VOL. 17 (2) JUNE 1997 61
AUSTRALIAN BIRD WATCHER 1997, 17, 61-70
The Juvenile Food-begging Calls and Associated Behaviour in the Lorikeets
by JOHN COURTNEY, 'Ashgrove', Swan Vale, via Glen Innes, N.S.W. 2370
Summary The juvenile food-begging calls of representative species of eight of the 11 genera of lorikeets were studied and compared with those of other Australian parrots and cockatoos. The food-begging call of most species is a uniform repeated hissing note. The calls of two genera, Charmosyna and Vini (incorporating Phigys), are a repeated tremulous trill. On the basis of their trilling food-begging calls, it is suggested that the Psittaculirostris fig-parrots may be related to Charmosyna and Vini.
Introduction The present paper, on the juvenile food-begging calls and begging postures of the lorikeets, is a continuation of a study covering these aspects in all groups of Australian parrots and cockatoos (Courtney 1996, 1997). The methods used and a discussion of food-begging calls were outlined in those papers. It is emphasised that the project, for all parrot groups studied, is simply a survey and qualitative description of these calls. Where differences in these calls between species or groups have been great or, conversely, where similarities have been undeniable even without statistical analysis, taxonomic conclusions have been drawn where appropriate. The lorikeets are a large assemblage of parrots, with 55 species in 11 genera, which occur widely through the Australasian Region and east into Polynesia, with one species extending north to the Philippines (Forshaw 1978). Six species in three genera occur in mainland Australia (Christidis & Boles 1994). On protein evidence, the lorikeets appear to be closely related to one another but their relationship to other parrots is obscure (Christidis et al. 1991). The food-begging calls of representative species of eight of the 11 lorikeet genera are described briefly and discussed. Possible taxonomic implications are discussed also, including a putative relationship between the Psittaculirostris fig-parrots and some lorikeet genera. For the purpose of this paper, Phigys is synonymised with Vini (following Low in press).
Methods Captive-bred young of five Australian species were reared by hand, and wild broods of some species were also studied. No Purple-crowned Lorikeets Glossopsitta porphyrocephala were available for study. Sample sizes of birds tape-recorded were: Musk Lorikeet G. concinna two broods of two chicks (B/2 X 2, unrelated) in the wild, B/2 x 1 in captivity; Little Lorikeet G. pusilla B/4 x 3 (unrelated) in the wild, B/4 x I in captivity; Varied Lorikeet Psitteuteles versicolor B/3 X 1 captive; Scaly-breasted Lorikeet Trichoglossus chlorolepidotus and Rainbow Lorikeet T. haematodus rubritorquis single, typical examples of captive young. The begging postures were noted and food-begging calls recorded on tape, which were then analysed using Audiograph 2.0 for Macintosh computer (see Courtney 1997). It is stressed that, from 11 years' prior experience of nestling parrots, the calls taped and analysed were typical of what were by then known to be the normal, species-specific calls; resulting audiographs presented here are representative examples for their species. Tape-recordings of the begging calls of captive-bred young of 10 species from six genera outside mainland Australia were also analysed (tapes provided by R. Low of England unless stated): Yellow streaked Lory Chalcopsitta scintillata B/1 x 2 (one brood from G. A. Smith of England); Dusky Lory Pseudeos fuscata B/ 1 x 2; Tahiti Blue Lory Vini peruviana Bll x 2 (unrelated); Red Lory Eos bomea B/2 X 1; and single, typical young of Duyvenbode's Lory Chalcopsitta duivenbodei, Black-capped Lory Lorius lory, Rainbow Lorikeet Trichoglossus haematodus (New Guinean form, race unknown), Goldie's Lorikeet Psitteuteles goldiei, Collared Lory Vini solitaria and Blue-crowned AUSTRALIAN 62 COURTNEY BIRD WATCHER
Lory V. australis (the last two species from G. Campbell, San Diego Zoo). A tape-recording of a single young Salvadori's Fig-Parrot Psittaculirostris salvadorii was provided by G.A. Smith. A composite cassette-tape of all recordings used has been lodged with the Australian National Wildlife Collection, CSIRO, Canberra. Comparison with other groups can be made by reference to the companion papers in this series (cockatoos: Courtney 1996; 'rose-tailed' parrots and others: Courtney 1997; platycercines: Courtney in press).
Results
Defensive behaviour in young When disturbed in the nest or (in captivity) when a nest-box is opened, nestling Australian lorikeets run rapidly over the floor of the nest and utter a loud buzzing call (pers. obs.), although many other species are silent or become so after 2-3 seconds (R. Low in litt.). This contrasts with the behaviour of other young parrots and cockatoos observed (except Budgerigars Melopsittacus undulatus), which sit in one place. Nestling Budgerigars behave like young lorikeets. Nestling Salvadori's Fig Parrots are also said to run rapidly about like lorikeets (G.A. Smith in litt.).
Food-begging posture All Australian lorikeets observed adopted a fairly horizontal stance and rapidly quivered their partly outspread wings in a manner reminiscent of young passerines. This behaviour is unlike that of any other species of parrot or cockatoo observed (Courtney 1996, 1997, in press).
Juvenile food-begging calls The lorikeets can be split into two categories based on the food-begging call: those species that utter a hissing note (most genera, Figures 1-12) and those that utter a tremulous trill ( Vini, Figures 13-16). Both types of call are of similar length and repeated at a similar rate, and do not change in basic pattern with age from downy to feathered young. In those species that hiss, audiographs reveal that the somewhat sharp-sounding, high-intensity repeated note uttered when young are hungry and being fed, begins with a structureless hiss abruptly giving way to a momentary high-energy, high-pitched 'blip' which precedes a structured hiss (Figures 1-12).. In lower intensity begging, as is typical of much calling, often only the structured hiss is given. The hissing call is similar among the various species, and the rate of repetition varies with degree of hunger. The length of note and emphasis in pitch among different species are clearly shown in the audiographs (Figures 1-12). The species and genera known to call in the hissing manner just described are as follow: Chalcopsitta (Duyvenbode's and Yellow-streaked Lories, Figures 1-2); Lorius (Black-capped Lory, Figure 3); Eos (Red Lory, Figure 4); Pseudeos (Dusky Lory, Figure 5); Trichoglossus (Rainbow and Scaly-breasted Lorikeets, Figures 6-8); Psitteuteles (Varied and Goldie's Lorikeets, Figures 9-10); and Glossopsitta (Musk and Little Lorikeets, Figures 11-12). A videotape of Musschenbroek's Lorikeet Neopsittacus musschenbroekii (provided by R. Low) shows that it also utters a hissing food-begging call; unfortunately the recording is too noisy to analyse by audiograph, but Low (in litt.) has subsequently confirmed from hand-reared birds that it hisses like Trichoglossus . Lorikeets of the genus Vini (including Phigys) occur well east of Australia and New Guinea ·in the far-flung Pacific islands. Tape-recorded food-begging calls of hand reared Tahiti Blue Lories (Figures 13-14) show for the first time that species of Vini utter a brief repeated trill instead of a hiss. A recording of a single two-week-old, parent-reared young Collared Lory consisted of some~what high-pitched tremulous VOL. 17 (2) JUNE 1997 V ocalisations of Lorikeets 63
Duyvenbode 's Lory 10.00
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Figure 1. Juvenile food-begging call of Duyvenbode's Lory Chalcopsitta duivenbodei.
Yell ow-streaked Lory
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Figure 2. Juvenile food-begging call of Yellow-streaked Lory Chalcopsitta scintillata.
Black-capped Lory 10.00 kHz
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Figure 3. Juvenile food-begging call of Black-capped Lory Lorius lory. AUSTRALIAN 64 COURTNEY BIRD WATCHER
Red Lory
Figure 4. Juvenile food-begging call of Red Lory Eos bomea.
Dusky Lory 10.00
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Figure 5. Juvenile food-begging call of Dusky Lory Pseudeos fusctita.
Rainbow Lorikeet (New Gu inea form) 10.00
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Figure 6. Juvenile food-begging call of Rainbow Lorikeet Trichoglossus haematodus, New Guinea for1n (race unknown). VOL. 17 (2) JUNE 1997 Vocalisations of Lorikeets 65
\Rainbow Lorikeet 10.00 kHz
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4.00 kHz !~ 3.00 kHz f-·'· ::./;.;;~~.-::-.., -;;-:-::;;;.. - ~ 2.00 kHz .
LOOkH' ..~~ ~ ~ .?24fSP!Qf24i¥Mii4& 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
Figure 7. Juvenile food-begging call of Rainbow Lorikeet Trichoglossus haematodus rubritorquis.
Scaly-breasted Lorikeet 10.00
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Figure 8. Juvenile food-begging call of Scaly-breasted Lorikeet Trichoglossus chlorolepidotus.
\Varied Lorikeet
Figure 9. Juvenile food-begging call of Varied Lorikeet Psitteuteles versicolor. AUSTRALIAN 66 COURTNEY BIRD WATCHER
Goldie ' s Lorikeet 10.00
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Figure 10. Juvenile food-begging call of Goldie's Lorikeet Psitteuteles goldiei.
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Figure 11. Juvenile food-begging call of Musk Lorikeet Glossopsitta concinna.
Little Lorikeet 10.00
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Figure 12. Juvenile food-begging call of Little Lorikeet Glossopsitta pusilla. VOL. 17 (2) JUNE 1997 V ocalisations of Lorikeets 67
ILOOkHz~.--.~--r-~~--.-.--.~--~~~--.-.--r~--.-.-~-.--.--r~ > Tahiti Blue Lory (1) 10.00 kHz
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Figure 13. Juvenile food-begging call of Tahiti Blue Lory Vini peruviana, first chick.
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Figure 14. Juvenile food-begging call of Tahiti Blue Lory Vini peruviana, second chick.
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Figure 15. Juvenile food-begging call of Collared Lory Vini solitaria (with some background hissing by an unidentified individual). AUSTRALIAN 68 COURTNEY BIRD WATCHER
11.00 Blue-crowned Lory 10.00
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Figure 16. Juvenile food-begging call of Blue-crowned Lory Vini australis.
Tahiti Blue Lory Yellow-st reaked Lory G7 -~ kHz ·-\53 - 2-
0 1 sec
Figure 17. Juvenile food-begging calls of Tahiti Blue Lory and Yellow-streaked Lory on Kay Sonagram using wide-band (300 Hz) filter.
Salvadori's Fig- Parrot
Figure 18. Juvenile food-begging call of Salvadori's Fig-Parrot Psittaculirostris salvadorii. VOL. 17 (2) JUNE 1997 Vocalisations of Lorikeets 69 trilling (Figure 15), which sounded much the same as 16-day-old chicks of the Blue crowned Lory (Figure 16). A chick of the Collared Lory was said to utter a curious note somewhat like the loud ticking of a clock, and agitated its whole body when begging for food (Bahr 1912 in Forshaw 1978). When analysed by Kay Sonagram using the wide-band (300Hz) filter, the food-begging trill of the Tahiti Blue Lory can be seen to be formed by a series of brief, individual high-pitched notes in the shape of an inverted 'V' or chevron, placed closely together (Figure 17). Young of Cho.nnosyna lorikeets (Papuan C. papou, Josephine's C. josefinae, Little Red C. pulchella and Red-flanked C. placentis) trill like those of Vini when begging for food (R. Low in !itt.). The food-begging call of the Whiskered Lorikeet Oreopsittacus aifaki has not been recorded, but the chicks are morphologically identical to those of Vini and Cho.rmosyna (R. Low in !itt.). The tremulous food-begging trill of Salvadori's Fig-Parrot sounds similar to the tremulous trill uttered by young of the Vini group of lorikeets, but is not identical when analysed by audiograph (Figure 18).
Discussion and conclusions The results indicate that the juvenile food-begging calls of lorikeets are very similar across genera but different from those of the cockatoos, 'rose-tailed' parrots and platycercines (see Courtney 1996, 1997, in press). The uniform juvenile food-begging call of the lorikeets that hiss suggests that the food-begging call is a highly conservative character that has not changed over the evolutionary history of the group. The results of this study also support biochemical evidence (Christidis et al. 1991) that the lorikeets constitute a tight monophyletic group. Within the lorikeets, one can recognise two apparently closely connected groups: those that hiss (Trichoglossus, Glossopsitta and relatives) and those that trill (Cho.rmosyna, Vini and probably Oreopsittacus), a conclusion also reached by R. Low (in litt.) on the basis of adult behaviour and chick morphology as well as fledgling food-begging calls. The begging calls of one lorikeet group (Vini) suggest a possible connection with the fig-parrots. Like the structure of the food-begging call of the Tahiti Blue Lory, even when unfiltered the begging notes of Salvadori's Fig-Parrot clearly consist of inverted Vs or chevrons. This, too, is the shape of the basic note or fundamental tone in the Indian Ring-necked Parrot Psittacula krameri and the Princess Parrot Polytelis alexandrae (Courtney 1997), and may reflect retention of a primitive trait. Lorikeets and fig-parrots also share another character. An underwing stripe that is permanent in all individuals regardless of age or sex occurs only in three groups: lorikeets (most genera), some fig-parrots, and the most 'primitive' barred platycercines (Budgerigar, Ground Parrot Pezoporus wallicus and Night Parrot P. occidentalis). However, in Charmosyna the underwing stripe is present only in juveniles, and only in some species (R. Low in !itt.). The Psittaculirostris fig-parrots also show lorikeet like plumage features including lanceolate cheek feathers and pointed tails; they resemble small lorikeets in some respects (see photographs in Coates 1985). Feather lice (Mallophaga) have taxonomic significance, because louse genera are often exclusive to particular bird genera (McClure & Ratanaworabhan 1973, Clay et al. 1985). Lorikeets host a genus, Psittaconirmus, found on no other parrots (Forshaw 1978), which suggests long isolation on lorikeets. Another genus, Pacifimenopon, is also exclusive to lorikeets; a third, Eomenopon, is found only on lorikeets, fig-parrots and pygmy-parrots Micropsitta spp. (G.A. Smith in litt., from British Museum [Natural History] collections). This evidence further supports the notion of a distant relationship between lorikeets and fig-parrots. AUSTRALIAN 70 COURTNEY BIRD WATCHER
Even though lorikeets appear to have radiated in relatively recent times, the typical hissing begging call bears no resemblance to the begging calls of other parrots, suggesting in tum that the ancestrallorikeet stock may not be of recent origin. The results presented here agree with the conclusions of Christidis et al. ( 1991) that most species of lorikeets appear to be closely related and possibly r~ently evolved, but that the group as a whole is difficult to place in the general assemblage of Australian parrots. From this study, one could hypothesise that the Papuan/Polynesian 'trilling' lorikeets (Vini and Chaniwsyna) represent an older group, retaining connections with the fig-parrots, and the Papuan/ Australian 'hissing' lorikeets represent more recent speciation in a 'sister' lineage. A fruitful avenue of research might be to investigate the genetic distance between the 'trilling' lorikeets and the fig-parrots.
Acknowledgements As this paper is a continuation of Courtney (1996, 1997), all acknowledgements in those papers apply equally here. Special thanks are due to Miss Rosemary Low of England for her sterling efforts in supplying so many tapes of the begging calls of lorikeets, for commenting on a draft of this paper, and for her information and preview of her lorikeet encyclopaedia text; and to Ms Gwendolyn Campbell of San Diego Zoo for her invaluable contribution of a tape of Vini solitario and V. australis. My gratitude also to George A. Smith of England for a tape of Psittaculirostris, and for information on feather lice. Mr A. C. Hunt of Attunga, N.S.W ., gave me unrestricted access to the lorikeets in his aviaries. Neville and Enid Connors of Grafton, N.S.W., kindly allowed me to tape-record young Varied Lorikeets in their collection. An anonymous referee commented on the manuscript. For all this I am most grateful. References Bahr, P.H. (1912), 'On a journey to the Fiji Islands, with notes of the present status of their avifauna, made during a year's stay in the group, 1910-1911' , Ibis 9th ser., 6, 282-314. Christidis, L. & Boles, W.E. (1994), The Taxonomy and Species ofBirds ofAustralia and its Tenitories, RAOU Monograph 2, Royal Australasian Ornithologists Union, Melbourne. --, Schodde, R., Shaw, D.D., & Maynes, S.F. (1991), 'Relationships among theAustralo-Papuan parrots, lorikeets, and cockatoos (Aves: Psittaciformes): protein evidence' , Condor 93, 302-317. Clay, T ., Hutson, A.M. & Baker, A.S. (1985), 'Ectoparasite', in Campbell, B. & Lack, E. (Eds), A Dictionary of Birds, 170-172, Poyser, Calton. Coates, B.J . (1985), The Birds of Papua New Guinea, vol. 1, Non-passerines, Dove, Brisbane. Courtney, J. (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 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 folgidus', Aust. Bird Watcher 17, 42-59. --(in press), 'The juvenile food-begging calls and associated aspects in the Australian " broad- tailed" (platycercine) parrots', Aust. Bird Watcher 17. Forshaw, J.M. (1978), Parrots of the World, 2nd edn, Lansdowne, Melbourne. Low, R. (in press), Encyclopedia of Lories and Lorikeets, Hancock, Canada. McClure, H.E. & Ratanaworabhan, N. (1973), Some Ectoparasites of the Birds of Asia, privately published, Bangkok. Received 9 October 1996 •