The Auk 118(2):472-483, 2001

AVIFAUNA OF A LOWLAND FOREST SITE ON ISABEL,

ANDREW W. KRATTER?4 DAVID W. STEADMAN,• CATHERINE E. SMITH, 2 CHRISTOPHERE. FILARDI,2 AND HORACE P. WEBB3 •FloridaMuseum of NaturalHistory, P.O. Box 117800, Universityof Florida,Gainesville, Florida 32611, USA; 2BurkeMuseum, University of Washington,P.O. Box 353010, Seattle, Washington 98195, USA; and 32048White Oak Circle,Kennesaw, Georgia 30144, USA

ABSTRACT.--Weprovide the first comprehensivedescription of a community from a lowland rainforestsite on a major island in the SolomonIslands. During two dry season visits (July 1997,June 1998) to the lower GarangaRiver valley on the island of Isabel,we recorded65 residentand 6 migrant speciesof . We documentrelative abundances, hab- itat preferences,and foraging guilds for the membersof the bird community.The Garanga River site sustainsall but 11 of the 76 speciesof landbirds known from Isabel. Of those11 ,four are small-islandor beachspecialists, three are montane,and four are of un- known status.Habitat heterogeneity,maintained largely by river dynamics,is a major con- tributor to aviandiversity at the site.The avifaunais dominatedby nonpasserines,especially parrots,pigeons, , and hawks.The flightlessrail Nesoclopeuswoodfordi, previously regarded as rare and threatenedwith extinction,was common.We recordedIxobrychusfiav- icollis,Falco severus, and Eudynamysscolopacea for the first time on Isabel.We alsodocumented occurrencein the lowlands of Micropsittafinschii, Collocaliaspodiopygia, Coracina caledonica, and Pachycephalapectoralis, four speciespreviously thought to be confinedto upperelevations on Isabel. The depauperateunderstory avifauna of the Garanga River site may be anthro- pogenicand couldbelie what otherwiseseems to be an intact avifauna.Received 27 December 1999, accepted13 December2000.

THE SOLOMON ISLANDS have one of the In 1997 and 1998,we surveyedbirds at a low- world's most distinctive avifaunas, with more land forest site on Isabel, the fourth largestis- "restrictedrange species"(species with ranges land in the Solomons. Although Isabel is <50,000 km 2) than any other Endemic Bird known to sustain a rich avifauna (73 resident Area in the world (Stattersfield et al. 1998). Al- speciesare listed in Mayr 1945 and Webb 1992; though analysesof intriguing patterns of dis- currentisland total is 76 species),its birdshave tribution and endemism among Solomon Is- yet to receive an in-depth investigation.Four land landbirds have figured prominently in subspeciesare restrictedto the island(Nesoclo- island biogeographyand community ecology peuswoodfordi immaculatus, Ninox j. jacquinoti, (Greenslade 1968, Diamond et al. 1976, Dia- Pitta a. anerythra,and Coracinacaledonica welch- mond and Mayr 1976,Diamond 1982), we lack mani).Isabel sharesmore speciesand subspe- even one thorough descriptionof a bird com- cies of birds with Choiseul than any other is- munity from a single site within the Solomon Islands. In fact, descriptionsof avifaunasfrom land. Those two islands (as well as Buka, single sites are rare anywhere in out- Bougainville, and the Florida Islands) were side of .In contrast,single-site de- connectedduring late Pleistocenelow sea-level scriptionsof avifaunasare commonplacefor stands (Wickler and Spriggs 1988). For exam- Neotropicalbirds (e.g. Karr 1990, Terborghet ple, all but two of the 45 speciesof birds re- al. 1990, and various papers in Remsen1997). corded in lowland Bougainvilleby Kaestner Thesepapers have been instrumental in under- (1987) also occur on Isabel. standingthe ecologicalprocesses that promote In this paper, we describethe avifaunaof Is- and maintain avian diversity;they alsoprovide abel's lower Garanga River valley, including a foundationfor identifying areasof high con- data on endemism,species diversity, foraging servationpriority. guilds, and habitat associations.We also dis- cussimportance of the area for conservation. E-mail: [email protected] Our specimen-baseddata (mass,molt, age,and

472 April 2001] Birdsof Isabel, Solomon Islands 473

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'•1• ARASH,OWN INMAP 8 o •"• . Isabel •t• ß a,a,a NewGeorgia •. • • Group • • • •

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' r 154 ø 156 ø 158 ø 160 ø

0 4 8 12 16 km r• ELEVATIONOVER600 m

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FIG. 1. The northern coastof Isabel,showing localities mentioned in the text. reproductiveactivity) are reported elsewhere the coastalplain has been cleared for gardens,vil- (Kratter et al. in press). lages,and coconut(Cocos nucifera) plantations. From near the mouth of the Garanga River, a coconutplan- STUDY AREA AND METHODS tationextends southeast for 3 km alongthe coastand for -1 km inland. Much of this plantationis untend- Isabel(3,995 km 2) is a long (199km) narrow island, ed and thus many native trees are becomingestab- in most places20-25 km in width, centeredon 8øS lished beneath the coconut palms. Overgrown old and 159øE(Fig. 1). The island is geologicallycom- gardensare scatteredalong the lower GarangaRiver, plex, rises to 1,250 m elevation, and is incised by upstream to near our camp. Several limestone out- many river systemsthat empty onto a coastalplain crops below our camp, however, are coveredwith of variable width. The Garanga River (Fig. 1) begins mature forest. A -4 ha clearing was felled during at -1,100 m near the island's center and flows 13 km our visit in 1997 about 1.5 km downstream from our north into the Pacific.The coastal plain along the camp.In 1998,several small housesand gardenshad lower reachesof the Garanga,-5 km wide, is a mo- been started, althoughno one was living at the site saicof river terraces,gentle rolling hills, and isolated yet. At and aboveour camp,the forestsare more con- limestone outcrops.The river is about 30 m wide tinuousand mature.Trails, made by huntersto track here, alternatingbetween pools (to -4 m deep) and wild pigs (Susscrofa), extend to at least a few kilo- rocky-bottomed shallow rills. At low water, the meters aboveour camp. banks are -2 m tall. During our dry seasonvisits, Riveredge vegetation was dominated by earlysuc- sandy and gravelly beacheslined most of the river, cessionalspecies, including Hibiscustiliaceus, Cam- althoughwater levelsrose above the beachesduring prospermabrevipetiola, Macaranga sp., Pipturusargen- heavy rains. Behind the sandy beach at the river's teus,and Ficussp. (small, nonstrangling),as well as mouth is a small lagoon (-200 x 50 m). nonwoody Musa sp., Zingibersp., Dioscoreasp., and The natural vegetationon mostof Isabelis tall low- ferns. The more mature forest reached a canopy land forest, with shorter montane forestsat higher height of 25-30 m, with emergenttrees to 40 m. We elevations. Successionalvegetation occurs where were unable to assess the relative abundance of tree floodplain forestsare disturbed by flooding.Some of speciesin the species-richmature forest, although 474 KRATTERET ^L. [Auk, Vol. 118 these genera were conspicuous:Aglaia, Calophyllum, demic to the archipelago.All four subspecies Ficus,Myristica, and Pomefla.Large fruiting banyans endemic to Isabel were recorded at our site. No (Ficus spp.) were common and attracted multiple introduced specieswere recorded. speciesof frugivorousbirds. The SolomonIslands As on other islands in the Solomons (Gal- sustain63 speciesof Ficus--more than any Pacificis- braith and Galbraith 1962, Hadden 1981) the land group excluding New Guinea (Corner 1967). The beachfront forest at the mouth of the Garanga Garanga River avifaunaon Isabelis dominated River featured Barringtoniaasiafica, Calophyllum ino- by nonpasserines,whether in speciesrichness, phyllum,Heritiera littoralis,and Terminaliacatappa, generic-leveldiversity, or abundance(Table 1). with occasionalthickets of Hibiscustiliaceus along the Hawks, columbids,psittacids, and kingfishers river'sbanks. Absent in the area was Australianpine eachhave five or six residentspecies. Of the 42 (Casuarinaequisetifolia), which forms swampy thick- speciesof nonpasserines,19 were commonand ets in northern Isabel and the nearby island of San three--all parrots--were abundant. Morning Jorge(A.W.K., C.E.E, and C.E.S. pers. obs.). flight surveys were dominated by psittacids Our visits (9-23 July 1997 and 22-29 June 1998) were during the dry season.The weather was mild (especially Cacatuaducorpsii and Chalcopsitta to hot and humid, with partly cloudyto cloudyskies. cardinalis)and columbids(especially Ducula We recordedrain on sevendays in 1997and five days rubriceraand Ptilinopusviridis). Chalcopsitta car- in 1998,including a seriesof stormsfrom 15-18 July dinalisalso was abundantin the coconutplan- 1997. During most early mornings (from 0630-0730, tation closeto the river mouth, and many of the GMT + 12 h), at least one of us (C.E.E) conducted birds flying over our forestsite may havebeen dawn surveys of birds flying over the river near commutingfrom an inland roostto feed on the camp. All observationsand specimensare from prolific nectarproduced by coconutpalms. The within 1 km of the river, usually within 2 km but oc- tiny parrot Micropsittafinschii was abundantin casionallyas much as4 km upstreamor downstream the forest interior. accounted for from camp.Most birds were collectedwith shotguns by local hunters, especiallyMark Hafe, an Isabel is- only 23 (35%) of the resident speciesand 14 lander,who was thoroughlyfamiliar with the plum- (28%) of the residentgenera. The richestpas- ages,vocalizations, and behaviorof all specieson Is- serine families were cuckoo-shrikes(five spe- abel. Limited mist-nettingin interior forest (four 12 cies)and starlings(four species).Eleven pas- m, 36-mm mesh nets on the west side of the river in serine specieswere considered common and 1997,and five on the eastside in 1998)supplemented two were abundant. the collection via hunting. A single 12 m, 60-ram Habitataffinities.--Most resident species were mesh net also was placed acrossthe Garanga River found in a variety of habitats (Table 1). Forest -250 m upstreamfrom our campin 1997.Specimens interior habitats were preferred by slightly are housedat the Florida Museumof Natural History (UF), University of Washington Burke Museum more species(26) than river-edgehabitats (24 (UW), and Solomon Islands National Museum. Tis- species).Beaches and river edges were pre- suespreserved in lysisbuffer are storedat UW. Spec- ferredby 11 species.The remainingfive species imen tag data include sex, reproductivecondition, were noted only in the air (swiftlets, Falcosev- molt scoring, soft part colors, subdermal fat levels, erus)or were infrequently encountered(Podar- stomachcontents, skull ossification,presence of bur- gus ocellatus). sa, and habitat. Many of the most commonspecies in succes- sional river-edge and interior forest habitats RESULTS were canopy species.Among migrant species, shorebirdswere found along the river, primar- Overview.--Werecorded 71 speciesof birds ily on sand and gravel beaches,and both mi- alongthe lower GarangaRiver (Table1). All but grant landbirdsused river-edge vegetation, al- six of thesewere found in both years. Of the 71 though Chrysococcyxlucidus was found as well species,65 are residents, two (Halcyonsancta, in the interior forest. Chrysococcyxl. lucidus)are migrant landbirds Mixed speciesfiocks.--Mixed-species flocks from the south, and four (Pluvialisfulva, Nu- were encounteredmore frequently in forestin- meniusphaeopus, Tringa nebularia, and Actitishy- terior than in other habitats. Pachycephalapec- poleucos)are northern migrant shorebirds. toralis,Rhipidura ruJ•frons, R. cockerelli,and Cor- Among the residentspecies, 18 (27%) are en- acinacaledonica were found only as part of the demic to the Solomon Islands, with an addi- forest-interior flocks. Most flocks also con- tional 24 (37%) representedby subspeciesen- tained Coracina lineata, C. tenuirostris, C. holo- April 2001] Birdsof Isabel, Solomon Islands 475 polia,Myiagra ferrocyanea, Monarcha castaneiven- (Schodde 1977, Thiollay 1994, Debus 1995). tris,M. barbata,and Zosteropsmetcalfii, although Previous records of A. imitator from Isabel were these specieswere not restricted to flocks.The based on observations or vocalizations (Webb dominant members of most flocks, whether in 1992, 1995). Although other melanisticspeci- river-edge vegetation or the forest interior, mens of Accipiterfrom Isabel have been as- were M. castaneiventris,M. barbata,Z. metcalfii, signedto A. albogularis(Mayr 1945),all pied or C. tenuirostris,and C. holopolia.Other species black specimensof Accipiterfrom Isabelin the (e.g.Mic. finschii, Nectarinia jugularis) occasion- American Museum of Natural History are A. ally joined flocks.Within tall forests,the flocks imitator(LeCroy et al. in press). usually kept well abovethe ground(•10 m) in HALIAEETUS SANFORDI. Endemic to the Solo- the leafy subcanopy.Monarcha barbata and M. mon Islands, this eagle is usually regarded as castaneiventrisoccasionally descended to with- rare or uncommon.We recorded H. sanfordi in a few metersof the ground. The mixed-spe- nearly daily at our inland site;it was more com- cies flocks on Isabel resemblethose reported mon alongthe coast,as reported for New Geor- from Bougainville(Diamond 1975). gia (Blaber1990). Most of our inland records Foragingguilds.--We categorized foraging were of solitarybirds flying just abovethe can- guilds (Table 2) by the predominant substrate opy. M. Hafe collecteda male harassingfruit- or microhabitatwhere we observedthe species doves (Ptilinopusviridis, P. superbus)in a fruit- foraging (for speciesthat sally, we noted the ing Ficus tree. This specimen (UF 40196) perch, not the substrate)and by its predomi- yielded the world'sfirst skeletonof H. sanfordi, nant food. Stomachcontents from specimens along with a spreadwing and study skin (mi- (housed at UF) and literature sources (e.g. nus bill). As in other speciesof Haliaeetus(Ol- Schodde1977) were used to identify common son 1982),the first two phalangesin the fourth prey. The five canopyguilds made up 57.8% of toe are fused. A. Kratter observed an individ- the avifauna and included the three richest ual flying -50 m above the water at low tide guilds (14 frugivores,12 insectivores,9 omni- along a reef edge; it abruptly stoopeddown vores).The only comparablyspecies-rich guild and pluckeda -0.8 m long eel with its talons. was that feeding on fish and aquaticinverte- FALCOSEVERUS. On 11 July 1997, A. Kratter brates (9 species,12.7% of total); four of these and D. Steadmansaw a rather small but stocky were migrant shorebirds.The three terrestrial falcon flying -100 m over the river, just up- guilds (totaling 9.9% of the avifauna)and the streamfrom camp.The underpartswere most- two understory guilds (4.2%) were depauper- ly rufous, with streaking on the breast. This ate, especiallyin the interior of mature forests bird almost certainly was E severus,which (seeabove). The four insectivoreguilds (terres- agreeswith the observedfield marks and is the trial, undergrowth, canopy, aerial) totaled only small falconknown in the Solomons.This 29.6% of the avifauna,followed closelyby fru- is the first record for Isabel for E severus,which givores (24.0%). appearsto be rare in the archipelago,although Selectedspecies accounts.--We present ac- it has been recordedfrom Buka, Bougainville, countsfor three speciesnot previouslyknown Choiseul, Gizo, Kulambangara, , from Isabeland severalothers with poorly doc- and (Mayr 1945, Sibley 1946, Cain and umented natural histories. Galbraith 1956, Schodde 1977, Gibbs 1996, J. M. IXOBRYCHUS FLAVICOLLIS. The first Isabel re- Diamond pers. comm.). cords of I. flavicolliswere collectedon 11 July AMAURORNIS[OLIVACEA] MOLUCCANUS. In 1997, 1997 (UF 39523, adult female) and 27 June 1998 M. Hale saw an individual of this rail, but was (UW 60298,immature male). This widespread unable to collect it. We failed again to obtain speciesis known from other major islands in this speciesin 1998,although M. Hafe collected the Solomons. a male in hill forest,-3 km southof the village ACC•,•rER•M•'A•'OR. On 24 June 1998, we mist- of Tirotongna (UF 40216; complete skeleton, netted a dark-morphAccipiter in the forestin- skin, and spreadwing; first specimenof A. mol- terior (UF 40173, reproductively active male; uccanusfrom Isabel).The long wing (chord141 full detailsin LeCroy et al. in press).Melanistic mm) and pale plumageof UF 40216clearly ally specimenshad not been confirmed before in it with A. m. nigrifronsof Choiseul, Bougain- this species, which is difficult to identify ville, , and the BismarckArchi- 476 KRATTERET AL. [Auk, Vol. 118

TABLE1. Relativeabundance, habitat selection,foraging guild, and specimensof birds from the Garanga River valley. Relativeabundance: m = nonresidentmigrant; a = abundant(recorded daily in largenum- bers); c = common(encountered regularly by all observerseach time in habitat); u = uncommon(en- countereddaily or almost daily in small numbers);r = rare (encounteredless than five times).Habitats, givenin decreasingorder of importancefor eachspecies: r = river, beaches;rev = river-edgevegetation; f = forestinterior; a = aerial.Foraging Guild (microhabitat/prey):microhabitats: AQ = waterand water edge;AE = aerial;B = branches;C = canopy;T = terrestrial;U = understory;prey: FR = fruit and other vegetativematerial; FI = fish and aquatic invertebrates;N = nectarivore;O = omnivore; I = insects(in- cludesall nonaquaticarthropods and other invertebrates;V = terrestrial vertebrates.* Taxonendemic to the SolomonIslands (* before species= speciesendemic; * before subspecies= subspeciesendemic; ** before subspecies= subspeciesendemic to Isabel).

Relativeabun- Speci- mens dance Foraging FAMILY species (97/98) Habitat guild 97 98 ARDEIDAE Egrettasacra sacra Pacific Reef-Egret c/u r AQ/FI 2 -- Butorides striatus solomonensis Striated Heron u r AQ/FI -- 1 Nycticoraxcaledonicus mandibularis Rufous Night-Heron c r AQ/FI i -- Ixobrychusfiavicollis *woodfordi Black Bittern u r, rev AQ/FI i 1 ANATIDAE Anassuperciliosa pelewensis Pacific Black Duck c r AQ/FR 6 -- PANDIONIDAE Pandionhaliaetus cristatus Osprey u r, a AE/FI -- -- ACCIPITRIDAE Avicedasubcristata *gurneyi Pacific Baza c rev, c C/I-V 7 -- Haliasturindus *fiavirostris Brahminy Kite u r, a AE/V 1 -- Haliaeetus*sanfordi Sanford's Fish-Eagle u r, f, a AE/V-Fi -- 1 Accipiter*imitator Imitator Sparrowhawk u f, a C/V -- 1 Accipiternovaehollandiae * rufoschistaceus Gray Goshawk c f, rev, a C/V-I 4 1 FALCONIDAE Falcoseverus Oriental Hobby r a AE/V -- -- MEGAPODIIDAE Megapodius[freycinetl eremita Dusky Scrubfowl u rev, f T/O -- 2 RALLIDAE Nesoclopeus*woodfordi **immaculatus Woodford's Rail c rev, f T/O 14 -- Amaurornismoluccanus *nigrifrons Rufous-tailed Bush-hen r/-- f T/O -- -- Porphyrioporphyrio samoensis Purple Swamphen c rev T-AQ/O 2 -- CHARADRIIDAE Pluvialis[dominicalfulva Pacific Golden Plover mu r AQ/FI -- -- SCOLOPACIDAE Numeniusphaeopus Whimbrel mu r AQ/FI -- -- Tringanebularia Common Greenshank mr r AQ/FI -- -- Actitishypoleucos Common Sandpiper mc/r r AQ/FI i -- COLUMBIDAE Macropygiamackinlayi *arossi Mackinlay's Cuckoo-Dove u rev C/FR 5 -- Chalcophapsstephani *mottoni Stephan's Dove u f, rev T/FR 1 -- Ptilinopuss. superbusSuperb Fruit-Dove u f, rev C/FR i -- Ptilinopusviridis lewisiiClaret-breasted Fruit-Dove c f, rev C/FR 2 6 Ducularubricera *rufigula Red-knobbed Imperial-Pigeon c f, rev C/FR 5 1 PSITTACIDAE Chalcopsitta*cardinalis Cardinal Lory a rev, f, a C/FR 9 -- Trichoglossushaematodus massena Rainbow Lorikeet c rev, a C/FR 3 -- Cacatua*ducorpsii Ducorps's Cockatoo a rev, f, a C/FR 5 1 Micropsittafinschii *nanina Finsch's Pygmy-Parrot a f, rev B/FR 14 3 Geoffroyusheteroclitus heteroclitus Singing Parrot u f, a, c C/FR Eclectus roratus solomonensis Eclectus Parrot c f, rev, a C/FR 2 1 CUCULIDAE ChrysococcyxI. lucidus Golden Bronze Cuckoo mu f, rev C-U/I 3 1 Cacomantis variolosus *addendus Brush Cuckoo r rev C/I i -- Eudynamysscolopacea *alberti Asian Koel u rev, f C/O i -- STRIGIDAE Ninox*jacquinoti **jacquinoti Solomon Islands Hawk-Owl c rev, f C/V-I 3 -- April 2001] Birdsof Isabel, Solomon Islands 477

TABLE 1. Continued.

Relative abun- Speci- mens dance Foraging FAMILY species (97/98) Habitat guild 97 98 PODARGIDAE Podargusocellatus *inexpectatus Marbled Frogmouth --/r rev? C/I-V -- -- APODIDAE Collocalia vanikorensis vanikorensis Uniform Swiftlet c a AE/I i -- Collocaliaesculenta *?becki Glossy Swiftlet c a AE/I Collocaliaspodiopygia reichenowi White-rumped Swiftlet r a AE/I -- -- HEMIPROCNIDAE Hemiprocnemystacea woodfordiana Moustached Treeswift c rev, a AE/I 2 -- ALCEDINIDAE Alcedoatthis *salomonensis Common c r AQ/FI 9 -- Ceyxlepidus *meeki Variable Kingfisher u f, rev U/I-V 2 3 Ceyxpusillus *bougainvilleiLittle Kingfisher --/r rev U/FI -- -- *leucopygia Ultramarine Kingfisher c rev U/I-V 7 2 Halcyonchloris *alberti c rev, f C/I-V -- 1 Halcyonsancta mu rev C/I-V 4 -- C ORACIIDAE Eurystomusorientalis *solomonensis Dollarbird u rev, f C/I i 1 BUCEROTIDAE Acerosplicatus *harterti Blyth's Hornbill c f, rev, a C/FR 4 -- PITTIDAE Pitta *anerythra**anerythra Black-faced Pitta r/-- f T/I HIRUNDIDIDAE Hirundotahitica subfusca Pacific Swallow c r AE/I 2 -- CAMPEPHAGIDAE Coracina caledonica**welchmani Melanesian Cuckoo-shrike --/r f C/O -- 2 Coracinalineata *nigrifrons Yellow-eyed Cuckoo-shrike u f C/O 6 2 Coracinapapuensis *perpillada White-bellied Cuckoo-shrike c rev C/O 4 3 Coracina tenuirostris *saturatior Slender-billed Cicadabird c f, rev C/O 5 1 Coracina*holopolia holopolia Solomon Islands Cuckoo-shrike u f, rev C/O 3 3 Rhipiduraleucophrys melaleuca Willie Wagtail c r T/I 7 2 Rhipidurarufifrons *commoda Rufous r f C/I 2 1 Rhipidura*cockerelli interposita White-winged Fantail r/u f C/I -- 3 MONARCHIDAE Monarcha *castaneiventriscastaneiventris Chestnut-bellied Monarch c f, rev C/I 10 6 Monarcha *barbata barbataBlack-and-white Monarch u/c f C-U/I 4 7 Myiagra*ferrocyaneaferrocyanea Steel-blue Flycatcher u f, rev C/I 2 2 PACHYCEPHALIDAE Pachycephalapectoralis *orioloides Golden Whistler u f C/O 4 3 STURNIDAE Aploniscantoroides Singing Starling r/-- rev C/FR -- -- Aplonis*grandis grandis Brown-winged Starling c rev,f C/FR 6 -- Aplonismetallica nitidus Metallic Starling c rev, f C/FR 10 -- Mino dumontiikreffti Yellow-faced Myna c rev, f C/FR 8 3 CORVIDAE Corvus*woodfordi vegetus White-billed Crow c f, rev C/O 4 1 NECTARINIIDAE Nectariniajugularisflavigaster Olive-backed Sunbird a rev, f C-U/N 12 -- DICAEIDAE *aeneum aeneum Midget c rev,f C-U/N 13 3 ZOSTEROPIDAE Zosterops*metcalfii **metcalfii Yellow-throated White-eye a f, rev C/O 11 2 MELIPHAGIDAE Myzomela*lafargei Scarlet-naped Myzomela r/c f C/N -- 1 478 KRATTERET AL. [Auk,Vol. 118

TABLE2. Foragingguilds of residentspecies. For def- other pairs would follow when the first duet inition of guilds, see text and Table 1. For species died off, and so on up and down the river. with multiple categoriesof microhabitator prey (Table1), the predominant(first) categoryis used. The three forms of N. woodfordiare N. w. im- maculatuson Isabel,N. w. woodfordion Guadal- Guild canal,and N. w. tertiuson Bougainville.An un- (microhabitat/prey) Number Percentage describedform of Nesoclopeusmay occur as Aerial/insects 5 7.0 well on New Georgia (Blaber1990). Our spec- Aerial / vertebrates 3 4.2 imens representtwo downy chicks;one im- Aerial/fish I 1.4 mature female and four immature males(bursa Aquatic/fish 9 12.7 Aquatic/ fruit I 1.4 present); and five adult females and one adult Terrestrial/fruit I 1.4 male (bursaabsent); the age of onemale could Terrestrial/insects 2 2.8 not be assessed.The immature and adult spec- Terrestrial/omnivore 4 5.6 imens showeda great deal of plumagevaria- Understory/ insects 2 2.8 Understory/fish I 1.4 tion, even within age/sex classes.Nine of the Branches/fruit I 1.4 12 nondownyspecimens had somewhite spot- Canopy/fruit 14 19.7 ting in the remiges, underwing coverts,and Canopy/insects 12 16.9 undertail coverts. The characters used to dis- Canopy/nectar 3 4.2 Canopy/ omnivore 9 12.7 tinguish N. w.immaculatus (larger, blacker, lack- Canopy/ vertebrates 3 4.2 ing white mottling on the underparts and white barring in the remiges, axillaries, and wing coverts;Mayr 1949) are negated by the variation in our specimens.Intrapopulation pelago,rather than A.m. ultimusof islandseast variation went unnoticed by Mayr (1949) be- of Isabel (Taylor1998). causeso few specimenswere availablethen. On the basisof vocalizations,Webb (1992) re- Previous authors describedN. woodfordias gardedA. moluccanusto occuron Isabel.How- "probably flightless."Men from Isabel stated ever, the calls he described ("wild duets con- that it never flies; this was confirmed in our sisting of alternatingshrieks overlapping in a specimens.The distal part of eachremix or rec- harmonic beat") sound similar to us to the calls trix is lax, with a weak rachis and noninter- of Nesoclopeuswoodfordi (see below). We did not locking barbs. Lacking aerodynamicfunction, hear the calls that Webb (1992) describedfor N. the flight featherscannot support the body woodfordi("unmusical series of metallicshrieks weight. The wing is small relativeto the leg. unvarying in pitch"), but theseare reminiscent Also, the sternal keel and the relative mass of of the distinctivegrowling shrieksdescribed the breastmuscles (pectoralis major and supra- for A. moluccanusand A. olivaceus(Lambert coracoideus) are much reduced. 1998). MICROPSITTA FINSCHII. In contrast to Webb's N•socLoe•us WOODFORDI.This rail has been de- (1992) report that M. finschii was confined to scribed as "extinct or in low numbers and en- montane areas on Isabel, we found this tiny dangered"(Diamond 1987), "obviously scarce" parrot to be one of the most abundant interior (Blaber1990), or "closeto extinction"(Taylor forest speciesalong the Garanga River. We 1998), even though the last sourcecited Webb foundthree other species (Collocalia spodiopygia, (1992),who regarded it as uncommonon Isabel Coracinacaledonica, Pachycephala pectoralis) at in old gardensand river edge habitats.In fact, our lowland site that previouslyhad beencon- this large flightlessrail was commonin early sidered as largely or wholly montanespecies successionalhabitats within 100 m of the river, on Isabel (Mayr 1945,Webb 1992). althoughwe alsofound it in matureforests. Elu- Aided by stiffenedrectrices, Micropsitta fin- sive and difficult to see when not calling, N. schiiusually hitched along trunks and larger woodfordivocalized infrequently(a pair near branches,from 4-20 m up in monospecific camp would duet only onceor twice per day), flocksof two to eightbirds. We observedbirds yet the loudnessand distinctivenessof its vo- entering arborealtermitaria at dusk, presum- calizationsmade detection easy. In mostlate af- ably to roost (seeJuniper and Parr 1998).The ternoonsor on cloudydays, this rail engagedin displaysin speciesof Micropsittaare not well boutsof duets:one pair would begin,and then knownbecause they are rarelykept in captivity April2001] Birdsof Isabel, Solomon Islands 479 and are inconspicuousin the wild. On 27 June pers. obs.), although we found four of those 1998, AWK observed a pair of M. finschii speciesnear sealevel alongthe GarangaRiver. perchedsideways on a 2 cm verticalvine, -6 m Thus, the relativelyhigh overall diversity of above the ground. The female was -20 cm birds on Isabel may result more from species above the male, and their bills were oriented in packing within habitats and speciesreplace- the same direction. The male hitched up the ments across habitats within the floristically stem and touched bills with the female. This rich lowlands,rather than from elevationalspe- processwas repeated several times over the cialization of species. next minute as the male chasedthe female up We found at least 86% of Isabel's resident the vine. At one point, the male flew 30 cm out speciesat the Garanga River site, indicating away from the vine and hoveredfor a few sec- that a singlelowland site dominatedby forest onds.Both birds gavea high-pitchedchattering may containalmost all specieson an island, continuouslyduring the display. even an island with elevations >1,000 m. The EUDYNAMYS$COLOPACEA. Except for an uncon- lower Garanga River has created a mosaic of firmedreport of a bird seenby J.Diamond (cit- older forests, successionalvegetation, rocky ed in Webb1992), E. scolopaceahad not beenre- and sandybeaches, and forestedges. The bird ported for Isabel. We recorded E. scolopaceacommunity reflects that habitatheterogeneity. daily in small numbers, especially in river- The erosional and depositional processesas- edge vegetation.Although difficult to see, its sociatedwith meanderingrivers are key con- distinctivesong was given frequently and often tributors to habitat heterogeneityat the land- at night (as reported by Blaber 1990).An im- scapescale elsewhere in tropicallowlands (e.g. mature (UW 58734), collectedon 10 July 1997, Robinsonand Terborgh1997). is the first specimenfrom Isabel. Understoryavifauna.--We were struckby how few birds we observedin the understoryof ma- DISCUSSION ture forests.Our mist-net efforts, though min- imal, supported these observations.A single Diversity.--Incontrast to islandsfurther east mist net acrossthe Garanga River captured in Polynesia(e.g. Steadman et al. 1990,Stead- Egrettasacra, two Anas superciliosa,Actitis hy- man and Freifeld 1998, Freifeld 1999), in Va- poleucos,and three Alcedoatthis in -30 net- nuatu(Bowen 1997), or on peripheralislands in hours in 1997. In the interior forest, 170 net- the Solomons (Diamond et al. 1976, Diamond hours in 1997 yielded a single bird (Monarcha 1984, Filardi et al. 1999), the bird communities barbata);in 1998, 182 net-hoursyielded eight of islands in the main chain of the Solomons birds, all nonpasserines(4 Ceyx lepidus,3 Mi- have more direct Papuan influencesand are cropsittafinschii, and 1 Accipiterimitator). Thus, more diverse(Diamond and Mayr 1976).Major 352 net-hours from nine nets at two mature for- islands of the Solomons, such as Isabel, are estsites captured only onepasserine and eight larger and more geologicallyand topographi- nonpasserines(2.6 birds/100 net-hours). In cally heterogeneousthan most islandsfurther contrast,most continentaltropical forest sites eastin Oceania.Increased diversity of birds on have diverse communities of terrestrial and un- larger or higher islandsin Oceaniais thought derstory species,especially passerines, which to result from non-overlappingdistributions are usually the easiestavifaunal component to amongecologically similar speciesacross ele- samplewith nets (Remsenand Good 1996). vational gradients,which may arise from com- Our observationsand capture rates indicate petitive interactions(Mayr and Diamond 1976, thatthe terrestrialand understory avifauna at the Diamond and Mayr 1976, Schodde1977). Al- GarangaRiver site is depauperatein both diver- thoughmontane areas of Isabelare not partic- sity and abundance.Terrestrial and understory ularly high or extensive,two strictlymontane guildstotaled only 16%of the residentlandbirds speciesare known, Charmosynameeki and Phyl- (Fig. 2), comparedto 31% acrossseven lowland loscopuspoliocephalus (Mayr 1945).Several other forestsites in New Guinea (Bell1982, Beehler and speciesare primarily montaneon Isabel(Nes- Mack in press).The only regularlyoccurring un- asio solomonensis,Collocalia spodiopygia, Pitta derstoryor terrestrialspecies that we did not net anerythra,Coracina caledonica, Pachycephala pec- were large non-passerines(Nesoclopeus woodfordi, toralis; Webb 1992, A.W.K., C.E.E, and C.E.S. Megapodiuseremita, Chalcophaps stephani) and one 480 KRATTERET AL. [Auk,Vol. 118

branches a) ISABEL aerial 2% na (106species) than Isabel,12 of the 50 species (65 spp.) 14% of birds recordedin prehistoriccultural depos- its no longerlive on the island (Steadmanet al. aquatic 9% 1999).Other possiblecauses for a low capture- rate of understorybirds seemless likely, such terrestrialas seasonal shifts in foragingheight or habitat 11% preference. 59% Speciesnot recorded.--Withour new records understory for Ixobrychusfiavicollis, Falco severus, and Eu- 5% dynamusscolopacea, Isabel and its offshoreislets are known to have 76 speciesof residentland b) NEW GUINEA and freshwater birds (previous summations (7 sites, average145 spp.) 1%6%2% from Mayr 1945, Webb 1992). One other spe- cies,Accipiter albogularis, has been listed as oc- 14% curring on Isabel (Mayr 1945, Webb 1992, 1997), although all Isabel specimensin the American Museum of Natural History appear to be A. imitator (see its account) and the few 600/, .... .-.-.-.- ... 17% sight recordsare controversial(Debus 1995). We found 65 of the 76 speciesknown from Isabel(86%) in the GarangaRiver valley. Of the 11 speciesnot recorded, we found five at other FIG. 2. Distribution of resident landbird species sites on Isabel in 1997 and 1998. Four of these among preferred microhabitatsat lowland forest sitesin (a) the SolomonIslands, along the Garanga (Island Imperial-Pigeon Ducula pistrinaria, River on Isabel (this study, Table 1); and (b) Papua Halcyons. saurophaga,Islet New Guinea(one site on the BrownRiver floodplain Monarch Monarchacinerascens impediens, and [Bell 1982] and six sites in the Lakekamu Basin Nicobar Pigeon Caloenasn. nicobarica)are [Beehlerand Mack in press]).For b the proportionof small-islandor beachspecialists that occuron the avifaunausing eachmicrohabitat was averaged isletsjust off the north coastof Isabel(see Fig. across the seven sites; microhabitat use for these sites 1). Another, the Fearful Owl (Nesasiosolomonen- is from the above citations, as well as Beehler et al. sis), we found in the mountains southeast of (1986). Buala (Fig. 1). Of the remaining six species,two (Island (Pitta anerythra),the latter two being Leaf-Warbler, Phylloscopuspoliocephalus, and rare (Table1). Missingfrom the site were those Meek's Lorikeet, Charmosynameeki) may be terrestrialand understorytaxa that eitherstill oc- confinedto montaneforests (Mayr 1945,Webb cur or onceoccurred on nearbylarge islandsin 1992), one is nocturnal (White-throated Eared- the Solomonsor Bismarcks(Mayr 1945,Diamond Nightjar, Eurostopodusmystacalis) and may not 1991,Steadman et al. 1999):a largespecies of Me- have been calling during our visits, and one gapodius,other flightless rails (Gallirallus,Porphy- (White-throated Pigeon, Columbavitiensis) is rio, Gallinula),columbids (Gallicolumba spp., Mi- known on Isabelfrom a Whitney SouthSea Ex- crogoura meeki), and passerines (Turdus pedition specimen,but has not beenseen since poliocephalus,Petroica multicolor). (Webb 1992). Reinwardtoenacrassirostris (Crest- An impoverishedunderstory bird commu- ed Cuckoo-Dove) has never been collected on nity is a widespreadphenomenon in Oceania, Isabel--the only record is a sight recordby muchof this resultingfrom prehistoric,anthro- Webb (1992), who reported that locals said it pogenic extinctions (Steadman 1995). The de- was common(but his given local name--fela-- pauperatenature of the understoryavifauna in is the sameas that given to us for Macropygia the GarangaRiver valley may alsobe anthro- mackinlayi).The other speciesnot definitelyre- pogenic,which would include predationby corded by us (DuchessLorikeet, Charmosyna non-native mammals. Even on New Ireland, margarethae)may have been observedin the which is larger, higher, and closer to New Garanga River valley in 1998,but could not be Guinea, and with a much richer modern avifau- confirmed. On two occasions, A. Kratter noted April 2001] Birdsof Isabel, Solomon Islands 481 flocks(2-6 birds) of very small psittacidsflying thoughthe Melanesiantradition of customland within the canopy giving two-note calls that can complicate issuesof land protection, that were less shrill than those of Trichoglossussystem of land managementhas resultedin a haematodus. fairly intact avifauna surviving on Isabel in Conservation.--TheGaranga River valley is a spite of many millennia of human occupation. prime candidatefor conservation.A protected Other than an expansionof commerciallog- area from the river mouth and adjacent off- ging or other intensiveland uses(e.g. oil palm shore islets and extendingalong the lowland plantations),the future of birdlife on Isabel is portions of the River would include popula- not bleak. tions of at least 91% of the Isabel'sspecies of birds.Incorporating highlands at the sourcesof ACKNOWLEDGMENTS the Garanga watershed, a linear distanceof only 8 km from the coast,would protect pop- Field researchwas supportedby the Universityof ulationsof all residentlandbird species,except Florida College of Liberal Arts and Sciences(grant perhaps the severalspecies of uncertain status. RDA-1-23 95-86) to D.W.S.and the Ornithology En- Protecting the lower parts of the watershed dowmentof the Universityof WashingtonBurke Mu- seum to C.E.F. and C.E.S. Museum-based research is of foremostimportance because lowland for- estsare the first to be destroyedin the Solomon was supported by the National ScienceFoundation Islands (Stattersfield et al. 1998) and across (grant EAR-9714819to D.W.S.).Mary LeCroy and Ja- red Diamond provided information that was instru- Oceania (Steadman 1995). Although mature mental to our analyses.Terry Taylor helped to pre- forestsare preferredby many speciesalong the pare specimens.Moses Biliki, JoeHorako, Lawrence Garanga River, successionalriverine habitats Foanaota,and Audrey Rusa kindly provided logis- are essentialfor others,including the flightless tical support and researchpermits from the national rail Nesoclopeuswoodfordi, a species of greatevo- governmentof the SolomonIslands. On Isabel,Amos lutionary importancethat is likely to remain Gigini, Mark Hafe, Austel Ruumana, and William, commonas long as riverine habitatsare intact. Veronica, and Roger Manehage supplied compan- Although we doubt that the GarangaRiver ionship,access to the site,and immeasurablehelp in avifauna is pristine, we found populationsof the field. For constructivecomments that improved many speciesin the samefamilies (megapodes, the manuscript,we thank B. M. Beehler,S. Debus,J. rails, columbids,and psittacids)that dominate M. Diamond, H. Ford, and T. K. Pratt. the anthropogenicextinction record in Oceania Noting that Diamond (1975) dedicateda paper on Bougainvillebirds (written in 1974)to Ernst Mayr on (Steadman 1995). Introduced speciesof birds his 70th birthday, we take this wonderful opportu- are absent.The GarangaRiver sitesustains one nity to dedicate our paper to ProfessorMayr on the species listed as "near-threatened" (Corvus occasionof his 97th birthday. All that we have done woodfordi),two listed as "vulnerable"(Haliaee- in the Solomon Islands rests on the foundation laid tussanfordi, Pitta anerythra),and two others(Ac- by thesetwo gifted men. cipiterimitator, Nesoclopeus woodfordi) listed as

"endangered" (Collar et al. 1994). At popula- LITERATURE CITED tion levels suggestedby our fieldwork, those classificationsmay be exaggeratedfor all five BEEHLER,B. M., AND A. L. MACK. 1999. Constraints species.The conservationstatus of birds in to characterisingspatial heterogeneity in a low- Oceania can be evaluated accurately only land forest avifauna in New Guinea. In Proceed- throughrigorous fieldwork. ings XXII International Ornithological Con- The Garanga River valley is particularly in- gress.(N. Adams and R. Slotow,Eds.) University viting for protection becausethe provincial of Natal, Durban, South Africa. governmentof Isabelowns its lower stretches, BEEHLER,B. M., T. K. PRATT,AND D. A. ZIMMERMAN. 1986.Birds of New Guinea. PrincetonUniversity therebysimplifying ownership issues. The pro- Press,Princeton, New Jersey. vincial governmentalso owns muchof FeraIs- BELL,H. L. 1982.A bird community of lowland rain- land (12 km ESE of the Garanga River mouth), forest in New Guinea. I. Compositionand den- where we found four of the speciesmissing sity of the avifauna.Emu 82:24-41. along the Garanga. Thus, a tract of land with BLABER,S. J. M. 1990. A checklist and notes on the at least91% of Isabel'sresident species of land- current statusof the birds of New Georgia,West- birds is already in government hands. A1- ern province,Solomon Islands. Emu 90:205-214. 482 KRATTERET AL. [Auk, Vol. 118

BOWEN,J. 1997. The status of the avifauna of Loru HADDEN, D. 1981. Birds of the North Solomons. Wau Protected Area, Santo, . Bird Conserva- EcologyInstitute Handbook 8. Wau,Papua New tion International 7:331-344. Guinea. CAIN, A. J., AND I. C. J. GALBRAITH.1956. Field notes JUNIPER,T., AND M. PARR. 1998. Parrots, a Guide to on birds of the eastern Solomon Islands. Ibis 98: the Parrots of the World. Yale University Press, 100-134. New Haven, Connecticut. COLLAR,N.J., M. J. CROSBY,AND A. J. STATTERSFIELD. KAESTNER,P. 1987. Some observations from lowland 1994. Birds to Watch 2. BirdLife Conservation Se- swamp forest in south Bougainville.Muruk 2: ries no. 4 34-38. CORNER,E. J. H. 1967. Ficus in the Solomon Islands KARR, J. R. 1990. The avifauna of Barro Colorado Is- and its bearing on the post-Jurassichistory of land and the PipelineRoad, Panama. Pages 183- . PhilosophicalTransactions of the 198 in Four NeotropicalRainforests (A. H. Gen- Royal Societyof London, SeriesB 253:23-159. try, Ed.). Yale University Press, New Haven, DEBUS,S. J. S. 1995.The morphsof SolomonIslands Connecticut. Accipiterspp.--A commenton Webb(1992). Emu KRATTER,A. W., D. W. STEADMAN, C. E. SMITH, AND 95:71-72. C. E. FILARDI.2001. Reproductivecondition, DIAMOND,J. M. 1975. Distributional ecology and moult, and body massof birds from Isabel,Sol- habits of some Bougainville birds (SolomonIs- omon Islands. Bulletin of the British Ornitholo- lands). Condor 77:14-23. gists'Club 121:in press. DIAMOND,J. M. 1982.Effect of speciespool size on LAMBERT,E R. 1998. A new speciesof Amaurornis speciesoccurrence frequencies: Musical chairs from the Talaud Islands, , and a review on islands.Proceedings of the National Acade- of of bush hens occurring from the my of SciencesUSA 79:2420-2424. Philippinesto Australasia.Bulletin of theBritish DIAMOND, J. M. 1984. The avifauna of Rennell and Ornithologists'Club 118:67-82. Bellona islands. Natural History of Rennell Is- LECROY, M., A. W. KRATTER, D. W. STEADMAN, AND land, British Solomon Islands 8:127-168. n. P. WEBB.2001. Accipiterimitator on Isabel Is- DIAMOND,J. M. 1987. Extant unlessproven extinct? land, SolomonIslands. Emu 101:in press. Or, extinct unless proven extant? Conservation MAYR, E. 1945. Birds of the Southwest Pacific. Mac- Biology 1:77-79. millan, New York. DIAMOND,J. M. 1991.A new speciesof rail from the MAYR, E. 1949. Notes on the birds of northern Mel- SolomonIslands and convergentevolution of in- anesia, 2. American Museum Novitates 1417. sular fiightlessness.Auk 108:461-470. MAYR, E., AND J. M. DIAMOND. 1976. Birds on islands DIAMOND, J. M., M. E. GILPIN, AND E. MAYR. 1976. in the sky: Origin of the montane avifauna in Species-distancerelation for birds of the Solo- northern Melanesia.Proceedings of the National mon Archipelago,and the paradoxof the great Academy of SciencesUSA 73:1765-1769. speciators.Proceedings of the National Acade- OLSON,S. L. 1982.The distributionof fusedphalan- my of SciencesUSA 73:2160-2164. ges of the inner toe in the Accipitridae.Bulletin DIAMOND,J. M., ANDE. MAYR.1976. Species-area re- of the BritishOrnithologists' Club 102:8-12. lationfor birdsof the SolomonArchipelago. Pro- REMSEN,J. V., JR.,ED. 1997.Studies in Neotropicalor- ceedingsof the National Academy of Sciences nithology honoring Ted Parker.Ornithological USA 73:262-266 Monographs,no. 48. FILARDI, C. E., C. E. SMITH, A. W. KRATTER,D. W. REMSEN,J. V., JR., AND D. A. GOOD. 1996. Misuse of STEADMAN, AND H. P. WEBB. 1999. New behav- data from mist-net captures to assessrelative ioral, ecological,and biogeographicdata on the abundancein bird populations.Auk 113:381- avifauna of Rennell, Solomon Islands. Pacific 398. Science 52:319-340. ROBINSON,S. K., AND J. TERBORGH.1997. Bird com- FREIFELD,H. B. 1999. Habitat relationshipsof forest munity dynamics along primary successional birds on Tutuila Island, American Samoa. Jour- gradientsof a Amazonian whitewater river. Or- nal of Biogeography26:1191-1213. nithologicalMonographs no. 48:641-672. GALBRAITH,I. C. J., AND E. H. GALBRAITH. 1962. Land SCHODDE,R. 1977. Survey of birds of southernBou- birds of Guadalcanal and the San Cristoval gainville Island, . Contribu- group, easternSolomon Islands. Bulletin of the tions to Papuasianornithology VI. Common- British Museum (Natural History), Zoology 9:1- wealth Scientific and Industrial Research 86. Organization, Melbourne,. GIBBS,D. 1996. Notes on Solomon Island birds. Bul- SIBLEY,C. G. 1946.The secondspecimen of the Ori- letin of the British Ornithologists' Club 116:18- ental Hobby from the SolomonIslands. Condor 25. 48:97. GREENSLADE,P.J. M. 1968.Island patternsin the Sol- STATTERSFIELD,A. J., M. J. CROSBY,A. J. LONG, AND omon Islands bird fauna. Evolution 22:751-761. D.C. WEGE. 1998. Endemic bird areas of the April2001] Birdsof Isabel, Solomon Islands 483

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