88 AUSTRALIAN FIELD 2005 , 22, 88-103 Nesting and Foraging by a Pair of Striped at Baradine, New South Wales

K.A. WOOD1 and ANDREW J. LEY2 17 Maralinga Drive, Ashmore, Queensland 4214 219 Lynches Road, Armidale, New South Wales 2350

Summary A breeding pair of Striped Honeyeaters Plectorhyncha lanceolata was observed at three nests at Baradine, northern New South Wales, for 43.3, 25 .8 and 42.5 hours overall in the nest-building, incubation and nestling phases respectively during spring 2003. The breeding home-range around one nest (nest 2) was 4.2 ha, dominated by mature White Cypress Pines glaucophylla. Both parents defended a small area within a radius of about 20m from the nest and were engaged almost equally in nest-building, incubation and feeding a single nestling. The mean rate at which both parents fed the nestling was 11.3 (range 4.3-16.7) visits/hour. The female's brooding effort was three times greater th an that of the male, but the male uttered most (88%) solo songs and performed most (if not all) territorial song-flights. Overall, there were at least twice the number of calls and displays during nest-building as during incubation or while feeding the young. Food, foraging, and use of feet, as we ll as flight and comfort behaviour, are also described.

Introduction In the family Meliphagidae, the Striped Plectorhyncha lanceolata is a medium-sized monotypic species (length 22 em, weight 40 g) with a spear-shaped bill, a black-and-white striped head and long buffy-white lanceolated feathers on the upper breast (Schodde & Tidemann 1986; Longmore 1991). It breeds throughout its range from northern Queensland, through New South Wales to northern Victoria and eastern South Australia (Blakers et al. 1984).1ts preferred habitat is semi-arid woodlands dominated by eucalypt, or cypress pine trees (Higgins et al. 2001). Some aspects of its breeding biology are known (see Higgins et al. 2001) but it seems that only one quantitative study has been undertaken. Moffatt et al. (1983) watched a single nest with young for 7 hours over 3 days near Meandarra, Queensland, and observed cooperative feeding of nestlings. Nevertheless, Higgins et al. (2001) concluded that Striped Honeyeaters usually breed as simple pairs with both sexes building the nest, incubating the eggs and feeding the young. The only reported systematic foraging study was in south-western New South Wales in July 1993, when Franklin & Alley (1995) watched non-breeding Striped Honeyeaters as they foraged in Black Oak and mallee woodland. In 21 minutes of cumulative sampling, Franklin & Alley (1995) found that Striped Honeyeaters spent the majority of time (80.8%) feeding in the canopy (2-10 m high) of woodland trees. They suggested that arthropods, rather than or lerp, were the main source of food. In order to further our knowledge of the Striped Honeyeater, we obtained sample data on a breeding pair at the showground at Baradine, NSW (30°57'S, 149°05'E), during 24 days in spring 2003. Observations were made of parent attending nests and foraging in the breeding home-range. The female and male were colour-banded about halfway through the study. In this paper, information is presented on some aspects of the breeding biology and foraging ecology. VOL. 22 (2) JUNE 2005 Striped Honeyeater Nesting and Foraging 89

Methods Three nesting attempts were observed (Table 1, Figure 1). The first nest was watched for 30 minutes on 11 September 2003 when it appeared to be completely built and was attended by two Striped Honeyeaters, but incubation had not commenced. Ten days later it looked dishevelled and was inactive apart from one visit by a Striped Honeyeater that collected some nest-material and carried it to a second nest 75 m away. Subsequently, nest 2 was watched for 30.5, 25.8 and 42.5 hours respectively in the nest-building, incubation and nestling phases. Given that we later determined that the first nest was located in the home-range around nest 2, it seems likely that the same pair built both nests. Three eggs were being incubated in nest 2 on 13 October but only one hatched (Table 1). The nestling was estimated to be c. 4 days old on 27 October and, since the nestling peri od is c. 17 days (Higgins et al. 2001), its projected fledging date was about 9 November (Figure 2). In the mornings of 7, 8 and 9 November, this nestling was fed by a parent but was absent at approximately 0900 h on 10 November (R. Nixon, pers. comm.). Because th e colour-banded parents, and not the juvenile, were found during 150 minutes of searching on the afternoon of 10 November (Table 1), we concluded that the young died on 9 or 10 November. At 1155 h on 11 November, the breeding pair was found again, carrying the nest-materials from nest 2 to a third nest. Located 440 m from nest 2, nest 3 was watched cumulatively fo r 12.3 hours on 4 days until 24 November. It was abandoned about 24 November when c. 70% constructed.

Foraging Foraging data were acquired by focal- sampling (Altmann 1974). Parent birds were followed opportunistically during foraging bouts which lasted > 1 minute. Tree species in which the Honeyeaters foraged were recorded, as well as the time spent foraging at va ri ous heights, estimated to the nearest metre. Both parents were followed and watched until they ceased foraging or were lost from sight. The foraging method (sensu Recher eta!. 1985) and food consumed were noted if possibl e. 'Prise' differs from 'probe' in that 'prising' involves leverage and sometimes breakage of the substrate (usually bark) to obtain hidden prey. Foraging in fo rm ation was obtain ed during the 33.2 hours that parent birds were followed away from nests (Table 1).

Displays and calls The following displays and calls were recorded when they occurred (see Wood & Ley 2005): 1. Wing-quivering by one of the pair 2. Mutual wing-quivering 3. Solo songs, usually while stationary 4. Duets, usually while perched 5. Response songs 6. Territorial song-flights 7. Alarm calls. If a display and call occurred together (e.g. wing-quivering with solo song), each behaviour was counted separately. Binoculars were used for nest-watches and to obtain foraging information, but two recording methods were employed. AJL dictated onto a hand-held tape-cassette-recorder and later transcribed the tapes onto a word-processor. KAW recorded obse1vations and information directly into fi eld books. Nest-watches were from unconcealed positions between 10 m and 20 m from nests. Most distances were measured with a portable odometer, driven by a rolling wheel. Otherwise they were stepped. Heights were calculated from measurements with a clinometer or 15-m extendable fibreglass tube. Times were taken from watches, set digitally to indicate hours, minutes and seconds. Ambient temperatures were obtained from the Meteorological Station at the New South Wales State Forests Office in Baradine. All times are Eastern Standard Time. Home-range and territory terminology foll ows Odum & Kuenzler (1955). The breeding home­ ra nge is the maximum area used by the breeding pair in the nest-building, incubation and nestling \0 Table 1 0 Observations of a breeding pair of Striped Honeyeaters at Baradine, NSW, in 2003.

Date Breeding stage Duration of observations (h) Notes (day/month) Nest-watches Other a

11/9 Nest-building 0.5 (KW) Nil Nest 1 seen-appeared complete. 21/9 Nest-building Nil 2.5 (KW) Nest 2 found at 1650 h. 22/9 Nest-building 12 (KW) Nil 23/9 Nest-building 7.5 (KW) Nil ~ 29/9 Nest-building 2.3 (KW) Nil 0 30/9 Nest-building 7.2 (KW) 4 (KW) No eggs at 0721 h. 0v 01/10 Nest-building 1.5 (KW) Nil No eggs at 0842 h. [-< 16/10 Incubation 3.3 (KW) 1.5 (KW) Light rain in early morning. 17/10 Incubation 4.7 (AL), 1 (KW) Nil Female banded orange at 0910 h. 27/10 Nestling 5.5 (AL) Nil One nestling c. 4 days old, one egg at 1645 h. 28/10 Nestling 8.4 (AL) 3.8 (KW) Male banded white at 0950 h. 29110 Nestling 6 (AL), 4.4 (KW) 0.5 (KW) Cold gusty wind all day. 30/10 Nestling 7.6 (KW) Nil One nestling but no eggs at 1510 h. 03/11 Nestling 4 (AL) Nil ...... 'T1 04/11 Nestling 6.6 (AL) Nil Nestling growth well advanced. til 10/11 Post-fledging Nil 2.5 (KW) Parents but not nestling seen. 11/11 Nest-building 4.3 (KW) 2.5 (KW) Nest 2 collected at 0945 h. Nest 3 found at 1155 h. G 0 12/11 Nest-building 2 (KW) Nil 13/11 Nest-building 5 (KW) 2.5 (KW) ...... ~6 Cil 24/11 ? 1 (KW) 0.5 (KW) Only male seen (not female). i..., 25/11 ? Nil 9 (KW) Only male seen (not female). Combined 111.6 33.2 Total o-12~ a Other observations away from the nest related to foraging, home-range boundaries and breeding behaviour. ~~ VOL. 22 (2) JUNE 2005 Striped Honeyeater Nesting and Foraging 91 ...... ·.·.·.·.·.·.·.·.·.· .·.·.·.·.·.·.·.·.·.·.·.·.·.·.· • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 0 ••••• • • 0 . 0 ••••••••••• •••••••••••••••• • ••••••••••••••• 0 • • 0 •••• 0. 0 •• • • • • • • • • • • • • • • • • • • • • • 0 ••• •• • •••••••• •••• ••••• ••••••••••••••••••• •• • • •• • • • • • • 0., ••••••. .0 . .••••••••• ...... •• • ••••••• • ••• • ••••••••••• • • •••••• ••• 0 • • ••••••• :~:~:~ :~ :~:~ :~:~:~: ~ .• .• ...... •• ••• 0 •••..• ••••..... 0 ...... • 0 • • •• •••.•••••• • •• ......

,...... ""'...... ------...... , ...... , .... " ... , ...... /// ·----,,, ' Racecourse ' :.: (treeless) ',,\, ' :. \ LEGEND * Nests 1, 2 & 3 - Home-range

D Buildings

100m 1-:-:·:-:1 Box woodland

Figure 1. Breeding home-range of a pair of Striped Honeyeaters around nest 2 at showground, Baradine, NSW, in spring 2003. Nests l and 3 were abandoned.

stages. To calculate the area, the furthermost points at which parent birds were seen in all directions were plotted to scale and joined by eye; the enclosed area was calculated using a su rvey software package. The core area of the home-range is that portion which is used most f r e q u e n t I y (>80% sightings), irrespective of geometry. Playback tapes were used to assist in determining home-range boundaries (Wood & Ley 2005). The defended territory is that portion of the home­ range around the nest that was defended from conspecific and interspecific individuals either by physical attack or displacement (McFarland 1984; Armstrong 1991).

Sexing and colour-marking the pair Parent birds were difficult to capture with a mist-net, even when playback tapes were used (Wood & Ley 2005) and a stuffed Fairy Prion Pachyptila turtur decoy was placed 50 em from the net. However, the female and male were eventually caught in nets on 17 and 28 October respectively (Table 1 ). The sexes are monomorphic in but females are smaller than males (Higgins et al. 2001). The female, colour-banded orange, was measured as follows (see Lowe 1989): head-bill = 41.1 mm, wing chord = 112 mm, tail = 102 mm, tarsus (without foot) = 31 mm, weight = 43 g. The male, colour-banded white, had the following measurements: head-bill = 43.2 mm, wing chord = 118 mm, tail = 105 mm, tarsus (without foot) = 31 mm, weight = 43.5 g. As a further aid to separate the sexes in the field, the distal1 em of the tail of the female was dyed purple and a 1-cm band in the middle of the tail of the male was dyed green.

A 11hropod samples Foliage from White Cypress Pine Callitris glaucophylla was taken from the lowest branches of several trees close ( < 40 m) to the nest and immediately placed in six-litre plastic bags. This fol iage was then sprayed with household insecticide and the bag sealed for about 10 minutes. Dead arthropods were collected and stored in a liquid consisting of 70% methylated spirits, 25 % water and 5% glycerine. Batches of arthropods were collected on 22 and 29 September from foliage that almost filled three and two plastic bags respectively. AUSTRALIAN 92 WOOD&LEY FIELD ORNITHOLOGY

NEST-BUILDING 15 d

------30 Sept.----- (a) nest appeared complete

EGG-LAYING 7 d

-- 10 October

INCUBATION 16 d

-r-20 October

-----(b) nestling estimated c. 4 days old

-r- 30 October NESTLING 17 d

----- (c) nestling growth well advanced

------9 November (fledged)

Figure 2. Estimated breeding chronology of a pair of Striped Honeyeaters at nest 2, Baradine, NSW, in spring 2003, based on observations (a)-(c) a nd assumed incubation and nestling periods of 16 and 17 days respectively (see Higgins eta/. 2001).

Proportion of wool in nest Combined wool and plant-down from the nest were weighed and then immersed in a solution of caustic soda. After the wool was dissolved, plant-down residue was collected in a filter, dried in an oven and weighed. The weight of wool was calculated by subtraction.

Results

Nests All three nests (Table 1) were suspended and attached by the rim at the leafy ends of thin drooping branchlets in tall (15- 17 m) White Box Eucalyptus albens or White Cypress Pine trees (Table 2). Nests 1 and 3 were > 7 m above ground whereas nest 2 was only 2.8 m above ground (Table 2). Nest 2 resembled a slightly urn­ shaped, small woollen bag c. 90 mm in diameter and 110 mm deep. Its rim was festooned and strongly fixed to six thin (1.2- 2 mm diameter) Callitris branchlets. VOL. 22 (2) JUNE 2005 Striped Honeyeater Nesting and Foraging 93

Table 2 Physical characteristics of nests of a pair of Striped Honeyeaters at Baradine, NSW.

Nest number 1 2 3

Nest height (m) 8.5 2.8 7.2 Nest-tree height (m) 15 16 16.5 Horizontal distance 2.2 2 3.7 of nest from trunk (m) Nest-tree species White Cypress Pine White Cypress Pine White Box

Using the lowest point of the festoon rim as reference, the inside diameter of the nest was 60 mm and the inside depth was 75 mm. Cohesion between the various components of nest 2 was substantial. When dismantled it consisted of 93 dry grass stems (25-140 mm x 0.75 mm diameter), 61 twisted grass rootlets (50-200 mm x 0.5 mm diameter), 170 white and 12 orange synthetic twines (75-250 mm x 0.5 mm diameter), 17 blue synthetic twines (75-500 mm x 0.5 mm diameter), nine lengths of cotton thread, nylon thread or white plaited yarn (130-330 mm x 0.25-1 mm diameter), fo ur round, prickly plant seeds (10 mm diameter), one white feather (20 mm long), one white spider's egg-sac (20 mm diameter), 6.3 g of white wool and 1.2 g of white plant-down. The grass stems were mostly branched, providing a gripping point, whereas the twisted grass rootlets gripped materials everywhere. The spider's egg-sac and prickly plant seeds were on the outside, but the wool and other components were distributed generally throughout the entire nest. Most long twines were woven around the attachment branch lets, into the rim and downward into the body of the nest. There did not appear to be any distinctive lining, but the thickness of wool and plant­ down was greater in the base than in the sides (c. 25 mm vs c. 15 mm).

Breeding home-range The breeding home-range around nest 2 comprised an oval-shaped core area of 3.5 ha and an annex of 0.7 ha to the east (total4.2 ha, Figure 1). The core area was used during all breeding stages but the annex protruding into box woodland seemed to be used only in the nestling stage (from 28 October). In the core area there were 110 mature White Cypress Pines (15-18 m tall), nine Eucalyptus box trees (18- 22 m) (mostly White Box) and 17 smaller trees (6- 10 m) (mostly White Cedar Melia azedarach, KurrajongBrachychiton populneus and Peppercorn Schinus areira ). The overall tree density was c. 40 trees/ha. No shrubs were growing in the core area. We were unable to define precisely the home-range around nest 3. In the central portion around the nest there were mostly tall (16- 22 m) White Boxes interspersed with a few White Cypress Pines of various ages and a few shrubs. Tree density was c. 70 trees/ha. The food resources and nesting materials in the home-range around nest 2 were shared with other . Closest nests were of White-browed Woodswallow Artamus superciliosus ( 4 m away), White-winged Triller Lalage sueurii (12m, 20m), Willie Wagtail Rhipidura leucopl11ys (35m), Noisy Miner AUSTRALIAN 94 WOOD&LEY FIELD ORNITHOLOGY

melanocephala (60 m), Yellow-rumped Thornbill Acanthiza chtysorrhoa (60 m) and Magpie-lark Grallina cyanoleuca (70 m). White-plumed Honeyeaters Lichenostomus penicillatus and Rufous Songlarks Cincloramphus mathewsi were also common, especially east of nest 1. There was no nesting association with as no Cracticus species were observed nesting within 100m of any of the three Honeyeater nests in Figure 1.

Building of nest 2 We estimated that nest 2 took c. 15 days to build (16-30 September). Visits by both adults were to inspect only, deliver material only or undertake construction (with or without delivery). By 22 September this nest had taken shape. During 19.5 hours of observation on 22 and 23 September (Table 1), the adults made 89 visits (14 inspection, 25 delivery, 50 construction) at a combined rate of 4.6 visits/h. The longest absence was 63 minutes. Inspection and delivery visits lasted < 1 minute whereas construction visits were between 1 and 14 minutes (median = 6 min., n = 50). Nest 2 appeared complete by 30 September. During 11 hours of observation on 29-30 September and 1 October, parents did not deliver additional material but made 13 short ( < 1 min.) inspection visits and another 8 visits to modify or maintain construction (median = 5 min., range = 1- 12 min.). The visi tation rate was less than on 22 and 23 September (1.9 vs 4.6 visits/h). Some materials for nest 2, particularly grasses, were gathered within a radius of 40 m from the nest-site, and on three occasions (21, 22 and 23 September) we saw one of the pair take wool or plant-down from nest l and carry it to nest 2. During long construction visits, the adults threaded and wove materials into the body of the nest and tied the rim to the supporting branch lets. When this nest was almost completed the visiting bird often sat inside, in the incubation position, with only the tip of the tail visible, and wriggled as if compacting the base and sides. Nest-building changeovers were frequent although both adults were seen together at nest 2 only seven times, mostly for about 15- 20 seconds (maximum 40 sec.).

Building of nest 3 Nest 3 comprised only a few pieces of wool or plant-down when it was found on 11 November. In 11.3 hours of observation on 11, 12 and 13 November, the adults made a total of 44 visits (12 inspection, 3 delivery, 29 construction) at a combined rate of 3.9 visits/h. The longest absence was 28 minutes. Construction visits were shorter than at nest 2 (median = 1 min., range 0.5- 5 min.). The male and female made 23 and 14 visits respectively (for seven visits the sex of the bird could not be determined). Both adults were at this nest together four times for 10 to 30 seconds. Otherwise, the behaviour of the sexes in building nests 2 and 3 was sim il ar. The materials used to build both nests were also similar, except that for nest 3 we did not see materials re-used from any other nests nearby and we once saw plant fibre stripped from the hanging case of a moth (fam ily Psychidae) and taken to the nest 25 m away.

Incubation It was estimated that fu ll-ti me incubation did not commence until about 7 days after nest 2 was completed (Figure 2). The presence of two adults near the nest at times of (mainly short) changeovers indicated that both sexes incubated. VOL. 22 (2) JUNE 2005 Striped Honeyeater Nesting and Foraging 95

Table 3 Feeding rate of a single Striped Honeyeater nestling at Baradine NSW, in spring 2003. M = male, F = female, US = sex undetermined.

Date Observation Number offe eding visits Total Feeding rate (day/month) period (h) visits/h) M F us Morning 28/10 2.4 12 10 22 9.2 29/10 6.1 27 14 41 6.7 30/10 5.2 34 27 61 11.7 04/11 6.6 42 54 96 14.5 Afternoon 27/10 5.5 33 29 6 68 12.4 28/10 6.0 55 39 94 15.7 29110 4.4 10 9 19 4.3 30!10 2.5 8 7 15 6.0 03/11 3.9 30 35 65 16.7 Total 42.6 251 224 6 481 ll.3

Sharing of incubation was confirmed in watches of 148 and 62 minutes after the female was colour-banded on 17 October, when she incubated in three bouts for 78 minutes overall and the unmarked male incubated in three bouts for 130 minutes (eggs uncovered for about 2 minutes). During 14, 15 and 16 October, before the female was banded, the incubation routine included some long ( >60 sec.) periods of absence during changeovers. Ten long changeovers ranged from 1 to 10 minutes (median = 5 min.). All long changeovers were after 1130 h, mostly after 1400 h, and were not related to abnormally high ambient tempe ratures (temperature range at 1500 h = 22.8-25.8°C). Overall , before the female was banded, 61 incubation bouts were recorded in eight watches. The so-called 'a' bird incubated for 558 minutes in 30 bouts (mean = 18.6 min.) and the 'b' bird incubated for 569 minutes in 31 bouts (mean = 18.4 min.). The total time off eggs during the 3 days before 17 October was estimated at 84.7 minutes, resulting in an incubation constancy of about 93% (1127/1211 minutes). Absences during short changeovers were mostly about 20 seconds (mean = 18.5, range 5-40 sec., n = 37).

Nestling stage The parents fed the single nestling almost equally (Table 3). Of 475 feedi ng visits by a parent of known sex, 251 (53%) were by the male and 224 (47% ) were by the female. Feeding rates varied widely between 4.3 visits/h in an afternoon watch of 4.4 hours on 29 October and 16.7 visits/h in an afternoon watch of 3.9 hours on 3 November. Nevertheless, there was only a slight difference between the overall morning and afternoon feeding rates (10.8 and 11.7 visits/h respectively). When morning and afternoon data were pooled, the mean feeding rate at which both parents fed the nestling was 11.3 visits/h. AUSTRALIAN 96 WOOD&LEY FIELD ORNITHOLOGY

The female's brooding effort was three times greater than that of the male (Table 4). The female brooded for 118 bouts compared with 39 bouts by the male (ratio 3.02) and for 945 minutes compared with 312 minutes (ratio 3.03). The mean brooding constancy during the 4 days 27-30 October was about 58% (1109/1922 minutes, Table 4), whereas between 3 and 4 November, it was only c. 23% (148/635 minutes). We were able to describe the food delivered to the nestling on only 42 of 481 visits (9% ). Mostly the item carried was very small and we could not see it being placed in the nestling's mouth. These 42 food items were 11 large black larvae (15-20 mm long), eight spiders, five insects, three white larvae or coccoid mealybugs (c. 8 mm long), one caterpillar, and 14 other items (10-15 mm long), some black and some white.

Displays and calls The breeding pair called throughout the breeding cycle but most frequently in the nest-building stage (Table 5). While nest-building, solo songs were given 3.4 times per hour compared with 1.2 and 1.6 times per hour in the incubation and nestling stages respectively. Territorial song-flights (0.9/h) and duets (0.43/h) were also performed at the highest rate in the nest-building phase. Displays were performed in all breeding stages (Table 5) but wing-quivering by one of the pair (n = 42) was much more common than mutual wing-quivering (n = 9). The most intense breeding interaction, involving mutual wing-quivering with duet, was seen twice in the nest-building stage, four times in the incubation stage and once in the nestling stage. Overall, there were twice as many breeding communications (calls and displays) in the nest -building stage as in the incubation or nestling stages (5.3 vs 2.6 and 2.6, Table 5). After the female was colour-banded, it was thought that only the male performed territorial song-flights. This suggestion was confirmed later, after the male was also colour-banded, when he performed all 14 song-flights observed. Ten song-flights by the male were in the nestling stage and four were during the time that nest 3 was being constructed. The male also uttered more solo songs than the female. After both parents were colour-banded, during 30.5 hours of observation in the nestling period (data included in Table 5), 43 solo songs (88%) were given by the male and six (12%) by the female. Otherwise, the sex of the birds singing solo was not known. Some solo songs were given from high 'sentry' branches of trees near the home-range boundary.

Nest -defence Both parents defended a small area around the nest (Table 6). Twenty-nine of the 35 chases seen (83%) were directed at intruders within a radius of 20 m. Occasionally, small passerines were ignored when close ( <8 m) to the nest. There were 18 chases by a parent of unknown sex, but after the sex of the parents was established by colour-banding, we saw nine chases by the male and four by the female. Only five chases were in the incubation stage compared with 14 and 16 respectively in the nest-building and nestling stages. Four chases were of intruding Striped Honeyeaters, three of which were during the nest-building stage (third conspecific repelled). Only three attacks involved simultaneous utterances of the short alarm call zeet. Table 4: Brooding constancy of a pair of Striped Honeyeaters at Baradine, NSW, in spring 2003.

Date Observation Number of brooding bouts (n) and Brooding Estimated age (day/month) pe1iod (min.) cumulative total of brooding time (min.) constancy% of nestling (days) (cumulative) Male Female n Minutes n Minutes

27/10 332 7 43 24 154 59.3 4 28/10 503 10 55 30 162 43.1 5 29/10 630 12 149 23 321 74.6 6 30/10 457 9 57 24 168 49.2 7 03/11 236 4 20 8.5 11 04111 399 1 8 13 120 32.1 12 Total 2557 39 312 118 945

Table 5: Number (n) and frequency (n/h) of different displays and calls of a pair of Striped Honeyeaters while breeding at Baradine, NSW, in spring 2003.

Display or call Breeding stage (and length of observation time, h) Nest-building (41.8)" Incubation (28.J)b Nestling (42.5) n n/h n nih n n/h

Wing-quivering (one bird) 20 0.5 9 0.3 13 0.3 Mutual wing-quivering 3 0.07 5 0.18 1 0.02 Solo song 142 3.4 35 1.2 68 1.6 Duet 18 0.43 7 0.25 15 0.35 Response song 2 0.05 1 0.04 3 0.07 Territorial song-flight 38 0.9 16 0.6 12 0.3 Total 223 5.4 73 2.6 112 2.6 a Pooled data from nest 2 (30.5 h) and nest 3 (11.3 h). b 2.5 hours of sampling were away from the nest (not nest-watching, see Table 1). AUSTRALIAN 98 WOOD&LEY FIELD ORNITHOLOGY

Table 6 Nest defence by breeding Striped Honeyeaters at Baradine, NSW, in spring 2003. Intruder bird species and distance (m) from nest 2 when intially chased by the Striped Honeyeaters during indicated breeding stage (total observation time in parentheses); M = male, F = female, BS = both sexes, US = unknown sex. Intruder species Chased by M F BS us Nest-building (30.5 h) Yellow-rumped Thornhill Acanthiza chryson-hoa 13 Striped Honeyeater Plectorhyncha lanceolata 4 13,35 Little Philemon citreogularis 40 Noisy Miner Manorina melanocephala 1,4 White-plumed Honeyeater Lichenostomus penicillatus 5 Willie Wagtail Rhipidura leucophrys 13 Black-faced Cuckoo-shrike Coracina novaehollandiae 6 White-winged Triller Lalage sueurii 5, 15,20,30

Incubation (25.8 h) Noisy Miner 30 White-winged Triller 3, 5, 6, 10

Nestling ( 42.5 h) Spiny-cheeked Honeyeater Acanthagenys rufogularis 1 Striped Honeyeater 6 Noisy Miner 4 4, 13 White-plumed Honeyeater 1, 3 White-winged Triller 4 5,6 Olive-backed Oriole Oriolus sagittatus 30 White-browed Woodswallow A11amus superciliosus 3,4,5,6 Australian Raven C01vus coronoides 50

Nest hygiene In 42.5 hours of observation over 6 days (Table 1), the nestling voided 29 faecal sacs (about one every 1.5 hours). All were carried at least 15m from the nest by a parent. Eighteen sacs ( 62%) were dropped in flight; the fate of the others was unknown although none was seen to be eaten by a parent. The male removed 16 sacs, the female 12; one sac was presented to a parent of unknown sex. All but two sacs were voided immediately after the nestling was fed.

Foraging The predominant tree species in which parent Honeyeaters foraged differed between home-ranges (Table 7). During 37 foraging bouts (1- 13 min.) in the home­ range around nest 2, adults spent 91 % of foraging time in White Cypress Pines compared with 8% in Eucalyptus box trees. Conversely, while building nest 3, the parents spent 83.5% of foraging time in Eucalyptus box trees and only 16.5% in White Cypress Pines (18 bouts, range 1- 11 min.). The height at which they foraged also differed between tree species (Figure 3). Whereas they spent a majority of VOL. 22 (2) JUNE 2005 Striped Honeyeater Nesting and Foraging 99

Table 7 Percentage of time that the breeding Striped Honeyeaters foragesd in various tree species at Baradine, NSW, in spring 2003.

Nest No. of Total T!·ee species no. foraging observation bouts time (min.) White Cypress Eucalyptus Other Pine box 2 37 169.5 91 8 1 3 18 72.5 16.5 83.5 0

time (56.4%) foraging at heights of 15-20 min Eucalyptus box trees, they foraged throughout the full range of heights in White Cypress Pines. Arthropods collected from White Cypress Pine foliage (Table 8) indicate food available to the Honeyeaters during the study period. However, we generally could not see the Honeyeaters capture prey or consume food because they searched mostly amid dense foliage of Callitris and Eucalyptus trees. Even when the bill was not hidden or obscured from view, the food taken was too small to identify. Occasionally they searched the insides of open (dead) White Cypress Pine cones without success. In 242 minutes of sampling, the only successful foraging manoeuvres seen were: gleaning (twice: an insect from a eucalypt branch and a spider from Callitris leaves), prising (twice: arthropods from under eucalypt bark), probing [three times: larva from moth's case (family Psychidae), twice, and small spiders (family Araneidae) from their compact 'nest', once], hawking (three times: a moth and two insects) and hovering (once: nectar taken from a flower of Patterson's Curse Echium plantagineum ). During the study period, only one eucalypt tree was in flower in the home-range around nest 2, a Fuzzy Box Eucalyptus

20-25

15-20 E .E _j------'1267 Ol 10-15 • Eucalyptus box "Q) ··········~~~··········13.4 56.4 (21 bouts, 74.5 min.) .r:::. Ol 1------'1234 White Cypress Pine c 5-10 D ·o, 22.1 (33 bouts, 166.5 min.) ~ 0 1------''33 LL 0-5 -

J16.8

0 10 20 30 40 50 60

Percentage of foraging time

Figure 3. Percentage of foraging time that the breeding male and female Striped Honeyeaters foraged at various heights in White Cypress Pine and Eucalyptus box trees at Baradine, NSW. Combined data for nests 2 and 3. AUSTRALIAN 100 WOOD&LEY FIELD ORNITHOLOGY

Table 8 Number of arthropods collected from two batches of leaves of White Cypress Pine at Baradine, NSW, in spring 2003 (see Methods).

Common name Order or Family First batch Second batch Total of m1hropod (22 Sept.) (29 Sept.)

Spider (Orb Weaver?) Araneidae 5 3 8 Plant bug Miridae* 3 4 7 Leafh opper Cicadell id ae * 2 1 3 M idge fl y Chironomidae 1 Nil 1 Beetl e Staphylinidae 2 Nil 2 'Round' bug Lestoniidae* Nil 1 1 Black aphid Aphididae* Nil 2 2 Psocopte ran Psocopte ra Nil 1 1 To tal 13 12 25

*Families in the Order H emiptera (bugs and leafhoppers)

conica adjacent to the nest-tree, but we did not see the Honeyeaters probing its flowers for either nectar or insects. On one occasion a parent defaecated a seed that was collected and identified as mistletoe, either Lysiana sp. or Amyema sp. (B. Wiecek in !itt. 19 December 2003).

Use of feet Striped Honeyeaters were observed to grip an object with a foot on 10 occasions. Five times prey was seen held against a branch while being dismembered with the bill; the prey items were insects (twice), an insect larva (once) and a spider (one arthropod was not identified). Twice, wool or plant-down was seen to be gripped against a branch and teased apart with the bill before being carried to the nest. Once, in the nest-building stage, a bird held a spikelet of live grass against a low branch with the foot while pulling the lower stem with the bill. Again in the nest­ building stage, a Honeyeater held a moth's case against a green leaf in a White Box to facilitate stripping of plant-fibre from the case before subsequent delivery to the nest. Lastly, a green White Box leaf was gripped with a foot to facilitate removal of a lerp or gall. The male held six of the objects mentioned, the female held one and the remaining three objects were held by a bird of unknown sex.

Flight and comfort behaviour In horizontal flights of> 50 m the Honeyeaters typically used undulating flight. However, in shorter flights of 20-30 m, particularly when reducing altitude, they often glided with upswept wings. Gliding flight was mostly seen as the Honeyeaters approached nest 2 from an adjacent tree. Parent birds often wiped their bills while foraging and handling nesting materials. They also preened themselves regularly during most days, sometimes in bouts of 2-3 minutes' duration. Allopreening was observed once in a bout of 5.5 minutes on 13 November when the parents perched on a 15-m high branch and both birds were oriented in the same direction. VOL. 22 (2) JUNE 2005 Striped Honeyeater Nesting and Foraging 101

Discussion The pair of Striped Honeyeaters at Baradine nested as a simple pair, with the male and female sharing the duties of nest-building, incubation and feeding the nestling almost equally. However, the female's effort in brooding the nestling was three times greater than that of the male, and the male performed most (if not all) territorial song-flights and uttered most (88%) solo songs. In contrast, the putative male and female Striped Honeyeaters studied at Meandarra, Queensland, made almost equal contributions to brooding ( 43% vs 57% total time) and were assisted by a third conspecific in feeding the nestlings (Moffatt et al. 1993). We found no evidence of cooperative care of the Baradine nestling. Differences between the Meandarra and Baradine pairs may be because of differences between their breeding systems or behaviour of the respective pairs. In most facultatively communally breeding species, the most frequent group is the simple male and female pair (Dow 1980; Rowley & Russell 1997). Some sex roles of the Baradine pair of Striped Honeyeaters were unlike the corresponding roles adopted by most non-cooperative honeyeaters. In many non­ cooperative species (Longmore 1991), including the Fuscous Honeyeater Lichenostomus fuscus (Dunkerley 1989), Regent Honeyeater Xanthomyza phrygia (Oliver 1998) and New Holland Honeyeater Phylidonyris novaehollandiae (Higgins et al. 2001), only the female builds the nest, incubates the eggs and broods the nestlings. Both sexes of the Striped Honeyeater participated in these duties at Baradine. Indeed, the Baradine pair adopted sex roles that are similar to the non­ cooperative Grantiella picta (Eddy 1961; Whitmore & Eller 1983; Tremont & Williams 1999). In the Painted Honeyeater, both sexes share duties of parental care although most of the brooding is by the female and territorial song-flights are by the male, as observed for the Striped Honeyeater at Baradine. This study provides new information about breeding habitat and foraging height of Striped Honeyeaters. The Baradine nest may have been located in habitat similar to a nest built in Callitris pine in Victoria (Chandler 1913) but a breeding home­ range of predominantly mature Callitris, as observed at Baradine (110/136 trees = 81% ), has not been previously reported (Higgins et al. 2001). Moreover, the Baradine pair foraged at all heights in White Cypress Pines (Figure 3) whereas canopy foraging was preferred by Striped Honeyeaters in mallee-oak woodland near Mungo in south-western New South Wales (Franklin & Alley 1995). Gleaning, probing, prising and hawking were observed at both Baradine and Mungo. Although we were unable to describe the food delivered to the nestling in the majority of visits to the nest, it is unlikely that nectar was carried because only one eucalypt was in flower in the home-range and we did not see parent Striped Honeyeaters probing its flowers. Of the 42 identified items of food given to the nestling, 41 (98% ) were spiders, insects or insect larvae. Spiders have been reported in the diet of Striped Honeyeaters in only one previous study, at Lake Cowal, NSW (Vestjens 1977; Higgins et al. 2001). Spiders, however, could be an important dietary component because they comprised 19% (8/42) of the items identified and represented 32% (8/25) of arthropods collected from samples of Callitris leaves (Table 8). As at Baradine, extraction of larvae from moths' cases also was observed at Inverell, NSW (Baldwin 1976). Our unsuccessful identification of most food taken by adult Honeyeaters while foraging is consistent with Franklin & Alley's (1995) general inability to determine AUSTRALIAN 102 WOOD&LEY FIELD ORNITHOLOGY

the nature of food taken near Mungo. As 17 of the 25 arthropods (68%) sampled at Baradine were only 1--4 mm long (Table 8), it is possible that the Honeyeaters captured arthropods (mostly hemipterans) that were too small to be seen with binoculars. Alternatively, it is possible that we did not see the birds licking exudate or honeydew from Callitris branches. Previously, Cleland (1919) noted a sugary exudate on branchlets of Callitris pines in the Pilliga that was attracting woodswallows. Furthermore, Paton (1980) cited a communication from N. Forde that honeyeaters sometimes 'fed on the honeydew of aphids living on Callitris'. The possibility that Striped Honeyeaters imbibe honeydew or a plant exudate when foraging along Callitris branches requires further investigation. The overall mean feeding rate at which both parents fed the nestling (11.3 visits/h) was similar to that for other honeyeaters near Armidale, NSW: Painted Honeyeaters (8.1 feeds/h, Tremont & Williams 1999), Red Wattlebirds Anthochaera carunculata (7.2 feeds/h, H.A. Ford unpublished, in Oliver 1998) and Noisy Philemon corniculatus (9.3 feeds/h; Ford unpublished, in Oliver 1998) but much lower than Regent Honeyeaters (23 feeds/h, Oliver 1998). Such feeding rates for non-social honeyeaters are probably dependent on biological and environmental factors such as number of nestlings, food size and type and food availability. Some behaviour types observed at Baradine have also been reported elsewhere. Allopreening, as seen in the present study, was also seen near Mungo (Franklin & Alley 1995), and use of feet to hold prey was observed there (Franklin & Alley 1995) and near Coffs Harbour, NSW (Ford 1992). In contrast, a nesting association with Grey Butcherbirds Cracticus torquatus has been observed previously (Higgins et al. 2001) but did not occur at Baradine. The Baradine Striped Honeyeaters took c. 15 days to build nest 2 whereas honeyeaters of similar size usually take about 8-10 days to build nests (Higgins et al. 2001 ). Finding wool and other suitable materials may contribute to protracted nest-building, as also may the time taken to weave the various materials into a durable nest at the extremity of a drooping branch. We were unable to find comparative data, but a mean duration of 6 minutes at the nest during construction visits may be significant. The size and shape of the bill of the Striped Honeyeater seems suited to weaving the wool and other materials together. Because most Australian honeyeaters make 'many attempts at breeding in a season, even after successfully raising young' (Higgins et al. 2001), it was not surprising to see the Baradine pair building nest 3, 1- 2 days after the disappearance of their young from nest 2. Performing displays and calling during all breeding phases (Table 5) prolonged the adults' sexual bonding and ensured that they were ready to recommence breeding after rearing their nestling in nest 2. As for the Baradine pair, most honeyeaters build a new nest for each breeding attempt rather than re-use an old nest (Higgins et al. 2001). Re-use of materials from old nests is well documented (Ley et al. 1997).

Acknowledgements We thank Kim Pullen and Michelle Michie of CSIRO Entomology for identifying arthropod samples and Ken Hill of the Royal Botanic Gardens, Sydney, for identifying some eucalypt pressings. Ron Nixon, Barry Johnston and Neil McDonald made us welcome at Camp Cypress, Baradine. David Drynan expedited colour-marking approval from Environment Australia. Andrew Lindsay, Australian Wool Testing Association, assisted with the separation of wool and plant­ down in nest 2. We also thank Damon Oliver and Hugh Ford for critically commenting on the manuscript. VOL. 22 (2) JUNE 2005 Striped Honeyeater Nesting and Foraging 103

References Altmann, J. (1974), 'Observational study of behaviour: Sampling methods', Behaviour 49, 227- 267. Armstrong, D.P. (1991 ), 'Aggressiveness of breeding territorial honeyeaters corresponds to seasonal changes in nectar availability', Behavioural and Ecological Sociobiology 29, l 03- 111. Baldwin, M. (1976), 'Striped Honeyeater and case moth', Canberra Bird Notes 3, 25. Blakers, M., Davies, S.J.J.F. & Reilly, P.N. (1984), The Atlas ofAustralian Birds, Royal Australasian Ornithologists Union, Melbourne University Press, Melbourne. Chandler, L.G. (1913), 'Bird life of Kow Plains (Victoria)', Emu 13, 33-45. Cleland J.B. (1919), 'The birds of the Pillaga Scrub, New South Wales', Emu 18, 272-285. Dow, D.D. (1980), 'Communally breeding Australian birds with an analysis of distributional and environmental factors', Emu 80, 121-140. Dunkerley, G.M. (1989), The Fuscous Honeyeater: Food Resources and the Bird Communitv, PhD thesis, University of New England, Armidale, NSW. Eddy, R.J. (1961), 'Twenty years of the Painted Honeyeater', Australian Bird Watcher 1, 122- 128. Ford, H.A. (1992), 'Striped Honeyeater Plectorhyncha lanceolata using feet to hold down prey', Australian Bird Watcher14, 190. Franklin, D.C. & Alley, J.C. (1995), 'Some observations of foraging and social behaviour of the Striped Honeyeater in south-western New South Wales',Australian Bird Watcher 16, 71- 74. Higgins, P.J., Peter, J.M. & Steele, WK. (Eds) (2001), Handbook ofAustralia n, New Zealand and Antarctic Birds, vol. 5, Oxford University Press, Melbourne. Ley, A.J., Oliver, D.L. & Williams, M.B. (1997), 'Theft of nesting material involvi ng honeyeaters (Meliphagidae)', Corella 21, 119-123. Longmore, W. (1991 ), Honeyeaters & Their Allies of Australia , Harper Collins, Sydney. Lowe, K.W. (1989), The Australian Bird Bander's Manual, Australian Bird & Bat Banding Schemes, Canberra. McFarland, (1984), 'Behaviour of White-cheeked Honeyeaters and Little Wattlebirds in Banksia integrifolia ',Australian Birds 19, 1-6. Moffatt, D., Whitmore, M.J. & Date, E.M. (1983), 'Communal breeding by Striped Honeyeaters', Emu 83, 202-203. Odum, E.P. & Kuenzler, E.J. (1955), 'Measurement of territory and home range size in birds', Auk 72, 128- 137. Oliver, D.L. (1998), 'The breeding behaviour of the endangered Regent Honeyeater, Xanthomyza pluygia, near Armidale, New South Wales', Australian Journal of Zoology 46, 153-170. Paton, D.C. (1980), 'The importance of manna, honeydew and lerp in the diet of honeyeaters', Emu 80, 213-216. Recl1er, H.F., Holmes, R.T., Schulz, M., Shields, J. & Kavanagh, R. (1985), 'Foraging patterns of breeding birds in eucalypt forest and woodland of southeastern A ustrali a',Australian Journal of Ecology 10, 399-419. Rowley, I. & Russell, E. (1997), Fa i1y Wi·ens and Grasswrens: Maluridae, Oxford University Press, Oxford, UK. Schodde, R. & Tidemann, S.C. (Eds) (1986), Reader's Digest Complete Book ofAust ralian Birds, 2nd edn, Readers Digest Services, Sydney. 1h~ mont , S. & Williams, M.B. (1999), 'Observations at nests of the Painted Honeyeater Gran tiel/a picta', Australian Bird Watcher 18, 49- 58. Vestjens, W.J.M. (1977), 'Status, habitats and food of vertebrates at Lake Cowal', Technical Memorandum 12, CSIRO, Canberra. Whitmore, M.J. & Eller, C.M. (1983), 'Observations at a nest of Painted Honeyeaters', Emu 83, 199- 202. Wood, K.A & Ley, A.J. (2005), 'Breeding displays and calls of the Striped Honeyeater at Baradine, New South Wales', Australian Field Ornithology 22, in press.

Received 26 April 2004 •