22 AUSTRALIAN FIELD ORNITHOLOGY 2005, 22, 22- 28 Breeding Behaviour of the near Armidale, New South Wales

S.J.S. DEBUS and G. LOLLBACK Division of Zoology, University of New England, Armidale, New South Wales 2351

Summary A pair of Restless Flycatchers inquieta was observed in eucalypt woodland near Arm idale, New South Wales, fo r 10.2 hours over 5 days during the nest-building (6 h) and early incubation phases ( 4.2 h) in spring 2000. Male and female had similar rates of nest-building visits (7 and 4.5 visits/h respectively) and contributions to incubation (60% and 46% of incubation time respectively) , covering the eggs for 52% of observation time. Male and female also called from the nest during both phases, including while incubating, at simi lar rates; the whistle and grate call types were given at similar rates by both sexes. Incidental observations of two pairs over 2 years, on nesting chronology, nest-sites, fl edgling behaviour and nest success (14%, n = 7), are given.

Introduction The Restless Flycatcher Myiagra inquieta is a common and familiar in rural Australia where remnant bushland remains, yet its biology is little documented. There is a brief study of its breeding behaviour (Roberts 1942), and there are unquantified and unsourced general statements on its biology in bird handbooks and field-guides: its nest, eggs, clutch-size and fledgling morphology are described; nest -building takes 4-7 days, incubation and nestling periods are about 14 days each, both sexes build the nest, incubate, brood and feed the nestlings, and pairs may be multi-brooded (Schodde & Tidemann 1986, Boles 1988). As the Restless Flycatcher is one of the declining in fragmented woodlands of the sheep-wheat belt in Australia (Cogger et al. 2003), there is a need for quantified information on its ecology as a basis for conservation. Furthermore, quantified information on its parental behaviour and breeding biology is of general ecological interest, with regard to the interplay between social monogamy, sexual dimorphism, sex roles and parental breeding investment, and the possibility of extra-pair paternity (Goodey & Lill1993, Tremont & Ford 2000). The aim of this study was primarily to quantify male and female contribution to nest-building and incubation in the Restless Flycatcher, a sexually almost monomorphic . Other behavioural observations, and data on nest success, are included.

Study area and methods The study site was Imbota Nature Reserve (formerly Eastwood State Forest), 10 km south­ east of Armidale (30°30' S, 151°40' E) on the New England Tablelands of New South Wales. This woodland patch of about 270 ha, including contiguous woodland on private property, has been described by Ford eta!. (1985, 1986), Tremont & Ford (2000) and NPWS (2002). The physiography and climate of the New England region are described elsewhere (Heatwole & Simpson 1986; Heatwole eta!. 1995, 2003). Two pairs of Restless Flycatchers in habited Imbota in 2000. The pair in territory A was observed for approximately 2 hours per day, between 0730 and 1600 h (standard time), on five VOL. 22 (1) MARCH 2005 Breeding Behaviour of Restless Flycatcher 23 non-consecutive days between 17 October and 3 November 2000 (total 10.2 h). The adults were initially sexed by the black Iores of the male versus grey Iores in the female (Schodde & Tidemann 1986); they were colour-banded by SD on 19 October, before the second observation day (20 October). Data on calling rates were collected on the four observation days after the could be positively sexed by their colour-bands. Two observers (GLandS. Teale) stood 20m from the nest with 8 x binoculars, one dictating events while the other scribed the details and times. The birds appeared unaffected by the proximity of observers. The fema le of the pair in territory B was also colour-banded in spring 2000. Nest-watches in territory A in 2000 were conducted from the late nest-building stage (final placing of plant fibres and cobweb on the cup) to early incubation stage. Incubation was inferred from prolonged sitting on the nest (>6 minutes without other activity), lack of nest-building or maintenance, and egg-turning behaviour by the sitting bird. Nest-building occurred on the first two and the fifth observation days (the last being a repeat nest after failure), and incubation was in progress on the third (24 October) and fourth (27 October) observation days. Nest-building days were pooled for analysis, as the same birds of pair A were involved at the two nests. Other nests in 2000 and 2001 were monitored opportunistically by SO. Vocalisations of the pair in territory A in 2000were quantified as discrete bouts of the repeated whistle calls (1 bout = 1 call), and as single grate calls (the monosyllabic harsh call), from the second observation day so that the individuals were accurately sexed on the basis of colour­ bands. Variations on the whistle calls (upslurred versus downslurred) and grate calls (abrupt versus slightly extended, cf. Buckingham & Jackson 1992) were not separated for the purpose of analysis. Paired t-tests were used to compare sex roles in nest-building, and to compare calling rates on the nest within and between the sexes according to call type.

Results

Territories and occupancy Imbota was occupied by two pairs of Restless Flycatchers in 2000, 1 km apart: territory A (o' red/yellow, 9 red/blue = dRY, 9 RB) and B (o' unhanded, 9red/green = o'U, 9RG). During 2001 9RG disappeared and 9RB moved to pair with aU; an unhanded pair (o' subsequently metal-banded= o'Me) then arrived, ousted mateless RY and occupied his territory. In August 2001 RY was seen once in Imbota, remote (1 km) from either territory, then disappeared. A pair (o'Me, 9U) continued to occupy territory A to autumn 2003, but the pair in territory B (o'U, 9RB) had one unsuccessful nesting attempt in spring 2001 then disappeared. In autumn 2001 the pair in territory A ranged out into surrounding paddocks 1 km from the nesting area.

Ten-itmy and mate defence The female of the pair in territory A (9RB) was captured during the nest­ building phase, and while she was being removed from the mist-n et the male (dRY) was captured when he defended her by swooping and 'scolding' the handler with the grate call. In 2001 the new male in territory A ( o'Me) was captured for banding when he vigorously attacked his reflection in the mirrors of a vehicle, and entered the vehicle through an open door (B. Chaffey pers. comm.).

Nesting chronology The pair in territory A (pair A) commenced breeding activity in September 2000, laid in early October, and had repeat attempts until December, with no re­ laying in January 2001 (Table 1). The pair in territory B (pair B) followed a similar pattern, but a first attempt in 2000 may have been missed and the pair disappeared before a second attempt in spring 2001. Pair B fledged a brood of four in early AUSTRALIAN 24 DEBUS & LOLLBACK FIELD ORNITHOLOGY

Table 1 Nesting chronology of two pairs of Restless Flycatchers in two territories (A and B) near Armidale, NSW (see page 23 for identity of individual birds of each pair).

Date (day.month) Event(s)

Pair A, 2000-01: 13.9 Nest-building; half-built (N1) 14.9 Nest storm-damaged, abandoned. Further attempt(s) over next 2 weeks not detected. 29.9 Pair selecting new nest-site 2.10 Sitting on completed nest (taken as N2) 11.10 Incubating 13.10 Nest vacant; adults selecting new site (N2 damaged 17.10) 17-22.10 Building and finishing new nest (N3) 24-27.10 Incubating 30.10 Broken eggs on ground (predation); building new nest (N4) 3.11 Building 8-17.11 Incubating 21.11 Eggs taken by predator 11-12.12 Building new nest (N5) 18.12.00-1.1.01 Incubating/brooding 2-5.1 Nest vacant (failed); no further nesting activity Pair B, 2000-01: 1.11 Selecting nest -site 8.11 Nest-building 13.11 Sitting; finishing nest 17-21.11 Incubating 26.11 Broken eggs on ground (predation); building new nest (N2) 7-22.12 Incubating/brooding 24-29.12 Nestlings visible 30.12.00-1.1.01 Near-fledged 2-3.1 1 fledgling out of nest 10.1 4 fledglings Pair A, 2001: 2.10 Nest-building (N1) 4.10 N1 destroyed, building new nest (N2) to 11.10 12.10 Sitting (laying?) 15-31.10 Incubating/brooding 2-8.11 Nestlings (2+) visible, half-grown 6.11 12.11 Nest empty (predation), adults selecting new nest-site (no further monitoring; no fledglings in summer) Pair B, 2001: 2-19.10 Incubating/brooding; no further activity, adults disappeared VOL. 22 (1) MARCH 2005 Breeding Behaviour of Restless Flycatcher 25

January 2001, asynchronously over 2+ days, suggesting either that incubation started before the clutch was complete or that the nest was too crowded for all four large chicks. Pair A fledged no young.

Nest-sites and nest-building Known nest-sites of pair A were in live Blakely's Red Gums Eucalyptus blakelyi (n = 6) or Manna Gum E. viminalis (n = 1) near or beside water, within about 50 m of an earth dam. The birds often perched on and foraged from fallen branches protruding from the water. Nest-sites were typically a fork in a horizontal live or dead branch, <1m beneath a near-horizontal live branch of similar orientation, in an open situation in the tree canopy. The pair appeared to nest in loose association with a pair of Magpie-larks cyanoleuca that had active nests in neighbouring Red Gums beside the dam, and with a pair of Willie Wagtails Rhipidura leucoph1ys that nested successfully in the same area. No interactions between any of these species were observed. Pair B nested in the crowns of live eucalypts [Broad-leaved Stringybarks Eucalyptus caliginosa (n = 2) or Yellow Box E. melliodora (n = 1)], in trees within 20m of that containing an active Little Eagle Hieraaetus morphnoides nest. A pair of Willie Wagtails also nested successfully near the Eagles, once in a dead mistletoe in the lower canopy of the Eagles' tree. No interactions between the two flycatcher species, or between either flycatcher species and the Eagles, were observed. The Restless Flycatchers re-nested almost immediately after failure at the egg stage (Table 1). Within 1-2 days of egg predation the adults were selecting a new site by testing several possible sites: squatting in an apparently suitable horizontal fork and calling. The pair started building on the favoured site, signalled by frequent squatting and calling on it, within a further day. One repeat nest was apparently complete, with the bird sitting on it, 4 days after site selection. The next repeat nest was partly built 4-5 days after the previous failure. Pair A was observed as the birds built their third and fourth nests of the season. The male visited the nest 60 times, 70% of visits being to build the nest, and the female 36 times, 78% to build, in 6 hours of observation (7.0 ± S.E. 2.0 and 4.5 ± S.E. 0.42 nest-building visits/h for male and female respectively), with no significant difference between the sexes (t2 = 1.01, P = 0.42).

Incubation and parental care The adults of pair A incubated for 52% of observation time ( 4.1 h), leaving the eggs uncovered for a total of 2 hours ( 48% ). The contribution of both sexes was similar on average, although the male incubated about twice as much as the female on the first observation day (24 October 2000, 2 days after nest completion) and the reverse applied on the second observation day (27 October). The male sat on the nest for 60% of incubation time, and the female for 46%. The male incubated for 18 stints averaging 4.3 minutes, with an average of 4.7 minutes between stints, and the female for nine stints averaging 5.7 minutes, with an average of24.6 minutes between stints. Minor nest-building or maintenance, of addition of cobweb to the inside of the cup, occurred during the incubation phase. At other nests of pairs A and B both sexes incubated; one incubation changeover was seen for pair B. For the only nest of pair A that reached the nestling stage before failure (2001), and for the successful nest of pair B (2000), both sexes AUSTRALIAN 26 DEBUS & LOLLBACK FIELD ORNITHOLOGY

Table 2 Calling rate, by call type, for male and female Restless Flycatchers on the nest in the nest-building and incubation periods (five observation days): mean number of calls per minute (standard error in parentheses).

Sex Whistle Grate Male 1.46 (1.06) 1.29 (0.30) Female 0.51 (0.24) 1.26 (0.64)

shared brooding and feeding of nestlings and feeding of fledglings. Colour-banding confirmed that only two birds attended the A nests, and only two birds (banded female and unhanded male) attended the B nests.

Calling rates Both birds of pair A made both types of call (whistle and grate) at the nest, in the nest -building and incubation phases, at an average rate of 3.4 calls per minute. During the observation period, the male and female made 2.7 and 1.8 calls per minute, respectively. Calling rates of each sex during the nest-building phase were similar (male 2.5 calls/min., female 2.3 calls/min.; t2 = 0.02, P = 0.99). Calling rates of each sex during the incubation phase were also similar (male 1.7 calls/ min., female 0.9 call/min.; t1 = 0.89, P = 0.54). Male and female both gave 1.3 grate calls per minute while on the nest. Similarly, there was no significant difference between the sexes in the rate of whistle calls given while on the nest (male 1.5 calls/min., female 0.5 call/min.; t1. = 0.99, P = 0.38). Additionally, there was no significant difference in the rate ot each call type for the male (t4 = 0.11, P = 0.92) or female (t = 1.36, P = 0.23). However, there were only three observation sessions in dw nest-building period and two in the incubation period, with high statistical variance, blurring any intersexual differences that may prove real with larger sample sizes (Table 2).

Fledglings At 1 week post-fledging, the four fledglings of pair B huddled together or within a few metres of one another in the same tree, in the canopy of live eucalypts within 50 m of the nest. At that stage they flew little, although competently, and did not descend to mist-net height (i.e. <3 m). By the end of their second week they flew frequently, ranged about 100m from the nest, and started to descend nearer ground-level ( <2 m) with their parents. Thereafter, they ranged at least 200 m from the nest before dispersing by autumn.

Breeding success Pair A had five known nesting attempts in spring 2000, all unsuccessful (one nest dislodged by wind, four had the eggs taken by a predator of unknown identity). Pair B had one unsuccessful attempt (eggs taken by a predator), then successfully fledged a brood of four in January 2001. Nest success in 2000was 14% (n = 7). Pair A reared no fledglings in 2001 or 2002, from an undetermined (but multiple) number of nesting attempts in each year. VOL. 22 (1) MARCH 2005 Breeding Behaviour of Restless Flycatcher 27

Discussion Our observations are consistent with previous knowledge of the Restless Flycatcher's basic biology (Roberts 1942, Schodde & Tidemann 1986, Boles 1988), this study providing some quantification of sex roles and confirmation of social monogamy. Male and female shared nest-building and incubation equally, although the clutch may not have been complete on the first observation day of incubation and the female may have later increased her share. An incomplete clutch may also explain the frequently uncovered eggs. The male called more frequently from the nest than the female did, although the difference was not statistically significant. Samples sizes (number of observation days and hours) were small and the study based mostly on one pair, so further quantified study is required. The results of this study were similar to those of Roberts (1942) for nest-building rates (7.8 nest-building visits/h for male and 8.5/h for female). However, Roberts recorded incubation stints averaging 17 minutes, with a suggestion of greater contribution by the female (longest stint, 31 min.) than the male (shortest stint, 5 min.), presumably when the clutch was complete and incubation fully established. Roberts (1942) also found that parental feeding rates to nestlings were about equal (male 13 feeds/h, female 11.5/h; observation period = 2 h). Breeding success of the Restless Flycatcher was low, nest success (14%) being similar to that of the Myiagra mbecula at lmbota (19%: Ford et al. 2001). The Restless Flycatchers at lmbota suffered frequent nest predation, and had frequent re-nesting attempts. High nest predation may be partly attributable to the open nesting situation, the amount of time spent off the eggs, and to frequent advertising of the site by calling from nests (which were easily found by humans). Calling from the nest may function as intraspecific territorial advertisement. Rapid turnover of nesting attempts, and nesting near aggressive species for possible protection, may be strategies for counteracting predation. Low nest success, and high rates of nest predation, are typical for declining woodland passerines (Ford et al. 2001). The breeding and calling behaviour of the Restless Flycatcher was similar to that described for the Leaden Flycatcher, whose whistling (siren or chewee) and harsh zeep calls are homologous with the former's whistle and grate calls respectively (see Tremont 1994, Tremont & Ford 2000). Leaden Flycatchers also call frequently from the nest. In the sexually dimorphic Leaden Flycatcher both sexes share nest­ building equally, females contribute more daytime incubation and brooding, and both share feeding of nestlings and fledglings equally. Leaden Flycatchers also frequently nest near larger, aggressive species (Marchant 1983, Tremont 1994). Breeding behaviour of the Restless Flycatcher was also similar to that of the confamilial , which is sexually monomorphic, socially monogamous, and biparental with equal shares in nest-based duties (Goodey & Lill 1993). The almost complete plumage monomorphism, equal biparentalism and high nest-failure rate of the Restless Flycatcher suggest that males of this species are unlikely to be free of parental or defensive duties sufficiently for extra-pair matings (see Tremont & Ford 2000 for related discussion on the Leaden Flycatcher). The high level of biparental investment needed for fledging success, to counter the high nest-predation rate, may leave little opportunity for extra-pair paternity. Males may also strongly defend or guard females, and repel intruding males. It is likely, therefore, that mated pairs of Restless Flycatchers are biologically as well as socially monogamous, although this suggestion requires testing. AUSTRALIAN 28 DEBUS & LOLLBACK FIELD ORNITHOLOGY

In some circumstances monogamy may be temporary, with females opportunistically changing mates for a better territory or likelihood of offspring, as happened with the original A female in this study. Whether long-term monogamy is the norm in this species also requires further study.

Acknowledgements This study was largely an undergraduate project by GL, supervised by Hugh Ford. We thank Shannon Teale for sharing the nest-watches, Jim Palmer and Kihoko Tokue for assistance with mist-netting, Andrew Ley for the loan of equipment, and Brian Chaffey for banding the new A male. The research was conducted under University of New England Ethics permit AEC 2000/0081, with authorisation from the NSW National Parks & Wildlife Service and the Australian Bird & Bat Banding Scheme. We also thank Hugh Ford, Steve Tremont, Geoff Price and the Handbook team at Birds Australia for comments on a draft, and Andrew Ley for editing.

References Boles, W.E. (1988), The Robins and Flycatchers ofAustralia, Angus & Robertson, Sydney. Buckingham, R. & Jackson, L. (1992), A Field Guide to Australian Birdsong, Cassette 8, Little Shrike·thrush to Hall's Babbler, Birds Observers Club of Australia, Melbourne. Cogger, H.G., Ford, H.A., Johnson, C.N., Holman, J. & Butler, D. (2003), Impacts of Land Clearing on Australian Wildlife in Queensland, World Wide Fund for Nature Australia, Brisbane. Ford, H.A., Barrett, G.W., Saunders, D.A. & Recher, H.F. (2001), 'Why have birds in the woodlands of southern Australia declined?', Biological Conservation 97, 71- 88. Ford, H.A., Bridges, L. & Noske, S. (1985), 'Density of birds in eucalypt woodland near Armidale, north-eastern New South Wales', Corella 9, 78-107. Ford, H.A., Noske, S. & Bridges, L. (1986), 'Foraging of birds in eucalypt woodland in north­ eastern New South Wales', Emu 86, 168- 179. Goodey, W. & Lill, A. (1993), 'Parental care by the Willie Wagtail in northern Victoria', Emu 93, 180-187. Heatwole, H. & Simpson, R.D. (1986), 'Faunal survey of New England. I. Introduction and general description of the area', Memoirs of the Queensland Museum 22, 107-113. Heatwole, H., de Bavay, J. & Webber, P. (2003), 'Faunal survey of New England. V The lizards and snakes', Memoirs of the Queensland Museum 49, 299- 325. Heatwole, H., de Bavay, J., Webber, P. & Webb, G. (1995), 'Faunal survey of New England. IV The frogs', Memoirs of the Queensland Museum 38, 229- 249. Marchant, S. (1983), 'Suggested nesting association between Leaden Flycatchers and Noisy Friarbirds', Emu 83, 119-122. NPWS (2002), Imbota Nature Reserve Draft Plan of Management, National Parks & Wildlife Service, Sydney. Roberts, N.L. (1942), 'Breeding activities of three common species', Emu 41, 185- 194. Schodde, R. & Tidemann, S.C. (Eds) (1986), Reader's Digest Complete Book ofAustra lian Birds, 2nd edn, Reader's Digest Services, Sydney. Tremont, S.M. (1994), Breeding Biology of the Leaden Flycatcher (Myiagra rubecula ), BSc Hons Thesis, University of New England, Armidale, NSW. Tremont, S.M. & Ford, H.A. (2000), 'Partitioning of parental care in the Leaden Flycatcher', Emu 100, 1- ll.

Received 14 April 2004 •