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UPLAND BIRD COMMUNITIES ON SANTO, VANUATU, SOUTHWEST PACIFIC PDF

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4 The Wilson Journal ofOrnithology 118(3):295—308, 2006 UPLAND BIRD COMMUNITIES ON SANTO, VANUATU, SOUTHWEST PACIFIC ANDREW W. KRATTER,1 JEREMY J. KIRCHMAN,23 AND DAVID W. STEADMAN1 — ABSTRACT. We surveyed indigenous landbirds at two upland, mostly forested sites in southwestern Santo, Vanuatu. One site (Wunarohaehare, 600-1,2502 m elevation) lies on the western, rain-shadowed slope of Mt. Tabwemasana. The other (Tsaraepae, 500-700 melevation) is 16 km to the south, on the southeastern, very wet slope ofPeakSanto. These are the richest single-sitebird communities yet surveyedin Vanuatu, with 30species ofresident birds recorded at each site, 27 ofwhich were common to both sites, including 6 species endemic to Vanuatu. Wejudged that 12 ofthe shared species were common atboth sites. The non-overlappingspecieswere a megapode, a parrot, and four understory passerines. We present new data on vocalizations for four species endemic to Vanuatu (Ptilinopus tannensis, Todiramphusfarquhari, Neolalage banksiana) or to Vanuatu plus New Caledonia (Clytorhynchus pachycephaloides). We found less seasonality in breeding than previously re- ported for Vanuatu. Most human impact at the sites today may be from non-native mammals (rats, cats, pigs, cows), along with low levels of hunting and forest clearing. Based on prehistoric bones from elsewhere in Vanuatu, we suspectthat formerly the sites on Santo may have supported additional speciesofmegapode, hawk, parrot, and starling. Received28July 2005, accepted 14 March 2006. The Republic ofVanuatu (12,195 km Fig. of surveys conducted decades ago (e.g., San- ; 1) consists of 12 islands >270 km2 and nearly derson et al. 1998, Gotelli and Entsminger 100 smaller ones in the tropical Pacific Ocean. 2001), little recent attention has been paid to Approximately 190,000 persons inhabit 70 is- gathering new data on intra- and inter-island lands (Lai and Fortune 2000) that range from variation in Vanuatu’s bird communities (al- active volcanoes to limestone islands to older, though see Bowen 1997). Bregulla (1992) geologically composite islands, such as Santo summarized information on identification, (MacFarlane et al. 1988, Nunn 1994). Anal- life-history, and distribution for each species yses of avian distributions in Vanuatu, based recorded from the island group, yet made it largely on collections made during the Whit- clear that much remains to be learned about ney South Sea Expedition on 31 islands in the basic biology ofVanuatu’s birds. Although 1926 and 1927 (e.g., Mayr 1934, 1941), have most biogeographic analyses ofinsular faunas been important in the development of evolu- (or floras) are based on lists of species from tionary theory (Mayr 1963) and the fields of an entire island, such lists typically contain island biogeography (MacArthur and Wilson species that seldom, if ever, interact because 1967) and community ecology (Diamond they are not syntopic. Especially on large is- 1975). Aside from the study by Scott (1946), lands such as Santo, the sets of species found field ornithology in Vanuatu lagged until the at single sites provide fertile grounds for anal- Percy Sladen expedition of 1971 focused on ysis. inter-island and altitudinal patterns of avian In 2002 and 2003, we made two trips to distribution across six islands in the archipel- Santo, Vanuatu’s largest (3,900 km2 andhigh- ago (Medway and Marshall 1975). Despite the est (1,879 m) island, home to eight)ofthe nine continued interest by ecologists in the results bird species endemic to the archipelago (Bre- gulla 1992). We surveyed birds at two mid- 1 FloridaMuseum ofNatural History, Univ. ofFlor- elevation rainforest sites, one each on the ida, P.O. Box 117800, Gainesville, FL 32611, USA. southeastern (windward) and western (lee- 2Dept, ofZoology,Univ. ofFlorida,Gainesville,FL ward) slopes of Santo’s rugged west-coast 32611, USA. mountain range. Our surveys were based on 30233C,urCruelntturaadldrEedsusc:atNieown CYeonrtker,StaAtlebaMnuys,eNumY,1R2o23o0m, sight/sound records, mist netting, tape-record- USA. ings, and specimens collected: skins with 4Corresponding author; e-mail: wings spread, skeletons, tissues, stomach con- [email protected] tents, and ectoparasites from the same indi- 295 296 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 118, No. 3, September2006 0 5 10 15 20 25km FIG. 1. Map of Espiritu Santo, Vanuatu, with an inset of Melanesia. Islands and island groups mentioned in the text are named. Sites of bird surveys conducted from 2002-2003 by the authors are indicated by the triangle (Wunarohaehare) and the square (Tsaraepae), and filled circles indicate sites surveyed by Bowen (1997; Loru Protected Area) and Medway and Marshall (1975; Nokovula, Apuna River, Hog Harbour). Asterisks = mountain peaks >1,400 m; dashed line = 600-m contour. METHODS vidual, along with data on habitat, molt, diet, and reproductive condition. Such information The island of Espiritu Santo (generally is a first step in the investigation ofecological, called Santo; Fig. 1) probably originated in morphological, and genetic differences among the Oligocene (ca. 25-30 mya) through vol- populations, and it is important for conserva- canism and tectonic uplift, although most of tion efforts that often focus on endemic taxa. its land formed during or since the Miocene In this paper, we present the results of our through these same processes (Mallick 1975, surveys at each site, focusing on Vanuatu’s Collot and Fisher 1989). Much of the island’s endemic and poorly known species. We also eastern half is flat or has rolling hills, with present comparisons with previous surveys at most land <300 m in elevation and very little sites elsewhere on Santo and in the Solomon of it above 600 m. The western half of Santo Islands. is dominated by a north-south trending moun- — Knitteret al. • SANTO, VANUATU BIRD COMMUNITIES 297 TABLE 1. Study sites and mist-netting effort on Santo, Vanuatu, 2002-2003. Site(latitude,longitude) Majorhabitats Nettingdates Elevation(m) No.nets Net-hr Wusi village (15° 22.7' S, Dry lowland forest, 22-27 Oct 2002, 0-50 8 165 166° 39.7' E) secondary scrub 4-5 Nov 2002 Wunarohaehare1* (15° 20.5' S, Humid premontane forest, 29 Oct-2 Nov 600-1200 18 337 166° 40.5' E) forest patches, grassy 2002 ridge Kerevalissy village (15° 35.7' S, Secondary lowland forest 3-6 and 14 Jun 200 5 14 166° 50.0' E) patches 2003 Tsaraepae3 (15° 32.7' S, Wet, primary, premontane 7-14 Jun 2003 500-700 15 575 166° 48.4' E) forest aPrimarystudysites. tain range that reaches its greatest height at much wetter than sites in the rain shadow Mt. Tabwemasana (1,879 m). Prevailing Wusi and Wunarohaehare. From 3 to 7 June, winds push moist air off the Pacific Ocean we surveyed a patchy secondary forest near across the eastern lowlands and into the east- Kerevalissy village (Fig. 1, Table 1), a land- or southeast-facing slopes of the main cordi- scape dominated by coconut plantations, —4 llera. Thus, the eastern and southern slopes of km north of the coastal village of Ipayato. the cordillera are humid with high precipita- From 7 to 14 June, we mist-netted (Table 1) tion, whereas the western slopes, which and observed birds on the southern slopes of plunge into the Pacific with little development Peak Santo at Tsaraepae (—500 m; denoted by ofa coastal plain, lie in a rain shadow and are the square in Fig. 1) and on nearby slopes up m relatively dry. to 700 elevation. Ridges in the lower ele- From 22 October to 5 November 2002, we vations had a broken canopy and were cleared (AWK, JJK) mist-netted and observed birds in of undergrowth, grazed by feral cattle (Bos d(Friyg.fo1r,esTtabalned1)s,crauvbililnagteheinvtihceinirtayinosfhWaudsoiw ftaa)u.rTush)e, aarnedab>r7o0w0semd bwyasfemraailnpliygstal(lSu(scasncorpoy- on the western coast 10 km west of Mt. Tab- 12-25 m) forest. wemasana, and in humid premontane forests Trees identified (by DWS) to genus includ- and grassy ridges from 600 to 1,250 m ele- ed Garuga (Burseraceae), Calophyllum (Clu- vation on the northern slope of Mt. Wunaro- siaceae), Elaeocarpus (Elaeocarpaceae), Her- haehare (denoted by a triangle in Fig. 1; Table nandia (Hemandiaceae), Ficus (Moraceae; at 1). At Wunarohaehare, figs (Ficus spp.) and least five species, some of them emergent), nutmegs Myristica spp.) are the dominant and Myristica (Myristicaceae); those we iden- ( sfrouniitainsgppt.re)esb.eTcroemeefceronmsm(oCynatahbeoavesp7p.0,0Dmicki-n t(iBfairerditnogtsopneicaiceesaei)ncalnuddeEdnBdaorsrpinegrtmounmiameedduulli-s a transitional habitat between the “high-stat- losum (Euphorbiaceae; often emergent). There ure lowland rain forest” and the “montane also were a number of unknown species, in- cloud forest” (described in Mueller-Dombois cluding various Myrtaceae and Rubiaceae. and Fosberg 1998). The weather at Wunaro- Also present were Pandanus spp. (Pandana- haehare is cool and moist in the morning, as ceae), tree-ferns (Cyathea spp.; Cyatheaceae), cloud cover descends below 600 m. By 10:00 and Dicksonia spp. (Dicksoniaceae). The edg- UTC + 11, however, the clouds dissipate and es included trees and shrubs of Macaranga the canopy receives direct sunlight. Short pe- spp. (Euphorbiaceae), lnocarpus fagifer (Fa- riods (<1 hr) of rain occur most afternoons. baceae), Ficus spp., Piper spp. (Piperaceae), From 3 to 14 June 2003, AWK, JJK, and Alphitonia spp. (Rhamnaceae), Pipturus spp. DWS worked on the southern slopes of Peak (Urticaceae), palms (Cocos spp.; Metroxylon Santo (also called Lairiri; 1,704 m), —16 km spp. [Arecaceae]), and thickets ofHibiscus til- south-southeast of Mt. Tabwemasana. This iaceus (Malvaceae), bananas (Musaceae), and area received the full precipitative effects of gingers (Zingiberaceae). moist air coming off the Pacific, and was The weather at Kerevalissy and Tsaraepae — 298 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 118, No. 3, September2006 in 2003 was extremely wet, with heavy rain- found on Santo. Nonetheless, cryptic species fall occurring on 11 ofour 12 days. On 9 days may have been missed if they were not vocal we estimated that the daily rainfall exceeded during our visits. 100 mm, including 6 days (5, 6, 8, 9, 12, and RESULTS 13 June) on which it probably exceeded 150 — mm. The excessive rain was due to an unusu- Diversity andcommunity composition. We ally late tropical storm that paused just north recorded 33 indigenous species oflandbirds at of Santo over the Banks and Torres islands. Wunarohaehare and Tsaraepae, with 27 spe- Because avian activity did not diminish no- cies common to both sites (Table 2). As is the ticeably during rains at Tsaraepae, we con- case across most of Oceania (Steadman 1997, ducted our sight/sound surveys and set mist 2006b), pigeons and doves (Columbidae) nets even during the very rainy weather. Vo- composed a large part of the avifauna; the calizations were tape-recorded on several days same seven species of columbids were found at each ofour two primary sites (Table 1), and at each site. We also recorded seven of the the original tapes were deposited in the Flor- eight species endemic to Vanuatu, failing to ida Museum of Natural History (UF) Sound record only Aplonis santovestris (see below). Archives. Birds were collected according to Six of the endemic species (all but Megapo- the stipulations of our permits from the Va- dius layardi) were recorded at both sites. nuatu Ministry of Lands, Environment Unit. Although three species of non-native birds Specimens were prepared as various combi- are widespread on Santo (Red Junglefowl, nations of round skins, complete or partial Gallus gallus; Common Myna, Acridotheres skeletons, and with spread wings. Stomach tristis; and Black-headed Munia, Lonchura contents and two tissue samples were taken malacca), the only one we recorded was G. from each specimen; one tissue sample is gallus, and it was uncommon (<5/day) atboth housed at UF and the other at the Louisiana sites. All three species were common in plan- State University Museum of Natural Science. tations and villages at elevations lower than All non-tissue material is housed at UF. As far those of Wunarohaehare and Tsaraepae. Con- as we know, neither tissue nor skeletal spec- tamination of the bird communities by non- imens of birds had been collected previously native species on Santo is minor (by Pacific in Vanuatu. The skeletal specimens ofthe Va- Island standards); however, both sites are nuatu endemics Ducula bakeri Ptilinopus heavily infested with non-native mammals. At , tannensis, Todiramphusfarquhari, Neolalage Tsaraepae, we noted feral cats (Felis catus), banksianci, Zosterops flavifrons, and Glycifo- pigs (Sus scrofa), and cows (Bos taurus); dogs hia notabilis (see Tables 2 and 3 for English (Canis familiaris) seemed to be confined to common names) are the first in the world’s villages. Inside our leaf house at Tsaraepae, DWS inventories. snap-trapped 10 rats (7 Rattus rattus, 3 In addition toAoWurKwork at the two primary R. exulans) in 3 nights, using only two traps. sites, JJK and collected and surveyed Although species richness was the same at birds in patchy forested sites near sea level on our two primary sites, composition of the the eastern coast of Santo for 2 days in Oc- landbird communities differed slightly. Me- tober—November 2002 and for 4 days in June gapodius layardi Charmosyna palmarum AWK , , 2003. In northern Santo, visited lowland and Clytorhynchus pachycephaloides were forests of the Vatte Conservation Area (near found only at Tsaraepae, although the latter Matantas; Fig. 1) from 17 to 19 November species was found in the dry forests nearWusi DWS 2002. visited Aore Island (Fig. 1) on (lower elevations than at Wunarohaehare). 15-16 June 2003, surveying (sight/sound The mound-building Megapodius layardi may only) birds in patches of tall (canopy 15—30 be absent from dry forests due to unsuitable m) lowland rainforest. soil conditions. Our failure to record Char- Although this was the first visit to Santo by mosyna palmarum at Wunarohaehare may all three authors. AWK and especially DWS have been aconsequence ofits nomadic habits have wide experience with the avifauna in (see C. palmarum species account, below). western Oceania. They know the vocalizations Three species with widespread distributions in and behaviors of all but one of the genera Oceania Lalage leucopyga Turdus polioce- , Kratteret al. • SANTO, VANUATU BIRD COMMUNITIES 299 — phalus, and Petroica multicolor were not re- (Table 4). We suspect, nevertheless, that the corded at Tsaraepae. The four passerine spe- difference between the two sites (87% versus cies found at only one of the two sites have 60% of species) does reflect seasonal trends been recorded on both sides of the cordillera more than inter-site variation. (Medway and Marshall 1975; Table 3), so their apparent absence at one site may be re- Selected Species Accounts lated to inadequate sampling. We note, how- We present ourfindings forspecies endemic ever, that our guides at Tsaraepae did not rec- to Vanuatu and for some others that are poorly ognize the illustration in Bregulla (1992) of known in Vanuatu or th—roughout their range. Turdus poliocephalus, suggesting that the lo- Megapodius layardi. The endemic Vanu- cal absence of this conspicuous species was atu Megapode was not recorded at Wunaro- genuine. The guides did not distinguish be- haehare, but, at Tsaraepae on 1 1 and 12 June, tween L. maculosa and L. leucopyga (Hakei three individuals were heard calling at an el- m language names forLalage were “vasoimoto” evation of 550 in the thick undergrowth and “losoloso,” which seemed to apply to ei- near an active incubation mound in a large ther species), so it is possible that the latter tract of forest. This was the only mound near species was present. Our guides did know Pe- Tsaraepae known to our guides. Another bird troica multicolor, however, and called it “pa- was observedmin a dense Hibiscus tiliaceus nopano.” thicket at 600 on 11 June. Single birds also We observed inter-site differences in the al- were seen twice in secondary forest patches titudinal ranges ofsome species. Two endemic on Santo’s eastern coast, and once near Ma- species characteristic ofthe highlands Ducula tantas. Villagers showed us eggs from an ac- ( bakeri and Glycifohia notabilis) were more tive mound near Matev—ulu on 16 June. common at Tsaraepae than at Wunarohaehare, Chalcophaps indica. This terrestrial dove where D. bakeri was not seen below 800 m. is widespread in Oceania, with the subspecies At Tsaraepae, D. bakeri was found regularly C. i. sandwichensis confined to New Caledon- m as low as 500 and locally in forest patches ia, the Santa Cruz Group, and Vanuatu. Abun- m as low as 200 along the trail south toward dant in disturbed forest and forest edge from m the coast. At Tsaraepae, the fantail, Rhipidura sea level to 400 elevation (lower than either spilodera, was scarce above 500 m, but at study site), the Emerald Dove was much less Wunarohaehare it was common up to 800 m. common in more mature forest near our two Some species associated with less forested primary study sites. In 337 net-hr at Wuna- habitats (Todiramphus chloris, Lalage macu- rohaehare, only one bird was netted at eleva- hliosgahe,rGeelreyvgatoinoensflaatvWoluantaerroalhiase)hawreer,ewfhoeurnedwaet tnieotn-shr>5at000-m5,0wmherneeaasrfWiuvesiw.erBeecneatutseediint s1e6l5- sampled open habitats up to 1,000+ m; atTsa- dom vocalizes and is rather furtive, mist-net- raepae, however, we did not find these species ting may yield better evidence ofthe Emerald at elevations above 550 m, which were almost Dove’s population density than auditory or vi- entirely forested. — sual data. The species is common in village Seasonality of reproduction. Our visit to gardens, where it often is lured with papaya Wunarohaehare during October-November {Carica papaya) into traps; stomachs of near- coincided with the reported breeding period ly all collected individuals contained seeds of for most species of birds in Vanuatu, which this non-native plant. The four birds taken generally is September-February (Bregulla near Wusi village included two males with en- 1992). Our visit to Tsaraepae took place dur- larged testes, an adult male (no bursa) with ing June, a month when Bregulla (1992) unenlarged testes, and an adult female (no found breeding activity for only 5 of the 33 bursa, convoluted oviduct) with slightly en- species we recorded (Table 2). We found less larged ova. The single bird from Tsaraepae evidence of marked seasonality in breeding, was a male with enlarg—ed testes. with signs of reproductive activity (enlarged Ptilinopus tannensis. Endemic to Vanua- gonads in specimens, active nests, or recently tu, the Tanna Fruit Dove was common (up to fledgedjuveniles) in 20 of23 species at Wun- 15 per day) at each site, especially in montane arohaehare and 12 of 20 species at Tsaraepae forests. This fruit dove was heard much more I 300 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 118, No. 3, September2006 TABLE 2. Summary of native bird communities at two sites (Wunarohaehare, 600-1,200 m; Tsaraepae, 500-700 m) on Santo, Vanuatu, surveyed in 2002-2003. E = endemic to Vanuatu, e = endemic to Vanuatu plus New Caledonia and/or the Santa Cruz Group. Relative abundance: c = common (encountered regularly by all observers), u = unc—ommon (encountered daily or almost daily in small numbers), r = rare (encountered fewer than five times), = not recorded. Foraging guild (microhabitat/prey): A = aerial, C = canopy, T = terrestrial, U = understory, F = fruit, G = granivore (seeds), I = insects and other invertebrates, N = nectar, V = vertebrates. Avian nomenclature follows Dickinson (2003), except that we do not recognize Aerodramus, which has been used for some species in Collocalia (but see Price et al. 2004). Relativeabundance Foraging Species Wunarohaehare Tsaraepae guild Megapodiidae Megapodius layardi, Vanuatu Megapode (E) T/F,G,I Accipitridae Circus approximans. Swamp Harrier A/V Columbidae Columba vitiensis leopoldi White-throated Pigeon T,U,C/F,G , Macropygia m. mackinlayi, Mackinlay’s Cuckoo-Dove U/F Chalcophaps indica sandwichensis, Emerald Dove T/G,I,F Ptilinopus tannensis, Tanna Fruit Dove (E) C/F Ptilinopus greyii. Red-bellied Fruit Dove (e) U,C/F Duculap. pacifica. Pacific Imperial Pigeon C/F Ducula bakeri, Vanuatu Imperial Pigeon (E) C/F Psittacidae Trichoglossus haematodus massena. Rainbow Lorikeet C/N,F Charmosynapalmarum Palm Lorikeet (e) C/N , Cuculidae Chrysococcyx lucidus layardi. Shining Bronze-Cuckoo C/I? Apodidae Collocalia esculenta uropygialis, Glossy Swiftlet A/I Collocalia v. vanikorensis, Uniform Swiftlet A/I Alcedinidae Todiramphusfarquhari. Chestnut-bellied Kingfisher (E) U/I,V Todiramphus chloris santoensis, Collared Kingfisher C/I,V Meliphagidae Glycifohia n. notabilis. White-bellied Honeyeater (E) C/N,I Myzomela cardinalis tenuis. Cardinal Honeyeater C/N, Acanthizidae Gerygoneflavolateralis correiae. Fan-tailed Gerygone U,C/I Artamidae Artamus leucorhynchus tenuis. White-breasted Woodswallow A/I Campephagidae Coracina caledonica thilenii, Melanesian Cuckoo-shrike U,C/F,I Lalage maculosa modesta, Polynesian Triller U,C/F,I Lalage leucopyga albiloris. Long-tailed Triller U/F,I Pachycephalidae Pachycephala [pectoralis] caledonica intacta. New Caledonian U/I Whistler (e) Petroicidae — Petroica multicolorambrynensis. Pacific Robin U,C/F,I Rhipiduridae Rhipidura fuliginosa] albiscapa brenchleyi. Gray Fantail U/I [ Rhipidura s. spilodera. Streaked Fantail T,U/I Kratter et al. • SANTO, VANUATU BIRD COMMUNITIES 301 TABLE 2. Continued. Relativeabundance Foraging Species Wunarohaehare Tsaraepae guild Monarchidae Neolalage banksiana, Buff-bellied Monarch (E) —C C U/I Clytorhynchuspachycephaloides grisescens Southern Shrikebill (e) U U/I , Myiagra caledonica marinae, Melanesian Flycatcher C C U,C/I Zosteropidae Zosteropsflavifrons brevicauda, Yellow-fronted White-eye (E) C C U,C/N,F,I Zosterops lateralis tropicus. Silver-eye c C U,C/N,F,I Turdidae — Turduspoliocephalus vanikorensis. Island Thrush c T,U/F,I — often than seen, although it called less fre- Ducula bakeri. The monotypic Vanuatu quently than the Red-bellied Fruit Dove. Con- Imperial Pigeon is endemic to seven islands trary to Medway and Marshall (1975) and in northern Vanuatu. Although rare or absent Bowen (1997), we found the Tanna Fruit in the lowlands of Santo, it was common at Dove above 500 m; it remained common up Tsaraepae, where two or three calling individ- to the highest continuous forests that we uals often were audible from many points on reached at both Wunarohaehare (800 m) and a forested ridge at —600 m, and we recorded Tsaraepae (700 m). The most common call as many as 20 on single days. It was less com- was a series (~10+) of low, upwardly inflect- mon on the disturbed slopes below 500 m, ing woot notes, spaced up to 2 sec apart. In- although we heard it in a forest patch adjacent frequently, it also gave a soft, single woot to Kerevalissy on 14 June. At Wunarohaehare, note. we found the Vanutau Imperial Pigeon only at We found the Tanna Fruit Dove breeding at elevations >800 m, where up to three indi- both sites. Bregulla (1992) reported its nesting viduals called in heavy forest cover on most status as poorly known, with previous evi- days. The birds taken at Tsaraepae were an dence reported only in April and May, a time adult female with enlarged ova and ajuvenile of little breeding activity among other land- male. They differed little in plumage, and both birds in Vanuatu. At Wunarohaehare, a nearly had Myristica spp. fruits in their crops and fledged nestling was found on the ground after stomachs. — a windy evening, and two males had enlarged Charmosyna palmarum The monotypic . testes and a female had enlarged ova. At Tsa- Palm Lorikeet is endemic to Vanuatu and the raepae, the one bird collected was a female Santa Cruz Group. We recorded this species with enlarged ova. — only twice (a flock of six on 8 June, a group Ptilinopus greyii. The monotypic Red- oftwo on 1 1 June), both times in a Ficus spp. bellied Fruit Dove is confined to New Cale- tree with large, fleshy fruits, in humid forest donia, the Loyalty Islands, and Vanuatu. The at 650 m on the main ridge at Tsaraepae. Al- species was abundant (<50/day) at both sites though more characteristic of montane than in heavily disturbed to mature forests and at lowland habitats, the Palm Lorikeet seems to all elevations. It vocalized throughout the day. undergo population fluctuations and has a pro- All specimens showed evidence of breeding: pensity to wander (Medway and Marshall at Wunarohaehare, these included a female 1975, Bregulla 1992). Its preferred foods with a ruptured follicle, another with enlarged (flowers and fruits) may have been scarce at ova, a male with enlarged testes, and a re- the time of our visits. cently fledgedjuvenile; atTsaraepae, the spec- Collocalia e—sculenta uropygialis and C. v. imens included two males with enlarged tes- vanikorensis. Each of these widespread tes, a female with enlarged ova, and two ju- swiftlets was common at Tsaraepae. The veniles. Glossy Swiftlet (C. esculenta uropygialis 20- ; 302 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 118, No. 3, September2006 TABLE 3. Indigenous birdsrecorded (+ = present, - = notrecorded) atsix sites on Santo, Vanuatu, 2002- 2003. English common names are provided for the species not included in Table 2. E = endemic to Vanuatu, e = endemic to Vanuatu plus New Caledonia and/or the Santa Cruz Group. Sources are Bowen (1997) for Loru Protected Area; Medway and Marshall (1975) for Apuna River, Hog Harbor, and Nokovula; and our own data for Wunarohaehare and Tsaraepae. For each site, the elevation (m) is included. LoruProtected ApunaRiver HogHarbor Wunarohaehare Tsaraepae Nokovula Species Area0-120m 100m 160m 600-1,250m 500-700m 1,120m Megapodius layardi (E) + + - - + - Falcoperegrinus. Peregrine Falcon + - - - + Circus approximans + + - + + + Galliraliusphilippensis Banded Rail + - - - - - Columba vitiensis , + + - + + - Macropygia mackinlayi + + + + + + Chalcophaps indica + + + + + + Ptilinopus tannensis (E) + + - + + — Ptilinopus greyii (e) + + + + + + Duculapacifica + + + + + - Ducula bakeri (E) - - - + + + Trichoglossus haematodus + + + + + - Charmosynapalmarum (e) - - - - + + Chrysococcyx lucidus - - - + + - Tyto alba. Barn Owl + - - - - - Collocalia esculenta uropygialis + + - + + + Collocalia v. vanikorensis - + - + + - Todiramphusfarquhari (E) + + + + + - Todiramphus chloris + - - + + - Glycifohia n. notabilis (E) - - - + + + Myzomela cardinalis - + + + + + Gerygoneflavolateralis - + + + + + Artamus leucorhynchus + — — + + + Coracina caledonica + + + + + + Lalage maculosa - - - + + - Lalage leucopyga - - - + - + PPeatcrhoyicceaphmualltaic[opleocrtoarmalbirsy]nceanlseidsonica (e) -+ -+ -+ ++ -+ ++ Rhipidura [fuliginosa] albiscapa + - - + + - Rhipidura spilodera + + + + + + Neolalage banksiana (E) + + + + + + Clytorhynchuspachycephaloides grise- scens (e) + + + - + — Myiagra caledonica + + + + + Cichlornis whitneyi, Melanesian Thick- - - - - - + etbird Zosteropsflavifrons (E) + + + + + + Zosterops lateralis + + + + Alponis zelandica. Rufous-winged Star- - - - - — + ling (e) Turduspoliocephalus vanikorensis + + + + Erythrura cyaneovirens. Red-headed Parrotfinch - - - - - + Total species 25 22 16 30 30 24 Total endemic species (E + e) 8 8 6 8 11 8 50/day) generally flew much closer to the noted at all sites visited on Santo. Despite our ground than the Uniform Swiftlet (C. v. van- careful observations of all swiftlets detected ikorensis; <20/day, except for loose flocks of on Santo, we did not record the White-rumped —400 that passed over on several mornings at Swiflet (Collocalia spodiopygia), which was Tsarapae, all flying west). Both species were unknown to our guides. H' Kratteret al. • SANTO, VANUATU BIRD COMMUNITIES 303 and no Wusi Group.reproductiveN=I + i + + + i I Z + 1 Z 1 + z z y+ + b|+ i z Cruz of + + + + + ZZZ + + + + + + + + + + + + + + + around juveniles; 1 1 1 Santa the condition elevations fledged and/or 0 On laotw Caledona ossification, recently I I orr*n-« 11 11 11 11 11 11 11 N<ON 11 11 11 o^CM 11 11 -—~cf NO | \I O°n). 0—r0t or CO CM CO skull cd 00 d collected pNleuwsodegfreeacnteisvtes, g*-sa <<cnNm <Orr~N-- c0n0 1 ^oTj- 1 1 1 t NO 1 1 ^>n 11 C-CMtM1 11 r- 11 00 | | dcm"0in Specimens Vanuatu tract, o no Fabricius, J20u0n3e. entdemoic oburfsa reproductive MmCM. OC0-'M0.8M— Co3M„ ON>nOn- I2CONM O^ OiCrMni„COCoOnOo. cCcN'oMdtO" 'OtN„• 0^0 OcCCcMMd)d„in d 11qdN——OI iinn O0n (r(T-NNnH Ot1qd—-!H CicC—MMnd„N0iTtnO and = of of (N NO I I co it co CM 00 11 CM CM 06 r- —i CM 1 't e 2002 presencecondition Vanuatu, to byof O 6~ tOn oo basis October-November o o endemic determined on no odn the Tt ft oo -t 't = on 00 T+ on co co in co o E cj2 £ £ ^ 2 status Vanuato, included. determined C0M0 ON—O ccOmm oNoO. ii—nn1 Octno o—o1 t°j°-. cfot NimOncOmNh-— (ooNcMo_OOnO. 0O0.Ot^-r)io(Noo^-(N ccod ncm Juvenile ) Santo, are — or sites names. (+ from study data common evidence •3 5Sa 3 montane specimen English .Ift w t3o 3f3t A4TvAiB.aLnE avKeirnlelvadalgisessy2a3fSTaonbelrdees Baprlteunremadadcigtne,g.information. 32.§f2t8:£S2f13t ^3»USSQ3gf32St.W3a3aSOo3C,3a'l3a35<SSS3.303i. -Q<Isfff3S3ttt. -^-Qfa*s3aSt fftt <f3t o -3t<3f3o3-t.3ftf«3tot J&_3ci 3-.•S3ff^f§29S2ttt **~§CsC£<ocs«*53i33 §iaP^„If.t i3.f2§fftItt;S.1l3.f<fftt3t.I.lw"1^^^eS>oJGsNfbfJtt -SrS0f“s8tSi3t'0ts"^63a|^N?-63••Ib^f5ff^f2cS02Ntttti..,"3^-§>f£2SCfC332t2ti-- 304 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 118, No. 3, September2006 — Todiramphusfarquhari. Endemic to San- from Tsaraepae also was molting, probably its to, Malo, and Malakula, the Chestnut-bellied first pre-basic molt. — Kingfisher was slightly more common in the Petroica multicolor ambrynensis. The wet forests near Tsaraepae (<5/day) than in subspecies ofPacific Robin from Santo, P. m. the dry forests of the western slope, although ambrynensis is one of 5 subspecies from Va- , we recorded up to six daily at Wunarohaehare. nuatu and 14 across Oceania. In the Solomons It was most common in high-canopy forests, and New Guinea, the Pacific Robin is restrict- but also persisted in forest patches, even near ed to montane forests. Although apparentlyre- Kerevalissy village. It ranged from the low- stricted to high-elevation forests (>500 m) on lands up to at least 800 m, overlapping the Santo, the Pacific Robin may be found at low- entire elevational range of its larger congener, er elevations elsewhere in Vanuatu. JJK found the Collared Kingfisher (T. chloris), which it to be common in lowland forests on the prefers more open habitat. The Chestnut-bel- rain-shadowed Dillon’s Bay area of western lied Kingfisher was very vocal at both sites, Erromango. On Efate, however, DWS found often singing throughout the day. The call is it in humid, mid-elevation forest (—350 m). In a series of ascending notes with decreasing addition to not finding the Pacific Robin at intervals, not the “monotonous single note” Tsaraepae (although our guides there knew of described by Bowen (1997). The two birds this species), no one has recorded it from any collected at Wunarohaehare, both at 600 m, lowland location on the wet (eastern) side of were adult males, one in non-reproductive Santo. Medway and Marshall (1975) recorded condition (testes 3 X 1.5 mm) and the other it at an elevation of 1,100 m on the eastern with somewhat enlarged testes (6X4 mm). flank of Mt. Tabwemasana, but we recorded Evidence ofreproductive activity at Tsaraepae robins (up to four daily) only in forest from included ajuvenile male (probably in first pre- 650 to 800 m near Wunarohaehare. The three basic molt, with heavy wing molt and mod- specimens were two adult males with enlarged erate body molt), and two adult females with testes and seminal vesicles, and an adult fe- convoluted oviducts but unenlarged ova. male that probably had nested recently (ova Stomachs contained the remains of large bee- not enlarged, but oviduct somewhat thickened tles (including Cerambycidae), large orthop- and convoluted). — terans, spiders, skinks, and—geckos. Neolalage banksiana. The Buff-bellied Glycifohia notabilis. The monotypic Monarch belongs to a monotypic genus en- White-bellied Honeyeater is endemic to Santo demic to Vanuatu. It occurs on most major and Malakula. With Dickinson’s (2003) place- islands south to Efate and was common at ment of this species in the genus Glycifohia both ofour primary study sites, with daily re- (previously cla—ssified as Phylidonyris), its cords of up to 25 at Wunarohaehare and 12 at only cong—ener the Barred Honeyeater (G. Tsaraepae. It was found most often in pairs or undulata) is endemic to New Caledonia. family groups in the undergrowth of forest Previously, both had been placed in the wide- patches or large tracts of forest, especially spread Australian genus, Phylidonyris. The where vine tangles or thickets ofHibiscus til- White-bellied Honeyeater occurred in similar iaceus dominate the understory, although abundance between 600 and 800 m at both some birds were found in forests with an open sites, usually in large tracts of forest. Often, understory. these birds congregated at flowering trees in The song ofthe Buff-bellied Monarch is ap- noisy groups of <15 individuals. Of four parently undescribed; Bregulla (1992) stated specimens (two from each site), only one was AthaWt,K“. . . it is said to have melodious song.” reproductively active, a male from Wunaro- tape-recorded a bird singing in scrubby haehare with enlarged testes. The other bird dry forest adjacent to Wusi village on the from this site, an adult female (no bursa; skull morning of 25 October. The song had a stut- 100% ossified), had minute ova, a straightovi- tering,jumbled beginning, then three rapid se- duct (probably had not yet bred), and its ries of reedy, high-pitched, whistled notes. wings, tail, and body were molting. An adult The first and last series consisted of three de- female from Tsaraepae had these same char- scending notes, whereas the second series acteristics. A young male (bursa 2X2 mm) consisted of only two descending notes: tee-

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