Terrestrial birds of the Indo-Pacific 361 Terrestrial birds of the Indo-Pacific

B Michaux Private Bag, Kaukapakapa, New Zealand

Key words: Indo-Pacific biogeography, fragmentation of east Gondwana, nightjars, Gallirallus philippensis

Abstract “ of the 265 species of land birds which are known from that part of New Guinea which is The avifaunas of New Zealand, New Caledonia, central Poly- opposite New Britain, only about 80 species nesia, New Guinea, Maluku and Sulawesi are discussed Dis- have a representative on New Britain In other tributional patterns within and between avifaunas are de- scribed and related to Mesozoic tectonic activity along the words, the 45 mile [70 km] stretch of water east Gondwana margin Spreading, rift formation, subduc- which separates the two islands has prevented tion, obduction and mobile arc systems were all important the crossing over of 70 percent of the New components of this activity Avian distributional patterns at Guinean species” family, genus and species levels are discussed in terms of rift- arc interactions within modern zones of active plate conver- gence Clearly there is more to colonisation than cross- ing barriers because most birds should be able to cross 70 km of sea Yet according to Mayr Introduction only 30% of the Huon peninsula avifauna has shown evidence of colonising New Britain Mayr I’m sure that Leon Croizat would have derived (1941) thought that 70% ‘sedentary’ species was some satisfaction at this coming together of biol- general for avifaunas within the Indo-Australian ogists and geologists to discuss matters of mutu- archipelago Poor colonising ability and a devel- al interest regarding the Indo-Pacific region For oped taxonomic and systematic base make birds biologists the Indo-Pacific is an evolutionary excellent biogeographical tools laboratory in which taxonomic diversification The avifaunas of New Zealand, New Guinea, has occurred on a dynamic and complex stage Fiji, Tonga, Samoa, Maluku and Sulawesi are For geologists it is a region where a modern described and broad patterns in the distribu- orogeny can be studied and its development tional data discussed These patterns link Indo- through time reconstructed The key question Pacific islands into a number of groupings for me is to what extent are distributional pat- These groupings do not always follow conven- terns consistent with the reconstructions provid- tion, for example Timor is linked to south ed by geologists? Maluku Island groupings based on avifaunal Even though most birds are highly mobile, a distributions are related to three broad geologi- component of an island’s avifauna is sedentary cal systems These are the Melanesian rift-arc What proportion of an avifauna is sedentary de- system, the Banda rift-arc system and the Sumba pends on several factors, including distance to ar- terrane The geological histories of these struc- eas that could be colonised Mayr (1941) dis- tures are described and differences in geological cussed the efficiency of sea barriers to bird coloni- interpretations discussed sation from New Guinea to New Britain and said: Finally, distributional patterns are interpreted

Biogeography and Geological Evolution of SE Asia, pp 361-391 Edited by Robert Hall and Jeremy D Holloway © 1998 Backhuys Publishers, Leiden, The Netherlands 362 B Michaux in the light of geologists’ reconstructions These islands such as New Caledonia, Vanuatu, the interpretations are necessarily general for two Solomon Islands, Fiji, Samoa and Tonga, or In- reasons Firstly, cladistic bird phylogenies of ap- donesia The genera Nestor and Cyanoramphus propriate scope are not available In most cases (Psittacidae) and the species Eudynamis the taxa used in describing patterns are either taitensis (Cuculidae) are endemic to the south- species or genera for which monophyly is likely, west Pacific but as yet undemonstrated cladistically How- Low diversity at family, generic and (with few ever, Sibley and Ahlquist (1990) have produced exceptions) specific levels is also a characteristic a systematic treatment which is used to examine of the New Zealand terrestrial bird fauna While the relationship between nightjar evolution and it is true that extinction has reduced taxonomic the Mesozoic and Tertiary history of the Indo- diversity (the families Dinornithidae, Emeidae, Pacific in some detail The second reason is that Pelagornithidae, Pelecanidae, Phasianidae, Ap- the geology of the Indo-Pacific is complicated tornithidae, Aegothelidae, Turnagridae and Cor- and many geological questions are yet to be re- vidae are no longer part of New Zealand’s en- solved Definitive answers to problems of Indo- demic fauna), it would seem that low diversity Pacific biogeography are not available at has always been a characteristic of New Zea- present, but this is precisely what makes the land’s terrestrial avifauna Bird families that one Indo-Pacific such an interesting area for re- might have expected to be part of New Zea- search land’s pre-European avifauna are missing For example, some self- or deliberately introduced members of the families Cracticidae (Australian Avifaunas Magpie), Hirundinidae (Welcome Swallow) and Turdidae (Blackbird and Song Thrush) have es- New Zealand tablished themselves in New Zealand Within families there is a general paucity of Extant and extinct New Zealand (Fig1: 1) terres- genera and species This point is illustrated by trial birds are listed in Table 1 The data came the family Columbidae which is represented in from Falla et al (1979), Fordyce (1982), Fuller New Zealand by a single species Hemiphaga (1987) and Turbott (1990) Species unknown to novaeseelandiae The genus Hemiphaga is now pre-European Maori (i e, lack a Maori name) confined to New Zealand, but an extinct sub- and self-introduced since European arrival are species, H novaeseelandiae spadicae, was not included in Table 1, because many of these found on Norfolk Island (Schodde et al , 1983) species appear to be associated with man-made This can be contrasted with the Columbidae in habitats The presence of these species may be New Caledonia, where there are six species in due to the availability of grassland habitats, in- five genera, or in Australia where there are 23 troduced food sources or nesting sites, and they species in eleven genera The exceptions to this have been excluded Table 1 shows two charac- pattern are the Anatidae, Rallidae and Phalacro- teristics of New Zealand’s terrestrial avifauna, a coracidae This latter family reaches its greatest high degree of endemism and low taxonomic specific diversity in New Zealand waters Cor- diversity morants have been included in this study be- Five endemic terrestrial avian families are cause four species (Phalacrocorax carbo, P sul- listed in Table 1 The Apterygidae (kiwi), cirostris, P melanoleucos, and P varius) are Acanthisittidae (New Zealand wrens) and predominantly estuarine or freshwater, but oth- Callaeatidae (New Zealand wattle birds, includ- er members of the family are marine or oceanic ing the extinct huia) are extant, while two fami- species lies of moa, the Emeidae and Dinornithidae, are extinct New Zealand thrushes were represented by a single extinct species — the piopio — re- New Caledonia garded by Turbott (1990) as a member of the Paradisaeidae, but better placed either in the The terrestrial avifauna of New Caledonia (Fig1: Turnagridae or Ptilonorhynchidae (Sibley, pers 2) and the Loyalty Islands is listed in Table 2 comm, 1996) Thirty five genera and 79 species The data for Table 2 are from Mayr (1945) Table listed in Table 1 are endemic Only seven spe- 2 shows that the number of avian taxa endemic cies are shared with Australia and three of these in New Caledonia is less than in New Zealand are also found elsewhere A further twenty spe- There are five endemic genera (Keast (1996) cies are shared with Australia, southwest Pacific records eight), 21 endemic species and a single Terrestrial birds of the Indo-Pacific 363

Fig 1 The Indo-Pacific region showing extent of continental crust and marginal oceanic basins (stippled) Avifaunas discussed in text: 1 = New Zealand, 2 = New Caledonia, 3 = Fiji, Tonga and Samoa, 4 = New Guinea, 5 = Maluku, 6 = Sulawesi Marginal basins: AS = Andaman Sea, sBS = south Banda Sea, CS = Coral Sea, CeS = Celebes Sea, sCSB = South China Sea, M = Makassar Strait, MB = Manus Basin, NC = New Caledonian Basin, NFB = North Fiji Basin, PO = SW Pacific, SB = Sulu Basin, SCB = Santa Cruz and Loyalty Basins, SFB = south Fiji Basin, SS = Solomon Sea, TS = Tasman Sea CP = Campbell Plateau, CR = Chatham Rise, LHR = Lord Howe Rise, NR = Norfolk Ridge, OJP = Ontong Java Plateau, IO = Indian Ocean Bo = Borneo, F = Fiji, nI = North Island, New Zealand, sI = South Island, New Zealand, J = Java, NG = New Guinea, S = Samoa, So = Solomons, T = Tonga, V = Vanuatu

endemic family — the Rhynochetidae — in New tralian avifauna Caledonia The number of endemic taxa has Within the non-endemic element of New Cal- been greater in the past Balouet and Olson edonia’s avifauna, distributional patterns are (1989) describe a large, flightless species similar to those discussed for New Zealand Sylviornis neocaledonica (Megapodiidae?) from Only three species are shared exclusively with Holocene deposits, together with eleven extinct Australia despite New Caledonia’s proximity to non-passerine species that have relatives on the Australian continent Thirty three species smaller islands off New Zealand, northern Mela- (44% of the total avifauna) have widespread dis- nesia and Asia Two extinct birds of prey have tributions, listed in Table 2 as Australia-south- been recorded by Thiollay (1993) New Zealand west Pacific-Indonesia Five of these species are and New Caledonia have few avian families, absent from Australia and ten are present in which indicates a long isolation from the Aus- New Zealand There is also a pronounced south- 364 B Michaux

Table 1 Terrestrial birds of New Zealand Bold entries are extinct taxa Note on flycatchers and warblers: in Table 1 and following, Rhipidura is included with other monarch flycatchers in the Monarchidae and Gerygone with other Australian wren- warblers in the Acanthizidae Sibley and Ahlquist (1990) include the former in the expanded family Corvidae and the latter in the family Pardalotidae

Family Endemic Shared with Australia Shared with Australia SW Pacific +SW Pacific/ New Guinea/Indonesia

Emeidae Anomalopteryx Megalopteryx Pachyornis Emeus Euryapteryx Dinornithidae Dinornis Apterygidae Apteryx Acanthisittidae Acanthisitta chloris Xenicus longipes Xenicus gilviventris Traversia lyalli Pachyplichas yaldwyni Pachyplichas jagmi Callaeatidae Callaeas cinerea Philesturnus carunculatus Heteralocha acutirostris Podicipedidae Poliocephalus rufopectus Podiceps cristatus Pelagornithidae Pelagornis miocaenus Pseudodontornis stirtoni Pelecanidae Pelecanus novaezealandiae Phalacrocoracidae Leucocarbo carunculatus Phalacrocorax varius Phalacrocorax carbo Leucocarbo chalconotus Phalacrocorax sulcirostris Leucocarbo onslowi Phalacrocorax melanoleucos Leucocarbo ranfurlyi Leucocarbo colensoi Leucocarbo campbelli Leucocarbo atriceps Stictocarbo punctatus Stictocarbo punctatus steadi Stictocarbo featherstoni Ardeidae Ixobrychus novaezelandiae Egretta alba Egretta sacra Botaurus poiciloptilus Threskiornithidae Platalea regia Anatidae Pachyanas chathamica Oxyura australis Anas gibberifrons Euryanas finschi Anas rhynchotis Anas superciliosa Cnemiornis gracilis Anas gracilis Hymenolaimus malacorhynchos Tadorna variegata Anas chlorotis Anas aucklandica Aythya novaeseelandiae Biziura delautouri Malacorhynchus scarletti Cygnus sumnerensis Mergus australis Accipitridae Circus eylesi Circus approximans Harpagornis moorei Haliaeetus australis Falconidae Falco novaeseelandiae Phasianidae Coturnix novaezealandiae Rallidae Fulica chathamensis Porzana tabuensis Fulica prisca Porzana pusilla Gallinula hodgeni Porphyrio porphyrio Capellirallus karamu Gallirallus philippensis Diaphorapteryx hawkingi Rallus pectoralis Gallirallus australis Rallus modestus Porphyrio mantelli Terrestrial birds of the Indo-Pacific 365

Table 1 Continued

Family Endemic Shared with Australia Shared with Australia SW Pacific +SW Pacific/ New Guinea/Indonesia

Aptornithidae Aptornis otidiformis Aptornis defossor Charadriidae Charadrius obscurus Charadrius bicinctus Anarhynchus frontalis Thinornis novaeseelandiae Scolopacidae Coenocorypha aucklandica Coenocorypha pusilla Coenocorypha chathamica Haematopodidae Haematopus unicolor Haematopus ostralegus Haematopus chathamensis Columbidae Hemiphaga novaeseelandiae Psittacidae Strigops habroptilus Nestor Nestor meridionalis Cyanoramphus Nestor notablis novaezelandiae Cyanoramphus unicolor Cyanoramphus auriceps Cuculidae Chrysococcyx lucidus Eudynamis taitensis Strigidae Sceloglaux albifacies Ninox novaeseelandiae Aegothelidae Megaegotheles novaezealandiae Alcedinidae Halcyon sancta Eopsaltriidae Petroica macrocephala Petroica australis Petroica traversi Monarchidae Rhipidura fuliginosa Sylviidae Bowdleria punctata Bowdleria rufescens Pachycephalidae Mohoua albicilla Mohoua ochrocephala Mohoua novaeseelandiae Acanthizidae Gerygone igata Gerygone albofronta Motacillidae Anthus novaeseelandiae Meliphagidae Notiomystis cincta Anthornis melanura Prosthemadera novaeseelandiae Zosteropidae Zosterops lateralis Turnagridae Turnagra capensis Corvidae Palaeocorax moriorum

west Pacific influence in the New Caledonian Fiji, Tonga and Samoa avifauna, with four genera, eighteen species and two subspecies linking New Caledonia exclu- The avifaunas of Fiji, Tonga and Samoa (Fig1: sively to other southwest Pacific islands 3) are listed in Table 3 The data are from Mayr The avifauna of Norfolk Island, which lies be- (1945) and Watling (1982) The terrestrial avifau- tween New Zealand and New Caledonia, has a na of these central Polynesian islands, like that number of southwest Pacific genera in common of New Zealand and New Caledonia, is depau- with New Caledonia It has links to New Zea- perate in avian families While species ende- land with the occurrence of the genera Hemi- mism is high, there are only six endemic genera phaga (Columbidae) and Nestor (Psittacidae) These are the Collared Lory (Phigys solitarius), The majority of species found on Norfolk Island Red-breasted Musk Parrot (Prosopeia tabuensis) are widespread As one would expect for such a and Kadavu Honeyeater (Xanthotis provocator) small island the endemic element is small, of Fiji and the Tooth-billed Pigeon (Didunculus Schodde et al (1983) listing only five endemic strigirostris) of Samoa An extinct species, Di- species (including two Zosterops spp) and nine dunculus sp nov has been reported from ‘Eua, endemic subspecies Tonga by Steadman (1995) The Silktail (Lam- 366 B Michaux

Table 2 Terrestrial birds of New Caledonia and the Loyalty Islands

Family Endemic Shared with Australia Shared with Australia SW Pacific +SW Pacific/ New Guinea/Indonesia

Podicipedidae Podiceps novaehollandiae Phalacrocoracidae Phalacrocorax melanoleucus Ardeidae Botaurus poiciloptilus Ardea novaehollandiae Butorides striatus Egretta sacra Nycticorax caledonicus Anatidae Anas rhynchotis Dendrocygna arcuata Anas superciliosa Anas gibberifrons Aythya australis Accipitridae Accipiter haplochrous Haliaster sphenurus Accipiter fasciatus Circus approximans Pandion haliaetus Falconidae Falco peregrinus Turnicidae Turnix varia Rallidae Tricholimnas lafresnayanus Gallirallus philippensis Porzana tabuensis Poliolimnas cinereus Porphyrio porphyrio Rhynochetidae Rhynochetos jubatus Columbidae Drepanoptila holosericea Chalcophaps indica Columba vitiensis Ducula goliath Ducula pacifica Ptilinopus greyii Psittacidae Vini diadema Trichoglossus haematodus Eunymphicus cornutus Cyanorhamphus novaezelandiae Cuculidae Cacomantis pyrrhophanus Eudynamis taitensis Chrysococcyx lucidus Tytonidae Tyto alba Tyto longimembris Aegothelidae Aegotheles savesi Caprimulgidae Eurostopodus mystacalis Apodidae Collocalia spodiopyga C s leucopygia Collocalia esculenta C e uropygalis Alcedinidae Halcyon sancta Hirundinidae Hirundo tahitica Campephagidae Coracina analis Coracina caledonica Lalage leucopyga Turdidae Turdus poliocephalus Sylviidae Megalurulus mariei Acanthizidae Gerygone flavolateralis Eopsaltriidae Eopsaltria flaviventris Monarchidae Rhipidura spilodera Myiagra caledonica Clytorhynchus pachycephalodes Pachycephalidae Pachycephala caledonica Pachycephala pectoralis Pachycephala rufiventris Artamidae Artamus leucorhynchus Sturnidae Aplonis striatus Corvidae Corvus moneduloides Meliphagidae Phylidonyris undulata Myzomela sanguinolenta Myzomela cardinalis Gymnomyza aubryana Lichmera incana Philemon diemenensis Gymnomyza Zosteropidae Zosterops minuta Zosterops lateralis Zosterops xanthochroa Zosterops inorta Estrildidae Erythrura psittacea Erythrura trichroa Terrestrial birds of the Indo-Pacific 367

Table 3 Terrestrial birds of Fiji, Tonga and Samoa (S) = Samoan endemics, (T) = Tongan endemics

Family Endemic Shared with Australia Fiji/Tonga/Samoa SW Pacific +SW Pacific/ New Guinea/Indonesia

Ardeidae Egretta sacra Butorides striatus Anatidae Dendrocygna arcuata Anas superciliosa Accipitridae Accipiter rufitorques Circus approximans Falconidae Falco peregrinus Megapodiidae Megapodius pritchardii (T) Rallidae Nesoclopeus poecilopterus Gallirallus philippensis Nesoclopeus Pareudiastes pacificus (S) Porzana tabuensis Pareudiastes Poliolimnas cinereus Porphyrio porphyrio Columbidae Ptilinopus luteovirens Ptilinopus perousii Ptilinopus porphyraceus Ptilinopus layardi Gallicolumba stairii Columba vitiensis Ptilinopus victor Ducula pacifica Ducula latrans Didunculus strigirostris (S) Psittacidae Charmosyna amabilis Vini australis Phigys solitarius Prosopeia tabuensis Prosopeia personata Cuculidae Cacomantis pyrrhophanus Eudynamis taitensis Tytonidae Tyto alba Tyto longimembris Apodidae Collocalia spodiopygia Alcedinidae Halcyon recurvirostris (S) Halcyon chloris Hirundinidae Hirundo tahitica Campephagidae Lalage sharpei (S) Lalage maculosa Turdidae Turdus poliocephalus Sylviidae Vitia ruficapilla Vitia Trichocichla rufa Eopsaltriidae Petroica multicolor Monarchidae Rhipidura personata Clytorhynchus vitiensis Rhipidura spilodera Rhipidura nebulosa (S) Mayrornis Mayrornis lessoni Clytorhynchus Mayrornis versicolor Clytorhynchus nigrogularis Myiagra albiventris (S) Myiagra azureocapilla Lamprolia victoriae Pachycephalidae Pachycephala flavifrons (S) Pachycephala pectoralis Artamidae Artamus leucorhynchus Sturnidae Aplonis atrifuscus (S) Aplonis tabuensis Meliphagidae Gymnomyza samoensis (S) Foulehaio carunculata Myzomela cardinalis Gymnomyza viridis Xanthotis provocator Myzomela jugularis Zosteropidae Zosterops samoensis (S) Zosterops lateralis Zosterops explorator Estrildidae Erythrura kleinschmidti Erythrura cyanovirens

prolia victoriae) of Fiji was placed by Mayr characteristic of the New Guinean avifauna dis- (1945) in ‘uncertain family’, although Sibley and cussed below The occurrence of megapodes Ahlquist (1990) regarded it as a monarch fly- (Megapodiidae: one endemic species in Tonga) catcher on DNA evidence The genus Foulehaio is also a characteristic of tropical Indo-Pacific is- (Meliphagidae) is endemic to Fiji and Samoa and lands was formerly found in Tonga (Steadman, 1995) No species is shared exclusively with Aus- Both the pigeon (Columbidae) and flycatcher tralia, but a number of widespread species (Aus- (Monarchidae) families are diverse in species, a tralia-southwest Pacific-New Guinea-Indonesia) 368 B Michaux

Table 4 Terrestrial birds of New Guinea that are either not found or are poorly represented in Australia X = presence; MP = Malay peninsula; WIs = western islands of New Guinea; CY = one species restricted to Cape York; numbers refer to the

number of species found in Australia; ? = unknown

N, C & SE New Guinea New SE & C N,

Bismarck Bismarck Archipelago

Andamans/Nicobars

Greater Sundas Greater

New New Caledonia

S New Guinea New S

Lesser Sundas Lesser

Africa/Eurasia

India/S China India/S

New New Zealand

Philippines

Micronesia

N Maluku N

Solomons

S Maluku S

Sulawesi

Australia

Vanuatu

Pacific SEA

Casuariidae Casuarius bennetti XX Podicipedidae Tachybaptus ruficollis XXX XXXXXXX XX Anatidae Dendrocygna guttata XX XXXXX Accipitridae Macheiramphus alcinus XXXX XX Circus spilonotus XXX X X XX Accipiter meyerianus XXX XX Falconidae Falco severus XX XXX XXXXX? Megapodiidae Megapodius XXXXXXXX XXXX 1 Rallidae Gymnocrex plumbeiventris XX X Gallirallus torquatus X X Sula X Columbidae Columba vitiensis XXXXXXXXXXX X Macropygia XXXXX XXXXXXX 1 M nigrirostris XXX M mackinlayi XXXX Reinwardtoena XXXXXX R reinwardtii XXX Chalcophaps stephani X XXXX Caloenas nicobarica XXXXXXXX X X Gallicolumba XX XX X XX G beccarii XXXX G jobiensis XXXX Ptilinopus MPXXXXXXXXXXXXXX 4 P wallacii XX P rivoli XXX X P solomonensis XXX P viridis XX X Ducula XXXXXXXXXXXXXXXX 1 D concinna XXX D pacifica XX D myristicivora XX D rufigaster X D pistrinaria XXX Gymnophaps XXXXXX Psittacidae Chalcopsitta XXX Eos XXX Trichoglossus Bali X X X X X X X X X X X 3 Lorius XXXXXX L hypoinochrous XX Charmosyna XXXXXXXX C rubrigularis XX C placentis XXXXXX Micropsitta XXXX Tanygnathus X X ? X X WIs T megalorynchos X X X WIs Alisterus amboinensis XXXX Loriculus XXXXXXX XXXX L aurantiifrons XXX Eclectus roratus XXXX XX CY Geoffroyus XXXXXXX CY Cuculidae Centropus XXXXXXX? XXXXXX 1 Strigidae Otus XXXXX? XXXXX Otus magicus XXXXX Terrestrial birds of the Indo-Pacific 369

Table 4 Continued

N, C & SE New Guinea New SE & C N,

Bismarck Bismarck Archipelago

Andamans/Nicobars

Greater Sundas Greater

New New Caledonia

S New Guinea New S

Lesser Sundas Lesser

Africa/Eurasia

India/S China India/S

New New Zealand

Philippines

Micronesia

N Maluku N

Solomons

S Maluku S

Sulawesi

Australia

Vanuatu

Pacific SEA

Hemiprocnidae Hemiprocne XXXXXX? XXXXXX Hemiprocne mystacea XXXXXX Apodidae Collocalia XXXXXX? XXXXXXXXXX 1 C esculenta XXX XXXXXXXX X Aerodramus vanikorensis X XXXX? XX XX CY A whiteheadi XXX Alcedinidae Tanysiptera XXXXX CY T galatea XXXX Halcyon saurophaga XX XX Ceyx lepidus X XXXXXX Alcedo XXXXXXXXXXXXXX Alcedo atthis XXXX X XXX XX M philippinus XXXXXXXX XXX Coraciidae Eurystomus orientalis XXXX? X? XXXXX X Bucerotidae Rhyticeros XXXXXXXXXXXXX R plicatus XXXXXX Pittidae Pitta erythrogaster XX XXXX ? P sordida XXXXX XX Hirundinidae Hirundo tahitica XXXX XXXXXXXXXX X Campephagidae Coracina morio XX X Lalage atrovirens XX Laniidae Lanius XXXXX X L schach XXXX X Turdidae Saxicola XXX XXXXX X X S caprata XXX XXXXX X X Turdus XXX XXXXX X XXXX X T poliocephalus XXXXXXXXXX Sylviidae Phylloscopus XXXXX X XXX XX P trivirgatus MPX XXXXX XX Monarchidae Monarcha cinerascens X XXXX XX Rhipidura XX XXX XXXXXXXXXXX3 Zosteropidae X X X X X X X X X X X X X X X X X X X 4 Nectariniidae Nectarinia sericea X XXXXX Sturnidae Aplonis XXXXXXXXXXXXXXXXX CY A cantoroides XXXX A mysolensis X XXX Mino dumontii XXXX Oriolidae Oriolus XXXXX XXXXXX 2 Dicruridae Chaetorhynchus X Dicrurus XXXXXXX? XXXXXX 1

form an important element of the central New Guinea Polynesian avifauna A southwest Pacific ele- ment forms a second important grouping Seven Bird life in New Guinea (Fig1: 4) is varied with species and one genus are endemic to Fiji + 708 species officially recorded (Beehler et al , Tonga + Samoa, while nine species and five 1986) Besides birds of paradise (Paradisaeidae: genera are endemic to the southwest Pacific 38 species) and bower birds (Ptilonorhynchi- Niue, which lies to the east of Tonga, shares six dae: 11 species), hawks and eagles (Accipitri- of these southwest Pacific endemics (Wodzicki, dae: 25 species), parrots (Psittacidae: 46 spe- 1971) cies), pigeons (Columbidae: 45 species), king- 370 B Michaux

Table 5 Terrestrial birds of Maluku S Maluku = Sula + Buru + Seram + Ambon + Kai + Timor; N Maluku = Morotai + Halmahera + Ternate + Bacan + Obi; * = S Maluku only; # = N Maluku only; (T) = Timor only;

Family Endemic Endemic Endemic S Maluku-S NG/Aust S Maluku N Maluku Maluku

Casuariidae Casuarius casuarius Podicipedidae

Phalacrocoracidae

Ardeidae Butorides striata moluccarum Ardea novaehollandiae Egretta intermedia

Threskiornithidae Threskiornis moluccus Anatidae Anas gibberifrons (T) Nettapus pulchellus Accipitridae Accipiter henicogrammus Accipiter erythrauchen Accipiter fasciatus

Falconidae Megapodiidae Megapodius wallacei Phasianidae Turnicidae Turnix maculosa Rallidae Habroptila wallacii Gallinula tenebrosa

Jacanidae Columbidae Ducula cineracea (T) Ducula basilica Ducula melanura Ducula concinna Gymnophaps mada Ptilinopus granulifrons Ducula perspicillata Ptilinopus regina Treron psittacea Ptilinopus hyogaster Ptilinopus wallacii Turacoena modesta (T) Ptilinopus monarcha Ptilinopus viridis Gallicolumba hoedtii Ptilinopus bernsteinii

Psittacidae Charmosyna toxopei Loriculus a amabilis Trichoglossus haematodus

Eos bornea Lorius garrulus Micropsitta keiensis Eos reticulata Cacatua alba Eos semilarvata Lorius domicella Cacatua moluccensis Cacatua goffini Prioniturus mada Tanygnathus gramineus Psitteuteles iris (T) Aprosmictus jonquillaceus(T) Cuculidae Centropus spilopterus Centropus goliath Centropus phasianinus Chrysococcyx crassirostris Cuculus heinrichi

Tytonidae Tyto sororcula Tyto novaehollandiae Strigidae Ninox novaeseelandiae

Aegothelidae Aegotheles crinifrons Caprimulgidae Apodidae

Hemiprocnidae Alcedinidae Halcyon lazuli Halcyon funebris Halcyon macleayii Halcyon diops

Meropidae Coraciidae Eurystomus azureus Bucerotidae Pittidae Pitta versicolor elegans Pitta maxima Campephagidae Coracina ceramensis Coracina parvula Coracina atriceps Coracina fortis Lalage aurea Coracina dispar Terrestrial birds of the Indo-Pacific 371

West = Sundaland/Asia; East = New Guinea/Australia/Pacific; Widespread = found either side of Wallace’s Line

N Maluku-NG West East Widespread

Tachybaptus novaehollandiae#

Tachybaptus ruficollis Phalacrocorax sulcirostris Phalacrocorax melanoleucos Ardea purpurea* Nycticorax caledonicus Ardea sumatrana Ixobrychus flavicollis australis Ardea alba modesta* Bubulcus ibis Egretta sacra

Dendrocygna a arcuata (T) Anas superciliosa Dendrocygna guttata* Tadorna radjah Aquila gurneyi Ictinaetus malayensis Accipiter meyerianus Haliaster indus Accipiter novaehollandiae Alanus caeruleus Aviceda subcristata Falco moluccensis Falco severus Megapodius freycinet Megapodus reinwardt Synoicus ypsilophorus (T)

Gymnocrex plumbeiventris Amaurornis phoenicurus* Amaurornis olivacus Gallirallus philippensis Rallina fasciata Rallina tricolor* Porphyrio porphyrio* Irediparra gallinacea* Ducula myristicivora Ducula rosacea Ptilinopus superbus Ducula bicolor Gymnophaps albertisii Ptilinopus cinctus Ptilinopus rivoli Caloenas nicobarica Ptinopus melanospila* Reinwardtoena reinwardtsii Chalcophaps indica Streptopelia bitorquata (T) Macropygia amboinensis Geopelia striata* Columba vitiensis Macropygia ruficeps (T) Macropygia magna* Treron pompadora* Eos squamata Eclectus roratus Tanygnathus megalorhynchos

Alisterus amboinensis Micropsitta bruynii* Charmosyna placentis Cacatua galerita* Geoffroyus geoffroyi

Centropus bengalensis Cuculus variolosus Eudynamis scolopacea Chrysococcyx crassirostris Scythrops novaehollandiae Tyto alba* Ninox connivens Otus magicus Ninox squamipila

Caprimulgus affinis * Caprimulgus macrurus Aerodramus fulciphagus (T) Collocalia esculenta Aerodramus infuscatus Aerodramus vanikorensis Hemiprocne mystacea Halcyon saurophaga Halcyon australasia* Ceyx lepidus Halcyon chloris Alcedo azurea Tanysiptera galatea Halcyon sancta Alcedo pusilla Alcedo atthis Merops superciliosus Eurystomas orientalis Rhyticeros plicatus Pitta erythrogaster Coracina novaehollandiae* Coracina tenuirostris Coracina papuensis Lalage atrovirens* Lalage leucomela 372 B Michaux

Table 5 Continued

Family Endemic S Maluku-S NG/Aust S Maluku N Maluku Maluku

Pycnonotidae Ixos affinis Turdidae Zoothera schistacea Zoothera dumasi Zoothera machiki Saxicola gutturalis (T) Timalidae Sylviidae Buettikoferela bivittata (T) Urosphena subulata

Acanthizidae Gerygone inornata (T) Muscicapidae Rhinomyias addita Ficedula buruensis Ficedula timorensis (T) Cyornis hyacinthina (T) Microecia hemixantha Monarchidae Rhipidura superflua Myiagra galeata Rhipidura dedemi Monarcha pileatus Rhipidura opistherythra Rhipidura fuscorufa Monarcha leucurus Monarcha mundus Monarcha loricatus Pachycephalidae Pacycephala orpheus (T) Pachycephala griseonata Pachycephala simplex

Dicaeidae Dicaeum vulneratum Dicaeum erythrothorax Dicaeum hirundinaceum

Nectariniidae

Zosteropidae Madanga ruficollis Zosterops atriceps Lophozosterops pinaiae Tephrozosterops stalkeri Zosterops buruensis Zosterops grayi Zosterops uropygialis Zosterops kuehni Heleia muelleri (T) Meliphagidae Lichmera monticola Philemon fuscicapillus Lichmera squamata Melitograis gilolensis Lichmera deningeri Lichmera flavicans (T) Lichmera notablis (T) Myzomela blassii Myzomela vulnerata (T) Philemon moluccensis Philemon subcorniculatus Philemon inornatus (T) Philemon citreogularis Meliphaga reticulata (T) Estrildidae Padda fuscata (T) Erythrura tricolor

Sturnidae Basilornis corythaix Aplonis crassa Oriolidae Oriolus bouroensis Oriolus phaeochromus Oriolus flavocinctus Oriolus forsteni Sphecotheres viridis Dicruridae

Artamidae Paradisaeidae Lycocorax pyrrhopterus Semioptera wallacei Corvidae Corvus validus Terrestrial birds of the Indo-Pacific 373

N Maluku-NG West East Widespread

Brachypteryx leucophyrs (T) Turdus poliocephalus* Zoothera peronii (T) Zoothera doherty (T) Zoothera andromedae (T) Pneopyga pusilla (T) Cettia vulcania (T) Megalurus timoriensis* Acrocephalus stentoreus* Orthotomus cuculatus Phylloscopus poliocephala Cisticola exilis* Bradypterus castaneus* Cisticola juncidis* Bradypterus seebohmi (T) Phylloscopus presbytes (T) Seicercus montis*

Eumyias panayensis Ficedula dumetoria* Ficedula hyperythra Ficedula westermanni*

Rhipidura rufiventris* Rhipidura leucophrys Rhipidura rufifrons* Myiagra ruficollis* Monarcha cinerascens* Monarcha trivirgatus Piezorhynchus alecto Pachycephala phaionotus Pachycephala pectoralis Pachycephala rufiventris Pacycephala leucogastra Dicaeum agile (T) Dicaeum maugi* Dicaeum sanguinolentum (T) Nectarinia solaris (T) Nectarinia jugularis

Zosterops montanus Zosterops atrifrons* Zosterops chloris

Lichmera argentauris Lichmera indistincta (T) Myzomela sanguinolenta Myzomela obscura# Philemon buceroides (T)

Lonchura quinticolor (T) Erythrura trichroa* Lonchura molucca Poephila guttata Lonchura punctulata* Amandava amandava Aplonis metallica Aplonis mysolensis

Dicrurus densus Dicrurus bracteatus Dicrurus hottentottus Artamus cinereus* Artamus leucorhynchus

Corvus macrorhynchos (T) Corvus orru Corvus enca* 374 B Michaux fishers (Alcedinidae: 22 species), cuckoos (Cuc- classed as north Maluku The island of Obi has ulidae: 21 species), Australian warblers (Acan- an uncertain biogeographical status (Duffels and thizidae: 20 species), flowerpeckers (Dicaeidae: Boer, 1990), but for the present is regarded as 10 species), whistlers (Pachycephalidae: 26 spe- part of north Maluku Southern Maluku is com- cies), flycatchers (Monarchidae: 30 species) and posed of the islands of Buru, Seram, Ambon and honeyeaters (Meliphagidae: 65 species) are all Kai White and Bruce (1986) argued for the in- highly diversified In addition to the Ptilo- clusion of Tanimbar in southern Maluku and rhynchidae there are six other endemic New Michaux (1994) for the inclusion of Timor Guinea/Australian bird families (Casuariidae, The data in Table 5 are arranged to demon- Maluridae, Climacteridae, Grallinidae, Cractici- strate several distributional patterns These pat- dae and Orthonychidae) Notable absences from terns involve an endemic element (endemic), the New Guinean avifauna are Old World fly- taxa distributed between south Maluku and con- catchers (Muscicapidae), trogons (Trogonidae), tinental Australia (S Maluku-S NG/Aust) and taxa barbets (Capitonidae), woodpeckers (Picidae), that link north Maluku to New Guinea (N bulbuls (Pycnonotidae) and broadbills (Eury- Maluku-NG) Many birds found in Maluku are laimidae) widely distributed Some of these are Asian or Although New Guinea’s avifauna shows Sundaic (West in Table 5), some Australasian strong similarity to that of Australia, particularly (East), while still others are found both east and to northern and northeastern regions, there are west of Maluku (Widespread) New Guinean taxa that are not found in Austral- White and Bruce (1986) discussed endemism ia or are poorly represented and often restricted in Maluku and noted that while there are few to Cape York Table 4 lists species from many endemic genera, species endemism is high Ta- bird families that are present in New Guinea and ble 5 lists 72 endemic species in south Maluku areas to the east and west, but absent from Aus- with three endemic, monotypic genera — tralia Many of these species are widespread Tephrozosterops, Madanga (Zosteropidae) and Examples of these taxa include a hornbill (Buce- Buettikoferela (Sylviidae) In north Maluku there rotidae) the genera Caloenas, Reinwardtoena, are 26 endemic species or subspecies and two Gallicolumba (Columbidae), Chalcopsitta, Eos, endemic genera — Lycocorax and Semioptera Lorius, Charmosyna, Tanygnathus, Micropsitta, (Paradisaeidae) Table 5 also lists eleven species Loriculus (Psittacidae), Otus (Strigidae), Hemi- endemic to Maluku as a whole procne (Hemiprocnidae), Alcedo (Alcedinidae), North and south Maluku have different rela- Lanius (Laniidae), Saxicola, Turdus (Turdidae) tionships with surrounding areas Wallace and Phylloscopus (Sylviidae) Other examples in (1880) noted that the fauna of north Maluku which there is a single Australian species, often shows many similarities with New Guinea, while restricted to Cape York or the Queensland the fauna of south Maluku is closer to that of coast, include Megapodius (Megapodiidae), continental Australia (including southern New Macropygia, Ducula (Columbidae), Geoffroyus Guinea) White and Bruce (1986) confirmed that (Psittacidae), Centropus (Cuculidae), Collocalia the avifauna of north Maluku has a New (Apodidae), Tanysiptera (Alcedinidae), Oriolus Guinean element, and listed 23 species they re- (Oriolidae), Merops (Meropidae), Aplonis (Stur- garded as New Guinean (White and Bruce, nidae), Nectarina (Nectariniidae) and Dicrurus 1986: 44) Apart from the nine species listed as N (Dicruridae) Maluku-NG in Table 5, a number of north Maluku endemics can also be classed as New Guinean These include Habroptila wallacii Maluku (Rallidae), Centropus goliath (Cuculidae), Hal- cyon diops (Alcedinidae), as well as various taxa The avifauna of Maluku (Fig1: 5), listed in Table from the families Columbidae, Psittacidae, 5, is based on the distributional data in Bemmel Meliphagidae and Paradisaeidae (1948), Bemmel and Voous (1953), Peters (1934- The south Maluku avifauna shows a relation- 86) and White and Bruce (1986) Distributional ship to cratonic areas of southern New Guinea data were cross-referenced with MacKinnon and and Australia (Table 5: S Maluku-S NG/Aust and Phillipps (1993) and Beehler et al (1986) entries marked * under East) The families Acan- Wallace (1880) recognised that the avifauna of thizidae, Meliphagidae, Monarchidae and Ae- the northern Maluku islands was distinct from gothelidae and the genera Gymnophaps, Ducu- that of the southern islands In this paper la, Gallicolumba (Columbidae), Charmosyna, Ternate, Halmahera, Morotai, and Bacan are Eos and Lorius (Psittacidae) illustrate this pat- Terrestrial birds of the Indo-Pacific 375 tern A second pattern in the south Maluku dis- genetically isolated, but also geographically iso- tributions links south Maluku to Sundaland or lated The island of Borneo, separated from Su- Asia (Table 5: West) Examples illustrating this lawesi by the Makassar Strait, is only 105 km pattern include the families Timaliidae, Sylvii- distant (reduced to 40 km during the Pleis- dae, Pycnonotidae and Muscicapidae and the tocene), yet many Bornean species are absent genera Treron (Columbidae), Brachypteryx, White and Bruce (1986) discussed the differenc- Saxicola, Zoothera (Turdidae) and Oriolus (Ori- es in avifauna between Borneo and Sulawesi olidae) They provided the following list showing the reduction between Borneo and Sulawesi (spe- cies numbers in brackets):Phasianidae (12), Sulawesi Strigidae (8), Podargidae (6), Trogonidae (6), Bucerotidae (8), Capitonidae (9), Picidae (17), A list of non-endemic birds of Sulawesi (Fig1: 6) Eurylaimidae (8), Aegithinidae (6), Pycnonoti- is provided in Table 6 The distributional data dae (23), Timaliidae (36), and Arachnothera (7) are from Walters (1980), White and Bruce (1986) The families Podargidae, Trogonidae, Capitoni- and MacKinnon and Phillipps (1993) Thirty six dae, Eurylaimidae, Aegithinidae and Pycnonoti- of 140 species can be classed as Asian Twenty dae are absent from Sulawesi five of these species extend from mainland Asia to Sulawesi, which is at the eastern extremity of their ranges A further eleven species extend to Geological histories and local avifaunas the Sula Islands and/or the Lesser Sundas Twenty five species are Australasian, with The New Zealand subcontinent Sulawesi at the western extremity of their ranges A further 26 species are endemic to Fig1 is a simplified version of the Plate-Tectonic Wallacea and the remaining 53 species cross Map of the Circum-Pacific Region (Southwest Wallace’s line Some 60% of the species listed in Quadrant) by AAPG (1981) Fig1 shows an ex- Table 6 are also found in the Philippines Other tensive area of predominantly submerged conti- areas that share species with Sulawesi include nental crust off the east coast of Australia (Fig1: the Greater Sunda islands (particularly Java, LHR + NR + nI + sI + CR + CP) Kamp (1986) western Sumatra and north Borneo), the Lesser referred to this area as the New Zealand subcon- Sundas and south Maluku tinent The New Zealand subcontinent is sepa- White and Bruce (1986) discussed the high rated from Australia by the Tasman and Coral degree of generic and specific endemism shown Seas (Fig1: TS and CS), which are underlain by by the Sulawesian avifauna Twelve of the en- oceanic crust Fig1 shows that the New Zealand demic genera are monotypic: Macrocephalon subcontinent has been disrupted in the north by (Megapodiidae), Aramidopsis (Rallidae), Cryp- the New Caledonian basin (Fig1: NC) Accord- tophaps (Columbidae), Cittura (Alcedinidae), ing to Wood and Uruski (1990) this basin repre- Meropogon (Meropidae), Cataponera, Hein- sents a failed rift underlain by thinned continen- richia (Turdidae), Malia, Geomalia (Timalii- tal crust dae?), Coracornis, Hylocitrea (Pachycephali- A series of small basins, composed of oceanic dae), Enodes and Scissirostrum (Sturnidae) crust, are shown in Fig1 (SFB, NFB, SCB, SS and There are two species belonging to the endemic MB) A summary of basin ages is given in Fig2 genus Myza (Meliphagidae) In addition, the The ages given in Fig2 are from Yan and genera Prioniturus, Tanygnathus (Psittacidae), Kroenke (1993) There are discrepancies in Streptocitta and Basilornis (Sturnidae) are re- dates quoted in Yan and Kroenke (1993) and stricted to Sulawesi + Philippines + S Maluku, AAPG (1981) When dates differ those from the genus Turacoena (Columbidae) to Sulawesi AAPG (1981) are indicated by horizontal bars in and Timor and the genus Penelopides (Bucerot- Fig2 A series of archipelagos and island groups idae) to Sulawesi and the Philippines are found on the Pacific side of these basins The number of endemic species in Sulawesi is (Fig1: T, S, F, V and So) Vanuatu (Fig1: V), the not known for certain Stresemann (reported in Solomons (Fig1: So) and Tonga (Fig1: T) are White and Bruce, 1986) concluded that 84 out of island arcs situated above subduction zones 220 species were endemic (31%), but White and These archipelagos and island groups straddle Bruce (1986) suggested that this figure was too the present boundary between the Pacific and high Walters (1980) listed 80 Sulawesian en- Indo-Australian plates The relationship be- demics The Sulawesian fauna is not only phylo- tween oceanic basins, the New Zealand subcon- 376 B Michaux

Table 6 Non-endemic terrestrial birds of Sulawesi X = present; X* = present in S Maluku; [X] = not present in Malay peninsula Greater Sundas Greater Lesser Sundas Lesser New Zealand New Guinea Philippines Australia S China S Maluku Eurasia Pacific Africa India SEA

Phalacrocoracidae Phalacrocorax melanoleucos ? XXX P sulcirostris X XXXX X Anhinga melanogaster X [X] X X Ardeidae Hydranassa picata ?X Ardea purpurea XXXXXXX X* A novaehollandia XXXXXX A speciosa [X] X X Egretta intermedia XXXXXX XXXXX Ixobrychus cinnamomeus XXX XXXX* Nycticorax caledonicus XXXXXXXX Ciconidae Ciconia episcopus X X [X] X X Mycteria cinerea [X] X Threskionithidae Plegadis perigrinus XX Platalea regia XX ? X Anatidae Anas gibberifrons XXX*XX?X A superciliosa X XXXXXX Dendrocygna guttata XX D arcuata XX X*XXX Netapus pulchellus XX*XX Accipitridae Aviceda jerdoni XX[X]XX Sula Elanus caeruleus XXX X XXX*X Macheiramphus alcinus XXXX Ichthyophaga humilis X XXX X* Haliaeetus leucogaster XX XXXX Butaster liventer X [X] X X* Ictineatus malayensis XXXX X Hieraatus kienerii X [X] X Circus assimilis X Pernis celebensis X Falconidae Falco moluccensis XXX F severus XXXXX Megapodiidae Megapodius cumingii XX Turnicidae Turnix suscitator XXXXXX Rallidae Gallinula chloropus X XXX G tenebrosa XXX*XX Gallirallus striatus XXXXX G torquatus X Sula X Rallina eurizonoides XX[X]XX Sula Porzana fusca XXXXXX P cinerea XXXXXXXX Gallicrex cinerea XXXXX Amaurornis phoenicurus XXXXX Fulica atra XXX[X]XX XXX Porphyrio porphyrio X X X XXXXXX Jacanidae Irediparra gallinacea XXXX*XX Charadriidae Charadrius peronii XXXX Burhinidae Burhinus giganteus [X] X X X X Columbidae Treron griseicauda X Sula T vernans XXXX Ptilinopus superbus XXX P melanospila XXXX* Ducula aenea X XXXXSula D pickeringi XX D luctuosa Sula D bicolor X X XXX Chalcophaps stephani X* X C indica X XX XXXX Macropygia amboinensis XXX M magna XX* Columba vitiensis XXXXX X Psittacidae Cacatua sulphurea X Tanygnathus megalorynchos XX*X T sumatranus XX Cuculidae Scythrops novaehollandiae X* X X Clamator coromandus X [X] X X C russatus XXX XX Surniculus lugubris XX XX X# Centropus bengalensis XXXXXXX Cuculus sepulcralis [X] X X X X* C merulinus XX XX Terrestrial birds of the Indo-Pacific 377

Table 6 Continued Greater Sundas Greater Lesser Sundas Lesser New Zealand New Guinea Philippines Australia S China S Maluku Eurasia Pacific Africa India SEA

Strigidae Ninox scutulata X XXXXX* Tyto capensis XXXXXX X XXX Caprimulgidae Eurostopodus macrotis XXXXX Caprimulgus affinis X [X] X X X X* C macrurus X XX XXXX Apodidae Collocalia salangana X C esculenta XXXXXX Aerodramus vanikorensis XX X Hirundapus giganteus X XXX H celebensis X Hemiprocnidae Hemiprocne longipennis XXX Sula Alcedinidae Alcedo meninting XXXXX Sula A atthis XXXXXXX XXX H coromanda XXXXX Sula Meropidae Merops philippinus ? [X] X X Coraciidae Eurystomus orientalis XXXXXXX X Pittidae Pitta erythrogaster XXXX P sordida XX XX X Campephagidae Lalage sueurii XXXX L nigra XXX Coracina tenuirostris XXXXX C morio XX*X Dicruridae Dicrurus bracteatus XXXX Oriolidae Oriolus chinensis XXXXXXSula Corvidae Corvus enca XXX X* C macrorhynchos XXXXXXXX Turdidae Saxicola caprata XXX[X]XXXXX Turdus poliocephalus XX X*X X T obscurus X Sylviidae Cisticola juncidis XXXXXXXXXXX C exilis XX[X]XXXXXX Culicicapa helianthea X Bradypterus castaneus X* Orthotomus sepium XX O cuculatus XXXX XX* Megalurus timorensis XX*XX Acanthizidae Gerygone sulphurea [X] X X X Monarchidae Culicicapa helianthea X Hypothymis azurea XXXXXX Muscicapidae Ficedula hyperythra XXXXXXX F westermanni XXXXXXX* Cyornis rufigastra XXX Eumyias panayensis XX* Muscicapa basilanica X M griseisticta XXXX XX Motacillidae Anthus novaeseelandiae X XXXXX XX X Pachycephalidae Colluricincla megarhyncha XX Sturnidae Aplonis minor XXX A mysolensis XX A panayensis X XXX Acridotheres javanicus ? X [X] X A fuscus XXX Nectariniidae Anthreptes malacensis XXX Sula Nectarinia sperata X XXX N jugularis X XXX X N aspasia X* X Aethopyga siparaja XXXXX Meliphagidae Myzomela sanguinolenta X* X X X Zosteropidae Zosterops montanus XXXX Z atrifrons X* X Z chloris XXX*X Ploceidae Erythrura trichroa XXXX E hyperythra XXXX Lonchura molucca XX L punctulata X XXXXXX L malacca XXXXX X# L pallida X Fringillidae Serinus estherae XX 378 B Michaux

struction of Yan and Kroenke (1993) this block started separating from Antarctica at 85 Ma (old- est magnetic anomaly 72 Ma) and moved north paralleling Pacific plate movement It united with the rest of the New Zealand subcontinent (Fig1: LHR + NR + nI) in the Oligocene (40 Ma) Where the intervening ocean between this block and the rest of the New Zealand subconti- nent was subducted, what (if any) movement occurred along the Campbell fault, and how the collision between the two blocks relates to the on-shore geology of New Zealand are not clear from Yan and Kroenke’s (1993) account According to Yan and Kroenke (1993) the Lord Howe Rise-Challenger Plateau-Norfolk Ridge-North Island, New Zealand block sepa- rated from the Australian margin about 85 Ma (oldest magnetic anomaly 78 Ma) and moved away in a northeasterly direction This implies that the boundary between the Indo-Australian and Pacific plates at the time was a west-dipping subduction zone east of the New Caledonian Ridge The Lord Howe Rise and New Caledo- nian basin quickly submerged to abyssal depths following thermal relaxation (Wood and Uruski, Fig 2 Age of marginal oceanic basins in the southwest Pa- 1990) New Caledonia and parts of New Zealand cific A = formation of South Island, New Zealand-Campbell are emergent segments of the New Caledonian Plateau-Chatham Rise block B = formation of the Rise today It is probable, given the tectonic ac- Melanesian rift system C = fragmentation of the Melanesian rift into a rift and arc system D = fragmentation of the tivity described above, that parts of this ridge Melanesian arc, Fiji moves away from Vanuatu have been emergent since the beginning of the Tertiary The New Caledonian Ridge is identified as the Melanesian rift in this paper The Melanesian rift is equivalent to the inner tinent and these outer islands is critical in the Melanesian arc of others, but this name is inap- interpretation of Pacific biogeography propriate (Polhemus, 1996) because the Norfolk The oldest basins are the Tasman Sea and Ridge may not be the remnant of an island arc, southwest Pacific basin (Fig1: TS and PO and but a rifted continental block Fig2) There are many published reconstruc- The Solomons, Vanuatu, Fiji, Tonga and Sa- tions of the New Zealand subcontinent prior to moa form the Melanesian arc The history of this their opening (Kamp, 1986) While all models arc system is complicated and poorly resolved show the New Zealand subcontinent adjacent to The arc may represent a single structure that has the Australian/Antarctic section of east been disrupted (Burrett et al, 1991), a multiple Gondwana at 100 Ma, there are differences in arc system presently juxtaposed by tectonic dy- the details of various reconstructions Some re- namics (Polhemus, 1996), or an arc system with constructions place the Chatham Rise and embedded microcontinents in Fiji With our Campbell Plateau (Fig1: CR and CP) adjacent to present state of knowledge such detail is sec- either the North Island, New Zealand (Fig 1: nI), ondary to knowing where the arc or arcs were or to both North and South Islands (Fig1: nI and formed There are two schools of thought on sI) (Borg and Depado, 1991; Kamp, 1986) In the origin of the Melanesian arc, both of which these reconstructions the Chatham Rise and have different implications for Pacific biogeog- Campbell blocks have always been adjacent to raphy the rest of the New Zealand subcontinent One hypothesis is that the Melanesian arc(s) In other reconstructions (Yan and Kroenke, was formed within the Pacific plate far from any 1993) the Chatham Rise-Campbell Plateau-South land This implies that the Melanesian arc biota Island block (Fig1: CR + CP + sI) was derived was derived by over-water colonisation (e g, from Marie Byrd Land (Antarctica) In the recon- Boer and Duffels, 1995) A second hypothesis is Terrestrial birds of the Indo-Pacific 379 that the Melanesian arc (or parts thereof) was be less favourable for a family generally found formed along the eastern edge of the in more equitable climes Melanesian rift between 50 and 40 Ma (Hall, Five other species within the genus are recog- 1998 this volume) The present position of this nised The Yellow-crowned parakeet (C arc system is seen as the result of opening the auriceps) is distributed throughout mainland marginal basins discussed above, which trans- New Zealand and a number of off-shore islands, ported the arc to the east The biogeographical although its numbers on the mainland had al- implication of this is that the Melanesian arc ready sharply declined by the turn of the cen- biota is original and derived from that of the tury (Hutton and Drummond, 1904) A subspe- Melanesian rift The composite rift-arc structure cies, forbesi, is recognised on Chatham, Pitt and is referred to here as the Melanesian rift-arc Mangere islands The Orange-fronted parakeet (C malherbi) is found in a few sites in the South Island, while C unicolor is restricted to the An- Implications for southwest Pacific biogeography tipodes Islands Two species, now extinct, were known from the Society Islands C ulietanus Craw (1988) interpreted New Zealand biogeog- was found on Raiatea and C zealandicus on raphy in terms of a “parallel arcs model” In this Tahiti (Fuller, 1987) model the western ‘arc’ (and its associated Three other parrot species are found in New biota) was related to cratonic Gondwana frag- Zealand, the kea (Nestor notabilis), kaka (Nestor ments and the eastern ‘arc’ to what were, prior meridionalis), and the kakapo (Strigops habrop- to the Rangitata orogeny in the early Cretaceous, tilus) The kea is restricted to the mountains of geosynclinal sediments This model generates a the South Island, while kaka subspecies were number of systematic hypotheses The biota as- once widespread An extinct subspecies of the sociated with cratonic Gondwana fragments kaka was found on Norfolk Island and nearby should show relationships to other cratonic ar- Phillip Island (productus) The kakapo is en- eas of Gondwana such as Australia and South demic to New Zealand where its continued America The relationship of the biota associ- long-term survival depends on populations es- ated with what I have termed the Melanesian tablished on predator-free islands Although its rift-arc system (equivalent to Craw’s (1988) east- relationship to other parrots is obscure, some ern arc) should show closer relationships to workers suggest that the closest relative of the other rift and arc fragments to the north and kakapo may be found among the Australian en- northeast The lack of appropriate systematic demics Pezoporus wallicus (ground parrot), P data makes an unequivocal test of such hypoth- occidentalis (night parrot) and Melopsittacus eses impossible at present, but the data pre- undulatus (budgerigar) (MacDonald and Slater, sented below are suggestive 1992) The New Zealand parrot fauna does not Although the New Zealand avifauna can be appear to be particularly Australian with respect generally characterised as depauperate, a to phylogenetic affinity Cyanoramphus no- number of families such as the Psittacidae and vaezelandiae is endemic to the New Zealand Phalacrocoracidae are diverse The genus Cy- subcontinent The genus Cyanoramphus links anoramphus (Psittacidae) is a southwest Pacific the New Zealand subcontinent with eastern endemic (Table 1) and well represented in the Polynesia The genus Nestor is endemic to areas New Zealand region The red-crowned parakeet forming the southern parts of the Melanesian rift (C novaezelandiae) is found from the Three and, with Cyanoramphus, forms part of what is Kings Islands in the north to the Auckland Is- termed here as New Zealand’s northern biota lands in the south It is now rare on the New The kakapo’s relationship to other parrots Zealand mainland, but was once common needs clarification throughout the country A number of subspe- One bird family that does show clear austral cies are recognised which are found in the Ker- relationships is the Phalacrocoracidae New Zea- madecs (cyanurus), Chathams (chathamensis), land’s cormorant fauna is very diverse with a Antipodes (hochstetteri), Macquarie (erythrotis greater number of species (14) being found in extinct), Lord Howe (subflavescens extinct), New Zealand waters than anywhere else in the Norfolk Island (cooki) and New Caledonia (sa- world (Falla et al , 1979) Three groups of cor- isetti) The distribution of this species is remark- morants are recognised on the basis of leg col- able, because it is found further south than any our and crest characteristics The first group of 4 other parrot and on Southern Ocean islands species (Phalacrocorax carbo, P varius, P where environmental conditions could hardly sulcirostris, and P melanoleucos) are character- 380 B Michaux ised by black legs and feet A crest may or may not be present They are fresh water, estuarine or coastal birds that nest in trees The four Phalacrocorax species are also found in Sumatra, southern Borneo, Java, eastern Indone- sia, New Guinea, Australia, and New Caledonia The second group have pink legs and feet and a single crest They are marine or oceanic species that nest on ledges This group are placed in the genus Leucocarbo by Turbott (1990) and are found around the coasts of New Zealand and its subantarctic islands The genus is also found in South America and Kerguelen Island The third group have yellow legs and feet and are double Fig 3 Strict consensus cladogram of five minimal length crested These species are found around New trees (length = 25, CI = 088, RI = 089) for New Zealand Zealand, Stewart Island and the Chathams They cormorants See text for details are placed in the genus Stictocarbo S gaimardi is a South American representative of the dou- ble-crested New Zealand species A cladistic analysis of some external and eco- and a sister group to Leucocarbo + Stictocarbo logical characters was carried out using PAUP The New Zealand cormorant fauna is a mixture (Swofford, 1991) The character matrix is shown of ‘northern’ and ‘southern’ groups The former in Table 7 The branch and bound option was are associated with the Melanesian rift-arc sys- used and multistate characters were treated as tem, while the latter are associated with the polymorphisms Five minimal length trees were Campbell plateau-Chatham Rise-South Island, found (length 25, CI = 088, RI = 089) Fig3 New Zealand block Although the above analy- shows the strict consensus tree rooted using sis was intended only as an interest exercise, the Anhinga as an outgroup (Sibley and Ahlquist, phylogenetic pattern within New Zealand cor- 1990) In Fig3 Phalacrocorax is monophyletic morants is congruent with geological patterns The high degree of endemism of the New Zealand avifauna suggests that post-rifting isola- Table 7 Characters and character-state matrix of New Zea- tion has been an important factor in its evolu- land cormorants tion The fossil record shows influxes of north- ern marine invertebrates and plants into New Zealand during the upper Eocene and in the ABCDEFGHI Phalacrocorax carbo 111100211 earliest Miocene (Fleming, 1980) Michaux P varius 102100211 (1989) interpreted the earlier event as a re- P sulcirostris 103100211 sponse to subsidence along the Norfolk Ridge P melanoleucos 10,11100121 The Miocene event coincided with the final Leucocarbo carunculatus 204211112 L chalconotus/onslowi 214211112 stages of the opening of the South Fiji basin and L campbelli 214200132 emplacement of the Northland allochthon L ranfurlyi/colensoi 214201132 Sibley and Ahlquist (1990) suggested separation L atriceps 215311112 of the New Zealand Wrens (Acanthisittidae) Stictocarbo punctatus/steadi 326400132 from their ancestor at 42 Ma and kiwi from an S featherstoni 327?00132 Emu-Cassowary clade at 45 Ma Both these Anhinga 201000101 avian families have Papuan phylogenetic affini- ties and their origin coincides with the Eocene A leg colour: 1 = black 2 = pink 3 = yellow/orange event described above Endemic elements of the B crest: 0 = absent 1 = one crest 2 = double crest New Zealand avifauna may date from 65 Ma C facial skin: 1 = yellow 2 = blue 3 = olive 4 = red (initial isolation after rifting), 40 Ma (Eocene in- 5 = brown 6 = green 7 = purple flux) and 22 Ma (Miocene event) D gular pouch: 1 = yellow 2 = red 3 = brown 4 = blue The avifaunas of New Caledonia, Norfolk Is- E caruncles: 0 = absent 1 = present land, Fiji/Tonga/Samoa and Niue (Tables 2 and F alar bar: 0 = absent 1 = present 3) show a strong Melanesian arc influence Bal- G iris colour: 1 = brown 2 = green lance et al (1982) suggested that the Melanesian H eye ring: 1 = blue 2 = brown 3 = purple arc separated from the Melanesian rift in the I habitat: 1 = fresh water 2 = marine New Caledonia-Tongan region prior to the Terrestrial birds of the Indo-Pacific 381

Fig 4 New Guinea terranes Dots = Melanesian arc terranes, stars = terranes from the Australian craton, slanted line = Melanesian rift terranes B = Bowutu terrane, F = Finisterre terrane, J = Jimi, Schrader, Benabena and Marum terranes, K = Kemum terrane, O = Owen Stanley terrane, R = Rouffaer terrane, S = Sepik terrane, Wa = Wandamen terrane Stipple = continental crust, arrows = opening basins, toothed lines = subduction zones Closed teeth represent subduction zones marking plate boundaries and open teeth intra-plate subduction zones

opening of the South Fiji basin at 35 Ma Accord- nated by shear (the ‘bacon slicer’ effect) with ing to Burrett et al (1991) the Melanesian arc fragments entering the shear zone transported was still a relatively coherent entity until 7 Ma, westwards (Fig4: filled circles) To the west of when the southern section (Vanuatu, the Fiji New Guinea, Melanesian arc fragments (or its Platform and the Tonga Ridge) became isolated lateral equivalents) appear to have been incor- from the Solomons The opening of the North porated onto the Philippine plate and rotated Fiji basin subsequently separated the Fiji Plat- clockwise to amalgamate with pre-existing form-Tongan Ridge-Samoa block from Vanuatu terranes in north Maluku and the southern Phil- (Fig2D) Tentative ages from which New Cale- ippines (Hall, 1987; 1996) donian endemism date are 65 Ma (initial separa- Hall et al ’s (1995: Fig9) reconstruction of the tion from Gondwana) and 35 Ma (separation of Philippine Sea plate showed it to the north of the Melanesian arc from the Melanesian ridge) Australia at 45 Ma and rotating clockwise By 25 and for Fiji/Samoa/Tonga from 35 Ma and 7 Ma Ma this rotation had brought the southern boundary of this plate in contact with northern New Guinea, at a time when Pigram and Davies New Guinea and the Melanesian rift-arc system (1987) and Lee and Lawver (1995) suggested that exotic terranes had started to collide with An interpretive summary of Pigram and Davies’ the Australian craton According to Hall et al (1987) account of the collision and amalgama- (1995) clockwise rotation continued until 5 Ma tion of a variety of terranes with the northern by which time the eastern boundary was to the margin of the Australian craton is shown in northwest of New Guinea Rangin et al (1990) Fig4 In this paper these terranes are interpreted have described kinematic motions that are con- as lateral extensions of the Melanesian rift-arc sistent with this model Lee and Lawver (1995) system The interaction at this margin was domi- placed the Sepik arc terrane (= Melanesian rift) 382 B Michaux to the northeast of Australia at 50 Ma General Ma The relationship between these cratonic westward translation of this terrane resulted in terranes of the Bird’s Head, their lateral equiva- collision and amalgamation with the Australian lents in the central ranges interpreted here as craton at 30 Ma Northern New Guinea terranes, parts of the Melanesian rift (Fig4: diagonal which Lee and Lawver (1995) link to the lines), and outer Banda rift-arc fragments (dis- Bismarcks and Solomon Islands (= Melanesian cussed later) is unclear arc), also entered the collision zone from the A number of general points emerge from a east to amalgamate between 10 Ma and 5 Ma study of the New Guinean avifauna Firstly, Clockwise rotation of the Philippine Sea plate many of New Guinean taxa are endemic Sec- has been an important factor in the develop- ondly, the similarity to Australia is not as marked ment of the New Guinea margin as one might expect, with many taxa in central In Fig4 the terranes that Pigram and Davies and northern New Guinea showing phylogenet- (1987) identified along the north coast have ic links to the east and west, rather than to Aus- been interpreted as Melanesian arc islands, now tralia in the south The majority of New Guinean incorporated as coastal ranges along the New taxa with phylogenetic links to Australia are Guinea margin While each may have had a found in southern New Guinea Within central unique geological history, they are viewed here and northern New Guinea taxonomic differenti- as members of a tectonic unit that docked after ation is strongly developed along an east-west 10 Ma New Britain and New Ireland are adja- axis cent parts of the Melanesian arc that are cur- Table 4, which lists New Guinean taxa absent rently sandwiched between the Manus basin, from Australia, shows that northern and central which is opening in a west-east direction, and a New Guinea are linked to the southwest Pacific subduction zone along the northern boundary via the Melanesian arc islands of the Bismarck of the Solomon Sea (Fig4) archipelago and the Solomons Northern New The amalgamation of the northern section of Guinean taxa also have phylogenetic links to the Melanesian rift with the Australian craton is north Maluku and the Philippines Asian influ- dated by Pigram and Davies (1987) from 25 Ma ences are strong in central New Guinea New in the west to 15 Ma in the east By 15 Ma the Guinean montane flora shows a pronounced east Papuan composite terrane had docked mixture of Asian and Austral taxa, being the forming southeast New Guinea The east only place in the world where southern beeches Papuan composite terrane formed somewhere and northern oaks exist together Asian Diptero- to the northeast of the Australian craton prior to carpaceae dominate hill forest while Araucaria its docking In Fig4 this terrane is shown com- is found in lower montane zones with oaks at posed of both rift and arc fragments (Michaux, higher levels and Nothofagus above 2000 m 1994) The Louisiade plateau (Fig4: star) and a The higher moss forests are dominated by the section of the southern coast around Port Myrtaceae and at higher levels still by Eurasian Moresby are parts, according to Pigram and genera such as Vaccinium and Rhododendron Davies (1987), of the Australian craton moved (Ericaceae) (Gressitt, 1982a) Similar patterns into their present position by the opening of the occur in other groups Four anuran families are Coral Sea (63-53 Ma) found in New Guinea (Menzies, 1975) The Lep- In the Bird’s Head region, terranes have been todactylidae and Hylidae are shared with Aus- identified by Pigram and Davies (1987) as frag- tralia and South America (Cracraft, 1980) The ments of the Australian craton (Fig4: star) The Ranidae and Microhylidae are characteristic of largest of these terranes is the Kemum terrane Indonesia and Asia Gressitt has discussed the which makes up most of the Bird’s Head south ‘Oriental’ character of New Guinea’s rich and di- of the Sorong fault It is composed of Palaeozoic verse beetle fauna in which similar patterns are basement with only a thin, incomplete Mesozoic also evident (Gressitt, 1982a, b) sequence Pigram and Davies (1987) reported An examination of birds shared by New that polar wandering curves for the Kemum Guinea and Australia shows that southern New terrane are identical to those of Australia until Guinea and the Port Moresby area in the south- their separation in the Cretaceous Stratigraphic east peninsula feature in 42 of 121 of such distri- and palaeontological evidence also indicate an butions (MacDonald and Slater, 1992) The ma- Australian origin for the Kemum terrane During jority of these species are endemic to Australia the Oligocene to Miocene the Kemum terrane and New Guinea, although five species are amalgamated with other Bird’s Head terranes, more widely found in New Caledonia, New Zea- before docking with the Australian craton at 10 land, Timor and Aru New Guinean species Terrestrial birds of the Indo-Pacific 383 shared with Australia are often, but not exclu- collision zone Hall (1987, 1996) has outlined sively, restricted to Cape York Southern New the geological history of Halmahera, which he Guinea is geologically part of the Australian linked to the clockwise rotation of the Philip- craton What is more surprising is the evidence pine Sea plate discussed previously According for southern southeast peninsular Cape York to to Hall (1987), Halmahera has a basement of be an area of endemism, sharing the same or oceanic crust imbricated with Cretaceous to closely related species including the Magpie Eocene sediments This basement complex, Goose (Anseranus semipalmata), which Sibley which can be traced north into eastern and Ahlquist (1990) place with the South Ameri- Mindanao, is interpreted by Hall (1987) as the can screamers in a sister group to all other remnant of a late Cretaceous to early Tertiary ducks, swans and geese Pigram and Davies’ fore-arc ridge During the late Eocene and early (1987) suggestion, that the southern coast of the Oligocene the east Halmahera basement under- southeast peninsula represented a part of the went strong deformation and uplift, resulting in Australian craton detached and moved north by the deposition of river conglomerates in the spreading in the Coral Sea at 63 Ma, seems to be Oligocene and Miocene Synchronous deforma- borne out by bird distributional data tion is recognised over an area stretching from Pigram and Davies’ (1987) discussion of the western New Guinea to the western Pacific, composite geological nature of New Guinea leading Hall (1987) to view the east Halmahera- also makes intelligible the pronounced east- east Mindanao terrane as a westward continua- west taxonomic differentiation reported for tion of a Papuan arc complex (= Melanesian anurans (Menzies, 1975), murid rodents (Taylor arc) et al , 1982) and birds of paradise (Gilliard, Hall’s reconstructions (1998 this volume) 1969) For example, various species and sub- show east Halmahera continued to be translated species of the Six-wired Bird of Paradise west on the Philippine Sea plate Eastward sub- (Parotia spp) appear to be associated with the duction of the Molucca Sea microplate beneath following terranes: east Halmahera was initiated at c10 Ma in re- P wahnesi with Finisterre (Fig4: F) sponse to collision of continental fragments with P sefilata with Kemum and Wandamen Sulawesi that stopped westward movement (Fig4: K and Wa) along splays of the Sorong fault By 3 Ma sub- P carolae with Rouffaer and Sepik (Fig4: R duction under east Halmahera led to the build- and S) ing of an active arc on east Halmahera At 1 Ma P lawesi exhibita with Jimi, Schrader, Marum, arc volcanism ceased temporarily before shifting and Benabena (Fig4: J) westwards, building a new arc on the eroded P l fuscior and P l lawesi with Owen Stanley remnants of this Pliocene arc to form west (Fig4: O) Halmahera Halmahera’s continued western mi- P l helena with Bowutu (Fig4: B) gration eventually closed the Molucca Sea and These taxa link the Bird’s Head terrane to brought the island to its present position Melanesian rift fragments The occurrence of P Metamorphosed continental rocks in central l helena on the Bowutu terrane and P wahnesi Bacan have been linked by Pb-Nd isotope ratios on the Finisterre terrane, parts of the Melanesian to northern Australia (Vroon et al , 1996) Conti- arc system, may be the result of dispersal of P nental basement on Obi may also be a detached lawesi from the Owen Stanley ranges during col- fragment of the Australian craton Ali and Hall lision On the other hand an ancestral taxon may (1995) have discussed the geology of Obi and have been widely distributed and present on the the islands of Bisa and Tapas which lie to the Bird’s Head terranes and Melanesian rift-arc sys- northwest South Obi has a Palaeozoic continen- tem Further investigation into the phylogenetic tal metamorphic basement, while that in north relationship between taxa and terrane analysis Obi is composed of Jurassic ophiolite The is- would be of interest to biogeographers and ge- lands of Bisa and Tapas include high-grade Pal- ologists aeozoic metamorphics which are linked by Ali and Hall (1995) to similar metamorphics on Ba- can, 50 km to the north According to Ali and Arc and rift systems in Maluku Hall (1995) these terranes are derived from the Australian craton On the basis of palaeomag- North Maluku (Morotai, Halmahera, Ternate and netic studies the Obi-Misool-Sula platform block Bacan) is situated near the southern end of the was further south in Cretaceous times Obi un- Philippine trench at the boundary of a complex derwent northward translation during the Creta- 384 B Michaux ceous, westward translation during the late been discussed These phylogenetic patterns are Eocene-Oligocene (30 Ma) and northwest trans- intelligible in the light of Hall’s (1987, 1996, 1998 lation (north Obi) and northeast translation this volume) description of Halmahera’s geolog- (south Obi) in the Neogene These results sug- ical history The east Philippine-Halmahera-New gest that Obi is a detached fragment of the Aus- Guinea connection is an important result of tralian craton and may explain its anomalous Hall’s (1987) study as it explains similar distribu- biogeographical position noted before tional patterns in micro-lepidoptera (Diakonoff, The islands of south Maluku are continental 1967), murid rodents (Groves, 1984), weevils of fragments derived from the Australian craton, the tribe Pachyrrhynchini (Gressitt, 1956) and overthrust by oceanic basement material other groups discussed in Holloway (1990) The (Hartono and Tjokosapoetro, 1986; Hamilton, endemic element of north Maluku’s avifauna in- 1988; Audley-Charles et al , 1988; Lee and dicates a degree of isolation from other parts of Lawver, 1995) The term Banda rift-arc is used the Melanesian arc and from New Guinea Ac- here to describe this structure In Seram, cording to Hall (1987) the Halmahera section of ophiolite material has been thrust onto the Melanesian arc may have collided with New Palaeozoic continental basement (Linthout et al, Guinea in the late Eocene or early Oligocene It 1996) Linthout et al (1996) suggested this oc- is possible that the origin of the genera Lycoc- curred as Seram was ‘squeezed’ between Aus- orax and Semioptera (Paradisaeidae) dates from tralia and the inner Banda arc Seram was trans- this time (c40 Ma) lated northwards where it entered the Terera Aiduna fault zone (south of and parallel to the Sorong fault) where counter clockwise rotation The avifauna of south Maluku and the Banda and westward translation brought it to its rift-arc system present position Linthout et al (1996) dated these events as 8 Ma or younger There seems little doubt, given the preceding In the Timor region a similar collision be- discussion of avian and geological links, that tween continental fragment(s) and the Banda south Maluku was once part of the Australian arc also commenced in the Miocene In Timor, craton and was derived from northwest Aus- the interaction between rift and arc has led to a tralia sometime in the Mesozoic The high chaotic jumble of continental slivers, ophiolites number of endemic species (Table 5) indicates and subduction related sediments as Australian that south Maluku and Australia have been iso- continental crust is thrust under the Banda arc lated from each other, but the paucity of en- Snyder et al (1996) have interpreted seismic demic genera is problematical and gravity profiles across the cratonic margin Polhemus (1995) has shown that the water- behind Timor These profiles suggest that this strider Aquarius lili from Timor is not closely margin is rifting and providing further continen- related to Australian species, but is a member of tal fragments for eventual inclusion into the col- an Asian/Sundaic clade The non-endemic bird lision zone Their reconstructions show rifting in data in Table 5 also link south Maluku to Asia the Joseph Bonaparte Gulf, which is situated and Sundaland One factor of potential impor- between the Kimberley and Sturt blocks of tance in understanding this link between south northwest Australia This rifting appears to be Maluku and Asia is the collision of Timor with detaching fresh continental crust (Sahal and the Banda arc in the Pliocene (since 5 Ma) and Ashmore platforms) from the craton The age of the resulting dispersal between Flores and the detachment of the original Banda rift frag- Timor discussed by Michaux (1994) A second ments is not clear Jurassic to early Cretaceous factor is the possible relationship between south rifting has been suggested by various authors, Maluku and the Sumba terrane which is dis- but this seems rather early on avian evidence cussed in the following sections

Sulawesi as a geological composite The avifauna of north Maluku and the Melanesian/west Papuan arc The geological development of Sulawesi has been discussed by a number of authors The northern island of Halmahera is part of an Hutchison (1989) recognised that the southwest arc system and related to fragments now incor- arm and central Sulawesi was a microcontinental porated into northern New Guinea Taxa shared fragment Hutchison (1989) envisaged that an by north Maluku and New Guinea have already island arc was amalgamated to this terrane, to- Terrestrial birds of the Indo-Pacific 385 gether with the continental Sula platform Table 6 confirm that the Sulawesian avifauna terrane, in the late-Miocene The northern arm has close links with the Australian and Indian of Sulawesi was also interpreted by Hutchison cratons Musser (1987) regarded Sulawesian (1989) as an island arc which became sutured to mammals, with their high degree of endemism the rest of Sulawesi towards the end of the Ter- and phylogenetic isolation from Sundaic taxa, as tiary more typical of an oceanic island than one sur- Hall (1996) illustrated a more complex history rounded by lands rich in mammal families The for Sulawesi In Hall’s (1996) model the central/ avifauna also shows a high degree of endemism southwest Sulawesi terrane is sutured to (c25%) as do other groups such as the butterfly Kalimantan as in Hutchison (1989) At 45 Ma the families Papilionidae (47%) and Pieridae (56%) north arm of Sulawesi is shown as an arc close (Jong, 1990) and cicadas (95%) (Duffels, 1990) to its present position From 45 Ma to 25 Ma the This degree of endemicity shown by Sulawesi’s east Philippine-Halmahera arc (= Melanesian biota points to Sulawesi’s isolation from other arc) was translated westwards in response to landmasses clockwise rotation of the Philippine Sea plate A key task is to understand the relationship of The approach of the Philippine-Halmahera arc the southwest Sulawesian microcontinental to the Northern Arm arc lead to its suture to terrane to other terrane/arcs in the region southwest and central Sulawesi at 25 Ma This Rangin et al (1990) united the east Borneo time also marks the collision of the eastern arms terranes with southwest Sumatra, east Java, of Sulawesi, which Hall (1996) derived from the southwest Sulawesi, the “west Philippine is- south At 20 Ma Australian continental crust was lands” and Sumba as a Gondwana continental thrust under the eastern arms Two further con- block, which they termed the Sumba terrane tinental terranes, Buton and the Sula platform, This terrane became sutured to the margin of became amalgamated at 10 Ma and 5 Ma respec- the Sunda shelf during the early Tertiary, as tively Hall (1996) illustrated the derivation of what remained of the Ceno-Tethys ocean the Buton terrane and Sula platform from the (Metcalfe, 1996) was subducted Rangin et al Bird’s Head terrane by westward translation (1990) regarded the ophiolite-bearing melanges along splays of the Sorong fault system found in Borneo, central Java, and Sumatra as evidence of this Tethys suture zone Michaux (1996) discussed a distributional pat- Implications for Sulawesi biogeography tern linking Sulawesi with Mindanao, north Bor- neo, east and southeast Kalimantan, Java, west Table 6 shows twenty five bird species that have Sumatra (and islands), the Andamans/Nicobars their most westerly occurrence in Sulawesi and Burma Distributions of this type avoid the These species, together with thirteen bats listed Malay peninsula and reach Asia (and sometimes in Cranbrook (1991) and Musser (1987) which beyond) via Burma To the east of Java, this dis- have a Sulawesi/Maluku/New Guinea distribu- tributional pattern extends to the Lesser Sunda tion, three marsupials of the genus Phalanger islands The hypothesis of a Sumba terrane (Musser, 1987) and microhylid frogs of the ge- clearly has relevance for understanding the con- nus Oreophyrne (Kampen, 1923) can be classed nections shown by the Sulawesian avifauna to as Australasian elements in the Sulawesian surrounding avifaunas from Mindanao, north fauna An explanation for the presence of these Borneo, the Lesser Sundas, west Sumatra, taxa in Sulawesi is that they (or their ancestors) Andamans and Nicobars were part of the Sula platform or Buton terrane One might speculate that the fragments de- fauna There are also eleven Asian/Sundaic bird scribed above represented a rift-arc complex(es) species which only extend to the Sula Islands of east Gondwana affinity, which was sutured to (Table 6) These species appear to have colo- the Sundaland margin at the end of the Creta- nised the Sula Islands from Sulawesi ceous or early Tertiary Subsequent tectonic The southwest microcontinental terrane was events have dispersed these fragments Some of originally attached to southeast Kalimantan, but them may have been incorporated into the became separated when the Makassar Strait Banda arc which has transported them back to- opened in the mid-Tertiary (McCabe and Cole, wards Australia, a possible source area from 1989) Hutchison (1989) suggested that this frag- which they were derived Any relationship be- ment was derived from the north, but there is no tween the Sumba terrane of Rangin et al (1990) support for this hypothesis from faunal analysis and the Banda rift-arc system alluded to earlier (Michaux, 1991, 1994) The distributional data in would stem from the similarity of their respec- 386 B Michaux

Fig 5 Geological structures discussed in the text Stippled = cratonic areas; diagonal lines = Melanesian arc; circles = Melanesian rift; dots = Banda rift-arc structure; vertical lines = Sumba terrane B = Bismarck archipelago, Bo = Borneo, F = Fiji, J = Java, nM = north Maluku, sM = south Maluku, NC = New Caledonia, NG = New Guinea, NZ = New Zealand, P = Philippines, S = Samoa, So = Solomons, Su = Sulawesi, T = Tonga, V = Vanuatu Numbers refer to areas for which geological histories are provided tive source areas Pacific groups has the potential to answer these and other questions As far as I understand the geological literature Discussion describing the breakup of Gondwanaland in the Mesozoic and the reassembly of drifted frag- Clarifying the relationship between terranes, rift- ments in Asia, Southeast Asia and Indonesia dur- arc complexes, and mobile arc systems is an ing the Tertiary, an analysis using ridged plates important task in unravelling Indonesian bioge- models is not always appropriate The breakup ography (Polhemus, 1996) In turn, how are of east Gondwana appears to have involved two these structures related to the Indian and Aus- qualitatively different fragmentation regimes In- tralian cratonic regions derived from east ternal rifting produced large cratonic areas Gondwana? What is the explanation for the ori- which act as rigid structures Marginal rifting gin of ‘Oriental’ elements in the New Guinean produced smaller, elongate rifts and/or micro- and Pacific avifaunas but their absence from continents which are often (always?) associated Australia? A proper analysis of widespread Indo- with island arcs As these products of breakup Terrestrial birds of the Indo-Pacific 387 interact in collision zones, cratonic areas can ei- Nightjar family is discussed as an example below ther grow at the margins through accretion (e g, northern Australia) or their margins can fracture producing new rifts/microcontinents (e g, west Nightjar evolution and the fragmentation of east and east Australia) Where major plates interact Gondwana (e g, along the Banda arc) the movement of continental and arc structures caught up in colli- Fig6 shows the distribution of four avian fami- sion zones between rigid plates appear fluid lies (nightjars) that Sibley and Ahlquist (1990) over a geological time scale An example of this place in the Strigiformes The Eurostopodidae is way in which the northern section of the Ban- (Fig6: E) and the cosmopolitan Caprimulgidae da rift-arc structure has been translated north are ‘sister’ taxa The range of the Eurostopodi- and west to its present position Other examples dae defines the present day extent of former would include the movement of arcs across parts of east Gondwana The restriction of this plate boundaries in the southwest Pacific and family to former east Gondwana fragments indi- the lateral movement of rift-arc fragments along cates that the origin and diversification of the transform faults or in response to oblique sub- Eurostopodidae are related to an ancestral taxon duction in Indonesia widely distributed in east Gondwana, and sub- Marginal rifting occurred along the entire east sequent fragmentation of its range followed by Gondwana margin from the Indian to the taxonomic diversification southwest Pacific sector According to Metcalfe A further ‘sister’ grouping identified by Sibley (1996) a series of rifts were detached from the and Ahlquist (1990) is between the families Po- Indian margin from late Devonian (east Asia and dargidae (Fig6: P) and Batrachostomidae (Fig6: Indochina) to early Cretaceous (Burma-west B) The Batrachostomidae are distributed on the Sumatra-east Kalimantan-west Sulawesi-Lesser Indian craton and Gondwana terranes of South- Sundas) The rifted fragments incorporated into east Asia and Sundaland, while the Podargidae the outer Banda arc islands of Timor, Tanimbar, are found on the Australian craton and Melane- Kai, Seram and Buru were detached from sian rift-arc fragments This implies that these northwest Australia probably in the early two families date from the Cretaceous at c90 Cretaceous This pattern of marginal rifting is Ma According to Sibley and Ahlquist (1990), the repeated to the east with the formation of Aegothelidae is the most primitive nightjar fami- continental fragments now in the Bird’s Head, ly The range of this family is cratonic Australia continental terranes in central New Guinea and plus the Melanesian rift-arc system (Fig6: A) If the New Zealand subcontinent Larger cratonic Sibley and Ahlquist (1990) are correct in their fragments (India and Australia) were detached placement of the Aegothelidae then its present and conveyed north into the Himalayan and distribution represents a relict Two other cen- Indonesian-Papuan collision zones Significant tral and south American families included in the lateral movement, development of marginal Strigiformes are ‘sister’ taxa and together form oceanic basins and complex plate margin the sister taxon to the Eurostopodidae plus responses have subsequently occurred in the Caprimulgidae These two families, the Nyctibii- Indonesian-Papuan collision zone dae and Steatornithidae, must date their origin In my view the birds of the Indo-Pacific re- to a time after south America had split from west gion show clear evidence that their distributions Gondwana between 118-96 Ma (Veevers, 1988) have been determined by the geological events described above Fig5 shows areas in the mod- ern Indo-Pacific region which share a degree of Widespread species — Gallirallus philippensis geological similarity and whose avifauna, or portion thereof, reflect this in their phylogenetic The interpretation of widespread species is a links Such a hypothesis is likely to be more ac- key element in any biogeographical analysis ceptable to botanists than to zoologists, who These species may be efficient dispersers, they tend to view mammalian and bird families as may be unidentified species complexes that al- relatively recent arrivals However, Hedges et al ready had widespread ancestors on east (1996) have recently contested this view and Gondwana in the Cretaceous, they may repre- suggested not only that bird orders diversified sent a single species that has not responded to much earlier at c100 Ma, but that the fragmenta- geological fragmentation of their original range, tion of continents has been an important factor or they are not monophyletic and their wide dis- in avian diversification The evolution of the tribution is an artefact 388 B Michaux

Fig 6 Distributions of four night-jar families A = Aegothelidae, B = Batrachostomidae, E = Eurostopodidae, P = Podargidae See text for explanation

Extreme evolutionary conservatism seems Kingfishers (Tanysiptera: Maluku-New Guinea- rather improbable given the time scales in- north Queensland) An African species, H volved As for widespread distributions being an senegalensis, is the outgroup to these Austro- artefact, Musser (1981) has speculated that the papuan species murid genus Rattus may not be monophyletic The occurrence of widespread, polymorphic and Beehler et al (1986) have separated the species composed of allopatric populations pro- Australian false babblers (Pomatostomatidae) vide conditions for investigating some of the from African and Asian babblers (Timaliidae) taxonomic possibilities outlined above and as a Sibley and Ahlquist (1990) discussed the rela- test for geological models Ripley (1977) dis- tionships between various Alcedinidae and cussed the taxonomy of the rail Gallirallus showed the genus Halcyon to be paraphyletic philippensis which is widespread across eastern H sancta, the Australian craton-Melanesian rift Indonesia, Australia and the Pacific Twenty four species, is most closely related to the monotypic subspecies of G philippensis are described by Melidora macrorrhina (New Guinea) and these Ripley (1977) and their distributions shown in taxa are in turn the ‘sister’ group to Paradise Fig7 In Fig7 the subspecies have been labelled Terrestrial birds of the Indo-Pacific 389

Fig 7 Distributions of the subspecies of the rail Gallirallus philippensis 1 = philippensis (Philippines and Sulawesi), 2 = pelewensis (Palau), 3 = anachoretae (Anchorite Island), 4 = admiralitatis and praedo (Admiralty Island), 5 = lesouefi (New Hanover), 6 = meyeri (Witu Island), 7 = reductus (Long Island), 8 = sethsmithi (Vanuatu and Fiji), 9 = ecaudatus (Tonga), 10 = goodsoni (Samoa), 11 = randi (central mountains, Irian Jaya), 12 = wahgiensis (central mountains, PNG), 13 = swindellsi (New Caledonia), 14 = norfolkensis (Norfolk Island), 15 = assimilis (New Zealand), 16 = dieffenbachii (Chathams), 17 = macquariensis (Macquarie Island), A = yorki (northern Australia, Cape York and southern New Guinea), B = australis (NSW and South Australia), C = mellori (southwest Australia), filled circle = andrewsi (Cocos Island), open circle = xerophilus (Gunung Api), star in circle = wilkinsoni (Flores)

to indicate the degree of relationship that might pected to show the closest relationship to be expected as a consequence of geological Melanesian rift taxa (Fig7: 11-17) These groups models discussed earlier The Melanesian rift should in turn show a sister group relationship and arc taxa are numbered The nominate spe- to the Australian craton taxa (Fig7: A-C) The cies, G p philippensis (Fig7: 1) has been in- position of the Sumba terrane taxa (in which the cluded with other Melanesian rift and arc taxa, Cocos subspecies andrewsi is included (Fig7: but its position is equivocal because Sulawesi closed circle) depends on the geological rela- and Mindanao can also be related to Sumba tionship of this terrane to south Maluku, Aus- terrane fragments in the Lesser Sundas tralia and the Melanesian rift-arc system Melanesian arc taxa (Fig 7: 2-10) might be ex- Acknowledgments 390 B Michaux

(Homoptera: Cicadoidea) In Insects and the rainforests It is a great pleasure to acknowledge John of Southeast Asia pp 63-72 Edited by W J Knight and J D Holloway Royal Entomological Society, London Palmer of Arnold Books, Christchurch for the Duffels, J P and Boer, A J de, 1990 Areas of endemism loan of some of the sources used in this study and composite areas in east Malesia In The plant diversi- Robert Hall, Hans Duffels, Arnold de Boer, ty of Malesia, pp 249-272 Edited by P Baas et al Kluw- Hubert Turner and Rod Hay provided valuable er, Dordrecht assistance in improving earlier drafts The help- Falla, R A, Sibson, R B and Turbott, E G 1979 Birds of New Zealand Collins, Auckland, 247 pp ful comments and suggestions of the referees, Fleming, C A 1980 The geological history of New Zealand Charles Sibley and Edward Dickinson, are grate- and its life Auckland University Press, Auckland, 141 pp fully acknowledged Fordyce, R E 1982 The fossil vertebrate record of New Zealand In The 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