92 Notornis, 2018, Vol. 65: 92‒108 0029-4470 © The Ornithological Society of New Zealand Inc.

Evidence for year-round breeding of in the , in the context of the Australasian and South Pacific regions

MICHAEL K. TARBURTON School of Science & Technology, Pacific Adventist University, PMB, Boroko, Papua New Guinea Current address: 3 Freda Court, Blackburn South, Victoria,

Abstract This paper summarises the variations in breeding periodicity in the Australasian - South Pacific region, includ- ing recent evidence contrary to previous knowledge. Birds shown to breed in the Samoan islands throughout the year are white-tailed tropicbird (Phäethon lepturus), white-rumped swiftlet (Aerodramus spodiopygius), buff-banded rail Galli( - rallus philippensis), brown noddy ( stolidus), white (Gygis alba) and wattled honeyeater Foulchaio( carunculata). In addition, crimson-crowned fruit-dove (Ptilinopus porphyraceus), blue noddy (Procelsterna caerulea), cardinal honeyeater (Myzomela cardinalis) and Polynesian triller (Lalage maculosa) breed in at least 9 months of the year. The Samoan whistler ( flavifrons) has been found breeding in 8 months and the Samoan starling (Aplornis atrifusca) in 7 months of the year. Tarburton, M.K. 2018. Evidence for year-round breeding of birds in the Samoan Islands, in the context of the Australasian and South Pacific regions.Notornis 65 (2): 92–108. Keywords breeding; Samoan islands

INTRODUCTION The characteristic breeding cycles for some Reproduction in birds, as in all , takes place groups of birds have been known for some time; at the time of the year when it is biologically most this is particularly the case for in which advantageous for them to do so (Serventy & Warham their colonial nature facilitates the collection of 1971). Breeding periodicity allows birds and other data regarding breeding seasonality. For instance, organisms to synchronize breeding attempts with it has long been known that albatrosses nesting the most favourable ecological conditions under in temperate locations breed every second year which to raise their offspring. In most of the world, even though the breeding site will have half the ecological conditions vary seasonally and so impose population breeding in it each year (Jouventin & an annual breeding schedule on their populations Dobson 2002; Ratcliffeet al. 2008). (del Hoyo et al. 1992). In the tropics however, Regional variations have also been reported seasons are not as distinct and annual breeding in seabirds. Serventy & Warham (1971) note that cycles may be longer than in temperate regions almost all species of seabirds in eastern and south- (Ricklefs 1966) or may not develop for a population eastern Australia restrict their breeding to the spring or for individuals (Pyle et al. 2016). and summer; whereas in north-western Australia, the predominant period of egg-laying is in the Received 16 June 2017; accepted 19 January 2018 autumn and winter, and in south-western Australia Correspondence: [email protected] spring and early summer egg-laying is the rule. In Australasia & South Pacific breeding seasons 93 the intervening region, from the Lacepede Islands plants, invertebrates and vertebrates have greater in the north to Shark Bay, there are both autumn- possibilities of reproducing all year (Ricklefs 1966; breeding and spring-breeding populations on the Weimerskirch 2007). To prevent overpopulation same islands. there can be mechanisms to restrict that breeding The incidence of tropical cyclones and strong period, to restrict the number of progeny or to winter winds, as well as upwelling ocean currents, reduce the number of breeders. each appear to have an effect on the seasonality Watling (2001) uses information from the of egg-laying in seabirds. Silver gulls (Larus Whitney Expedition and his own observations to novaehollandiae) in Western Australia are double suggest that there is a distinct seasonal pattern in brooded with an interval of about 4 months both breeding and moult for many south Pacific between layings (Nicholls 1964). When silver gulls species. He points out that this seasonality is are taken from south-eastern Australia where they found in the majority of land birds in and , are single brooded and held in captivity in south- but less clearly in . Because the Samoan western Australia, they become double brooded in climate is almost aseasonal, it is tempting to agree synch with the local gulls (Nicholls 1964). In New with Dhondt (1976), that breeding is not restricted Zealand, the nitrogen-rich upwellings off Kaikoura to 1 part of the year. Dhondt’s assertion was based facilitate and control the timing and success of the on his finding that buff-banded rails (Gallirallus breeding of red-billed gulls (Larus novaehollandiae) philipensis) and Polynesian trillers (Lalage maculosa) and Hutton’s shearwaters Puffinus( huttoni)(Sherley were breeding in 6 to 8 months of the year. He 1992; Mills et al. 2008). During the breeding season, only visited a white-rumped swiftlet (Aerodramus the relative availability of New Zealand krill spodiopygius) colony once, but combining this with (Nyctiphanes australis) was correlated positively Ashmole’s data (1963), he determined breeding in with the Southern Oscillation Index and the April, June and November, albeit in different years. frequency of occurrence of north-east winds which Ashmole (1963) was able to make the generalisation induce upwelling. Incursions of warm oceanic that it is obvious that breeding is not restricted to 1 water, often induced by westerly winds which are part of the year. She found banded rails breeding characteristic of El Niño years, were associated in March and August and white-rumped swiftlets with reduced euphausiid availability and reduced breeding in February, April, June and November breeding success (Mills et al. 2008). Banks (1984) noted that even though 1,300 Less is known about the breeding seasons of specimens have been taken from , tropical seabirds; but what is known often shows the annual cycle remains incompletely known sub-annual breeding periodicity, rather than even for those few species with large samples of annual or biennial periodicity. Tropical seabirds specimens. Largely this is due to the limited timing known to breed every 6 months include swift of collecting. Probably about the same number of (Sterna bergii) on Aldabra Atoll (Diamond & specimens have been taken from Samoa (formerly Prys-Jones 1986) and white terns (Gygis alba) on Western Samoa) with similarly minimal information Christmas Island (Pacific) (Ashmole 1968). Those about the breeding seasons of the birds that breed breeding every 7 to 8 months include white-tailed there. tropicbirds (Phaeton lepturus) on Ascencion Island In 1985, the U.S. Fish & Wildlife Service (Stonehouse 1962) and Aldabra Atoll (Prys-Jones & organised a large survey of forest birds in American Peet 1980), and bridled terns (Sterna anaethetus) on Samoa, but at the end of that they still claimed “little Aldabra Atoll (Diamond & Prys-Jones 1986). Those is known about life histories of forest birds” (Engbring breeding every 9 to 10 months include the Christmas & Ramsey 1989). shearwater (Puffinus nativitatis) on Pitcairn Islands It has been shown that on Savaii, the largest (Brooke 1995), and Audubon’s shearwater (Puffinus island in the Samoan Archipelago, that the purple lherminieri) and swallow-tailed gulls (Creagrus swamphen (Porphyrio porphyrio), flat-billed furcatus) on the Galapagos Islands (Snow 1965; kingfisher Todiramphus( recurvirostris) and Samoan Snow & Snow 1967). In Fiji, it has been shown that flycatcher (Myiagra albiventris) do not inhabit while the colony of black noddies (Anous minutus) that island above 1,200 m (Pratt & Mittermeier on Vatu-i-Ra is occupied by birds throughout the 2016). What factor(s) restrict these birds? They year (Tarburton 1978), breeding takes place at 9 to also conclude that the Samoan woodhen (Galinula 10-month intervals, and individual birds breed each pacifica), a flightless rail, has been extinct for second cycle (Tarburton 1987). over 100 years and that recent “sightings” are Small islands in the tropical South Pacific have mis-identifications. Steadman (2006) estimated less seasonal climate variation than most other that between 500‒1,600 species of flightless rails places on earth (Steadman 2006). This means that inhabited Pacific islands prior to the first human along with the reliable year-round rainfall (except colonization. However, not a single species survives on a small part of large islands such as Savaii) on inhabited islands outside of New Zealand (Pratt 94 Tarburton

Fig. 1. Location of Samoan islands.

& Mittermeier 2016). It is too late to find out if they Non-breeding indicators from specimens could or did breed all through the year. or captured birds are not recorded because it is Since I was to live on Upolu in Samoa for 4 too uncertain as to whether the individual was years it was decided to start filling in some of the immature or in post-breeding condition. From blanks in the breeding cycles of Samoan birds. looking at skins in all Australian and New Zealand Accordingly, this paper collates information from Museums as well as live birds and the literature, I literature sources and field observations to provide see there is also considerable variation in observer an updated account of bird breeding activity on the ability with regard to judging the age and seasonal Samoan islands. status of skins – particularly in species new to most observers. It is also clear that in those species now known to breed all year that birds not in breeding MATERIALS AND METHODS condition do not preclude others from being in Breeding data from the literature were compiled breeding condition. and added to my own field observations, made Names and sequence of birds are taken between January 1994 and December 1997, while from Christidis & Boles (2008) or if not included locating and visiting 38 caves on the Samoan islands there, from the IOC Checklist (http://www. of Savai’i, Upolu and Tutuila, as well as doing over worldbirdnames.org/ioc-lists/master-list-2/). 400 1-hour censuses of movements around the entire coast of Upolu. To make headway on the bush birds, 35 trips RESULTS were made up the walking track from Vailima, past The location of the 8 main islands mentioned in the Robert Louis Stevenson memorial to the lookout this paper are shown in Fig. 1. The actual months on the west face of Mt Vaea, specifically to observe of breeding for each species for which data was evidence for breeding. Mt Vaea is close to the capital gained is shown in Table 1. Full details regarding of Samoa, Apia, on the island of Upolu. These walks the types of data used as evidence for breeding and were made between 1 July 1995 and 19 April 1997. the source of these observations are provided in One to 7 walks were made each month. Thus, while Appendix 1. not evenly spread over the study period, the walks Birds shown to breed in the Samoan Islands did add many data points to this paper. throughout the year (Table 1) are white-tailed Key indicators used to determine breeding tropicbird, white-rumped swiftlet, buff-banded rail, included anatomical features (i.e. enlarged testes or brown noddy, white tern and wattled honeyeater ovaries), nests with contents (i.e. eggs, nestlings or (Foulchaio carunculata). In addition, crimson- brooding adults), breeding behaviours (i.e. seabirds crowned fruit-dove (Ptilinopus porphyraceus), blue landing in forests, courtship behaviour, copulation) noddy (Procelsterna caerulea), cardinal honeyeater or the presence of recently fledged birds. (Myzomela cardinalis) and Polynesian triller (Lalage Australasia & South Pacific breeding seasons 95

Table 1. Months in which breeding has been observed in Samoan birds. * = historical observation, # = data from this study.

Common Name Scientific Name J F M A M J J A S O N D Red-tailed tropicbird Phaethon rubricauda * * * * * * * White-tailed tropicbird Phaethon lepturus * * # * * * * # # * * * Crimson-crowned fruit- Ptilinopus porphyraceus * * * * * * * * * * dove Many-coloured fruit-dove Ptilinopus perousii * * * * * * * Pacific imperial pigeon Ducula pacifica * * * * * Tooth-billed pigeon Didunculus strigirostris * White-rumped swiftlet Aerodramus spodiopygius * * # * # * # * * # * * Tahiti petrel Pseudobulwaria rostrata * * * * Herald petrel Pterodroma heraldica * * * * Audubon’s shearwater Puffinus lherminieri * * * * * Christmas shearwater Puffinus nativitatis * * Great frigatebird Fregata minor * * * Lesser frigatebird Fregata ariel * * * Masked booby Sula dactylatra * * * * * * Brown booby Sula leucogaster * * * * * * * * Red-footed booby Sula sula * * * * * Eastern reef egret Egretta sacra * * * Purple swamphen Porphyrio porphyrio * * * * * Buff-banded rail Gallirallus philippensis * * * * * * * * * * * * Spotless crake Porzana tabuensis * Brown noddy Anous stolidus * * # * * * * * # * * * Black noddy Anous minutus * * * * * * * * White tern Gygis alba * * # * # # * * # * * * Blue noddy Procelsterna cerulea * * * * * * * * * Spectacled tern Onychoprion lunatus * * Bridled tern Onychoprion anaethetus # # # Sooty tern Onychoprion fuscatus * * * Little tern Sternula albifrons * Black-naped tern Sterna sumatrana # # Blue-crowned lorikeet Vini australis * * * Eastern barn owl Tyto javanica # * * Flat-billed kingfisher Todirhamphus recurvirostris * Collared kingfisher Todirhamphus chloris * * * * * * * * Wattled honeyeater Foulehaio carunculatus * * # * # * * # # * * * Cardinal myzomela Myzomela cardinalis * * * * * # # * * # Mao Gymnomyza samoensis * Polynesian triller Lalage maculosa * * * * # * * * * # * Samoan triller Lalage sharpei * # * # Samoan whistler Pachycephala flavifrons # # # # # # # # Samoan fantail Rhipidura nebulosa # * # # Samoan flycatcher Myiagra albiventris * (broadbill) 96 Tarburton

Table 1 cont. Months in which breeding has been observed in Samoan birds. * = historical observation. # = data from this study.

Common Name Scientific Name J F M A M J J A S O N D Fiji shrikebill Clytorhynchus vitiensis * * Pacific robin Petroica pusilla * * # * * # Samoan white-eye Zosterops samoensis * Red-vented bulbul Pycnonotus cafer * # # # # # * # Island thrush Turdus poliocephalus * * * Polynesian starling Aplonis tabuensis * * * * # Samoan starling Aplonis atrifusca * * * * * * * Jungle myna Acridotheres fuscus # Common myna Acridotheres tristis * * # # # Red-headed parrot-finch Erythrura cyaneovirens # # #

maculosa) breed in at least 9 months of the year. recorded (Sekora 1974 & 1976 in Amerson et al. The Samoan whistler (Pachycephala flavifrons) has 1982). This is further supported by the recording of been found breeding in 8 months and the Samoan breeding birds in May 1974 (Amerson et al. 1982). Starling (Aplornis atrifusca) in 7 months of the year. A sub-annual breeding cycle could also explain the breeding dates given for the brown noddy in American Samoa by Amerson et al. (1982), whereas DISCUSSION they interpret their data as probably signalling Muse & Muse (1982) predicted that the white-tailed breeding throughout the year. Separate colonies tropicbird, the buff-banded rail, the Pacific robin may also not be synchronised with each other so we (Petroica multicolour) and cardinal honeyeater will need to be aware of the dangers of mixing data from be found to breed throughout the year. This paper different sea-bird colonies. adds credence to each of these suggestions, though Bridled terns, breeding on Penguin Island in only the first 2 are fully confirmed. The same Western Australia breed out of the tropics then authors also claimed that the Polynesian triller migrate to “winter” in the tropics in the Celebes bred throughout the year but gave no evidence for Sea (Dunlop & Johnstone 1994). However, Samoan the statement. This study found Polynesian trillers bridled terns stay in the tropics to breed on the 2 breeding in 2 of the remaining 3 months – so are stacks off O-Le-Pupu Pue National Park (Tarburton now almost certain to be found breeding in all 12 2001). months of the year. This study has also shown that 4 The concept of individual birds not participating unpredicted species nest in each month of the year: in breeding even though the colony might be white-rumped swiftlet, brown noddy, white tern breeding is also worth consideration. Tarburton and wattled honeyeater. (2009) has shown that white-rumped swiftlets With seabirds, there is the possibility in Polynesia breed throughout the year (contra Whitmee 1875) at of having a breeding period of less than 12 months. most colonies in Samoa, yet individual pairs in that This non-annual breeding regime has been found colony usually breed only once a year. If food levels in black noddies in Fiji where the colony has an 8 are continually reasonably high but insufficient for to 10-month breeding cycle (Tarburton1987), with all to breed all the time, then pairs can share them individuals breeding every 20 months. This means throughout the year. This is in contrast to 1 pair of the colony has eggs every 10 months and can look buff-banded rails on Upolu producing 5 broods in deserted diurnally at the time they had eggs the 1 year (Robinson 1995). Even though the swiftlets previous year. At sunset however, the noddies that in Samoa breed all year, there is an obvious peak in are not on migration to the Solomon Islands will September. mass offshore and come ashore to roost each night Amerson et al. (1982) suggest many-coloured of the year. A similar regime is possible for sooty fruit-doves (Ptilinopus perousii) breed in the terns (Onychoprion fuscatus) on Rose Island where southern winter, but from this paper it is now clear eggs were observed in October 1975, whereas in that they breed in all 4 seasons. The same authors October the following year 4,300 fledglings were also suggest that the collared kingfisher “probably Australasia & South Pacific breeding seasons 97 breeds in the southern summer”. This paper shows Del Hoyo, J.; Elliott, A.; Sargatal, J. (eds.) (1992). Handbook that to be true – but this kingfisher also breeds up to of the birds of the world, Volume 1. Barcelona, Lynx at least June – the Austral winter. Edicions. Turning from seabirds to landbirds, Pyle et Dhondt, A. 1976. Bird observations in Western Samoa. al. (2015) found in American Samoan birds that Notornis 23: 29‒43. while there was broad overlap between moult and Diamond, A.W.; Prys-Jones, R.P. 1986. The biology of breeding at the population level, there was very terns nesting on Aldabra Atoll, Indian Ocean, with little at the individual level. They suggest that particular reference to breeding seasonality. Journal of while moult may take precedence over breeding, Zoology, London 210: 527‒548. that tropical landbirds appear to be capable of Dunlop, J.N.; Johnstone, R.E. 1994. The migration of suspending moult to breed when environmental bridled terns Sterna anaethetus breeding in Western conditions shift to favour successful reproduction. Australia. Corella 18: 125‒129. Much further study is needed to understand why Dunmire, W.W. 1960. Some 1960 bird observations in so many Samoan bird species are breeding all year Samoa and Fiji. Elepaio 20: 76‒78. round. DuPont, J.E. 1972. Notes from Western Samoa, including I would be very wary of claiming too much for the description of a new parrot-finch (Erythrura). the family of crested terns I observed in Vaiusu Wilson Bulletin 84: 375‒376. Bay 27 July 1997. The 2 adults were accompanied Engbring, J.; Ramsey, F.L. 1989. A 1986 survey of the forest by a juvenile with 75% of the body covered with birds of American Samoa. Honolulu, U.S.D.I. Fish & brown flecking. The juvenile concerned was very Wildlife Service. like illustration #4 on Plate 36 in Higgins & Davies Green, R.H. 1965. Western Samoan bird notes. Elepaio 26: (1996). This does not prove breeding in Samoa as 19‒21. banded birds in Australia have been found up to 575 Higgins. P.J.; Davies, S.J.J.F. (eds.) 1996. Handbook of km from their natal colony 19 days after fledging, Australian, New Zealand and Antarctic birds: Volume 3, and over 1,000 km away within 7 weeks of fledging snipe to pigeons. 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Source Morrell & Aquilani 2000 Amerson Amerson Amerson Amerson Armstrong 1932 Armstrong 1932 Yaldwyn 1952 Clapp & Sibley 1966 Clapp & Sibley 1966 Banks 1984 Banks 1984 Banks 1984 Dunmire 1960 Banks 1984 Dunmire 1960 Clapp 1968 This study This study This study This study This study This study This study This study Armstrong 1932 Location Rose Rose Swain’s Rose Rose Upolu Upolu Upolu Tutuila Tutuila Tutuila Tutuila Tutuila Tutuila Ofu Ta’ü Swains Upolu Tutuila Upolu Upolu Savai’i Upolu Upolu Upolu Upolu x x x D x x x N x x x x x x x O x x x x S x x x x x A J x x x x J x x x x x x x x x Month x x x x x x x x x M x x x x A x x x M x x x x F J x x x x x x Evidence* Nt NB Fl E Nl Nl NS Fla Fl OE OE Nl, TE Fl Fl Fl Fl Oe FLa FLa PC SbL FLa NP Nl E E Specific evidences for breeding given for species recorded in this study and in literature, covering named Samoan islands. *Key to evidence: ASol = Species Red-tailed tropicbird Red-tailed tropicbird Red-tailed tropicbird Red-tailed tropicbird Red-tailed tropicbird White-tailed tropicbird White-tailed tropicbird White-tailed tropicbird White-tailed tropicbird White-tailed tropicbird White-tailed tropicbird White-tailed tropicbird White-tailed tropicbird White-tailed tropicbird White-tailed tropicbird White-tailed tropicbird White-tailed tropicbird White-tailed tropicbird White-tailed tropicbird White-tailed tropicbird White-tailed tropicbird White-tailed tropicbird White-tailed tropicbird White-tailed tropicbird White-tailed tropicbird Crimson-crowned fruit-dove Appendix 1. Appendix AFF = adult adult soliciting; flying with food; BP = brood patch; CNM = carrying nest material; Cop = copulation; E = egg; ES = egg shells; EOvi = egg in oviduct; F = - not coral island major over flying FLa = seabird seen; Fl = fledgling(s) fledgling; = feeding FFl in bill; fish with = flying FF begging; fledgling = FB departing; fledgling adult = NB nest; tending parents = NA reported; “nesting” = N juvenile; = Juv burrow; in heard = Hd enlarged; not gonads = GnE enlarged; gonads = GE diurnal; bar, PC enlarged; ovary = OE nest; = Nt site; one to trips repeated = NS prospecting; nest = NP nestling; = Nl behaviour; defence nest = NDf building; nest = NBl brooding; = pair courting; R runner; RB runner, in black down; SbL seabird landing forest/tree; TE testes enlarged. 100 Tarburton 1982 Banks1982; 1984 1982 1982 1982 1979 et al. et al. et al. et al. et al. et al. Source Armstrong 1932 Banks 1984 Banks 1984 Clapp & Sibley 1966 Ripley & Birckhead 1942 This paper Ashmole 1963 Amerson Clapp & Sibley 1966 Banks 1984 Banks 1984 Banks 1984 Amerson Whitmee 1875 Banks 1984 1876 Layard Ashmole 1963 Armstrong 1932 Whitmee 1875 Banks 1984 Banks 1984 Armstrong 1932 Armstrong 1932 Dhondt 1976 Ollier Amerson Engbring & Ramsey 1989 Bellingham & Davis 1988 Tarburton 2009 Amerson Amerson Engbring & Ramsey 1989 Location Upolu Tut uila, Manu’a Tut uila, Manu’a Tutuila “Samoa” Upolu Unspecified Tutuila Tutuila Tut uila, Manu’a Tut uila, Manu’a Tut uila, Manu’a Tutuila Upolu Tutuila Upolu Not given Upolu Upolu Ta’ü Ofu Upolu Upolu Upolu Upolu Tutuila Tutuila Upolu Upolu Ta’ü Ta’ü Ta’ü x x x x x x x D x x x x x x N x x x x O x x x x x x x S x x x x x x x A J x x x x x x x x J x x x x x x x Month x x x x x M x x x x x A x M x x x x x x F J x x x x Evidence* Nl Ge Juv TE Nl NB Nl TE TE NL GE TE TE, OE E OE TE R Juv Nl Nt Nt E Nl BP E, N, E E, Nl NB N E, Nl TE, PC SbL Fla,Hd, N Species Crimson-crowned fruit-dove Crimson-crowned fruit-dove Crimson-crowned fruit-dove Crimson-crowned fruit-dove Crimson-crowned fruit-dove Crimson-crowned fruit-dove Crimson-crowned fruit-dove Crimson-crowned fruit-dove Many-coloured fruit-dove Many-coloured fruit-dove Many-coloured fruit-dove Many-coloured fruit-dove Many-coloured fruit-dove Pacific pigeon Pacific pigeon Pacific pigeon Tooth-billed pigeon Tooth-billed pigeon White-rumped swiftlet White-rumped swiftlet White-rumped swiftlet White-rumped swiftlet White-rumped swiftlet White-rumped swiftlet White-rumped swiftlet White-rumped swiftlet White-rumped swiftlet White-rumped swiftlet White-rumped swiftlet Tahiti petrel Tahiti petrel Tahiti petrel Appendix 1. cont. Australasia & South Pacific breeding seasons 101 1982 1982 1982 1982 1982 1982 1982 1982 1982 1982 1982 1982 1982 1990 1990 1990 et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. et al. Source Engbring & Ramsey 1989 Engbring & Ramsey 1989 Pyle Pyle Pyle Amerson Banks 1984 Amerson Engbring & Ramsey 1989 Engbring & Ramsey 1989 Amerson Amerson Amerson Amerson Amerson Banks 1984 Amerson Amerson Banks 1984 Banks 1984 Banks 1984 Child 1979 Amerson Amerson Amerson Engbring & Ramsey 1989 Engbring & Ramsey 1989 Armstrong 1932 Banks 1984 Amerson Location Tutuila Olosega Ta’ü Ta’ü Ta’ü Ta’ü Ta’ü Ta’ü Ta’ü Tutuila Ta’ü Rose Rose Rose Rose Rose Rose Rose Nu’utele, Ofu Rose Ofu Fanuatapu Olosega Ofu(Nu’utele) Rose Tutuila Olosega Ofu(Nu’utele) Rose Tutuila Olosega Ofu(Nu’utele) Rose Tutuila Tutuila Olosega Nu’utele? Upolu, Off E Rose Rose x x x x D x x N x x x x x x x x x O x x S x x x x x A J x x x x x x x x J x x x x x x x x Month x x x x x x M x A x x M F J x x Evidence* NB Fla, N Fla, N Fla, Nl OE SbL Nt SbL Nl FLa, N SbL Nt Nl E Nt E, Nl E NL TE, OE E, TE TE Nl Nl E Nl Nl Nl Nl, E E E, NlNt, Species Tahiti petrel Tahiti petrel Herald petrel Herald petrel Herald petrel Herald petrel shearwater Audubon’s shearwater Audubon’s shearwater Audubon’s shearwater Audubon’s Christmas shearwater Great frigatebird Great frigatebird Lesser frigatebird Lesser frigatebird booby Masked booby Masked booby Masked Brown booby Brown booby Brown booby Brown booby Brown booby Brown booby Brown booby Brown booby Brown booby booby Red-footed booby Red-footed booby Red-footed Appendix 1. cont. 102 Tarburton 1982 1982 pers. comm. et al. et al. Source Amerson Banks 1984 Amerson Banks 1984 Banks 1984 Banks 1984 Banks 1984 Banks 1984 Banks 1984 Armstrong 1932 Armstrong 1932 Clapp & Sibley 1966 Banks 1984 Banks 1984 Green 1965 Dhondt 1976 Dhondt 1976 Dhondt 1976 Dhondt 1976 Dhondt 1976 Robinson 1995 Reed 1980 This study This study This study This study Banks 1984 Steve Brown Clapp & Sibley 1966 Clapp & Sibley 1966 Dhondt 1976 Clapp 1968 Clapp 1968

Location Tutuila, Poa (off Tutuila) (off Tutuila, Poa Rose Rose Ofu Ofu Tutuila Tutuila Ofu,Tutuila, Tau, Ofu,Tutuila, Tau, Upolu Upolu Tutuila Tutuila Tutuila Savai’i Upolu Upolu Upolu Upolu Upolu Upolu Savai’i Upolu Upolu Upolu Upolu Ta’ü Nu’utele, Upolu Tutuila Tutuila Tutuila Swain’s Swain’s x x x x x x x x x D x x x x x x N x x x x x O x x x x x x S x x x x A J x x x x x x J x x x x x Month x M x x x x x A x x x x x x M x x x x x F J x x x x x Evidence* Nt E E, Nt TE TE OE E GE N, E RB Juv OE OE, TE R R R, RB ES E Juv Cop RB R RB TE E R R, GE N Nt, NDf N NB Nl OE, BP

Species Red-footed booby Red-footed Eastern reef egret Eastern reef egret Eastern reef egret Purple swamphen Purple swamphen Purple swamphen Purple swamphen Purple swamphen Buff-banded rail Buff-banded rail Buff-banded rail Buff-banded rail Buff-banded rail Buff-banded rail Buff-banded rail Buff-banded rail Buff-banded rail Buff-banded rail Buff-banded rail Buff-banded rail Buff-banded rail Buff-banded rail Buff-banded rail Buff-banded rail Buff-banded rail crake Spotless Brown noddy Brown noddy Brown noddy Brown noddy Brown noddy Brown noddy Appendix 1. cont. Australasia & South Pacific breeding seasons 103 1982 1982 1982 1982 1982 1982 1982 1982 1982 et al. et al. et al. et al. et al. et al. et al. et al. et al. Source Clapp 1968 Clapp 1968 Clapp 1968 Amerson Amerson Amerson Amerson Amerson This study This study This study This study This study This study This study This study This study Clapp 1968 Clapp 1968 Clapp 1968 Banks 1984 Banks 1984 Amerson Amerson Amerson Amerson Engbring & Ramsey 1989 Clapp 1968 Clapp 1968 Clapp 1968 Banks 1984 Banks 1984 Banks 1984

Location Swain’s Swain’s Swain’s Tutuila Ofu Ta’ü Rose Swain’s Upolu Tutuila Upolu Upolu Savai’i Upolu Upolu Upolu Upolu Swain’s Swain’s Swain’s Swain’s Rose Tutuila Ta’ü Swains Rose Ta’ü Swain’s Swain’s Swain’s Tutuila Rose, Ta’ü Rose x x x x x x x D x x x x x x N x x x x x x O x x x x x S x x x x x A J x x x x x x J x x x x x x Month x x x x x x M x x x x x x x A x x x x x M x x x x x x F J x x x x Evidence* EOvi Nl, N BP, GE Nl Nl Nl N Nl FLa FLa SbL CNM FLa NB E NL PC NB NB NB, NL NB NB N N N N, E N E, BP BP NS NB GE, NB E, Nt

Species Brown noddy Brown noddy Brown noddy Brown noddy Brown noddy Brown noddy Brown noddy Brown noddy Brown noddy Brown noddy Brown noddy Brown noddy Brown noddy Brown noddy Brown noddy Brown noddy Brown noddy Black noddy Black noddy Black noddy Black noddy Black noddy Black noddy Black noddy Black noddy Black noddy Black noddy White tern White tern White tern White tern White tern White tern Appendix 1. cont. 104 Tarburton 1982 1982 1982 1982 1982 1982 1982 1982 1982 et al. et al. et al. et al. et al. et al. et al. et al. et al. Source Child 1979 Dhondt 1976 Amerson Dunmire 1960 Dunmire 1960 This study This study This study This study This study This study This study This study This study This study This study Clapp & Sibley 1966 Banks 1984 Banks 1984 Dhondt 1976 Amerson Amerson Amerson Amerson Amerson Amerson McAllan & Hobcroft 2005 Amerson This study This study Banks 1984 Amerson

Location Upolu, Upolu, Rose Tutuila Ta’ü Upolu Upolu Upolu Upolu Tutuila Tutuila Upolu Savai’i Upolu Upolu Savai’i Tutuila Tutuila Tutuila, Aunu’u, Ofu Tutuila Tutuila Aunu’u Ofu (Nu’utele) Ta’ü Olosega Tutuila Nu’uloa & Nu’utele Sand, Rose Upolu Upolu Rose Rose x x x x D x x x x x x N x x x x x x x x x x O x x x x x S x x x x x x A J x x x x x J x x x x x x x Month x x x x x x M x x x x x x A x x x x x M x x x x F J x x x x x x x Evidence* AFF, NBl AFF, FF N, E SbL PC FLa FF SbL PC FLa FF Nl FLa NB E PC Nl NB GE NB N Nt N N N N N Fl E, NB Juv OE E Species White tern White tern White tern White tern White tern White tern White tern White tern White tern White tern White tern White tern White tern White tern White tern White tern noddy Blue noddy Blue noddy Blue noddy Blue noddy Blue noddy Blue noddy Blue noddy Blue noddy Blue noddy Blue noddy Blue Spectacled tern Bridled tern Bridled tern Sooty tern Sooty tern Appendix 1. cont. Australasia & South Pacific breeding seasons 105 1982 1982 1982 et al. et al. et al. Source Amerson This study This study This study Banks 1984 Banks 1984 Banks 1984 Banks 1984 Armstrong 1932 This study Armstrong 1932 Clapp & Sibley 1966 Banks 1984 Banks 1984 Banks 1984 Ammerson Armstrong 1932 Banks 1984 Dhondt 1976 Dhondt 1976 Dhondt 1976 Dhondt 1976 Engbring & Ramsey 1989 This study This Study This study Banks 1984 Banks 1984 Dhont 1976 Dhont 1976 Amerson Engbring & Ramsey 1989 Location Upolu Tut uila, Manu’a Rose Upolu Upolu Upolu Tutuila Tutuila Ofu Ta’ü Upolu Upolu Upolu Tutuila Olosenga Tut uila, Manu’a Tut uila, Manu’a Tutuila Upolu Upolu Upolu Upolu Ofu? Upolu Upolu Upolu Tutuila Tutuila Upolu Upolu Tutuila Tutuila x x x x x D x x x x x x N x x x x x x O x S x x A J x x x x x J x x x x x x x x x Month x x x M x x x A x x x x x x M x x x x x x F J x x x x x x Evidence* Fl GE Fl Juv FFl F, Juv GE (4) GnE 64) GnE OE NS F F Nl Nl GE, Nl Nl Nl NBl CNM BP FFl FB FB, FFl NDf FB TE Fl BP Juv GE FB Species honeyeaterWattled honeyeaterWattled Sooty tern Little tern tern Black-naped tern Black-naped Blue-crowned lorikeet Blue-crowned lorikeet Blue-crowned lorikeet Eastern barn owl Eastern barn owl Eastern barn owl kingfisher Flat-billed Collared kingfisher Collared kingfisher Collared kingfisher Collared kingfisher Collared kingfisher honeyeaterWattled honeyeaterWattled honeyeaterWattled honeyeaterWattled honeyeaterWattled honeyeaterWattled honeyeaterWattled honeyeaterWattled Cardinal myzomela Cardinal myzomela Cardinal myzomela Cardinal myzomela Cardinal myzomela Cardinal myzomela Appendix 1. cont. 106 Tarburton 1979 1979 1979 et al. et al. et al. Source This study This study This study This study This study Mayr 1932; BanksMayr 1932; 1984 Dhondt 1976 Dhondt 1976 Dhondt 1976 Dhondt 1976 Dhondt 1976 Ashmole 1963 Ollier Ollier This study This study This study This study This study This study This study Dhondt 1976 Ollier Reed 1980 This study This study This study Dhondt 1976 This study This study Reed 1980 Beck,Banks fide 1984 Location Upolu Upolu Upolu Upolu Upolu Tutuila Upolu Upolu Upolu Upolu Upolu Unspecified Upolu Upolu Upolu Upolu Upolu Upolu Upolu Upolu Upolu Upolu Upolu Savaii Upolu Upolu Upolu Upolu Upolu Upolu Savai’i Ta’u x x D x x N x x x x x x x x x O x x x x x S x x x x x x x x x x x x A J x x x x x x x J x x x x x Month x x x M x x A x x x x M x x x x F J x x Evidence* FB, FFl Nl Fl CNM, NS NS TE AFF Nl JB, FFl JB E Nl NB FFl FFl Fl FB Nl AFF NBl Nt, NS NBE, N, FFl E, N FB, FFl FFl, FB FB FFl FB FFl NBl Nl, E Species Cardinal myzomela Cardinal myzomela Cardinal myzomela Cardinal myzomela Cardinal myzomela Mao triller Polynesian triller Polynesian triller Polynesian triller Polynesian triller Polynesian triller Polynesian triller Polynesian triller Polynesian triller Polynesian triller Polynesian triller Polynesian triller Polynesian triller Polynesian triller Polynesian triller Polynesian Samoan triller Samoan triller Samoan triller Samoan triller Samoan whistler Samoan whistler fantail Samoan fantail Samoan fantail Samoan Samoan broadbill shrikebillFiji Appendix 1. cont. Australasia & South Pacific breeding seasons 107 1982 1982 1979 et al. et al. et al. Source Engbring & Ramsey 1989 Banks 1984 Armstrong 1932 Muse & Muse 1982 Muse & Muse 1982 Dhondt 1976 Ollier Reed 1980 This study This study Beck, Armstrong fide 1932 Goodman in 1977 Dhont 1969 This study This study This study This study This study Armstrong 1932 Armstrong 1932 Green 1965 Armstrong 1932 Clapp & Sibley 1966 Banks 1984 Child 1979 Reed 1980 This study Armstrong 1932 Clapp & Sibley 1966 Amerson Amerson DuPont 1972 Dhondt 1976 Dhondt 1976 Location Ta’u Ofu Upolu Upolu Upolu Upolu Upolu Savai’i Upolu Upolu Savai’i Upolu Upolu Upolu Upolu Tutuila Upolu Upolu Upolu Upolu Upolu Tutuila Tutuila Tamaligi,Upolu Savai’i Upolu Upolu Tutuila Tutuila, Olosega Tutuila, Olosega Savai’i Upolu Upolu x x D x x x x N x x x x O x x x S x x x x x x x A J x x x x x x J x x x x x Month x x M A x x M x x F J x x x x Evidence* E Nl E Fl E FFl Fl Nl FFl NBl GE N E Fl PC FFl FFl Fl TE E Fl OE TE, BP Nl BP FB, FFl FFl TE TE GE, Ny NBl AFF CNM Species shrikebillFiji Fiji shrikebillFiji Pacific robin Pacific robin Pacific robin Pacific robin Pacific robin Pacific robin Pacific robin Pacific robin Samoan white-eye bulbul Red-vented bulbul Red-vented bulbul Red-vented bulbul Red-vented bulbul Red-vented bulbul Red-vented Island thrush Island thrush Island thrush starling Polynesian starling Polynesian starling Polynesian starling Polynesian starling Polynesian starling Polynesian starling Samoan starling Samoan starling Samoan starling Samoan starling Samoan starling Samoan starling Samoan Appendix 1. cont. 108 Tarburton 1979 et al. Source Ollier Ollier Engbring & Ramsey 1989 This study This study This study This study This study Dhondt 1976 Dhondt 1976 This study This study This study This study This study This study Location Upolu Tutuila Upolu Upolu Upolu Upolu Upolu Upolu Upolu Upolu Upolu Upolu Upolu Upolu Savai’i x x x x D N x x x O x x x x S x x A J x x x x J x x Month M A x M x F J Evidence* NA AFF CNM NBl NS E Nl FFl Juv NDf Fl Nl E, NB FFl Nl Species starling Samoan Samoan starling Samoan starling Samoan starling Samoan starling Samoan Jungle myna Jungle myna Common myna Common myna Common myna Common myna Common myna Red-headed parrot-finch Red-headed parrot-finch Red-headed parrot-finch Appendix 1. cont.