 1895 or 1896 was successful, and Hindwood mentions a claim that this introduction was undertaken by a person connected with the Melanesian Mission, which was founded in New Zealand and established a church and training centre on Norfolk Island in 1867.

In his summary of introductions made in Australia, Long points out that birds from New Zealand were brought to Victoria and Tasmania between 1863 and 1874. Ryan (1906) reported that six birds were liberated in Victoria in 1863, and at Gembrook a larger number was released in 1872 and another 40 in 1874; they thrived for two years, when many nests were found and young birds observed, but in one season all disappeared. Jenkins suspected that more than 260 birds may have been liberated in Victoria in 1873–1874 (in Long 1981). Escaped cagebirds rather than later introductions probably are responsible for sightings reported from various parts of Victoria in the late 1930s, and noted by Chisholm (1950). The only introduction listed by Long for South Australia is the liberation of one pair at Pewsey Vale by the South Australian Acclimatisation Society in 1879–1880. Also occurring in 1880, and attributed to Jenkins, are the earliest releases recorded in New South Wales, where birds were liberated in the Liverpool Plains district, at Bathurst and in the Blue Mountains (in Long 1981). In the 1936–1937 Report from the New South Wales branch of the Royal Australasian Ornithologists Union, Cooper (1938) advised that because of objections an application to import California Quail into the state was rejected by the Federal Government under the Customs Act. Tarr (1950) attributed to Keith Hindwood a report that birds were released in the Prospect district, near Sydney, in 1944, and breeding occurred, but no further information was available. The only report of these quail being introduced in Queensland is mention by Chisholm (1919) of birds having been released at Eton Vale on the Darling Downs in the early 1870s, probably by Robert Ramsay, a State Treasurer and member of the Legislative Council, but they ‘did not long survive’. ‘A fair number of American Quail’ were released on Rottnest Island, Western Australia, in 1876 by Sir Harry Ord, State Governor, and presumably these were California Quail, but these birds were said to have been exterminated by feral cats (see Storr 1965).

Apart from Norfolk Island, the only established feral population in Australia is on King Island in Bass Strait, Tasmania, where the introduction of birds, reportedly also from New Zealand, occurred in 1930, as reported by Frith (1973), or in about 1920, as reported by Green (1989). Disappearance of populations from elsewhere in Tasmania is not documented, but Tarr (1950) noted that in the south they formerly were plentiful on Huon Island and near Bridgewater. Frith pointed out that these quail were brought to King Island as a game species to replace the native Brown Quail Synoicus ypsilophorus, which was declining because of settlement and a lack of management. The introduction flourished, the birds becoming sufficiently numerous for an open season to be declared, but during the acclimatisation period island residents became attached to ‘the quaint crested birds’ and a public outcry against hunting prevented further open seasons.

HABITATS AND STATUS Leary (2016) reports that California Quail are abundant throughout their range in New Zealand, being most numerous in North Island from Northland to Waikato and in inland Bay of Plenty and coastal Wairarapa, and in South Island in Nelson, Marlborough and northern Canterbury, including Banks Peninsula, but are uncommon or absent in Southland and West Coast, including the Southern Alps, in South Island, and on the Chatham Islands. Williams (1952) noted that, although recorded up to 1800 m, they normally are rare above 900 m, and they are tolerant of high rainfall, small numbers being found in places with an annual rainfall of 2500 mm, but favouring areas where it is less than 1520 mm. Williams also pointed out that they tend to avoid areas where a mean annual temperature of less than 10°Celsius is accompanied by a mean annual rainfall exceeding 750 mm, this combination having thwarted repeated attempts to acclimatise them in Southland, and a mean annual rainfall above 2500 mm is responsible for their absence from cleared lands in the Southern Alps. They probably no longer occur on the Chatham Islands, the last record being of two sightings at Te Matarae, on the main island, in December 1996 (Miskelly et al. 2006).

Noting that forested and intensively farmed lands are avoided by California Quail in New Zealand, Williams identified favoured habitats as those providing an interspersion of cover and open spaces, and two natural plant associations meeting these requirements are Leptospermum heathlands and Poa or Festuca grasslands, the latter only when in association with shrubs like matagouri Discaria toumatou and Rubus or Muehlenbeckia creepers. The heathlands are more common in North Island, and the grasslands more common in South Island, where at one time flourished large populations of the extinct New Zealand Quail Coturnix novaezelandiae. New habitats have been created by burning, grazing or introducing exotic trees and shrubs in native grasslands, by the partial clearing of native forests, the invasion of wide riverbeds by introduced weeds, such as gorse Ulex europaeus, lupin Lupinus arboreus and broom Cytisus scoparius, and the establishment of Pinus plantations criss-crossed with wide firebreaks, and wherever the ground is relatively clear introduced herbs and grasses become a ready food source. Expanding on these comments, Leary confirms uncultivated open shrubland as the preferred habitat, and notes that rough, scrubby margins of rivers, inlets, forests, roads and rural gardens are frequented, with examples of good habitats being areas vegetated principally with Pteridium bracken, tussock grasses, matagouri, gorse, blackberries Rubus hybrid, Coriaria tutu and lupins.

Williams reported that on Cairnmuir Station, on the banks of the Kawarau River, near Cromwell, central Otago, the population of California Quail was under observation at various times for more than a year. Much of the land was poor tussock grassland with rare trees on hillsides and ridges, but along the banks of local streams were groves of manuka Leptospermum scoparium and matagouri with occasional elders, willows and briars. Near to the homestead, and surrounded on all sides by tussock grassland, were grass and lucerne paddocks with pines, presumably Pinus radiata, planted on three sides of one paddock and assorted fruit trees on the fourth side. Numerous small coveys of quail frequented the property, and in the winter of 1950 a total of 120 birds occupied a range of 14 ha – a density of 8.6 birds per ha, and in summer they ranged farther afield to occupy 40 ha – a density of 3.0 birds per ha.

Conceding that numbers of California Quail had declined since their introduction, Williams stressed that the role of predation, especially following the introduction of mustelids, almost certainly has been overestimated, and quality of habitat is more important in regulating numbers than is predation, hunting or disease. Initially, the partial clearing of native forest created much suitable habitat, but subsequent more intense cultivation and grazing of partially cleared land, or alternatively its reversion to second-growth forest following cessation of farming, rendered habitats unsuitable for California Quail. Dynamics of the population on Kapiti Island, North Island, were cited as an example of this effect of changes in land-use practices. Initially, the island supported a whaling station and the grazing of sheep and cattle, resulting in the modification of the original forest to dry, open country, where California Quail flourished after their introduction in 1893 but, after declaration of the island as a sanctuary in the early 1900s, browsing mammals were eliminated and a steady decline of the quail population coincided with gradual return of the original dense vegetation cover. Leary also concedes that, despite their continued overall abundance, numbers apparently have declined since the 1920s, when coveys of up to 100 birds were recorded frequently in Canterbury, where now it is unusual to see a covey of more than 20 birds. Birds and their eggs are taken by rats, cats and mustelids, eggs are taken by introduced hedgehogs Erinaceus europaeus, birds are poisoned by baits, especially poisoned grain laid for rabbits, and adults, chicks and nests are destroyed by farm machinery or by fires during clearing operations (Williams 1952; Oliver 1955).

California Quail are well established and very common on Norfolk Island, where undoubtedly they benefited from the near total clearance of native forest. Smithers and Disney (1969) recalled that, during a visit made to the island in November–December 1968 to observe terrestrial and freshwater birds, these quail were found to be common in more open habitats, such as pastures with thickets, open wet flats with nearby cover and more open areas of regenerating forest. In early December 1978, in a survey of birds undertaken by members of the Royal Australasian Ornithologists Union (RAOU) during an annual congress and field outing held on the island, California Quail were found to be widespread and very common, being recorded in 79 of 191 grid squares of 500 m × 500 m, or 39 per cent of the land area, with a preference for the same habitats as reported previously together with clearings within remnant stands of native vegetation (Schodde et al. 1983). In the 1970s and 1980s, during frequent visits to Norfolk Island to monitor numbers of Cyanoramphus parakeets, and again in 2007, I became familiar with these