121 AUSTRALIAN FIELD ORNITHOLOGY 2008, 25 , 121–131 Breeding Territories and Breeding Success of the fascinans in South-eastern

K.A. WOOD1, NICCI THOMPSON 2 and ANDREW J. LEY 3 18 Kalamunda Street, North Lakes, Queensland 4509 221 Welcombe Avenue, Toowoomba, Queensland 4350 319 Lynches Road, Armidale, 2350

Summary Breeding territories and breeding success of the Jacky Winter Microeca fascinans were investigated in the 2005–06 breeding season in 140 hectares of fragmented eucalypt woodland in south-eastern Queensland. The breeding density was eight territories in 140 hectares (one pair per 17.5 ha) and the mean territory area of a breeding pair was 1.7 hectares (range 1–3.1 ha, n = 8). All territories comprised similar proportions of box woodland and open box woodland, characterised by patches of mature or sapling box trees, isolated live trees, dead trees (standing and fallen), logs and stumps. Of 23 , 21 (91%) were in forks consisting of a pair of dead branchlets near the extremity of a long live branch in eucalypt trees estimated to be >160 years old. The sample of nests is small, but the study suggests that woodland without aged eucalypt trees might not provide the species’ preferred -sites. Three nests failed before incubation was observed. Of the remaining 20 nests with eggs, five nests produced seven fledglings, resulting in a nesting success of 25%, an estimated breeding success of 17.5%, and productivity of 0.88 young per territory in 2005–06. The post-fledging dependence period for one juvenile was ~9 weeks.

Introduction The Jacky Winter Microeca fascinans is a small, sexually monomorphic robin (length 12–14 cm, weight 14–18 g) that breeds mostly in the eastern states of , in eucalypt woodland west of the Great Dividing Range (Blakers et al . 1984; Schodde & Tidemann 1986; Boles 1988). It builds a small, open cup- shaped nest (Boles 1988; Beruldsen 2003) in an exposed situation, as do some other members of the family Petroicidae (Schodde & Mason 1999). Because its numbers are believed to be declining (Robinson 1991; Recher et al . 2002; Barrett et al . 2003) and its conservation is of concern (Ford et al . 2001; Olsen et al . 2005), the species has a high priority for research (Ford et al . 1995). Some aspects of its breeding are known. For example, the incubation period is between 16 and 18 days (Marchant 1985a; Keast 1994), the nestling period is between 15 and 20 days (Keast 1994; Higgins & Peter 2002), and only the female incubates eggs (Higgins & Peter 2002). There are no published studies of the species’ breeding territoriality or breeding success (see Higgins & Peter 2002). Accordingly, we investigated the characteristics of Jacky Winter breeding territories and estimated its breeding success in 140 ha of fragmented eucalypt woodland in south-eastern Queensland in the 2005–06 breeding season.

Study area and methods

Study site The study site (Figure 1; 28°22 aS, 151°30 aE) was part of the sheep-grazing property Glenelg (3660 ha) in the Parish of Moynalty, Queensland. Glenelg is ~8 km south of Gore on AUSTRALIAN 122 WOOD, THOMPSON & LEY FIELD ORNITHOLOGY

Figure 1. Study area at Glenelg , south-eastern Qld, showing nests 1 to 23 built by Jacky Winters between September 2005 and February 2006. Breeding territories (T1–T8) were determined around particular nests indicated in Table 1. Angophora –box and Angophora –Red Gum forests are shown cross-hatched as a-b and a-t respectively. Sub-areas X, Y and Z are described in text.

the Cunningham Highway, between Warwick and Inglewood. The landscape is fragmented eucalypt woodland with patches of remnant native vegetation growing mostly on the hills and along major streams such as Macintyre Brook. The study area was partly cleared for wool production ~160 years ago (French 1990). During this study, River Sheoak Casuarina cunninghamiana and River Paperbark Melaleuca trichostachya dominated the banks of the Brook, and mature Rough-barked Apples Angophora floribunda were abundant on alluvial flats beside the Brook, in association with mature Yellow Box melliodora and Grey Box E. microcarpa or mature Forest Red Gum E. tereticornis (see Figure 1). Isolated mature Yellow Box and Grey Box trees were present on the fertile ridges, interspersed with patches of box regrowth mostly 30–40 years old, between 3 and 5 m tall. Tumbledown Gum E. dealbata and some White Cypress-pine Callitris glaucophylla were present on less fertile stony ridges. Throughout the site, dead VOL. 25 (3) SEPTEMBER 2008 Jacky Winter Breeding Biology 123 trees and stumps were scattered about, standing and lying on the ground, sometimes with vegetation debris. No shrub layer existed, but a variety of native and introduced grasses and herbs was present.

Methods The study site was first visited for 9 h on 16 and 17 December 2004 (KW & NT) when Jacky Winters were observed and preliminary site information was obtained. It was visited again and searched for Jacky Winters for 5 h on 19 May 2005 (KW, NT & AL) and 15 h on 25, 26 and 27 May 2005 (KW). During a short inspection in early September, Jacky Winters were vociferous, and the detailed study commenced the following week. Subsequently, the site was visited every week from 15 September 2005 (week 1) to 2 February 2006 (week 21) except week 18 (no visits). Visits were typically midweek on three consecutive days (weeks 2, 3, 4, 5, 7, 8, 9 and 10), two consecutive days (weeks 1, 6, 11, 12, 13, 14, 15 and 19), or on a single day in weeks 16, 17, 20 and 21. The number of observers and observer-hours each week is shown in Figure 2 (overall, 431 observer-hours). The boundaries of the site were determined in mid September 2005. It was mapped using survey information on the 1955 parish map, measurements with a portable rolling- wheel odometer, and trigonometric calculations. Landmarks such as fences, gates, gullies and particular trees were plotted to an accuracy of ~5 m. Height of nests above ground and tree heights were measured if <15 m, and calculated trigonometrically, using direct measurements and angles measured with an inclinometer, if >15 m. We attempted to locate all nests by direct searching, aided by binoculars and behavioural cues of breeding . From weeks 1 to 17 the entire area was searched; thereafter we visited only territories numbered T1, T4, T5, T6, T7 and nest 23 (Figure 1). Incubating Jacky Winters were presumed to be female. Birds presumed to be male called much more frequently than presumed females did, and regularly uttered the territorial call peter-peter- peter (see Marchant 1985a; Keast 1993, 1994). After nests were located, attending birds were followed regularly until they were lost, to determine their extent of movement in the breeding territory. To confirm the attachment of birds to a nest, one observer watched the nest continuously while the other followed the birds that visited. Presumed females tended to stay in the vicinity of nests; presumed males tended to move elsewhere throughout the territory. Territorial calls were often uttered from tops (maximum 21 m) of preferred trees on territory boundaries. Calls from these locations regularly elicited counter-calling by presumed males from adjacent territories. Most observer time (~80% overall) was spent searching for nests and following birds that attended nests but, during some site visits, survey information, vegetation measurements and foraging data were also obtained. Nests were numbered sequentially as they were found. Their status was determined at least twice each visit, mostly on different days. For nests lower than 7.5 m (10 nests), inspections of contents were made with a mirror on a pole. For the other 10 nests with eggs, it was assumed that two eggs were laid, on the basis that 92% of 101 clutches in the Birds Australia Nest Record Scheme comprised two eggs (Higgins & Peter 2002). Incubation was inferred from prolonged sitting (>8 minutes) by the presumed female and lack of other activity that was indicative of any other phase in the breeding routine. Assessments of the status of nests were checked against known incubation and nestling periods of 16–18 days and 15–20 days, respectively. Failed nests disappeared, were abandoned or were found damaged. They were considered severely damaged if >50% of the original nest appeared to be destroyed. Nesting success was calculated as the percentage of nests with eggs that produced fledglings. Breeding success was the percentage of eggs that produced fledglings. The male from nest 5 was colour-banded on 22 September 2005 (week 2). He was sexed by the condition of the cloaca and subsequently verified as being a male by behaviour thought to be typical of males (see Higgins & Peter 2002). Foraging terminology follows Recher & Holmes (1985). Territory terminology follows Odum & Kuenzler (1955). The breeding territory is the maximum area used by the breeding pair in the nest-building, incubation and nestling stages. To calculate the area, the most distant points at which parent birds were seen in all directions from the nest were plotted and joined by eye; the enclosed area was calculated by graphical method. AUSTRALIAN 124 WOOD, THOMPSON & LEY FIELD ORNITHOLOGY

Table 1 Characteristics of eight breeding territories (containing closely studied nests) used by Jacky Winters at Glenelg , south-eastern Qld, between September 2005 and February 2006. Territory number 1 2 3 4 5 6 7 8

Containing nest no. 4 8 5 1, 6, 11 3 2, 7 16 10 Total area (ha) 1.05 1.24 3.1 1.64 1.19 1.02 1.84 2.2 Area of box woodland (%) 49 37 31 56 47 64 61 41 Area of open box woodland 1 (%) 51 63 69 44 53 36 39 59 PVC of box woodland 2 (%) 50 75 60 60 45 70 65 60 No. mature box patches 0 0 0 0 2 3 0 3 Area of mature box patches (m 2) 0 0 0 0 450 810 0 820 No. sapling box patches 2 3 5 2 0 2 4 0 Area of sapling box patches (m 2) 850 700 1550 1300 0 650 400 0 No. live isolated trees large box 3 18 12 19 18 30 13 103 76 large other 0 21 7 15 8 6 2 0 small 130 246 240 480 57 82 153 152 No. dead trees standing large 2 0 3 3 1 1 0 1 small 8 1 4 15 21 21 35 8 No. fallen dead trees large 6 2 6 0 0 0 2 3 small 10 0 4 3 4 4 8 10 No. logs large 4 2 3 6 6 1 4 3 small 11 34 58 98 41 52 111 52 No. stumps 17 5 12 12 4 2 8 7 No. sheep fences 1 0 1 1 1 1 1 1

1Percentage vegetation cover (PVC) <10% 2PVC of all trees, including box saplings >4 m high 3Includes Tumbledown Gum Eucalyptus dealbata

Areas of vegetation were classified as woodland or open woodland, depending on whether the percentage vegetation cover (PVC) of trees was greater or less than 10%, respectively (sensu Specht 1970). Trees and logs were classified as large or small depending on whether they were larger or smaller than about half the size of the tallest trees (~20 m) and largest logs (~8 m long × 0.6 m diameter), respectively. Vegetation areas within breeding territories were calculated from measurements made with a portable rolling-wheel odometer.

Results

Breeding territories Eight breeding territories were determined around the nests listed in Table 1. VOL. 25 (3) SEPTEMBER 2008 Jacky Winter Breeding Biology 125

Table 2 Physical characteristics of 23 nest-sites of the Jacky Winter at Glenelg , south-eastern Qld, September 2005 to February 2006.

Median (m) Range (m)

Height of nest above ground 8 3.2–11.2 Tree height 15 8.8–19.5 Height of vegetation over nest 4.5 0.3–13.5 Horizontal distance of nest from trunk 3.9 1.5–9.1 Horizontal distance to extremity of branch 0.7 0.3–2.4

The extent of new or adjusted territories that were occupied in the latter part of the breeding season around other nests was not determined. Distances between the centres of adjacent territories were: T1–T2 = 1000 m, T2–T3 = 270 m, T3–T4 = 550 m, T4–T5 = 400 m, T5–T6 = 250 m, T6–T7 = 250 m and T7–T8 = 190 m. The mean territory area was 1.7 ha (range 1–3.1 ha, n = 8). The breeding density was one pair per 17.5 ha (eight territories in 140 ha). All territories consisted of box woodland (mean = 48% of area) and open box woodland (mean = 52% of area). They were characterised by two to five patches of either mature box trees or sapling box trees, as well as mature isolated live trees, dead trees (standing and fallen), logs and stumps (Table 1). Territories 7 and 8 contained numerous large, isolated box trees. No nests were found in sub-areas X, Y or Z (see Figure 1), despite the presence of Jacky Winters. Sub-area X was open woodland. Sub-area Y was a combination of woodland and open woodland, and calling Jacky Winters, presumed to be males, were present, mostly in September and October. Noisy Miners Manorina melanocephala and other honeyeaters were generally absent from sub-areas X and Y. Sub-area Z was woodland and was occupied by one or two colonies of Noisy Miners. Jacky Winters were generally absent from other sub-areas such as the Angophora –box and Angophora –Red Gum forests (Figure 1).

Nests Twenty-three nests were constructed progressively within territories (Figures 1 and 2). Typically, a new nest was built when a former nest failed or was successful. Because only one individual was colour-banded, it is not known if replacement nests were used by the same pair of birds. Certainly, the yellow-banded male that attended nest 5 from week 2 to week 4 was not seen in attendance at adjacent nest 21 from week 13 to week 15, and not seen in attendance at nearby nest 13 in week 8. Adult birds additional to the primary male and female were sometimes seen in the nest-tree, but there was no regularity in their presence. All nests were in almost-horizontal forks of mature eucalypt trees, close (median 0.7 m) to the extremity of the branch (Table 2). Eight were in Grey Box, eight in Yellow Box, three in Forest Red Gum and four were in Tumbledown Gum. Most if not all nest-trees were judged as being alive ~160 years ago when the site was initially cleared. Twenty-one nests were in forks consisting of a pair of dead branchlets near the extremity of a much longer live branch. The two remaining nests had one of the forked branchlets dead, and the other alive. Four nests (1, 6, 11 and 19) were in the same Yellow Box, and two nests (13 and 21) were in the same Forest Red Gum. Some inter-nest distances within the same AUSTRALIAN 126 WOOD, THOMPSON & LEY FIELD ORNITHOLOGY . Failed nests are shown are nests Failed . ed, ed, DIS = disappeared. L , south-eastern Qld, monitored s) are displayed in italics below below italics in displayed are s) is shown as 1F or 2F. The number is shown as 1F or 2F. k 1) to 2 February 2006 (week 21). Nest Nest 21). (week 2006 February 2 to 1) k Glenelg t , fledgling(s) = fledgling(s) , n , nestling stage = stage nestling , l , incubation = incubation , U weekly (except week 18) from 15 September 2005 (wee 2005 September 15 from 18) week (except weekly as as A = abandoned, PD = partly damaged, SD = severely damag The number of birds that fledged from successful nests number. each week of observer-hours and number of observers (in parenthese (in observers of number and observer-hours of building = building Figure 2. Figure Breeding status of 23 nests of the Jacky Winter a

territory were: nest 4–nest 15 = 25 m, 15–17 = 31 m, 8–20 = 40 m, 5–13 = 50 m, 9–11 = 45 m, 3–14 = 22 m, 2–7 = 40 m, 7–12 = 17 m and 16–22 = 38 m. Nest 23 was 105 m from nest 2 in T6, and is likely to have belonged to T6, as nest 22 in T7 fledged a juvenile at about the same time (Figure 2).

Breeding success Of 23 nests, three failed before incubation was observed (nests 2, 6 and 18: Figure 2). Three other nests were abandoned, five nests were partly damaged, three were severely damaged, four disappeared and five nests produced a total of seven fledglings (Figure 2). of eggs or nestlings was not observed, but White- plumed Lichenostomus penicillatus or Fuscous L. fuscus Honeyeaters were seen at five damaged nests, inspecting remnant nesting materials. The following avian VOL. 25 (3) SEPTEMBER 2008 Jacky Winter Breeding Biology 127

Table 3 Nesting and breeding success of the Jacky Winter at Glenelg , south-eastern Qld, between September 2005 and February 2006. Clutch-size assumed to be two eggs where eggs not seen.

N nests N nests N eggs N N nests with N Hatching Nesting Breeding built with eggs laid nestlings fledglings fledglings success success success 23 20 40 16 5 7 40% 25% 17.5% nest-predators (from Major et al . 1996; Taylor & Ford 1998; Higgins 1999; Higgins & Peter 2002) were present at the site: Laughing Kookaburra Dacelo novaeguineae , Noisy Friarbird Philemon corniculatus , Grey Shrike-thrush Colluricincla harmonica , Grey Butcherbird Cracticus torquatus , Pied Butcherbird C. nigrogularis , Australian Magpie C. tibicen , Pied Currawong Strepera graculina and Torresian Crow Corvus orru . Overall, breeding success was 17.5% (seven fledglings from 40 eggs: Table 3). Nesting success was 25% (5/20). Of 18 nests that failed, most failures (44%) were in November (Figure 2). Of the five successful nests, two nests (40%) succeeded in December (Figure 2). From the locations of successful nests and sequence of events in the various territories (Figures 1 and 2), it is likely that the seven juveniles fledged from five nests occupied by five different pairs. Productivity was 0.88 young per pair (seven fledglings from eight territories).

Parental care of fledglings We recorded parental care of fledglings from nest 3 during the nest-building and incubation stages of replacement nest 14 (Figure 2). The two nestlings from nest 3 fledged between 7 and 12 October. On 13 October (study week 5), these two fledglings were watched for ~1 h while they were fed by parent birds in trees <8 m from the nest. In week 6, during 3 h of observation on successive days, only one fledgling was fed in the tree-canopy, leading us to conclude that the other had been killed or had died. The surviving fledgling once flew with a parent ~70 m circuitously, returning above nest 3 to the same clump of trees. During 66 minutes of observation in week 7, this juvenile was again fed by both parents in the canopy and was seen to make flights of ~25 m and 70 m with parent birds. Three watches (80, 20 and 60 min.) were conducted on successive days in week 8 when new nest 14 was found ~25% complete (Figure 2). For the first time, we saw the juvenile fly to logs on the ground and glean insects twice. For 30 minutes during one watch, the juvenile perched on the new nest-branch, 10–15 cm away from nest 14, and was fed there, once by the female and once by the male. In week 9, the female laid the first egg between 1145 h on 10 November and 0820 h on 11 November. During three watches (120, 30 and 220 min.) on successive days, the juvenile gleaned insects from foliage four times, snatched prey from foliage twice, ground-pounced twice and hawked successfully twice. It ranged to 120 m from nest 14 while following the male and begging for food. It was fed occasionally by both parents during these watches. During 80 minutes of observations in week 10, the female incubated two eggs in several successive stints while the juvenile foraged with the male, sometimes receiving food after begging and sometimes capturing prey itself. During 30 minutes of observation in week 11, the juvenile foraged successfully by hawking, ground-snatching and hovering. In week 12, nest 14 was found empty AUSTRALIAN 128 WOOD, THOMPSON & LEY FIELD ORNITHOLOGY and abandoned. During 40 minutes of observation, the juvenile fed itself 10 times (four ground-pounces, three foliage-snatches, two bark-snatches and once by pouncing after hawking), and was fed once by a parent. It was beginning to moult into immature plumage (see Boles 1985). The territory was not visited in week 13, but in week 14 the juvenile was still in moult and did not receive food from either parent while it was watched for 10 minutes. In summary, the juvenile was probably dependent for ~9 weeks after fledging (study weeks 5 to 14) when it was moulting into immature plumage.

Discussion Our finding of a mean territory size of 1.7 ha is consistent with territory areas of similarly-sized robins in other parts of Australia. At Nimmitabel, NSW, Flame Robins phoenicea (12–16 g) occupied a mean territory area of 1.79 ha and Scarlet Robins P. boodang (12–14 g) held 3.2 ha (Robinson 1990a). At Moruya, NSW, in wet eucalypt forest, the australis (18–23 g) occupied a mean territory area of 1.3 ha (Marchant 1987). However, unlike those of the Eastern Yellow Robin (Marchant 1987) or the (Robinson 1992), territories of the Jacky Winter were not contiguous. Jacky Winters held territories by regular sustained calling (peter-peter …) usually from the tops of particular tall trees (17–20 m: KAW unpubl. data). Territorial advertisement by Jacky Winters was similar to that of the Rufous Whistler Pachycephala rufiventris (McDonald 2001), a species whose territories are also non-contiguous (Bell & Ford 1987; Bridges 1994). The overall breeding density of Jacky Winters, about one pair per 17.5 ha, is comparable to that of the Eastern Yellow Robin (one pair per ~30 ha) and Scarlet Robin (one pair per ~25 ha) at a 270-ha woodland site near Armidale, NSW (Debus 2006a). Habitat selection by some Australian has received attention in recent years (Oliver 2000; Cousin 2004), but there is only scant knowledge of the preferred habitat of the Jacky Winter. The present study showed that breeding territories comprised similar proportions of box woodland and open box woodland (mean = 48% and 52% of area, respectively). These territories were characterised by patches of box trees and isolated live trees, dead trees (standing and fallen), logs and stumps (Table 1). Additionally, the study showed that Eucalyptus –box woodland, which appeared to be a healthy landscape, was not entirely occupied by Jacky Winters. Indeed, Jacky Winters were generally absent from the small Angophora –box and Angophora –Red Gum forests within the woodland. Similarly, the study showed that nests were not built (or did not survive) in some sub-areas where Jacky Winters were present (sub-areas X, Y, Z: Figure 1). Sub-area X may have been unsuitable for breeding because it was woodland with only a few trees, and in sub-area Z, Jacky Winters might have been discouraged from nesting because they experienced aggression from Noisy Miners (Dow 1977; Barrett et al . 1994). However, the reason for the absence of nests from sub-area Y is not known. This sub-area consisted of habitat which appeared to be similar to the habitat in other territories, Noisy Miners were generally absent, and calling Jacky Winters, presumed to be males, were seen there on most visits between September and December. Breeding success of Jacky Winters in this study was based on a small sample of only 20 nests and therefore may not be representative of the species. Nevertheless, the calculated value of 17.5% was higher than that of the Scarlet Robin at Nimmitabel (7.8%), but lower than breeding success of the at Nimmitabel (22.2%) and Eastern Yellow Robin at Moruya (28%) (see Table 4). VOL. 25 (3) SEPTEMBER 2008 Jacky Winter Breeding Biology 129

Table 4 Height of nest above ground, nesting and breeding success of some Australian robins (Mayfield method nest-success values from Debus 2006a), ns = not stated. Source: 1 = this study; 2 = Robinson 1990b; 3 = Debus 2006a,b; 4 = Marchant 1985b.

Species Location Mean or median Nesting Breeding Source height of nest success success above ground (m) (%) (%)

Jacky Winter Inglewood, Qld 8 25 17.5 1 Scarlet Robin Nimmitabel, NSW 7.2 10.3 7.8 2 Armidale, NSW 7.2 8 ns 3 Flame Robin Nimmitabel, NSW 4 25 22.2 2 Eastern Moruya, NSW 3 32 28 4 Yellow Robin Armidale, NSW 2.7 24 ns 3

Nesting success of the Jacky Winter in the present study (25%) was higher than that of the Scarlet Robin at Nimmitabel (10.3%) and Armidale (8%), but comparable to the nesting success of the Flame Robin at Nimmitabel (25%) and Eastern Yellow Robin at Armidale (24%) (see Table 4). Avian predation and pilfering of materials from constructed nests were assumed to be the most important causes of failure, although predation by mammals is also possible (Fulton 2006). Known avian predators (from Major et al . 1996; Taylor & Ford 1998; Higgins 1999; and Higgins & Peter 2002) that were moderately abundant at the site were the Pied Currawong, butcherbirds, Laughing Kookaburra, Grey Shrike-thrush and Noisy Friarbird. Two Lichenostomus honeyeaters, White-plumed and Fuscous, are known pilferers of nest-material (Ley et al . 1997), and were seen inspecting five damaged nests. All 23 nests were placed in almost-horizontal forks of mature eucalypt trees, close (median 0.7 m) to the extremity of the branch (Table 2). Twenty-one (91%) were in forks consisting of a pair of dead branchlets near the extremity of a long live branch, and most if not all nest-trees were considered to be at least 160 years old. The sample of nests is small, but nest-site selection at Glenelg suggests that woodland without very old eucalypt trees might not provide nest-sites preferred by Jacky Winters. Further work is required elsewhere to determine the ages of other nest-trees used by this species. Open-cup nests were placed in exposed positions at a median height of 8 m above the ground, and nestlings depended on crypsis to avoid detection. Although this height was similar to that for Scarlet Robin nests (7.2 m) near Nimmitabel and at Armidale, the nesting success of Jacky Winters at Glenelg was higher than that of Scarlet Robins at either location (Table 4). Conversely, Eastern Yellow Robins at Moruya and Armidale and Flame Robins at Nimmitabel nested closer to the ground (3 m, 2.7 m and 4 m, respectively, above the ground), with more vegetation cover, and their corresponding nesting successes were higher than Jacky Winters at Glenelg (Table 4). Jacky Winters in the present study may have had greater nesting success than Scarlet Robins at Nimmitabel and Armidale because they defend their nests against avian predators, whereas these other robins generally do not (S. Debus pers. comm.). In the Australian robins, few studies have reported the pattern of nest establishment and replacement through time and space. Such a pattern, as AUSTRALIAN 130 WOOD, THOMPSON & LEY FIELD ORNITHOLOGY determined for the Jacky Winter in the present study (Figures 1 and 2), is preliminary because the sample of nests is small and some territories were not mapped. However, our observations of the birds holding the eight territories, T1 to T8, were sufficient to determine the boundaries and times of occupancy of these particular territories. This information might assist in a better understanding of the process of predation (see Tokue & Ford 2006). The post-fledging dependence period for one juvenile (~9 weeks) was similar to a post-fledging dependence of 8 weeks for the Eastern Yellow Robin and 6 weeks for the Scarlet Robin (Debus 2006b). Marking of individuals and greater observer effort are recommended in future studies.

Acknowledgements We are grateful to Jan and Max Fitzgerald for allowing us to perform this study on their property. Stephen Debus and referees Graham Fulton and Hugh Ford made helpful comments to improve the manuscript. Licences to band and release birds were obtained from the Australian and Bat Banding Scheme and Queensland Parks and Wildlife Service.

References Barrett, G., Ford, H.A. & Recher, H.F. (1994), ‘Conservation of woodland birds in a fragmented rural landscape’, Pacific Conservation Biology 1, 245–256. Barrett, G., Silcocks, A., Barry, S., Cunningham, R. & Poulter, R. (2003), The New Atlas of Australian Birds , Royal Australasian Ornithologists Union, Melbourne. Bell, H.L. & Ford, H.A. (1987), ‘Fidelity to breeding-site in four migratory species near Armidale, New South Wales’, Corella 11 , 1–5. Beruldsen, G. (2003), Australian Birds, Their Nests and Eggs , author, Brisbane. Blakers, M., Davies, S.J.J.F. & Reilly, P.N. (1984), The Atlas of Australian Birds , Melbourne University Press, Melbourne. Boles, W.E. (1985), ‘Bird in the hand: Jacky Winter Microeca leucophaea ’, Corella 9, 66. Boles, W.E. (1988), The Robins and Flycatchers of Australia , Angus & Robertson, Sydney. Bridges, L. (1994), ‘Territory and mate fidelity in a migratory population of the Rufous Whistler Pachycephala rufiventris ’, Emu 94 , 156–165. Cousin, J.A. (2004), ‘Habitat selection of the (Eopsaltria griseogularis ) in a Wandoo woodland, ’, Emu 104, 229–234. Debus, S.J.S. (2006a), ‘Breeding and population parameters of robins in a woodland remnant in northern New South Wales, Australia’, Emu 106, 147–156. Debus, S.J.S. (2006b), ‘Breeding biology and behaviour of the Scarlet Robin Petroica multicolor and Eastern Yellow Robin Eopsaltria australis in remnant woodland near Armidale, New South Wales’, Corella 30 , 59–65. Dow, D.D. (1977), ‘Indiscriminate interspecific aggression leading to almost sole occupancy of space by a single species of bird’, Emu 77 , 115–121. Ford, H.A., Barrett, G. & Recher, H. (1995), ‘Birds in a degraded landscape—safety nets for capturing regional biodiversity’, pp. 43–50 in Saunders, D.A., Craig, J.L. & Mattiske, E.M. (Eds), Nature Conservation 4: The Role of Networks , Surrey Beatty, Sydney. Ford, H.A., Barrett, G.W., Saunders, D.A. & Recher, H.F. (2001), ‘Why have birds in woodlands of southern Australia declined?’, Biological Conservation 97 , 71–88. French, M. (1990), A Pastoral Romance—The Tribulation and Triumph of Squatterdom , University of Southern Queensland Press, Toowoomba, Qld. Fulton, G.R. (2006), ‘Identification of nest predators with remote cameras and artificial nests in extensive old-growth woodland of south-western Australia’, Corella 30 , 35–39. Higgins, P.J. (Ed.) (1999), Handbook of Australian, New Zealand and Antarctic Birds , vol. 4, Oxford University Press, Melbourne. Higgins, P.J. & Peter, J.M. (Eds) (2002), Handbook of Australian, New Zealand and Antarctic Birds , vol. 6, Oxford University Press, Melbourne. Keast, A. (1993), ‘Song structures and characteristics: Members of a eucalypt forest bird community compared’, Emu 93 , 259–268. VOL. 25 (3) SEPTEMBER 2008 Jacky Winter Breeding Biology 131

Keast, A. (1994), ‘The annual cycle in a vocalisation context: A comparison of the Eastern Yellow Robin Eopsaltria australis and Jacky Winter Microeca leucophaea ’, Emu 94 , 230–238. Ley, A.J., Oliver, D.L. & Williams, M.B. (1997), ‘Theft of nesting material involving honeyeaters (Meliphagidae)’, Corella 21 , 119–123. Major, R.E., Gowing, G. & Kendall, C.E. (1996), ‘Nest predation in Australian urban environments and the role of the Pied Currawong, Strepera graculina ’, Australian Journal of Ecology 21 , 399–409. Marchant, S. (1985a), ‘Nesting of the Jacky Winter’, Australian Birds 20 , 22–24. Marchant, S. (1985b), ‘Breeding of the Eastern Yellow Robin’, pp. 231–240 in Keast, A., Recher, H.F., Ford, H.A. & Saunders, D. (Eds), Birds of Eucalypt Forests and Woodlands , Surrey Beatty, Sydney. Marchant, S. (1987), ‘Territorialism and co-operative breeding of the Eastern Yellow Robin Eopsaltria australis ’, Corella 11 , 6–14. McDonald, P.G. (2001), ‘The function of vocalisations and aggressive behaviour used by male Rufous Whistlers Pachycephala rufiventris ’, Emu 101, 65–72. Odum, E.P. & Kuenzler, E.J. (1955), ‘Measurement of territory and home range size in birds’, Auk 72 , 128–137. Oliver, D.L. (2000), ‘Foraging behaviour and resource selection of the Regent Honeyeater Xanthomyza phrygia in northern New South Wales’, Emu 100, 12–30. Olsen, P., Weston, M., Tzaros, C. & Silcocks, A. (2005), ‘The state of Australia’s birds’, Supplement to Wingspan 15 , 1–32. Recher, H.F. & Holmes, R.T. (1985), ‘Foraging ecology and seasonal patterns of abundance in a forest avifauna’, pp. 79–96 in Keast, A., Recher, H.F., Ford, H.A. & Saunders, D. (Eds), Birds of Eucalypt Forests and Woodlands , Surrey Beatty, Sydney. Recher, H.F., Davis, W.E. & Calver, M.C. (2002), ‘Comparative foraging ecology of five species of ground-pouncing birds in Western Australian woodlands with comments on species decline’, Ornithological Science 1, 29–40. Robinson, D. (1990a), ‘The social organisation of the Scarlet Robin Petroica multicolor and Flame Robin P. phoenicea in southeastern Australia’, Ibis 90 , 78–94. Robinson, D. (1990b), ‘The nesting ecology of sympatric Scarlet Robin Petroica multicolor and Flame Robin P. phoenicea populations in open eucalypt forest’, Emu 90 , 40–52. Robinson, D. (1991), ‘Threatened birds in : Their distributions, ecology and future’, Victorian Naturalist 108, 67–77. Robinson, D. (1992), ‘Why do Flame Robins migrate? A comparison between the social and feeding ecologies of the Flame Robin Petroica phoenicea and Scarlet Robin P. multicolor ’, Corella 16 , 1–14. Schodde, R. & Mason, I.J. (1999), The Directory of Australian Birds: Passerines , CSIRO, Melbourne. Schodde, R. & Tidemann, S.C. (Eds) (1986), Readers Digest Complete Book of Australian Birds , 2nd edn, Readers Digest Services, Sydney. Specht, R.L. (1970), ‘Vegetation’, in Leeper, G.W. (Ed.), The Australian Environment , 4th edn, CSIRO, Melbourne. Taylor, L.N.H. & Ford, H.A. (1998), ‘Predation of artificial nests in a fragmented landscape on the New England Tablelands of New South Wales’, Wildlife Research 25 , 587–594. Tokue, K. & Ford, H.A. (2006), ‘Influence of food and nest predation on the life histories of two large honeyeaters’, Emu 106, 273–281.

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