AUSTRALIAN 42 BIRD WATCHER

AUSTRALIAN BIRD WATCHER 1997, 17, 42-59

The Juvenile Food-begging Calls and Related Behaviour in the Australian 'Rose-tailed' Parrots Alisterus, Aprosmictus and Polytelis; and a Comparison with the Eclectus Parrot Eclectus roratus and Pesquet's Parrot Psittrichas fulgidus

by JOHN COURTNEY, 'Ashgrove', Swan Vale, via Glen Innes, N.S.W. 2370

Summary The juvenile food-begging calls, food-begging postures and unique 'chick-to-parent' contact calls of the Australian King-Parrot Alisterus scapularis, Red-winged Parrot Aprosmictus erythropterus, Regent Parrot Polytelis anthopeplus, Superb Parrot P. swainsonii and Princess Parrot P. alexandrae are described, and compared where applicable with those of other Australian parrots. In addition, audio graphs of the begging calls in the Eclectus Parrot Eclectus roratus and Pesquet' s Parrot Psittrichas fulgidus are compared. It is concluded that the members of the Alisterus, Aprosmictus and Polytelis group are tightly monophyletic. It is proposed that because of this, their unique and distinctive rose­ pink tail colouration ought to be incorporated into a descriptive common name for the group to emphasise their distinctiveness, henceforth to be referred to as the 'rose-tailed' parrots to distinguish them from others such as the 'broad-tailed' parrots. The begging call of the Eclectus Parrot is different from that of the 'rose-tailed' parrots. However, this call in Pesquet's Parrot is so similar in all respects to that of the Australian King-Parrot that relationship between Pesquet's and the 'rose-tailed' parrots must be suspected. Introduction The present paper is a continuation of the study outlined by Courtney (1996) covering the juvenile food-begging calls of all groups of the Australian parrots and cockatoos. The methods used and a discussion of food-begging calls were fully outlined in that paper. The present paper has two aims: to place on record descriptions of the distinctive juvenile food-begging calls and related data of the parrot genera Alisterus, Aprosmictus and Polytelis as a contribution to the life-history of each species; and to explore the taxonomic possibilities presented by comparing these calls between species and with other species in nearby regions. This involves the principle (Courtney 1986, 1996) that the 'juvenile food-begging call is closely similar in closely related species, and bears little or no resemblance in distantly or unrelated species respectively, thus indicating relationships'. Historically, the parrots of the genera Alisterus, Aprosmictus and Polytelis have long been recognised as related and are non-controversial in this respect, except for the anomaly of some authors (e.g. Brereton 1963, Condon 1975, McAllan & Bruce 1988) in grouping the Cockatiel Nymphicus hollandicus with them as well. Holyoak (1972), Smith (1972, 1975), Courtney (1974), Hornberger (1980), Adams et al. (1984) and many others more recently, listed by Christidis & Boles (1994), have produced overwhelming evidence that the Cockatiel is basically a cockatoo and, as the juvenile food-begging call of the Cockatiel bears no resemblance to those of the 'rose-tailed' parrots (cf. Courtney 1996), that species is not considered part of the polytelitine group. Polytelis and Alisterus have been shown to be closely related to each other and are claimed to form a 'sister' group to the platycercines or 'broad-tailed' parrots (Christidis, Schodde et al. 1991). However, those authors did not examine relationships within the polytelitine group. The genera Alisterus and Aprosmictus have sometimes been merged (Anon. 1926, Storr 1984) and sometimes kept separate (Brereton 1963, Lendon 1973, Condon 1975); although the latter course is currently followed, the matter requires reappraisal (Christidis & Boles 1994). Conversely, the monotypic genus Spathopterus North 1895 has been proposed for the Princess Parrot Polytelis VOL. 17 (1) MARCH 1997 Vocalisations of Juvenile 'Rose-tailed' Parrots 43 alexandrae but has not gained acceptance (see Anon. 1926, Condon 1975, Christidis & Boles 1994). Peters (1937) gave it subgeneric recognition, with subgenus Polytelis for the Regent Parrot P. anthopeplus and Superb Parrot P. swainsonii. However, all three species of Polytelis seem not particularly close to one another (Lendon 1973). This matter also requires reappraisal. The large black-and-red Pesquet's Parrot Psittrichas fulgidus from New Guinea is discussed here too. Its relationship to the parrots of the Australian Region has been pbscure for more than a century, ironically not because of a lack of similarities, but because it shares many features with diverse groups which themselves appear not closely related (cockatoos, lorikeets, platycercines, nestorines, loriculines), thus confusing the issue. Its taxonomic position has been disputed, with no suggestion that it might be related to the 'rose-tailed' parrots (see Smith 1975, from which some of the following historical account is taken). Although Salvadori (1891) expressed doubts as to the true position of Pesquet's Parrot, he was certain that Reichenow (1881) was mistaken in placing it with the cockatoos. Thompson (1899) considered it to have no affinity with other parrots, and von Boetticher (1959) isolated it as a 'primitive' monotypic family. Peters (1937), Verheyen (1956) and Brereton (1963) followed Salvadori's reluctant lead and placed it next to the African Grey Parrot Psittacus erithacus and Madagascan black parrots Coracopsis. Although Thompson (1899) stated that Pesquet's Parrot is the only parrot lacking an opening in the interorbital plate, this peculiarity also occurs in the Palm Cockatoo Probosciger aterrimus (Smith 1975). The skull of Pesquet's Parrot is otherwise like that of Lorius lories (1orikeets), particularly in its long, narrow outline (Smith 1975). Dyck (1977) concluded that Pesquet's Parrot is a cockatoo, because it lacks 'Dyck' -texture (blue or green) in its plumage. Forshaw (1978) indicated that despite his previous belief that Pesquet's Parrot is close to the Eclectus Parrot Eclectus roratus (Forshaw 1973), new data from captive breeding had led him to suspect a relationship with the black-cockatoos Calyptorhynchus. Hornberger (1980) found that the drinking method of Pesquet's Parrot was like that of the hanging-parrots Loriculus; she placed both in monotypic subfamilies on morphological grounds. Low (1986) has drawn attention to the lory-like structure of the feathers of the hind-neck in Pesquet's Parrot, which are elongated and shaft-streaked like those of Chalcopsitta lorikeets from New Guinea. Pesquet's Parrot does possess many features common to cockatoos, lorikeets and (from this study) the king-parrot group. However, unlike these birds (all of which possess a 'primitive' A-1 carotid artery formula), Pesquet's Parrot has the 'advanced' A-2-s carotid arrangement which, in the Australian Region and nearby, occurs only in a large advanced group within the platycercines and in the New Zealand Nestor parrots (see Glenny 1957, 1959; Smith 1975). From genetic studies, Christidis, Shaw & Schodde (1991) speculated that the karyotype (chromosome arrangement) of Pesquet's Parrot could be derived from that of the Indian Ring-necked Parrot Psittacula krameri. The only other likely relatives of the Australian 'rose-tailed' parrots in the geographical vicinity are those that were grouped with them by Smith (1975), and referred to by him as the 'coral-billed' parrots (after von Boetticher 1959). These include the Eclectus Parrot and its presumed close relative the Red-cheeked Parrot Geoffroyus geoffroyi, both from Australia and New Guinea, and the non-Australian ring-necked parrots of the genus Psittacula, which are widespread in Asia with one species ranging as close to Australia as Bali. All these species (including the polytelitines), grouped as the Psittaculini, were aligned with the lorikeets by Smith (1975) and with the psittacines by Hornberger (1980). On biochemical and/or genetic AUSTRALIAN 44 COURTNEY BIRD WATCHER

grounds Eclectus and Geoffroyus are closely related to each other and form a 'sister' group to the fig-parrots Cyclopsitta spp., but are distant from Psittacula; and Psittacula has no close links with Australo-Papuan parrots such as the polytelitines (Christidis, Schodde et al. 1991; Christidis, Shaw & Schodde 1991). The results of the present study are used to investigate these taxonomic questions.

Methods As for all the parrot groups in this series of studies (cockatoos: Courtney 1996; lorikeets and platycercines: Courtney 1997 and in prep.), many broods of most species were hand-reared and their begging calls described in written form over a period of 11 years 1952-1963. In that period many broods were also observed closely in aviaries. When sound-recording equipment became available, the study was repeated on several hand-reared broods of all available species by tape-recording what were by then known to be their typical, species-specific begging calls. Representative examples of each species' typical begging call were then used to generate audiographs characteristic of each species. The begging calls of two unrelated captive broods ofeach of the five mainland Australian species in the 'rose-tailed' group were tape-recorded, except for the Red-winged Parrot Aprosmictus erythropterus in which only one brood (of two young, i.e. B/2 x 1) was studied (Australian King­ Parrot Alisterus scapularis: B/1 x 2; Princess Parrot: B/1 X 2; Regent Parrot: B/3 x 2; Superb Parrot: B/2 x 2). A comparison of the tapes, and audiographs generated, revealed little individual (intraspecific) variation and constant interspecific differences; the sample audiographs presented herein are representative of their species. Many other broods of species in the group were observed in aviaries, with no significant intraspecific variation in begging calls noted. 'Out-group' comparisons are provided by the Eclectus Parrot and African Grey Parrot audiographs herein, and by the extensive array of audiographs provided for most species in the other Australian parrot groups, in the other papers in this series (cockatoos: Courtney 1996; lorikeets and 'broad-tailed' or platycercine parrots: Courtney 1997 and in prep.). Tape-recordings of two broods of young Pesquet's Parrots (each B/1), two broods of young Eclectus Parrots (B/1 x 1, B/2 x 1) and one brood of African Grey Parrots (B/2) were received from Miss Rosemary Low, who bred and hand-reared the Pesquet's Parrots in 1995 and the others in 1980-81. Regrettably, no extensive samples of begging-calling in the nearby Asian genus Psittacula could be obtained. An attempt to rear young of the Indian Ring-necked Parrot failed because the young were remarkably wild and intractable, refusing to call. A brief tape-recording of one individual of the race P. krameri manilensis was obtained from Mr George A. Smith of England. The methods used have been described in Courtney (1996). However, for the present paper, audiographs were produced on a Macintosh LC 11 personal computer using the software program 'Audiograph 2.0' by Ralph Slater Sutherland, Canberra. The sound recordings were fed into the computer by line from a Sony WM D6C Professional Walkrnan cassette tape-recorder. Although there is no time restriction in this electronic program, the audiographs produced were mostly edited to 2.4 seconds in length for comparison with the familiar sonagrams in use for decades. Specifications for the print-outs (Figures 1-16) are: sample rate 1:1 (22.26 kHz); frequency range 0-11.13 kHz; resolution high (256 points vertically = 43.4631 Hz/point); width 512 points/53406 samples = 4.69 ms/point (total2.3999 s); sound level displayed = - 33 db (palest shading) to 0 db (darkest shading). A tape-recording of all calls used in the present paper has been lodged with the Australian National Wildlife Collection, CSIRO, Canberra.

Results In the Alisterus-Aprosmictus-Polytelis group, four main aspects were detected and examined. These were what appeared to be a 'chick-to-parent' contact call used in conjunction with the begging call (in Australian parrots a trait unique to the group), the juvenile food-begging call, juvenile food-begging posture, and the rose-pink edging to the inner webs of the lateral rectrices (again a feature unique to this group). Of these, only the juvenile food-begging call is universal to all parrots and, as the most important feature, is discussed first. Other minor features of chicks (e.g. bill colour, aggressive stance etc.) are discussed also.

Food-begging calls All begging calls of the five Australian 'rose-tailed' parrots followed a similar VOL. 17 (1) MARCH 1997 Vocalisations of Juvenile 'Rose-tailed' Parrots 45

pattern in which the basic note was brief and somewhat high-pitched; the notes were repeated rapidly and were strung closely together in lengthy uninterrupted bursts of loud calling which drifted up and down in tonal inflection, with brief pauses between the bursts of calling (Figures 1-5). This calling was unlike that of any other mainland Australian parrot, but was also typical of Pesquet's Parrot nestlings (Figures 6-7), whose calling strongly resembled that of the Australian King-Parrot (Figure 2). The brief tape-recording of the Indian Ring-necked Parrot (Figure 10) resembled the begging call of the Princess Parrot (Figure 1), but was not of sufficient duration to 'show length of calling bursts or whether the tone drifts up and down. The multiple audiographs, from different individuals of a given species (Figures 1 and 2), exemplify the negligible intraspecific differences and constant (though minor) interspecific differences in calls of the five Australian 'rose-tailed' parrots. More importantly, the audiographs of all five species (Figures 1-5) emphasise the interspecific similarity, as noted above, of the calls across this natural group (and inter-group differences, cf. lorikeets and platycercines: Courtney 1997 and in prep.). The main distinguishing feature in the begging call between the different species of 'rose-tailed' parrots is simply repetition rate. This is fastest in the Princess Parrot with the mean number of basic notes about 10 per second; Australian King-Parrot 5 per second; Red-winged Parrot 4.5 per second or slower; and Regent and Superb Parrots 2.25 notes per second. It would seem that the faster these notes are delivered, the sharper they sound. On sonagram, the sharper-sounding notes (Princess and Indian Ring-necked Parrots) are pointed in shape like an inverted 'V' or chevron (Figure 17), whereas the less sharp notes (the other parrots studied) are flatter at the apex. The Princess Parrot has the chevron shape in the fundamental tone of the begging note, which has five accompanying harmonics. These extend to 9 kHz, with most energy in the 2nd harmonic at a little above 3 kHz (Figure 1). The Australian King-Parrot begging call sounds like a slower, less sharp version of the Princess Parrot, but on audiograph the fundamental tone is rounded at the apex rather than chevron-shaped (Figure 2). This species has 10 accompanying harmonics covering the same range as the five harmonics of the Princess Parrot reaching to 9 kHz, with most energy in the 4th harmonic at 3 kHz. The two strongest harmonics, the 4th and 5th, are often quite chevron-shaped and presumably this makes the call resemble that of the Princess Parrot, particularly as they occur at the same frequency. In the Red-winged Parrot (Figure 3), the fundamental tone is flatter at the apex than that of the Australian King-Parrot, and is a little longer on the time scale, reflecting the slightly slower repetition rate. There are multiple harmonics, many too faint to document. In many respects, the begging call of the Red-winged Parrot seems to fall between those of the Australian King-Parrot on one hand, and Regent and Superb Parrots on the other. In the Regent Parrot (Figure 4) and Superb Parrot (Figure 5), the fundamental tone is flat in shape, not formed like a chevron as in the Princess Parrot or rounded as in the Australian King-Parrot. Again (reminiscent of the Red-winged Parrot) there are multiple harmonics, many too faint to count. When large and at the fledging stage, Superb Parrots tend to introduce a quaver into the begging call, but this may represent only the first stage of a gradual change-over to adult-type calling in this particular species. There is no quaver present in the basic begging note of downy young Superb Parrots, or at any stage in the Regent Parrot. The structure of the basic note (from two large nestlings) in the begging call of Pesquet's Parrot (Figures 6-7) is similar to that of the Australian King-Parrot, with a distinct rounded shape in the fundamental tone. The pattern of delivery of these AUSTRALIAN 46 COURTNEY BIRD WATCHER

Princess Parrot (a)

0.00 s 0.20 s 0.40 s 0.60 s 0.80 s 1.00 s t .20 s 1.40 s 1.60 s 1.80 s 2.00 s 2.20 s

Princess Parrot (bll 10.00 kHz

9.00 kHz

8.00 kHz

0.00 s 0.20 s 0.40 s 0.60 s 0.80 s 1.00 s 1.20 s 1.40 s 1.60 s 1.80 s 2.00 s 2.20 s

Princess Parrot (ell 10.00 kHz

9.00 kHz

8.00 kHz

7.00 kHz .;, .;,. ::r,;"": "' ~ ·,. -

Figure 1. Juvenile food-begging calls of nestling Princess Parrots: (a) with some adult calls in the background; (b) and (c) different, unrelated individuals. VOL. 17 (1) MARCH 1997 Vocalisations of Juvenile 'Rose-tailed' Parrots 47

Australian King-Parrot (a) 10.00 kHz

9.00 kHz

8.00 kHz

7.00 kHz

6.00 kHz

O.OOkHzL-~~--~~--~~~--~~--~~~~~~--~~--~~~--~~--~~~ 0.00 5 0.20 s 0.40 s 0.60 s 0.80 s 1.00 s 1.20 s 1.40 s 1.60 s 1.80 s 2.00 s 2.20 s

11.00 kHz.---,---,--...,--.,.---.,---,---,--.,---,---..---,..--~.,--~-~-.--~-,-~-....--.,.--~-.--, Australian King-Parrot (b) 10.00 kHz

9.00 kHz

0.00 s 0.20 s 0.40 s 0.60 s 0.80 s 1.00 s 1.20 s 1.40 s 1.60 s 1.80 s 2.00 s 2.20 s Figure 2. Juvenile food-begging calls of nestling Australian King-Parrots: (a) and (b) different, unrelated individuals.

ll.OO kHz.,---,--..--...,----.,.---..--,--,--...,----,---..--,..---,~.,---,----,---,..---~-,--..--...,--.,.--..--,..--, Red-winged Parrot 10.00 kHz

9.00 kHz

8.00 kHz

7.00 kHz

6.00 kHz

5.00 kHz ~"'; · .-..-: g_..-::. ~

4.00 kHz

0.00 s 0.20 s 0.40 s 0.60 s 0.80 s 1.00 s 1.20 s 1.40 s 1.60 s 1.80 s 2.00 s 2.20 s Figure 3. Juvenile food-begging call of nestling Red-winged Parrot. AUSTRALIAN 48 COURTNEY BIRD WATCHER

Regent Parrot 10.00 kHz

9.00 kHz

8.00 kHz

7.00 kHz

6.00 kHz

5.00 kHz

4.00 kHz

3.00 kHz

2.00 kHz

1.00 kHz

0.20 s 0.40 s 0.60 s 0.80 s 1.00 s 1.20 s 1.40 s 1.60 s 1.80 s 2.00 s 2.20 s Figure 4. Juvenile food-begging call of nestling Regent Parrot.

11 .00 kHz ,.-~~~~-~.,-~~~-~~~~-~~~~~-~.,-~-----, Superb Parrot 10.00 kHz

9.00 kHz

8.00 kHz

7.00 kHz

6.00 kHz

5.00 kHz ir.

.:.--r . k~ .~___f' 0.00 kHz L--'--'-~~-~~-'-~-~~~~~-~-'--'-~~-~~-'-~-'------' 0.00 s 0.20 s 0.40 s 0.60 s 0.80 s 1.00 s 1.20 s I .40 s 1.60 s 1.80 s 2.00 s 2.20 s Figure 5. Juvenile food-begging call of large feathered young Superb Parrot, close to fledging. notes follows that of the Australian group just described, especially in the tonal inflection which drifts slowly up and down during bursts of rapid calling, making the begging call in advanced nestlings sound remarkably similar to that of the Australian King-Parrot. Chicks of Pesquet's Parrot develop 'second' down (unlike cockatoos) and lack the typical cockatoo Juvenile Food-swallowing Vocalisation (R. Low in litt.), thus making any link with the cockatoos unlikely (cf. Reich!!now 1881, Dyck 1977, Forshaw 1978). The juvenile food-begging call of the African Grey Parrot (Figure 8) was found not to resemble that of Pesquet' s Parrot. The notes were delivered much more slowly, were evenly repeated, and had a downward-sliding quality, all of which was reminiscent of the begging call of the Eclectus Parrot. The juvenile food-begging call of the Eclectus Parrot sounds like a brief, sharp downward-sliding whistle (Figure 9), which is repeated at a much slower rate than the notes of the 'rose-tailed' parrots and Pesquet's Parrot. The notes are given at intervals of 2 seconds or less and are usually regularly repeated. There are no bursts VOL. 17 (1) MARCH 1997 Vocalisations of Juvenile 'Rose-tailed' Parrots 49

Pesquet's Parrot 10.00 kHz

9.00 kHz

8.00 kHz

7.00 kHz '1'-.

6.00 kHz . &: 5.00 kHz }

4.00 kHz 3.00 kHz 71 ~ 2.00 kHz ~

Figure 6. Juvenile food-begging call of Pesquet's Parrot, chick (no. 1) 32 days old. 11.00 kHz.-----r----r-.,---,---,.-.,----,---,.-,--r----.-,--r----.-..--,---,-..---.----.r--..--.----.,.--, Pesquet's Parrot - older nestling 10.00 kHz

9.00 k.Hz

8.00 kHz

7.00 kHz

0.20 s 0.40 s 0.60 s 0.80 s 1.00 s 1.20 s 1.40 s 1.60 s 1.80 s 2.00 s 2.20 s Figure 7. Juvenile food-begging call of Pesquet's Parrot, chick (no. 2) 39 days old. 11.00 kHz.-----r----r-.,---,---,....,.-.,----,---,.-,--r----.-,--r----.-..--,---,-..---.----.-..--.----.,.--, African Grey Parrot 10.00 kHz

9.00 kHz

8.00 kHz :..~- 7.00 kHz ~,-:;J. 6.00 kHz ~ ~--~ ,_~~~~ -· 5.00 kHz

4.00 kHz ~~ ~~-

3.00 kHz

2.00 kHz 1.00 kHz ~ II§ O . OOk Hz L-~~~~~~~~~~-~~~~~~-~~~-~~~-~~-·~~~~~ 0.00 s 0.20 s 0.40 s 0.60 s 0.80 s 1.00 s 1.20 s 1.40 s 1.60 s 1.80 s 2.00 s 2.20 s Figure 8. Juvenile food-begging call of the African Grey Parrot. AUSTRALIAN 50 COURTNEY BIRD WATCHER

11.00 kHz r--r--,--r-----.-~-,--,-~----,-~--,--p-~-..----,----,--r-----.-r--,--,---,-----,,--, Eclectus Parrot l 10.00 kHz :r 9.00 kHz rl 8.00 kHz ~-- - 7.00 kHz -~ .. ~ " ~ J,.J 6.00 kHz ~ ); I "!!-

5.00 kHz

4.00 kHz

3.00 kHz 2.00 kHz ~ 1.00 kHz r~ 0.00 kHz'----'---:--'----'--:-'-""'-~--'--'-'-"'--'---'---'--'--'-"-_...-..__--'---:_.____._-:-'_..__~_,____.__,____, 0.00 s 0.20 s 0.40 s 0.60 s 0.80 s 1.00 s 1.20 s 1.40 s 1.60 s 1.80 s 2.00 s 2.20 s Figure 9. Juvenile food-begging call of the Eclectus Parrot.

Indian Ring-necked Parrot 10.00 kHz

9.00 kHz

8.00 kHz

7.00 kHz

Figure 10. Juvenile food-begging call of the Indian Ring-necked Parrot. of calling which drift up and down in tonal inflection, as in the 'rose-tailed' parrots and Pesquet's Parrot. The calls are unlike those of the 'rose-tailed' parrots but somewhat like the begging call of the African Grey Parrot. The begging call of the Red-cheeked Parrot has yet to be obtained but, from a videotape by John Young of a pair at a nest with young, this call seems similar to that of the Eclectus Parrot (pers. obs.). No begging call ofthe fig-parrots Cyclopsitta is available for analysis, but this call would appear to differ from that of Eclectus and Geoffroyus. Bohner (1986) described the begging call of the red-browed race of the Double-eyed Fig-Parrot C. diophthalma macleayana as a loud, drawn-out chattering, with emphasis on the first note. In contrast, Forshaw (1978) described it as a constant twittering. A recording of the Indian Ring-necked Parrot showed that the basic tone and attendant harmonics were chevron-shaped to an extreme degree (Figure 10), more so than in the Princess Parrot. In common with the 'rose-tailed' parrots and Pesquet's Parrot, the individual begging notes were sharp-sounding, rapidly repeated and evenly VOL. 17 (1) MARCH 1997 Vocalisations of Juvenile 'Rose-tailed' Parrots 51 spaced, but were delivered only in a brief burst. It is not known whether longer bursts are made, or if they drift up and down in tonal inflection.

'Chick-to-parent' contact call Unlike all other parrots and cockatoos studied (except Pesquet's Parrot), the chicks of all species of the Alisterus-Aprosmictus-Polytelis group uttered another call in addition to the begging call which was totally different. This was interpreted as a _'chick-to-parent' contact call, for it was given when their keeper entered the room and the chicks became aware that they were about to be fed. In the Australian King­ Parrot (Figure 11) and Princess Parrot (Figure 12), the call sounds like a two-syllable whistle, usually uttered several times before the beginning of begging calling and mostly not used again during feeding. The Regent, Superb and Red-winged Parrots differ in that the call is a quavering whistle, and usually uttered at intervals in amongst the begging calls instead of at the beginning. There is an interspecific difference in that the Red-winged Parrot utters a piercingly shrill quavering whistle at very frequent intervals while begging (Figure 13); the Superb Parrot frequently utters a less shrill and very long quavering whistle (Figure 14); and the Regent Parrot utters a seldom­ used duller-sounding quavering note which is very short in duration (Figure 15). No other Australian parrot chicks were noted to use a 'contact call' in association with the food-begging call. However, the recording of two young Pesquet's Parrots showed that they, too, uttered a vocalisation before begging-calling which was totally different from the begging call and sounded like a cork being removed slowly from a bottle by twisting (Figure 16). In this respect Pesquet's Parrot resembles the 'rose­ tailed' parrots. However, this may prove to be a nest-defensive call rather than a contact call.

Juvenile food-begging posture When begging for food, the young of the Alisterus-Aprosmictus-Polytelis group bobbed their heads up and down in an irregular way, on out-stretched necks. This action is similar to that of young of the platycercines, the Cockatiel and the Gang­ gang Cockatoo Callocephalon fimbriatum but not of lorikeets or the other cockatoos (pers. obs.).

Rose-pink margins to rectrices In plumage, members of the Alisterus, Aprosmictus and Polytelis group share a common characteristic, the striking rose-pink edging to the inner webs of the lateral rectrices, apparently not possessed by any other parrot (e.g. see the descriptions in Forshaw 1978). This feature is present in all individuals regardless of age or sex in the Princess Parrot. It is present in adult female and juvenile Superb, Regent and Red-winged Parrots but is lost by moult in the adult male, except that some adult male Red-winged Parrots retain pink on the tips (pers. obs.). It is present to a lesser degree in the Australian King-Parrot, being confined to an area near the tip of the tail in most females and juvenile males and is lost in adult males (Lepschi & Courtney 1992). Two subspecies of the Amboina King-Parrot Alisterus amboinensis (A.a. amboinensis and A.a. buruensis) have prominent rose-pink edging on the inner webs of the rectrices, whereas four other alleged subspecies (one on mainland New Guinea) have no pink and much shorter tails. The only other New Guinea species in this genus, the Green-winged King-Parrot A. chloropterus, has no pink either (Forshaw 1978, Smith 1979). The tail of the Timor Red-winged Parrot Aprosmictus jonquillaceus has greenish-yellow tips instead (Forshaw 1978; see Smith 1979 for distributions of species and subspecies in the complex). AUSTRALIAN 52 COURTNEY BIRD WATCHER

11.00 kHz ,--~----.--.---,--..---,-----.--.---,--..----,--~~----.--.---,--..---,-----.--.---,--..----,-----, Australian King-Parrot 10.00 kHz

9.00 kHz

8.00 kHz

7.00 kHz

6.00 kHz

5.00 kHz

4.00 kHz

3.00 kHz

2.00 kHz

1.00 kHz

O.OO kHzL-~~~~~==~~~~~~-~~-~~~-~~-~~~~~~-~-L~ 0.00 s 0.20 s 0.40 s 0.60 s 0.80 s 1.00 s 1.20 s 1.40 s 1.60 s 1.80 s 2.00 s 2.20 s Figure 11. Chick-to-parent contact call, Australian King-Parrot (large feathered nestling about to fledge).

11.00 kHz ,--~----.---.,-.---,--..---,-----,-~--....-..----,--.--~----.--.---,--..---,-----.~~--....-..----,-----.,

~ Princess Parrot 10.00 kHz ~ 9.00 kHz f 8.00 kHz i, 7.00 kHz i

6.00 kHz !),t ~ ~ ~ ~i' · ~~ " .. 5.00 kHz s·_ ,. • • L::. t-" ~~ --': ~ -~ 4.00 kHz i ;·~ - ~ . ~ .-f*'"· : ~· ::\ . 3.00 kHz ~ ! . ~ - ' .. ' .. . 2.00 kHz ~ 1.00 kHz f - · : : •

O.OOkHzL-~~-~s-~~-~~~-~~-~~~~~~~-~~-~~~-~~-~~ 0.00 s 0.20 s 0.40 s 0.60 s 0.80 s 1.00 s 1.20 s 1.40 s 1.60 s 1.80 s 2.00 s 2.20 s Figure 12. Chick-to-parent contact call, Princess Parrot (pin-feathered nestling).

Red-winged Parrot 10.00 kHz

9.00 kHz

8.00 kHz

7.00 kHz

6.00 kHz

5.00 kHz ~

4.00 kHz ,

3.00 kHz~ 2.00 kHz L

1.00 kHz ~

O . OOkHz L-~~-~~-~~~-~~-~~-~~~-~~-~~~-~~-~~~ 0.00 s 0.20 s 0.40 s 0.60 s 0.80 s 1.00 s 1.20 s 1.40 s 1.60 s 1.80 s 2.00 s 2.20 s Figure 13. Chick-to-parent contact call, Red-winged Parrot (small feathered nestling). VOL. 17 (1) MARCH 1997 Vocalisations of Juvenile 'Rose-tailed' Parrots 53

Superb Parrot 10.00 kHz

9.00 kHz

8.00 kHz

7.00 kHz

6.00 kHz

0.00 s 0.20 s 0.40 s 0.60 s 0.80 s 1.00 s 1.20 s 1.40 s 1.60 s 1.80 s 2.00 s 2.20 s Figure 14. Chick-to-parent contact call, Superb Parrot (large feathered nestling).

11.00 kHz r-~--,-~~-~--,---,-~-,---,----,--,.---,---,-~--,--~--,---,--,----.--~--,---, Regent Parrot 10.00 kHz

9.00 kHz

8.00 kHz

7.00 kHz

6.00 kHz

Figure 15. Chick-to-parent contact call, Regent Parrot (small feathered nestling; second chick giving juvenile begging call in background).

Pesquet's Parrot 10.00 kHz

9.00 kHz

8.00 kHz

7.00 kHz

6.00 kHz ~t~~$~ ':·~~: .. ~ : ...... 5.00 kHz --~~ ;; 4.00 kHz , , ,!j~i~ ~: ; 3.00 kHz " ~"'-:':''' 2.00 kHz

1.00 kHz

Figure 16. Chick-to-parent contact call, Pesquet's Parrot (chick 32 days old). AUSTRALIAN 54 COURTNEY BIRD WATCHER

Indian Ring-necked Princess Parrot s7-• Parrot N 6- 2 s• }I\\ I \ >. !\ f\\\ "I <: "Q) , I ' r\ \ ! ~ v \I' 0' 4- "Q) ...'" 3- j .·~ 2- ~A~ i ~ ,tl , ~lJl , t ,I, I /,, 1- i \ I ~ /\ ; \

0 sec 1 sec Figure 17. Comparison of juvenile food-begging call of Indian Ring-necked Parrot and Princess Parrot, using Kay Sonagram and wide-band (300 Hz) filter.

Bill colouration of nestlings Nestlings of the Australian King-Parrot may have pink bills, the bill colour of young fliers (see Lepschi & Courtney 1992). The mandibles of nestling Princess Parrots were dull brownish pink, whereas those of Superb, Regent and Red-winged Parrots were yellowish with an infusion of orange. The bills of the latter two become orange much earlier in life than that of the Superb Parrot. It would appear that loss of yellow and increase of orange comes progressively with age, leading to coral-red in adults. In the Princess Parrot and Australian King-Parrot the pink deepens to coral-red in adults.

Feather colouring of juveniles Two aspects of this seem worth recording. Up to the age of 10 months a male Red-winged Parrot had pinkish red on the secondary wing-coverts. From this age to over a year, these were replaced with slightly deeper red ones. Until then, the similarity in these feathers between Red-winged and Regent Parrots was striking. Two Superb Parrots less than four months old moulted the dull olive-green feathers on the crown and replaced them with pale bluish-mauve ones strongly reminiscent of the Princess Parrot. Change to this colour did not occur in the Regent Parrot.

Defensive behaviour in young Nestling King-, Red-winged, Princess and Superb Parrots frequently, and Regent Parrots infrequently, uttered a constant 'growling' sound if their nest-box was touched. This defensive vocalisation of nestlings, apparently to deter nest predators, is unique to the group and comparable with the intermittent 'churring' call and head-waving display of nestling platycercines, constant buzzing call of lorikeets (pers. obs.), and snort-hissing and swaying of cockatoos (Courtney 1974) when they sense a potential nest predator. Small feathered nestling Superb Parrots were aggressive in an unusual way. They behaved in a manner not seen in any other parrot, by lunging at an intruding hand and striking powerfully with one and sometimes both wings, which were kept partly VOL. 17 (1) MARCH 1997 Vocalisations of Juvenile 'Rose-tailed' Parrots 55

folded. Contact was made with the underside of the bend of the wing, and the three or more hits were delivered too swiftly for the human eye to count. Similar behaviour was encountered in an incubating Squatter Pigeon Geopfu:Jps scripta (pers. obs.). This behaviour must have arisen through a special need to repel nest intruders.

Discussion and conclusions The present paper and others in the series (Courtney 1996, 1997 and in prep.) show that, on the basis of juvenile calls and associated behaviour, the Australian 'rose- . tailed' parrots (Alisterus, Aprosmictus and Polytelis) constitute a tight group. The calls are similar among the species of the group, but different from those of other parrot groups. The present study failed to provide any evidence that the 'rose-tailed' parrots are part of the core platycercine ('broad-tailed' parrot) group, and thus supports the biochemical and genetic data on that point (Christidis, Schodde et al. 1991; Christidis, Shaw & Schodde 1991). In captivity, members of the genera Alisterus, Aprosmictus and Polytelis frequently hybridise within the group, but only with extreme rarity do these species hybridise with other parrots. Because of this, many authors (e.g. Lendon 1973; Smith 1975, 1979; Forshaw 1981) consider them monophyletic and more closely related to one another than to other parrots, a conclusion supported by biochemical and genetic data (Christidis, Schodde et al. 1991; Christidis, Shaw & Schodde 1991). This conclusion is borne out by the present study, in which the three-factor combination of distinctive juvenile food-begging call, use of a 'chick-to-parent' contact call and unique rose­ pink edging to the inner webs oflateral rectrices of these genera provides convincing evidence that all are descended from a single common ancestor, which had these characteristics. However, the variation in these features within the group, and the nature of its distribution among the species, suggest that generic interrelationships may be more complex than previously recognised. A comparison of the juvenile food-begging calls shows a notable difference in repetition rate and structure between the Princess Parrot on the one hand, and Regent and Superb Parrots on the other, with the Australian King-Parrot and Red-winged Parrot in between. A possible interpretation is that the Princess and King-Parrot share an 'ancestral' type of begging call, and that the begging call changed in the common ancestor of the Regent and Superb Parrots (after the Princess Parrot had split off). This suggests a wide genetic gap within Polytelis approaching that of subgenera at least, and there are other differences as well. Like the king- and red-winged parrots of Australia, New Guinea and the Indonesian islands, the Princess Parrot has a blue rump which contrasts strongly with the colour of the body. This is lacking in the Regent and Superb Parrots which instead have rumps the colour of their bodies. Before feathering, Princess Parrot chicks give the appearance of lacking 'second' down, which is more developed in the others. The dull brownish-pink bill colouration in Princess Parrot chicks differs from the yellowish to yellowish orange of Superb and Regent Parrot chicks respectively. Though the general similarity in the juvenile food-begging calls, 'chick-to-parent' contact-calls, lack of a blue rump and bill colouration of chicks suggests that the Regent and Superb Parrots are genetically closer to each other than either is to the Princess Parrot, there is a mixture of differences in all three species which supports Lendon's (1973) view that 'none are particularly closely related'. For example, the Regent Parrot lacks the brilliantly coloured irides possessed by all other 'rose-tailed' parrots and used by them in courtship display. In addition, some of the combinations of similarities (and differences) between any two of the three Polytelis species are present in the king-parrots on the one hand, and the red-winged parrots on the other. The Superb AUSTRALIAN 56 COURTNEY BIRD WATCHER

(female) and Princess Parrots share reddish thighs with the Australian King-Parrot. This feature is lacking in the Regent Parrot and also in the Red-winged Parrot. The Regent Parrot has pinkish-red secondary wing-coverts as do juvenile Red-winged Parrots, lacking in all the other species. The quavering whistling 'chick-to-parent' contact call uttered mainly in amongst the begging call is a shared trait in Superb, Regent and Red-winged Parrots and differs in manner from the 'two-syllable' whistle of the Princess Parrot and Australian King-Parrot, which invariably is uttered several times before begging and not thereafter. Juvenile bill colouration also follows the same pattern, with the former three species having yellowish orange whereas the latter two are basically pink in the mandibles. Despite the differences, the Polytelis genus holds together well in the long graduated shape of the tail and in the flocking contact calls of adults which seem similar (cf. Buckingham & Jackson 1988, 1990). The differences between some species of Polytelis, which in tum equate to similarities in these features to either the Australian King-Parrot or conversely to the Red-winged Parrot, argue against the merging of Alisterus in Aprosmictus. Certainly these two genera seem close on anatomical grounds (Smith 1979), but even here there must be a difference in the tarsi (with Australian King-Parrots scratching 'over the wing' and Red-winged Parrots scratching 'under the wing'; pers. obs.). Other than the similarity in the juvenile food-begging calls between the two (which tends towards the Princess Parrot in the Australian King-Parrot, and to the Regent Parrot in the Red-winged Parrot), there are no other outward similarities between the king- and red-winged parrots that are not shared also with at least one member of Polytelis. Even the oft-quoted similarities in tail length and pattern of flight do not hold exactly (see Lendon 1973). Perhaps the most significant difference between the Australian King-Parrot and Red-winged Parrot is in the form of the sound and frequency of utterance of the 'chick-to-parent' contact call, a difference which is the widest in the group, with the three species of Polytelis coming in between the two extremes. In many respects Aprosmictus seems no closer to Alisterus than to Polytelis. The present study therefore supports the status quo for all genera as maintained in Christidis & Boles (1994), and the arrangement of Peters (1937) for two subgenera in Polytelis. The genera Alisterus, Aprosmictus and Polytelis are often referred to collectively as the 'long-tailed' parrots (e.g. Christidis & Boles 1994), presumably to distinguish them from others such as the so-called 'broad-tailed' parrots or platycercines. Clearly this is a misnomer, as a perusal of measurements given by Forshaw (1978, 1981) will show. These demonstrate that the 'broad-tailed' parrots Platycercus, Bamardius, Purpureicephalus, Northiella, Psephotus, Cyanoramphus, Eunymphicus, Melopsittacus, Neopsephotus, Neophema and Pezoporus all have tails much longer than their combined head and body measurement, with the only exceptions being Lathamus, Prosopeia and, significantly, the former Geopsittacus (now merged with Pezoporus on molecular data: Christidis & Boles 1994). In the 'long-tailed' parrots, the three species of Polytelis and the Green-winged and Amboina King-Parrots have tails longer than their head and body, but the Australian King-Parrot, Red-winged Parrot and Timor Red-winged Parrot all have tails considerably shorter than their head-body measurement, so on balance the term 'long-tailed parrots' for the group has no validity. It is proposed, instead, that the members of this group be known as the 'rose-tailed parrots', to distinguish them from the 'broad-tailed parrots' and other groups and to emphasise their distinctiveness. Such is shown not only by their rose-coloured tails but also by their uniquely shaped pineal body, shared only with the Cockatiel, and which is different from that of other cockatoos and all other major groups of Australian parrots (Renzoni & Watters 1972). / The three distinct types of pineal organ in the Australian Psittaciformes are: (1) VOL. 17 (1) MARCH 1997 Vocalisations of Juvenile 'Rose-tailed' Parrots 57

wedge-shaped (Cacatua, Calyptorhynchus); (2) elongated, tubular (Nymphicus, polytelitines); (3) extremely long, tubular (lorikeets, Melopsittacus, platycercines; Renzoni & Watters 1972). It is thought that type 2 is the 'primitive' one, with types 1 and 3 evolving from it (Renzoni & Watters 1972, Courtney 1974, Adams et al. 1984). This interpretation suggests that the 'rose-tailed' lineage is the primary one for Australian parrots, reaching back to the split with the cockatoos and forwards in time to the lorikeets and 'broad-tails' . Therefore, the karyotype of Alisterus is not necessarily highly derived as thought by Christidis, Shaw & Schodde (1991), but the basic one from which the others are derived instead. The chromosomal data (diploid numbers etc.) of Christidis, Shaw & Schodde (1991) appear to reinforce the data on pineal organs that the connection between cockatoos and 'rose-tailed' parrots is closer than that between cockatoos and 1orikeets/'broad-tails'. Thus, the high diploid figure for Alisterus (76, cf. 76-80 for cockatoos, 72 for Cockatiel and (70 for lorikeets and platycercines) supports the notion of an ancient primary lineage for its group. The similarity among the species of 'rose-tailed' parrot in proteins (Christidis, Schodde et al. 1991) and begging calls (this study), implying recent differentiation within the group, does not contradict the notion of a possibly ancient lineage for the group's ancestor. In considering the other taxonomic questions, a major conclusion from this study is that Pesquet's Parrot is probably related to the 'rose-tailed' parrots, on the basis of its similar juvenile food-begging call and possession of a possible chick-to-parent contact call. This apparent connection was hitherto unsuspected by parrot systematists. The red on the ventral surface, vestigial red on the head of the male and the begging call of Pesquet's Parrot are so similar to those features of the Australian King-Parrot that, were it not for certain anatomical differences (Glenny 1957, 1959; Smith 1975; Hornberger 1980), it would be tempting to conclude that Pesquet's is close to the 'rose-tailed' group. The red forewings of Pesquet's Parrot are also similar to those ofthe adult male Red-winged Parrot, and both have much black in the plumage. The evidence presented herein suggests that Pesquet's and the 'rose-tailed' parrots must have had a common ancestor. The apparent derivation of the Pesquet' s Parrot karyotype from the Psittacula type (Christidis, Shaw & Schodde 1991) may reflect the 'ancestral' condition of the latter, shared with divergent other parrot groups. The present study failed to provide any indication that the Eclectus Parrot is closely related to the 'rose-tailed' parrots or to Pesquet's Parrot, or that the African Grey Parrot is related to Pesquet's; the respective begging calls are not alike (other than a slight resemblance between those of the Eclectus and African Grey Parrots). The biochemical evidence agrees that Eclectus is not closely related to the polytelitines (Christidis, Schodde et al. 1991). Limited evidence on begging calls suggests that Eclectus and Geoffroyus are closely related to each other but somewhat apart from Cyclopsitta, again in agreement with the biochemical evidence (cf. Christidis, Schodde et al. 1991). Christidis & Boles (1994) listed Cyclopsitta immediately after Eclectus and Geoffroyus, the systematic arrangement advocated by Courtney (1974). Finally, the Asian Psittacula parrots have been thought to be related to the 'rose­ tailed' parrots (e.g. Immelmann 1968, Forshaw 1978, Hornberger 1980), and included with them (and the Eclectus Parrot) as the 'coral-billed' parrots by Smith (1975), on morphological and behavioural grounds. The biochemical and genetic evidence suggests that Psittacula is not closely related to Eclectus or the polytelitines (Christidis, Schodde et al. 1991; Christidis, Shaw & Schodde 1991). However, hybrids have been recorded between Psittacula and the Princess Parrot and Eastern Rosella Platycercus eximius (G.A. Smith in litt.), so a genetic connection is certain if perhaps somewhat distant. [Another hybrid of interest, in view of the inability of Christidis, Schodde AUSTRALIAN 58 COURTNEY BIRD WATCHER et al. (1991) to place the lorikeets, is one between a male Rainbow Lorikeet Trichoglossus haematodus and a female Australian King-Parrot (Silva 1984).] The similarity in the basic note of the begging call between Psittacula and the Princess Parrot (Figure 17) therefore suggests retention of an ancestral character. Better tape­ recordings of Psittacula begging calls may help to resolve the issue. In view of the uniform morphology of Psittacula, notably a long graduated tail exceeding the head­ body length in all species, the name 'long-tailed parrots' is best restricted to that genus. Such usage recognises ti'J.at group's distinctiveness, and its possible link with the 'rose­ tailed' parrots.

Acknowledgements As the present paper is a continuation of Courtney (1996), the acknowledgements there apply equally here. In addition, I owe special thanks to the following people. My daughter Julianne Farrell of Sydney obtained for me and set up the computer used in this study. Mr Ederic Slater and Dr Peter Fullagar of Canberra spent much time over two days teaching me the rudiments of using the Audiograph program. In 1984 Dr R. Schodde read and commented on the preliminary draft of the present paper, and gave me invaluable literature. Mr A.C. 'Sandy' Hunt of Attunga bred and lent me the young 'rose-tailed' parrots used in this study. Mr G.A. Smith of Peterborough, England, sent me the recording of the Indian Ring-necked Parrot and discussed many aspects of parrots with me. Last but not least I thank Miss Rosemary Low ofNottinghamshire, England, for her sterling efforts in breeding, hand­ rearing and tape-recording the Eclectus, African Grey and Pesquet's Parrots and making these recordings available to me. I am also most grateful to the editor, Mr Stephen Debus, for his infinite patience and unstinting assistance with this manuscript, and the assistant editor, Mrs Julia Hurley, for her care and thoroughness during the production process. Two referees commented on a draft.

References Adams, M., Haverstock, P.R., Saunders, D.A., Schodde, R., & Smith, G.T. (1984), 'Biochemical systematics of the Australian cockatoos (Psittaciformes: Cacatuinae)', Aust. J. Zoo/. 32, 363-377. Anon. (1926), Official Checklist of the Birds ofAustralia, Royal Australasian Ornithologists Union, Melbourne. Bohner, F. (1986), 'The Red-browed Fig-Parrot', Aust. Avicult. 40, 95-98. Brereton, J. LeG. (1963), 'Evolution within the Psittaciformes', Proc. Xllllnternatl Omithol. Congr. 499-517. Buckingham, R. & Jackson, L. (Eds) (1988), A Field Guide to Australian Bird Song, Cassette 4, Sooty Tern to Superb Parrot, Bird Observers Club of Australia, Melbourne. --& --(1990), A Field Guide to Australian Bird Song, Cassette 5, Regent Parrot to Masked Owl, Bird Observers Club of Australia, Melbourne. Christidis, L. & Boles, W.E. (1994), The Taxonomy and Species ofBirds ofAustralia and its Territories, RAOU Monograph 2, Royal Australasian Ornithologists Union, Melbourne. · --, Schodde, R. , Shaw, D.D. & Maynes, S.F. (1991), 'Relationships among the Australo-Papuan parrots, lorikeets, and cockatoos (Aves: Psittaciformes): protein evidence', Condor 93, 302-317. --,Shaw, D.D. & Schodde, R. (1991), 'Chromosomal evolution in parrots, lorikeets and cockatoos (Aves: Psittaciformes)', Hereditas 114, 47-56. Condon, H.T. (1975), Checklist of the Birds of Australia: Non-passerines, Royal Australasian Ornithologists Union, Melbourne. Courtney, J. (1974), 'Comments on the taxonomic position of the Cockatiel', Emu 74, 97-102. --(1986), 'Age-related colour changes and behaviour in the Northern Funereal Black-Cockatoo Calyptorhynchus funereus funereus', Aust. Bird Watcher 11, 137-145. --(1996), 'The juvenile food-begging calls, food-swallowing vocalisation and begging postures in Australian cockatoos', Aust. Bird Watcher 16, 236-249. --(1997), 'The juvenile food-begging calls and associated behaviour in the lorikeets', Aust. Bird Watcher 17 (2), in press. Dyck, J. (1977), 'Feather ultrastructure of Pesquet's Parrot Psittrichas fulgidus', ibis 119, 364-366. Forshaw, J.M. (1973), Parrots of the World, 1st edn, Lansdowne, Melbourne. -- (1978), Parrots of the World, 2nd edn, Lansdowne, Melbourne. -- (1981), Australian Parrots, 2nd edn, Lansdowne, Melbourne. Glenny, F.H. (1957), 'A revised classification of the Psittaciformes based on the carotid artery arrangement patterns', Ann. Zool. , Agra 2, 47-56. --( 1959), 'Specific and individual variation in reduction of the clavicles in the parrots', Ohio J. Sci. 59, 321-322. Holyoak, D.T. (1972), 'The relation of Nymphicus to the Cacatuinae;, Emu 12, 77-78. VOL. 17 (1) MARCH 1997 Vocalisations of Juvenile 'Rose-tailed' Parrots 59

Hornberger, D.G. (1980), 'Funktionell-Morphologische Untersuchungen zur Radiation der Enilihrungs­ und Trinkmethoden der Papageien (Psittaci)', Bonner Zoologische Monographien 13, 1-192. Imrnelmann, K. (1968), Australian Parakeets, 2nd rev. edn, A. Ziemsen, Wittemberg. Lendon, A.H. (1973), Neville W. Cayley's Australian Parrots in Field and Aviary, Angus & Robertson, Sydney. Lepschi, B.J. & Courtney, J. E. (1992), 'Sexual dimorphism in immature Australian King-Parrots', Aust. Bird Watcher 14, 301-304. Low, R. (1986), Parrots, Their Care and Breeding, 2nd edn, Blandford Press, Poole. McAllan, I.A.W. & Bruce, M.D. (1988), The Birds of New South Wales- A Working List, Biocon Research Group, Sydney. -Peters, J.L. (1937), Checklist of Birds of the World, Harvard Mus . Comp. Zool. , Cambridge, Massachusetts. Reichenow, A. (1881), 'Conspectus Psittacorum. Systematische Uebersicht aller bekannten Papageienarten', J. Om., Lpz. 29, 1-49, 113-177, 225-289, 337-398. Renzoni, A. & Watters, P.A. (1972), 'Comparative observations on the pineal body of some Australian parrots', Aust. J. Zool. 20, 1-15 . Salvadori, T. (1891), Catalogue of Birds in the British Museum, Vol. 20, Psittaci, British Museum (Natural History), London. Silva, T. (1984), 'Some birds seen on a visit to Australia, New Zealand and Fiji', Avicult. Mag. 90, 108-112. Smith, G.A. (1972), 'Nesting and nestling parrots' , Avicult. Mag. 78, 155-165. -- (1975), 'Systematics of parrots', Ibis 117, 18-68. -- (1979), Lovebirds and Related Parrots, Inkata Press, Melbourne. Storr, G.M. (1984), 'Revised list of Queensland birds', Rec. West. Aust. Mus. suppl19. Thompson, D' A. W. (1899), 'On characteristic points in the cranial osteology of the parrots', Proc. Zool. Soc. Lond. 1899, 9-46. · Verheyen, R. (1956), 'Analyse du potential morphologique et projet d'une nouvelle classification des Psittaciformes', Bull. lnst. r. Sci. nat. Belg. 32 (55), 54 pp. von Boetticher, H. (1959), Papageien, A. Ziemsen, Wittenberg Lutherstad. Received 10 May 1996 •