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

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902 M. Heads Figure 19 Zosterops atriceps (hatched) subspecies on Morotai, Halmahera, and Bacan. Z chloris has subspecies (solid line) on Aru, Kai, Seram, Schildpad, Halmahera; S Moluccas, Tanimbar, Torres Srait Is. (albiventris), and others around Sulawesi, Lombok and islands off SE Borneo. Figure 21 The distribution of Aquilaria ®laria (Thymelaeaceae). Figure 20 Pachycephala phaionota, with two subspecies: one on Aru Is., N Moluccas, Western Papuan Islands and Geelvink Bay islets, the other endemic to Majau I. (N Moluccas). The genus ranges north to Thailand. The Moluccas have an important biogeographical connection to the south, for example Lichmera Cabanis reaches its northern limit in the Moluccas. To the north, the Moluccas link New Guinea with the Philippines. Examples are Amaurornis olivaceus (Meyen) and the tree Aquilaria ®laria (Oken) Merr. (Ding Hou, 1960; now also known from the Sepik region) (Fig. 21). Finally, the northern Moluccas are an important eastern boundary for many Indian Ocean groups, such as the land snail family Clausiliideae (Szekeres, 1980), the sylvine warbler Bradypterus Swainson (Africa ± northern Moluccas), the bat family Megadermatidae (Africa ± northern Moluccas) and the palm subtribe Oncospermatinae J.D. Hooker (Uhl & Drans®eld, 1987; map 43) (Seychelles has four genera, Sri Lanka ± northern Moluccas has one genus). Craw et al. (1999, Fig. 4.4) mapped vicariant butter¯y genera in, respectively, SE Asia east to Sulawesi, and the Moluccas east to Australasia. Summarizing, the biogeographical boundary, or node, between Sulawesi and the Moluccas (`Weber's line') may not be as well-known as its over-famous neighbour, the break between Sulawesi and Borneo (`Wallace's line'), but it is probably of similar signi®cance. As shown, a break between the Moluccas and Sulawesi occurs in many plants and animals, and this same break in Paradisaeidae cannot be interpreted as an anomaly. The northern Moluccas have several different standard biogeographical connections with north and south New Guinea, recalling the different character recombinations in Semioptera. De Boer (1995a) wrote that the cicadas of the Moluccas show ®ve distinct patterns of distribution, each indicating different relationships between parts of the Moluccas and different areas. Like Paradisaeidae and the other birds mapped above, the cicadas Cosmopsaltria StaÊl, Diceropyga StaÊl, Aedeastria de Boer (Fig. 22), Gymnotympana StaÊl (Fig. 23) and Baeturia StaÊl are all mainly in New Guinea, but each has a few species in NE Queensland and outlying endemics in the Moluccas. Discussing these genera, de Boer (1995a) cited three volcanic island arc systems in the the West Paci®c, one of which, the Halmahera arc, comprises eastern Mindanao (Philippines), the northern Moluccas and Waigeo Island, and possibly connects with the Mariana and Palau/Yap arcs. The Halmahera arc formed far to the east of its present position on a fracture of the Paci®c plate ± it might have ended close to the Papuan Peninsula. Since the early or middle Miocene the arc has swept 2000 km westwards past the north-western edge of New Guinea into the Moluccas region (Daly et al., 1991; Honza, 1991). The geology of the northern Moluccas region is highly complex and controversial; one interpretation is shown in Fig. 24. Ó Blackwell Science Ltd 2001, Journal of Biogeography, 28, 893±925
Biogeography of birds of paradise 903 arcs from eastern PNG. This would also account for the Moluccas ± New Britain/NE PNG disjunctions seen in Accipiter, Ninox and others. South of Halmahera, but still in the northern Moluccas, Obi Island (where Lycocorax pyrrhopterus obiensis Bernstein is endemic) and Bacan Island (where Semioptera w. wallacii Gould is endemic) comprise a separate microcontinent distinct from Halmahera. The origin of the southern Moluccas (Buru and Seram) is still subject to controversy, but the microcontinent they form is of Australian origin (de Boer, 1995a) and is quite distinct from that of the northern Moluccas. Figure 22 The distribution of Aedeastria (Homoptera) species. Figure 23 The distribution of Gymnotympana (Homoptera) species. Western Papuan Islands Seven species of birds of paradise are indigenous to the Western Papuan Islands, lying between the New Guinea mainland and the Moluccas. Several mainland species occur on Salawati and Misool and Manucodia ranges to Gebe Island. Cicinnurus respublica (Bonaparte) and Paradisaea rubra Daudin are striking species both endemic to Waigeo and Batanta; birds endemic to these islands are discussed below as examples of ophiolite terrane endemism. In stark contrast, Zosterops Vigors & Hors®eld, supposedly a lover of small islands, is `strangely enough' (Rand & Gilliard, 1967) totally absent from Waigeo, Batanta, Misool and Salawati, although Z. chloris extends right up to the approaches of Misool on the tiny Schildpad Islands (Fig. 19). This absence of Zosterops from Misool and the others is indeed striking, and inexplicable by dispersal as movement, as the genus has ®ve species in the Moluccas, nine in New Guinea, and there is an endemic genus of Zosteropidae on each of Seram and Buru. The Western Papuan Islands also form an eastern limit for widespread Indian Ocean groups such as Strophanthus DC. (Apocynaceae): West Africa to western Papuan Islands (Beentje, 1982). Figure 24 Geology of the Moluccas region. Tectonic elements (subduction zones, faults, volcanic arcs) as heavy lines, arrows indicate motion on strike-slip faults (from Axelrod & Raven, 1982 after Hamilton, 1977). de Boer (1995a,b) and de Boer & Duffels (1996a,b) proposed that the north Moluccan cicadas are derived from ancestral forms which travelled on fragments of volcanic The Vogelkop The Vogelkop, like the Huon and Papuan Peninsulas and major biogeographical nodes in general, is an important centre of biological endemism, absence and disjunction. The Kemum terrane in the central Vogelkop originated as a detached portion of Gondwana (Pigram & Davies, 1987) and other component terranes of the Vogelkop are also of interest. The Arfak Mountains are a major centre of endemism, for example, four out of twenty-two New Guinea species of Rapanea Aubl. (Myrsinaceae) are endemic there, at the Anggi Lakes (Sleumer, 1986). In the birds of paradise Astrapia nigra (Gmelin) is restricted to the Arfak and Tamrau Mountains, and its af®nities are with the Huon Peninsula species A. rothschildi Foerster, 1500 km to the east (Fig. 25). Similarly, within the black, blue-eyed Parotia species (Fig. 26), P. se®lata of the Vogelkop and nearby Wandammen Peninsulas may well be the sister species of P. wahnesi of the Huon Penisula and the Adelbert Mountains. Both 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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