Birds of paradise, biogeography and ecology in New Guinea: a review

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900 M. Heads Figure 14 Ceyx (ˆ Alcedo, ˆ Alcyone) azureus at its western limit: subspecies in N Moluccas, Tanimbar, Aru Is., Western Papuan Is. and New Guinea. There are three other races in NES Australia and Tasmania. C. pusillus also ranges from Solomons, Queensland west through New Guinea to Halmahera and Obi (C. p. halmaherae). Figure 16 Monarcha at its north-western limit (hatched line). Monarcha pileatus Salvadori, with subspecies on Halmahera, Buru, and Tanimbar. M. trivirgatus (Temminck), also disjunct in Queensland and the Louisiade Is. (arrow), M. julienae (Ko®au), and M. brehmii (Schlegel) (Misool, Biak I.) with af®nities to Manus and Bismarck Archipelago species (arrow). Figure 15 The six species of Eos: E. histrio P.L.S.MuÈ ller (Talaud Is.), E. squamata (Boddaert): Obi (obiensis), Maju I. (atrocaerulea), N Moluccas (riciniata), Western Papuan Is. (nominate). E. cyanogenia Bonaparte (Geelvink Bay islands), E. reticulata (S.Muller): Tanimbar and Kai Is., E. bornea Linnaeus (S Moluccas, Kai Is.), E. semilarvata Bonaparte (Seram). The related Chalcopsitta and Pseudeos occur on mainland New Guinea. Artamus leucorhynchus leucopygialis Gould) also have a NE Australia ± New Guinea ± Moluccas distribution more or less the same as that of Paradisaeidae. In beetles, Chrysomelidae subfamily Hispinae has twentyone genera in New Guinea, fourteen of which range west only to the Moluccas (Gressitt, 1982a). Likewise in frogs, Microhylidae subfamily Asterophryinae is endemic to New Guinea and the Moluccas (Richards et al., 2000), and the genus Nyctimestes Stejneger (Hylidae) has a range practically identical to that of the birds of paradise: New Guinea (most species), the Milne Bay islands, northern Australia, and the northern Moluccas (Zweifel, 1958; Tyler, 1968). The lizard genera Eugongylus Fitzinger and Carlia Gray also range through New Guinea west to the Moluccas (Greer, 1974). Many characteristic plants of Australasian and Melanesian rainforests also have a north-western limit at the Moluccas. For example, in the seed plants, Himatandraceae (comprising Galbulimima F.M. Bail.) is in NE Australia, New Guinea, and the Moluccas (Croft, 1978) like the Paradisaeidae. A group of four genera in Cunoniaceae: Spiraeanthemum A. Gray (including Acsmithia Hoogland), Gillbeea F. Muell., Aistopetalum Schlechter and Brunellia Ruiz & Pav., forms a basal clade in the family, sister group to the rest, and occurs in Central America, Polynesia, NE Queensland, New Guinea and the Moluccas (Hufford & Dickison, 1992). Other plant taxa which break off range westward at the Moluccas include the fern Leptopteris Presl (van Balgooy, 1966a,b), the conifers Decussocarpus de Laub. sect. Decussocarpus (de Laubenfels, 1984) and Papuacedrus Li (Li, 1953), the monocots Pandanus sect. Maysops St John (Stone, 1992), Cominsia Hemsl., Helmholtzia F. Muell. (Johns & Hay, 1981), Drymophloeus Zipp. (Zona, 1999), Gulubia Becc. (Drans®eld, 1993), Glossorhyncha Ridley (Smith, 1979±1996), and Pterostylis R.Br. (de Vogel, 1975a), and the dicots Levieria Becc. (Philipson, 1986), Pararistolochia Hutch. & Dalz. (re-appearing in Africa) (Parsons, 1996), Rhyticaryum Becc., Polyporandra Becc. (Sleumer, 1971), Hollrungia K. Schum. (de Wilde, 1975), Hugonia sect. Durandea Planch. (van Balgooy, 1993), Dubouzetia Pancher ex Brongn. & Gris (Coode, 1987), Schleinitzia Warb. (de Vogel, 1975b; as Prosopis insularum (Guill.) Bret.), Archidendron ser. Stipulatae (Mohl) Nielsen, A. ser. Pendulosae (Mohl) Nielsen, and A. ser. Morolobiae (Kosterm.) Nielsen (Nielsen et al., 1984), Pullea Schlechter (Hoogland, 1979; cf. Hufford & Dickison, Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 893±925
Biogeography of birds of paradise 901 1992), Flindersia R.Br. (Roos, 1984), Jagera Blume, Sarcopteryx Radlk., Sarcotoechia Radlk. (the last three from Adema et al., 1994), Burckella Pierre (Pennington, 1991), Eucalyptopsis White (White, 1951; Johns, 1980), Lindsayomyrtus Hyland & Steen. (Hyland & van Steenis, 1973), Octamyrtus Diels (Scott, 1979a), Mastixiodendron Melchior, Cyclophyllum Hook.f. (Smith, 1979±1996), Tecomanthe Baillon (van Steenis, 1977), and such characteristic, widespread New Guinea species as Schuurmansia henningsii K.Sch. (Kanis, 1978). Like most important centres of endemism, the Moluccas have several far-¯ung biogeographical af®nities. For example, four species of Accipiter (Accipitridae) show disjunct connections between the Moluccas and New Britain which do not involve mainland New Guinea: A. erythrauchen G.R. Gray (Fig. 17) of the northern Moluccas is similar to A. brachyurus (Ramsay) of New Britain, just as A. henicogrammus (G.R. Gray) of the northern Moluccas is related to A. luteoschistaceus Rothschild & Hartert of New Britain (Wattel, 1973). The butter¯y Eurema candida is also disjunct between Moluccas/Timor and the Bismarck Archipelago/Solomon Islands (Parsons, 1999). Similarly, the fern genus Christensenia Maxon. is disjunct between west Malesia/Moluccas and New Ireland ± Solomons (Johns & Bellamy, 1981). Also in ferns the closely related Christella perpubescens (Alston) Holttum group has one species in each of the following: the Moluccas; Waigeo Island and disjunct in the Solomons; Biak island; and New Ireland, all on limestone (Holttum, 1976, 1981). In the Araceae, Spathiphyllum commutatum Schott (Araceae) occurs in the Philippines, Palau, Sulawesi, and the Moluccas, and is disjunct from there to New Britain and Bougainville, being absent from mainland New Guinea. It is replaced in Manus, New Ireland and SE New Guinea by two further species in the genus, the remainder are all in tropical America (van Steenis, 1961; Hay, 1990). Spathiphyllum is replaced on mainland New Guinea outside the Papuan Peninsula by the more or less closely allied Holochlamys Engl. This Moluccas ± New Britain connection is related to similar disjunctions. For example, Ninox connivens Latham (Fig. 17) is in the northern Moluccas, eastern PNG and Australia, bypassing Irian Jaya, and a similar range (northern Moluccas; Sepik) is held by the tree Dictyoneura acuminata Blume ssp. acuminata, with the gap in Irian Jaya ®lled by D. acuminata ssp. microcarpa J. Dijk (van Dijk in Adema et al., 1994). Other examples were noted by Michaux (1994). An even greater disjunction across the north of New Guinea is shown in conifers. Dacrydium magnum de Laub. occurs only in the northern Moluccas (Obi), and on islands east of New Guinea ± the Louisiades (Sudest Island) and the Solomon Islands (de Laubenfels, 1988), and Podocarpus spathoides de Laub. has a very similar distribution: northern Moluccas (Morotai), disjunct to the Louisiades (Rossel) and Solomon Islands, with an additional disjunct record in the west on the Malay Peninsula (Mount Ophir ˆ Gunong Ledang). In many taxa the Moluccas populations are involved with those of the Aru Islands, with the af®nity skirting New Guinea to the south-west and not present on the mainland. Three examples are illustrated: Hemiprocne mystacea con®rmata Stresemann (Fig. 18) is on the Moluccas and Aru Islands; other races occur in New Guinea, Bismarck Archipelago and the Solomon Islands. Zosterops chloris Bonaparte (Fig. 19) ranges on the Banda Sea islands, islets off SE Borneo, and also on the Torres Strait islands, but is not on the New Guinea mainland. Z. c. chloris is restricted to the Moluccas, Kai and Aru Islands. Z. atriceps Gray is endemic in the northern Moluccas. Pachycephala phaionota (Bonaparte) (Fig. 20) surrounds the Vogelkop to the west, north and south: it is on the northern Moluccas, Aru Islands, the western Papuan Islands and also small islands in Geelvink Bay. Figure 17 Accipiter erythrauchen G.R.Gray: Bacan, Halmahera, Morotai, Obi (nominate), Seram, Buru (ceramensis (Schlegel)). This species is similar to A. brachyurus of New Britain (arrow). A. henicogrammus: Bacan, Halmahera, Morotai; related to A. luteoschistaceus of New Britain. Ninox connivens, disjunct in E New Guinea (east of Merauke and Karkar Island) and Australia. Figure 18 Hemiprocne coronata (Sulawesi), H. mystacea with subspecies on Moluccas and Aru Is., and in New Guinea, Bismarck Archipelago, Solomon Islands. Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 893±925
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Birds of paradise, biogeography and ecology in New Guinea: a review

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