152 AUSTRALIAN FIELD 2005,22, 152- 157 Breeding Displays and Calls of the Striped at Baradine, New South Wales

K.A. WOOD1 and ANDREW J. LEY2 1 7 Maralinga Drive, Ashmore, Queensland 4214 2 19 Lynches Road, Armidale, New South Wales 2350

Summary Displays and calls of the Striped Honeyeater Plectorhyncha lanceolata were recorded during 144.8 hours of observation of a breeding pair at Baradine, northern New South Wales, in spring 2003. Wing-quivering displays by one of the pair and mutual wing-quivering displays were observed; solo and duet singing were heard. Territorial song-flights were observed during each of the 24 days that the were studied. Phrases of calls uttered during solo and duet singing were different from phrases sung during territorial song-flights. Calls other than songs that were recognised were an alarm call, a contact warble near the nest and a food-begging call of the nestling. Responses to calls that had been taped by other workers are described.

Introduction Communication between birds consists of visual and vocal displays that enable messages to be conveyed to breeding partners, cohorts, rivals and offspring (Rowley 1982). Among Australasian perhaps the most striking displays are those of the so-called 'arena birds', the bowerbirds (Cooper & Forshaw 1977; Diamond 1986) and the Superb Lyrebird Menura novaehollandiae (Smith 1988). Less spectacular are the displays of Australian (family Meliphagidae). Although noisy and acrobatic, no species of honeyeater prepares an arena for display or courtship. Eight species, however, are known to perform a congregation display known as the corroboree (see Pyke & O'Connor 1989) and at least 13 species, mainly in the genera Lichenostomus and Phylidonyris, perform song-flights (Higgins et al. 2001 ). Except for the Noisy Miner melanocephala (Dow 1975), comprehensive knowledge about honeyeater communication is generally lacking because the displays and calls of most species 'have not been methodically studied and described' (H.A. Ford in Higgins et al. 2001). With respect to the Striped Honeyeater Plectorhyncha lanceolata only Lord (1956) and Longmore (1991) mentioned a wing-quivering display; only Longmore (1991) and Pizzey & Knight (1997) stated that the species performs song-flights; and only Slater et al. (1986), Schodde & Tidemann (1986) and Longmore (1991) reported duet-singing. In this paper, we describe displays and calls of a breeding pair of Striped Honeyeaters at Baradine, New South Wales (30°57'S, 149°5'E), in spring 2003. Responses to calls that had been taped by other workers are also described.

Methods From early September to late November 2003, a pair of breeding Striped Honeyeaters was watched during all nesting stages for a total of 144.8 hours during 24 days (Table 1 in Wood & Ley 2005). Three nesting attempts were made but most observations were at nest 2 (the only nest which produced young) or within the breeding home-range around nest 2 (subsequently calculated = 4.2 ha) (see Wood & Ley 2005). The observations were part of a larger study on breeding and foraging (Wood & Ley 2005) during which the female and male were colour- VOL. 22 (3) Striped Honeyeater Breeding SEPTEMBER 2005 Displays and Calls 153 banded and sexed by measurements on 17 and 28 October respectively. The number of times each display or call was given by each parent in the various nesting phases is presented elsewhere with other breeding information (Wood & Ley 2005). Playback of tape-recorded calls was used to assist with catching the parent birds, determination of the home-range around nest 2 and location of colour-banded Honeyeaters near nest 3. Tapes used were those of Crouch & Crouch (1977), Gillard (1988) and Buckingham & Jackson (1995) selected randomly. Taped calls were mostly played from a hand-held speaker, with the operator sometimes hidden, or from the fixed speaker in a car with the door open and the operator standing beside the vehicle. Because we were conscious of possible harmful effects on the breeding pair (see McClure 1984; Geering 1998), the number of playback trials was minimised. Each trial was terminated after three or four repeated calls or whenever a response was noticed. A total of 28 playback trials was conducted: 11 on 30 September when the pair was building the second nest and 15 after 10 November, when the same pair was attending nest 3. The other two trials were on 17 and 28 October, when playback was used to possibly assist with mist-netting the pair near nest 2, which was located 2.8 m above ground. Calls were verbalised into syllables and phrases. Verbalisations considered were chi1p chilp cheny cheny (North 1909), chirp chi1p cheny cheny chirp (Officer 1975; Hill 1976), cher cher cheny cheny chip chip (Stewart 1976) cher cher cheny cheny (Slater eta!. 1986; Simpson & Day 1989), cherree cherree chirrarip, free wheat peeler peeler (Pizzey & Knight 1997) and quirrip quarreep quirrip quarreep quirrip quarreep (Morcombe 2000). Other verbalisations were devised for calls that were difficult to transliterate as above. In this paper, hyphens are shown between syllables to assist with pronunciation; spaces between syllables indicate short intervals of time.

Results

Wing-quivering by one of the pair Quivering of both wings simultaneously, by one of either sex, was observed daily but there were numerous variations in performance. Sometimes the amplitude of wing displacement was exaggerated, sometimes the duration of quivering was sustained. Mostly, this display was given by a Honeyeater upon landing on a branch either near the nest (<20m) or near the breeding partner.

Mutual wing-quivering Although observed much less frequently than wing-quivering by a single bird, mutual wing-quivering had similar variations in execution. In the majority of instances the presumed male and female performing the display were less than 1 m apart. Mostly, they were near the nest ( <20 m). This display was often initiated by the landing of one partner close to the other. The most impressive mutual display involved sustained and exaggerated wing-quivering by both birds facing each other closely on the same branch (bills < 10 em apart) while singing a duet.

Solo song, usually while stationmy Songs delivered by a single bird were commonly heard, and were mostly sung while the bird was perched on a branch of a tree. Occasionally these songs were uttered by an incubating or brooding bird or by a parent flying about in the vicinity (within 50 m) of the nest below the tree-tops. Solo songs varied in length from a particular phrase sung once to a cheerful rollicking melody of the same phrase repeated three to five times. We found that individual phrases of the solo song were best verbalised as either chim chee-a-ree or chim chim cheree of duration of about 1 second (Table 1). AUSTRALIAN 154 WOOD&LEY FIELD ORNITHOLOGY

Table 1 Phrases recognised in 56 songs given by breeding Striped Honeyeaters (sex not determined) at Baradine, NSW.

Number of times recognised

Phrase During a song-flight As a solo song Chee-a-ree chree chirrup 15 Nil Chee-a-ree chirrup 5 Nil Chim chee-a-ree Nil 16 Chim chim cheree Nil 19 Chee-a-rim cheree Nil

Duet, usually while perched Duets were given infrequently but contained the same verbalised phrases as solo songs (Table 1). Mostly, the two birds giving the duet were perched close to each other ( < 10 m) but occasionally they were perched farther apart (60 m maximum). Rarely, one bird was stationary and the other was in flight near the nest. Only a few duets were totally synchronised; usually they were started by one parent, with the other joining the rendition soon after commencement. As with solo songs, duets varied in duration to a maximum of about 5 seconds if the same phrase was repeated five or six times. Spatially, more duets were within 100 m of the nest than farther away and few, if any, were delivered when one singer was close (<20m) to the home-range boundary.

Response song Occasionally one partner uttered phrases of a solo song and the partner responded by uttering phrases immediately afterwards. It was sometimes difficult to differentiate between a response song and a poorly synchronised duet.

Territorial song-flight Song-flights were observed over the territory and home-range during each day of the study. A majority of songs uttered in flight were sustained, rich in timbre and seemed to rise slightly in pitch and volume. Phrases were best verbalised as either chee-a-ree chree chirrup or chee-a-ree chirrup (Table 1). Usually six to eight phrases were sung in succession (song duration c. 7-9 seconds). Flights with solo song were often horizontal over distances of 40-50 m about 15- 20 m above the tree-tops; the compass bearings of these were distributed randomly over the breeding territory. Some were above the nest in various directions, others were above and roughly parallel to the territory boundary. Some were in a sweeping arc rather than in a straight line. In the most impressive, and less frequent, song­ flights, the displaying bird gained height rather than flew horizontally. The angle of ascent was c. 30-40° with the launching point usually near the top of a tree. In ascending territorial song-flights, the displaying bird sang crescendo until it was some 50-60 m above the tree-tops, when it descended steeply in silence and perched in trees below. The launching point for a few ascending song-flights was VOL. 22 (3) Striped Honeyeater Breeding SEPTEMBER 2005 Displays and Calls 155

a branch about 10m above the ground, with the ascent terminating just above the trees (tree height 22m).

Contact warble The male and female sometimes communicated by soft ( = low-volume) contact notes while near ( < 15 m) the nest in the nest-building stage. Typically, the contact warble was given by both parents almost simultaneously when they were together at the nest, when one was at the nest and the other was approaching with nesting material or when both were flying towards the nest. In the last instance, one or both parents may have been carrying nesting material. The contact warble was difficult to verbalise, consisting of a variety of notes given for a duration of about 5-10 seconds.

Alarm call The only alarm call heard was a short harsh note verbalised as zeet. It was used by parent birds when dive-bombing other avian species near the nest and when dive-bombing a person inspecting the contents of the nest.

Food-begging call of nestling The begging call of the single nestling was just audible a few metres from the nest about 5 days after hatching. It comprised a series of single notes given when the parent birds were nearby. Reminiscent of bleating or whimpering, these peep begging calls could be heard by listening intently from a distance of about 10m when the nestling was 8 days old. The begging-call rate on that day was about 30- 40 peeps per minute during the three bouts of begging observed.

Response to playback of taped calls Of 28 playback trials, 18 elicited a response. If the location of the speaker was clearly within the breeding home-range, the usual response was for one of the breeding pair to immediately fly towards it, just above the trees, and land in the top of a nearby tree. The responding bird often flew 40-50 m (c. 90 m maximum) to reach the speaker and sometimes landed 20-30 m beyond it (maximum flight = 120m). Occasionally the responding bird searched for the source of the playback after landing and sang a solo song. Five responses involved counter-singing or song-flights by Striped Honeyeaters in adjacent territories. One trial brought three Striped Honeyeaters together, and physical conflict occurred between two of them. In the ten trials where no response was detected, we concluded that the speaker was probably located close to (within 20 m) the home-range boundary and the breeding occupants were > 100 m away, or else the speaker was outside the breeding home-range of nest 2. Of 15 trials conducted after both parents were colour-banded, only four failed to elicit a response. In the eleven successful trials, the male responded four times, both parents responded once, and the colour of the band on the responding bird was not identified on six occasions. In the trials on 17 and 28 October, when playback was used as a possible aid to mist-netting, the speaker was 3m from the nest and the operator was hidden. The net was 3m high and about 1m from the nest. Both parents responded immediately to the played call by flying into the nest-tree and calling repeatedly (possibly AUSTRALIAN 156 WOOD&LEY FIELD ORNITHOLOGY

duetting) while moving about in an agitated manner. They moved about above the net, at a height of 3-5m, and gave animated calls similar to those recorded by Buckingham & Jackson (1995) and by Crouch & Crouch (1977), when Striped Honeyeaters in South Australia responded 'enthusiastically' to playback. While calling, the Honeyeaters peered about as if searching for a conspecific intruder but were not drawn to the speaker itself. The female and male were mist-netted 60 and 80 minutes respectively after cessation of playback on 17 and 28 October respectively.

Discussion Interpretation of the above displays and calls is necessary to provide an understanding of the reproductive biology of the Striped Honeyeater. We propose that the wing-quivering observed was pair-bonding, courtship or solicitation. Perhaps in its mildest form, when given by one of the breeding pair, it was pair­ bonding whereas mutual wing-quivering could have been solicitation. In passerines only a very small number of solicitations lead to copulation (Wood 2000; Higgins et al. 2001), an activity which we did not see during this study. However, in most of the honeyeaters that have been studied comprehensively, copulation follows wing­ quivering (e.g. Regent Honeyeater Xanthomyza phrygia, Bell Miner Manorina melanophrys, Yellow-tufted Honeyeater Lichenostomus melanops and New Holland Honeyeater Phylidonyris novaehollandiae: Higgins et al. 2001 ). In the Noisy Miner, copulation is rarely observed but does not follow a display involving wing movement (Dow 1975). The possibility that wing-quivering by the Striped Honeyeater was a greeting display, as suggested for some honeyeater species (Higgins et al. 2001), was unlikely because most greeting displays are by colonially nesting species during changeovers at their nests (Marchant & Higgins 1990). Song-flights were interpreted as territorial or home-range announcements. Firstly, they seemed to be directed at conspecific rivals in adjacent territories. Secondly, counter-singing by rival Striped Honeyeaters in adjacent territories was regularly noted. Lastly, because song-flights were performed by one bird, without any interaction from the breeding partner, we did not interpret them as a pair­ bonding or courtship activity. These considerations provide the basis for our assessment that these flights were probably territorial song-flights. The function of duetting in songbirds is equivocal but probably serves to maintain pair-bonds through individual recognition or to advertise the existence of a strong pair-bond and thereby discourage conspecific rivals from entering the territory (Farabaugh 1982 and references therein). It could be argued that synchronised duets from breeding partners perched next to each other are more effective in advertising a strong bond and territory ownership than non­ synchronised duets from partners that are separated. Given that song-flights were used to advertise the territory at Baradine, that only a small proportion of duets was synchronised and that the breeding partners giving the duet were occasionally perched apart (60 m maximum), it is possible that the primary function of duetting by the Baradine pair was to maintain pair-bonding through individual recognition. These duets, however, may also serve to discourage conspecific rivals from entering the breeding home-range.

Acknowledgements We thank Ron Nixon, Barry Johnson and Neil McDonald for making us welcome at Camp Cypress, Baradine. David Drynan expedited colour-marking approval from Environment VOL. 22 (3) Striped Honeyeater Breeding SEPTEMBER 2005 Displays and Calls 157

Australia. We also thank referees Hugh Ford and Damon Oliver for critically commenting on the manuscript.

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Received 26 April 2004 •