Geografi Tanaman di Bioregion Wallace
Transcription
Geografi Tanaman di Bioregion Wallace
GEOGRAFI TANAMAN DI BIOREGION WALLACE Johny S. Tasirin & Martina A. Langi TROPICAL PLANT CURRICULUM PROJECT Kerjasama UNIVERSITAS SAM RATULANGI Dan TEXAS A&M UNIVERSITY 2012 J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 1 DISCLAIMER This module is made possible by the generous support of the American people through the United States Agency for International Development (USAID). The contents are the responsibility of Texas A&M University and Sam Ratulangi University as the USAID Tropical Plant Curriculum Project partners and do not necessarily reflect the views of USAID or the United States Government. Kata Pengantar Puji Syukur Kehadirat Tuhan Yang Maha Esa, atas hikmat dan pertolonganNya sehingga penyusunan modul ini dapat terselesaikan. Terima kasih disampaikan kepada USAID dan Universitas TEXAS A&M atas dukungan dana bagi penyusunan modul ini. Modul ini ditujukan sebagai bahan ajar Keanekaragaman hayati. Modul ini dapat digunakan oleh tenaga pengajar (dosen) maupun mahasiswa sebagai bahan acuan untuk memperkaya pengetahuan terkait Keanekaragaman hayati tumbuhan dan mengenal keanekaragaman tumbuhan di daerah tropis. Penyusunan Modul ini masih banyak kekurangan. Untuk itu kritik dan saran yang membangun sangat kami butuhkan untuk penyempurnaan modul ini. Terima kasih, Penyusun J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 3 Deskripsi Ekoregion di Kawasan Wallacea 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. Halmahera rain forests Banda Sea Islands moist deciduous forests Buru rain forests Lesser Sundas deciduous forests Seram rain forests Sulawesi lowland rain forests Sulawesi montane rain forests Sumba deciduous forests Timor and Wetar deciduous forests Mindanao-Eastern Visayas rain forests Mindanao montane rain forests Mindoro rain forests Luzon montane rain forests Luzon tropical pine forests Luzon rain forests Palawan rain forests Sulu Archipelago rain forests J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 4 1. Halmahera rain forests Halmahera, Indonesia Photograph by Vincent Roelofs Southeastern Asia: Islands of Halmahera, Moratai, and Obi in Indonesia 10,400 square miles (26,900 square kilometers) -- about the size of Maryland This ecoregion comprises the original "Spice Islands." The tropical islands that constitute the complex and mountainous terrain of the Halmahera Rain Forests are an important part of the region known as Wallacea, which contains a very distinctive fauna representing a mix of Asian and Australasian species. This small ecoregion contains an astounding twenty-six bird species, including four monotypic genera, which are found nowhere else in the world. Although there is some exploitation by logging and mining companies, extensive blocks of habitat still cover all the islands, and nearly 80 percent of its original forest still intact. Location and General Description This ecoregion represents the moist forests on Halmahera, Morotai, Obi, Bacan, and the other nearby Maluku Islands in the Relatively northeastern Indonesian Archipelago. Tropical and Stable/Intact Subtropical Moist Based on the Köppen climate zone system, Broadleaf Forests this ecoregion falls in the tropical wet climate zone (National Geographic Society 1999). The geologic history of these islands is a very complex mixture of inner volcanic island arcs, outer volcanic island arcs, raised coral reefs, and fragments of continental crust. Halmahera is a product of a collision between two islands approximately 1-2 million years ago. The eastern half of the island was part of an outer arc on the Philippines tectonic plate and consists of sedimentary and intrusive igneous rocks. The western half of Halmahera and Morotai was part of an inner arc consisting of volcanic materials. Bacan is a mixture of volcanic inner island arc and some crustal materials (Monk et al. 1997). The natural vegetation of these islands was tropical lowland evergreen and semievergreen forest (Stattersfield et al. 1998). Most of the remaining habitat in this ecoregion is semi-evergreen rain forest and includes eight characteristic dipterocarp species: Anisoptera thurifera, Hopea gregaria, H. iriana, H. novoguineensis, Shorea assamica, S. montigena, S. selanica, and Vatica rassak. Volcanic soils and good aspect combine to produce almost optimal growth conditions. Most of the trees reach 30 m or more and carry thick-stemmed lianas and woody and herbaceous epiphytes. Rattans that grow to 130 m and other epiphytes are common in old-growth forests. The most luxuriant rain forests occur in northwest Morotai and north Halmahera, as opposed to the south arm of Halmahera, which is in the rain shadow of north Halmahera and Bacan. Low, shrubby vegetation is found in poor soil conditions on patches of ultrabasic rocks (Monk et al. 1997). J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 5 Biodiversity Features Overall diversity is low in this ecoregion, but overall endemism is moderate to high when compared with that of other ecoregions in Indo-Malaysia. This ecoregion falls within the Wallacean biogeographic zone, and thus exhibits a mixture of Asian and Australian fauna. Together with Seram, Buru, and the Banda Sea Islands, this island group forms part of a bioregion with perhaps the highest levels of bird endemism for its size anywhere in the world and the highest number of endemic birds of any area in Asia. The mammal fauna is depauperate, containing only thirty-eight species with both Asian and Australasian affinities (cuscuses), but includes eight ecoregional endemics (table 1). The Obi cuscus (Phalanger rothschildi) is considered vulnerable (IUCN 2000). Table 1. Endemic and Near-Endemic Mammal Species. Family Species Phalangeridae Phalanger ornatus* Phalangeridae Phalanger rothschildi* Phalangeridae Phalanger sp.* Pteropodidae Pteropus chrysoproctus Pteropodidae Pteropus personatus* Pteropodidae Nyctimene minutus Muridae Melomys obiensis* Muridae Rattus sp.* An asterisk signifies that the species' range is limited to this ecoregion. The ecoregion supports approximately 223 bird species, including 43 ecoregional endemic species (table 2). The ecoregion corresponds with the Northern Maluku EBA. There are four endemic monotypic genera: Habroptila, Melitorgrais, Lycocorax, and Semioptera. These species include the invisible rail (Habroptila wallacii), white-streaked friarbird (Melitograis gilolensis), paradise-crow (Lycocorax pyrrhopterus), and the standardwing (Semioptera wallacii). Of the forty-three restricted-range species found in this ecoregion (and EBA), an astounding twenty-six are found nowhere else in the world. Five vulnerable species, four of which are found nowhere else, are found in the ecoregion: invisible rail (Habroptila wallacii), caranculated fruit-dove (Ptilinopus granulifrons), chattering lory (Lorius garrulus), and white cockatoo (Cacatua alba) J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 6 (Stattersfield et al. 1998). Table 2. Endemic and Near-Endemic Bird Species. Family Common Name Species Accipitridae Moluccan goshawk Accipiter henicogrammus* Accipitridae Rufous-necked sparrowhawk Accipiter erythrauchen Megapodiidae Moluccan scrubfowl Megapodius wallacei Megapodiidae Dusky scrubfowl Megapodius freycinet Rallidae Invisible rail Habroptila wallacii* Scolopacidae Moluccan woodcock Scolopax rochussenii* Columbidae Scarlet-breasted fruit-dove Ptilinopus bernsteinii* Columbidae Blue-capped fruit-dove Ptilinopus monacha* Columbidae Grey-headed fruit-dove Ptilinopus hyogastra* Columbidae Carunculated fruit-dove Ptilinopus granulifrons* Columbidae White-eyed imperial-pigeon Ducula perspicillata Columbidae Spice imperial-pigeon Ducula myristicivora Columbidae Pink-headed imperial-pigeon Ducula rosacea Columbidae Cinnamon-bellied imperial-pigeon Ducula basilica* Psittacidae Moluccan hanging-parrot Loriculus amabilis Cacatuidae White cockatoo Cacatua alba* Loriidae Violet-necked lory Eos squamata Loriidae Chattering lory Lorius garrulus* Cuculidae Moluccan cuckoo Cacomantis heinrichi* Cuculidae Pied bronze-cuckoo Chrysococcyx crassirostris J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 7 Cuculidae Goliath coucal Centropus goliath* Strigidae Moluccan hawk-owl Ninox squamipila Aegothelidae Moluccan owlet-nightjar Aegotheles crinifrons* Alcedinidae Blue-and-white kingfisher Todirhamphus diops* Alcedinidae Sombre kingfisher Todirhamphus funebris* Coraciidae Purple roller Eurystomus azureus* Pittidae Ivory-breasted pitta Pitta maxima* Meliphagidae Olive honeyeater Lichmera argentauris Meliphagidae White-streaked friarbird Melitograis gilolensis* Meliphagidae Dusky friarbird Philemon fuscicapillus* Pachycephalida Drab whistler Pachycephala griseonota Monarchidae White-naped monarch Monarcha pileatus Monarchidae Moluccan flycatcher Myiagra galeata Corvidae Long-billed crow Corvus validus* Paradisaeidae Paradise-crow Lycocorax pyrrhopterus* Paradisaeidae Wallace's standardwing Semioptera wallacii* Oriolidae Halmahera oriole Oriolus phaeochromus* Campephagidae Moluccan cuckoo-shrike Coracina atriceps Campephagidae Halmahera cuckoo-shrike Coracina parvula* Campephagidae Pale-grey cuckoo-shrike Coracina ceramensis Campephagidae Rufous-bellied triller Lalage aurea* Zosteropidae Cream-throated white-eye Zosterops atriceps* Dicaeidae Flame-breasted flowerpecker Dicaeum erythrothorax An asterisk signifies that the species' range is limited to this ecoregion. J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 8 The world's largest bee-the rare, 4-cm Wallace's giant bee Chalocodoma pluto-is also found on Bacan, Tidore, and Halmahera. Wallace discovered this species in 1858, and it was thought to be extinct until 1981, when it was recollected. This ecoregion also has conservation importance for butterflies and includes Troides aesacus, which may be the most primitive member of the T. priamus species group (Whitten and Whitten 1992; K. Monk, pers. comm., 2000). Current Status The rich volcanic soils of Ternate, Tidore, and nearby islands have been aggressively cultivated for cloves and other spices for centuries (Stattersfield et al. 1998). From the 1920s through the 1970s, commercial logging and enforced cultivation depleted the forests of Halmahera and Morotai (Monk et al. 1997). On Morotai, large tracts of lowland rain forest were cultivated with papaya (Carica papaya) during World War II (Monk et al. 1997). Currently, the wet evergreen lowland forests in the northwest of Halmahera are exploited by logging companies, primarily for the valuable damar trees (Agathis) (Whitten and Whitten 1992). The eastern forests are threatened by pulp plantations, especially using local transmigrants. Extensive habitat blocks still cover all the islands, with only small areas near the coast cleared for human settlements (Monk et al. 1997). The seven protected areas cover 4,880 km2 (18 percent) of the ecoregion area (table 3). Three protected areas are greater than 1,000 km2 in area, and the average size is 697 km2. Table 3. WCMC (1997) Protected Areas That Overlap with the Ecoregion. Protected Area Area (km2) IUCN Category Waya Bula 830 PRO Lolabata 1,210 PRO Gunung Gamkonora 110 PRO Ake Tajawi 1,200 PRO Saketa 1,100 PRO Gunung Sibela 300 PRO Pulau Obi 130 PRO Total 4,880 Ecoregion numbers of protected areas that overlap with additional ecoregions are listed in brackets. J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 9 Types and Severity of Threats With nearly 80 percent of its original forest still intact, the Halmahera Rain Forests [AA0106] ecoregion is largely free of intense habitat conversion threats. However, as the forests are lost on other Indonesian islands, there is an increasing potential for commercial forestry operations to move to Halmahera. A mining company, PT Halmahera Mineral (NHM), has already obtained an exploration license for Bacan and "neighboring islands" to look for gold and other minerals. A Canadian mining company has a license to mine nickel near Ake Tajawi on Halmahera (K. Monk, pers. comm., 2000). Justification of Ecoregion Delineation The Sula Islands were included within the Sulawesi Lowland Rain Forests, and the Aru islands in the Vogelkop-Aru Lowland Rain Forests. Buru Island, identified as a distinct subunit (13c) by MacKinnon (1997) and as an EBA (Stattersfield et al. 1998), was delineated as a distinct ecoregion, the Buru Rain Forests. Seram, the larger island to the east of Buru, was also delineated as an ecoregion: Seram Rain Forests. The larger Halmahera Rain Forests ecoregion includes Obi Island, which MacKinnon (1997) recognized as a separate subunit (13b) from Halmahera Island (subunit 13a). We created the Banda Sea Islands Moist Deciduous Forests by combining the islands in the Kai and Tanimbar archipelagos, which were distinguished as a biogeographic unit by Monk et al. (1997). The primary vegetation on the islands in both these archipelagos is moist deciduous forests and semi-evergreen forests, whereas the vegetation in the other, nearby large islands (Seram and Aru) is evergreen rain forests (Monk et al. 1997). References References for this ecoregion are currently consolidated in one document for the entire Indo-Pacific realm. Indo-Pacific Reference List This text was originally published in the book Terrestrial ecoregions of the Indo-Pacific: a conservation assessment from Island Press. This assessment offers an in-depth analysis of the biodiversity and conservation status of the Indo-Pacific's ecoregions. For more general information on this ecoregion, go to the WildWorld version of this description. All text by World Wildlife Fund © 2001 2. Banda Sea Islands moist deciduous forests J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 10 Banda Islands, Indonesia Photograph by Anasia-Cruise Southeastern Asia: Small islands scattered across the Banda Sea in Indonesia Tropical and Subtropical Moist Broadleaf Forests 2,900 square miles (7,500 square kilometers) -about half the size of Hawaii The Banda Sea Islands Moist Deciduous Forests are found on small islands scattered across the Banda Sea and are part of the region known as Wallacea, which contains a distinctive fauna representing a mix of Asian and Australasian species. Active volcanoes are found on the Banda Islands, whereas other parts of the ecoregion represent portions of the Australian continent that have been torn off. The islands contain a remarkable twenty-one bird species found nowhere else on Earth. The forests in this ecoregion are still largely intact, but although many of these islands are tiny and uninhabitable, the bird populations are seriously threatened by accidentally released rats and cats and by the removal of their eggs for sale by fishers traveling through this area. Vulnerable Location and General Description This ecoregion represents the moist deciduous and limestone forests of Tanimbar, Kai, Banda, and smaller island groups in the Banda Sea, part of the eastern Indonesian Archipelago. Based on the Köppen climate zone system, this ecoregion falls in the tropical wet climate zone (National Geographic Society 1999). Geologically, the islands have a mixed history. The Banda Islands are part of the inner arc, whereas the rest of the ecoregion is part of the outer arc. The inner arc islands are a result of the subduction and partial melting of the Indo-Australian tectonic plate below the Eurasian plate. The inner arc islands represent young volcanoes that have coalesced with lava and sediment. The volcanically active Mt. Api is found in the Banda Islands, which represent the ruins of a very large volcano that erupted in prehistory. The basement rock of the outer islands, on the other hand, is composed of actual continental margin from the Australian plate that has not been subducted. These outer islands are less than 4 million years old. The resulting surface geology consists of complex sedimentary and metamorphic rocks: uplifted coral reefs over complex basement rocks (Monk et al. 1997). In the south the forest biogeography of the Moluccas differs from that associated with the classic dipterocarp forests of Borneo or Sumatra. Northern Maluku has relatively similar dipterocarp forests. Many of the dipterocarp species have been replaced by dominants more typical of the Australo-Melanesian area. The forests of this ecoregion are varied but include evergreen rain forest (Kepulauan Kai), semi-evergreen rain forest, moist deciduous forest, and dry deciduous forest (Monk et al. 1997). Biodiversity Features The mammal fauna consists of twenty-two species with both Asian and J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 11 Australasian affinities, and three species are endemic (table 1) (Flannery 1995). The Moluccan mouse-eared bat (Myotis stalkeri) is endangered, whereas the dusky pademelon (Thylogale bruinii) and brown-bearded sheathtail bat (Taphozous achates) are considered vulnerable (IUCN 2000). The dusky pademelon is the only macropodid (kangaroo) found in the Banda Sea islands (Kai), although it is also found in the Aru Islands and the Trans Fly of New Guinea (Flannery 1995). Table 1. Endemic and Near-Endemic Mammal Species. Family Species Macropodidae Thylogale bruinii Vespertilionidae Myotis stalkeri* Emballonuridae Taphozous achates* An asterisk signifies that the species' range is limited to this ecoregion. Together with Halmahera, Buru, and Seram islands, this ecoregion lies within an area with perhaps the highest levels of bird endemism for its size anywhere in the world and the highest number of endemic birds of any area in Asia. Even the smallest, uninhabitable islands are significant as breeding sites for large numbers of seabirds such as frigatebirds, tropicbirds, boobies, terns, and smaller species (Whitten and Whitten 1992). Manuk Island and Mt. Api (north of Wetar), two nature reserves in the Banda Sea, are the breeding and roosting sites for millions of seabirds. Active volcanoes, they are probably the greatest bird islands left in all southeast Asia (Whitten and Whitten 1992; Monk et al. 1997). This ecoregion contains more than 225 species of terrestrial birds, of which forty-three are endemic or near endemic (table 2). This ecoregion corresponds with the Banda Sea Islands EBA, which contains forty-one restricted-range species, and includes eighteen species that are found nowhere else on Earth (Stattersfield et al. 1998). Only one of these species, the Damar flycatcher (Ficedula henrici), is considered threatened. Table 2. Endemic and Near-Endemic Bird Species. Family Common Name Species Megapodiidae Tenimbar megapode Megapodius tenimberensis* Megapodiidae Forsten's scrubfowl Megapodius forstenii Columbidae Dusky cuckoo-dove Macropygia magna Columbidae Wallace's fruit-dove Ptilinopus wallacii J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 12 Columbidae Pink-headed imperial-pigeon Ducula rosacea Cacatuidae Tanimbar cockatoo Cacatua goffini* Loriidae Red lory Eos bornea Loriidae Blue-streaked lory Eos reticulata* Loriidae Olive-headed lorikeet Trichoglossus euteles Cuculidae Green-cheeked bronze-cuckoo Chrysococcyx rufomerus* Cuculidae Pied bronze-cuckoo Chrysococcyx crassirostris Cuculidae Kai coucal Centropus spilopterus* Strigidae Moluccan hawk-owl Ninox squamipila Tytonidae Lesser masked-owl Tyto sororcula Alcedinidae Cinnamon-backed kingfisher Todirhamphus australasia Acanthizidae Rufous-sided gerygone Gerygone dorsalis Meliphagidae White-tufted honeyeater Lichmera squamata Meliphagidae Banda myzomela Myzomela boiei* Meliphagidae Black-faced friarbird Philemon moluccensis Eopsaltriidae Golden-bellied flyrobin Microeca hemixantha* Pachycephalida Drab whistler Pachycephala griseonota Pachycephalida Wallacean whistler Pachycephala arctitorquis* Rhipiduridae Cinnamon-tailed fantail Rhipidura fuscorufa* Rhipiduridae Long-tailed fantail Rhipidura opistherythra* Monarchidae White-naped monarch Monarcha pileatus Monarchidae Black-bibbed monarch Monarcha mundus* Monarchidae White-tailed monarch Monarcha leucurus* J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 13 Monarchidae Moluccan flycatcher Myiagra galeata Oriolidae Buru oriole Oriolus bouroensis Campephagidae Kai cuckoo-shrike Coracina dispar* Turdidae Slaty-backed thrush Zoothera schistacea* Turdidae Orange-banded thrush Zoothera peronii Turdidae Fawn-breasted thrush Zoothera machiki* Sturnidae Tanimbar starling Aplonis crassa* Muscicapidae Cinnamon-chested flycatcher Ficedula buruensis Muscicapidae Damar flycatcher Ficedula henrici* Zosteropidae Great Kai white-eye Zosterops grayi* Zosteropidae Little Kai white-eye Zosterops uropygialis* Sylviidae Timor stubtail Urosphena subulata Sylviidae Tanimbar bush-warbler Cettia carolinae* Estrildidae Tricolored parrotfinch Erythrura tricolor Dicaeidae Ashy flowerpecker Dicaeum vulneratum Dicaeidae Red-chested flowerpecker Dicaeum maugei An asterisk signifies that the species' range is limited to this ecoregion. Current Status This ecoregion consists of a chain of small islands. The forests in this ecoregion are still largely intact, with only about 20 percent of the habitat being lost. The island of Yamdena in the Tanimbars represents a fairly large block of undisturbed habitat. There are five protected areas that cover 1,500 km2 (27 percent) of the ecoregion (table 3). However, the largest of the protected areas is still in a proposed state. Table 3. WCMC (1997) Protected Areas That Overlap with the Ecoregion. Protected Area J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea Area (km2) IUCN Category 14 Kai Besar 290 PRO Pulau Nuswotar 40-75 I Jamdena 1,130 PRO Pulau Nustaram 30 I Pulau Lucipara 20 ? Pulau Angwarmase 10 I Gunung Api (north of Wetar) 1 ? Total 1,521 Ecoregion numbers of protected areas that overlap with additional ecoregions are listed in brackets. Types and Severity of Threats Although many of these islands are tiny and uninhabitable, the bird populations are seriously threatened by predatory rats and cats that have been accidentally released onto these islands and by the removal of their eggs for sale by fishers traveling through this area (Whitten and Whitten 1992). On inhabited islands, such as Manuk Island Nature Reserve, small-scale agriculture poses another threat as farmers oust both tree-nesting and ground-nesting birds (Whitten and Whitten 1992). Justification of Ecoregion Delineation The Sula Islands were included within the Sulawesi Lowland Rain Forests and the Aru Islands in the Vogelkop-Aru Lowland Rain Forests. Buru Island, identified as a distinct subunit (13c) by MacKinnon (1997) and as an EBA (Stattersfield et al. 1998), was delineated as a distinct ecoregion, the Buru Rain Forests. Seram, the larger island to the east of Buru, was also delineated as an ecoregion: Seram Rain Forests. The larger Halmahera Rain Forests includes Obi Island, which MacKinnon (1997) recognized as a separate subunit (13b) from Halmahera Island (subunit 13a). We created the Banda Sea Islands Moist Deciduous Forests by combining the islands in the Kai and Tanimbar archipelagos, which were distinguished as a biogeographic unit by Monk et al. (1997). The primary vegetation on the islands in both these archipelagos is moist deciduous forests and semi-evergreen forests, whereas the vegetation in the other, nearby large islands (Seram and Aru) is evergreen rain forests (Monk et al. 1997). References References for this ecoregion are currently consolidated in one document for the J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 15 entire Indo-Pacific realm. Indo-Pacific Reference List J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 16 3. Buru rain forests Satellite view of Buru Island, Indonesia Photograph by USGS The Buru Rain Forests are located on the small mountainous tropical island of Buru in the Banda Sea, part of the region known as Wallacea, which contains a distinctive fauna representing a mix of Asian and Australasian species. There are ten bird species in this ecoregion that are found nowhere else on Earth, including a monotypic bird genus. Although the northern portions of the island have been degraded by repeated burning and the coastal lowlands have been cleared, the remaining forest forms two large, contiguous blocks, current threats appear to be low, and the conservation outlook is relatively stable. Location and General Description This ecoregion represents the moist forests in the island of Buru. Based on the Köppen 3,300 square miles Southeastern climate zone system, this ecoregion falls in (8,600 square Asia: Island of kilometers) -the tropical wet climate zone (National Buru in about half the size Indonesia Geographic Society 1999). Buru is part of Hawaii remnant crustal fragment, probably from Tropical and the Australian continent, and part of the Vulnerable Subtropical Moist volcanic Inner Banda Arc. Consequently, Broadleaf Forests the surface geology of Buru is complex, consisting of older metamorphic schists and gneiss, younger volcanics, and recent alluvium (Monk et al. 1997). The natural vegetation of the island was tropical lowland evergreen and semi-evergreen rain forests (Stattersfield et al. 1998). The dominant tree species in this moist forest are the dipterocarps, Anisoptera thurifera, Hopea gregaria, H. iriana, H. novoguineensis, Shorea assamica, S. montigena, S. selanica, and Vatica rassak (Monk et al. 1997). In oldgrowth forests, the larger trees grow to more than 30 m in height and tend to be covered with thick-stemmed lianas and other epiphytes. Open forest, woodland, and savanna are also found in this ecoregion, with some being natural but most originating from human activity (Flannery 1995). The fire-resistant paper bark tree (Melaleuca cajuputi) is common and grows in nearly monotypic stands in dry areas (Whitten and Whitten 1992). The steep limestone cliffs in the northwestern part of the ecoregion are covered by mixed forests that include Shorea spp. (Monk et al. 1997). Exposed ridges between 1,800 and 2,000 m above sea level are characterized by stunted Dacrydium novo-guineense (Monk et al. 1997). Biodiversity Features J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 17 Overall richness and endemism in this ecoregion are low to moderate when compared with those of other ecoregions in Indo-Malaysia. Being in the Wallacean biogeographic zone, the ecoregion contains a mixture of Asian and Australian fauna. The mountainous areas of this island are largely unexplored and may contain many undiscovered species (Flannery 1995). The known mammal fauna of Buru consists of at least twenty-five species, including four near endemics (table 1). Two of these species are globally threatened: the vulnerable Seram flying-fox (Pteropus ocularis) and lesser tube-nosed fruit bat (Nyctimene minutus) (IUCN 2000). Table 1. Endemic and Near-Endemic Mammal Species. Family Species Pteropodidae Pteropus chrysoproctus Pteropodidae Pteropus ocularis Pteropodidae Nyctimene minutus Suidae Babyrousa babyrussa An asterisk signifies that the species' range is limited to this ecoregion. The bird fauna consists of 178 species, including twenty-nine endemic or near-endemic species (table 2). The ecoregion corresponds with the Buru EBA and contains twentyeight restricted-range bird species, ten of which are found nowhere else on Earth. Six of these species are considered vulnerable: Moluccan scrubfowl (Megapodius wallacei), blue-fronted lorikeet (Charmosyna toxopei), black-lored parrot (Tanygnathus gramineus), Buru cuckoo-shrike (Coracina fortis), streaky-breasted jungle-flycatcher (Rhinomyias addita), and rufous-throated white-eye (Madanga ruficollis), which represents a monotypic genus (Stattersfield et al. 1998). Table 2. Endemic and Near-Endemic Bird Species. Family Common Name Species Accipitridae Rufous-necked sparrowhawk Accipiter erythrauchen Megapodiidae Forsten's scrubfowl Megapodius forstenii Megapodiidae Moluccan scrubfowl Megapodius wallacei Columbidae White-eyed imperial-pigeon Ducula perspicillata J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 18 Columbidae Long-tailed mountain-pigeon Gymnophaps mada Psittacidae Buru racquet-tail Prioniturus mada* Psittacidae Black-lored parrot Tanygnathus gramineus* Loriidae Red lory Eos bornea Loriidae Blue-fronted lorikeet Charmosyna toxopei* Strigidae Moluccan hawk-owl Ninox squamipila Tytonidae Lesser masked-owl Tyto sororcula Meliphagidae Buru honeyeater Lichmera deningeri* Meliphagidae Wakolo myzomela Myzomela wakoloensis Meliphagidae Black-faced friarbird Philemon moluccensis Pachycephalida Drab whistler Pachycephala griseonota Rhipiduridae Cinnamon-backed fantail Rhipidura superflua* Monarchidae White-naped monarch Monarcha pileatus Monarchidae Black-tipped monarch Monarcha loricatus* Monarchidae Moluccan flycatcher Myiagra galeata Oriolidae Buru oriole Oriolus bouroensis Campephagidae Buru cuckoo-shrike Coracina fortis* Campephagidae Pale-grey cuckoo-shrike Coracina ceramensis Turdidae Moluccan thrush Zoothera dumasi Muscicapidae Streaky-breasted jungleflycatcher Rhinomyias addita* Muscicapidae Cinnamon-chested flycatcher Ficedula buruensis Zosteropidae Buru white-eye Zosterops buruensis* Zosteropidae Rufous-throated white-eye Madanga ruficollis* J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 19 Sylviidae Chestnut-backed bush-warbler Bradypterus castaneus Dicaeidae Flame-breasted flowerpecker Dicaeum erythrothorax An asterisk signifies that the species' range is limited to this ecoregion. Buru's butterflies include a large number of endemics and are therefore accorded highest conservation priority. Pifridae has 25 percent of the local species unique to Buru, and Papilionidae 7 percent (Vane-Wright and Peggae, in press). Current Status The coastal lowland forests have been cleared, and the northern and northeastern portions of the island now contain monsoon forest, gallery forest, and savannas as a result of repeated burning (Stattersfield et al. 1998). However, the remaining upland forest forms two large, contiguous blocks. Most of this forest is a mosaic of primary and secondary forest as a result of shifting cultivation (Monk 1997; Stattersfield et al. 1998). The two protected areas-of which one is greater than 1,000 km2-cover 17 percent of the ecoregion (table 3). Commercial logging on Buru intensified during the 1970s, but much of the island is still under extensive forest cover. Table 3. WCMC (1997) Protected Areas That Overlap with the Ecoregion. Protected Area Area (km2) IUCN Category Gunung Kelpat Muda 1,380 PRO Waeapo 50 PRO Total 1,430 Ecoregion numbers of protected areas that overlap with additional ecoregions are listed in brackets. Types and Severity of Threats Current threats to this ecoregion are low, causing its conservation status to remain vulnerable. Commercial logging and shifting cultivation are the primary threats to the remaining habitat. Justification of Ecoregion Delineation The Sula Islands were included within the Sulawesi Lowland Rain Forests and J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 20 the Aru Islands in the Vogelkop-Aru Lowland Rain Forests. Buru Island, identified as a distinct subunit (13c) by MacKinnon (1997) and as an EBA (Stattersfield et al. 1998), was delineated as a distinct ecoregion, the Buru Rain Forests. Seram, the larger island to the east of Buru, was also delineated as an ecoregion: Seram Rain Forests. The larger Halmahera Rain Forests includes Obi Island, which MacKinnon (1997) recognized as a separate subunit (13b) from Halmahera Island (subunit 13a). We created the Banda Sea Islands Moist Deciduous Forests [AA0102] by combining the islands in the Kai and Tanimbar archipelagos, which Monk et al. (1997) distinguished as a biogeographic unit. The primary vegetation on the islands in both these archipelagos is moist deciduous forests and semi-evergreen forests, whereas the vegetation in the other, nearby large islands (Seram and Aru) is evergreen rain forests (Monk et al. 1997). References References for this ecoregion are currently consolidated in one document for the entire Indo-Pacific realm. Indo-Pacific Reference List J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 21 4. Lesser Sundas deciduous forests Rinca Island, Indonesia Photograph by © WWF-Canon/Michel TERRETTAZ Southeastern Asia: Lesser Sundas Islands, Indonesia 15,200 square miles (39,400 square kilometers) -- about twice the size of Massachusetts The Lesser Sundas Deciduous Forests are found on a string of volcanic islands. They stretch across the Java Sea between Australia and Borneo. It is part of a unique biogeographic region known as Wallacea, which contains a very distinctive fauna representing a mix of Asian and Australasian species. These distinctive seasonal dry forests harbor unique species, including the Komodo dragon, the largest lizard in the world, and seventeen bird species found nowhere else on Earth. A combination of shifting agriculture and humancaused fires has significantly reduced the amount of natural forest in this ecoregion. Location and General Description This ecoregion represents the semievergreen dry forests in the Lesser Sunda Islands. It extends east from the islands of Tropical and Lombok and Sumbawa to Flores and Alor in Subtropical Dry Critical/Endangered Broadleaf the Indonesian Archipelago. Rinjani Forests volcano on Lombok is the highest mountain in the ecoregion, at 3,726 m. The Lesser Sundas are an inner volcanic island arc, created by the subduction and partial melting of the Australian tectonic plate below the Eurasian plate. The islands represent tertiary and quaternary volcanoes that have coalesced with lava and sediment. There is actually a geologic discontinuity between Lombok and Sumbawa, on the Sunda Arc, and the rest of the islands, part of the Banda Arc. With the exception of Komodo, which is Mesozoic, most of the islands were built during two pulses in the Tertiary (Mio-Pliocene) and Quaternary (recent) (Monk et al. 1997). This ecoregion is separated from Bali and Java to the west by Wallace's Line, which marks the end of the Sunda Shelf. With an average annual rainfall of 1,349 mm, this region is the driest but also the most seasonal in Indonesia. Based on the Köppen climate system, this ecoregion has a tropical dry climate zone (National Geographic Society 1999). This distinctive climate has given rise to a vegetation that is strikingly different from that of the rest of the archipelago. Much of the natural habitat is composed of monsoon forests and savanna woodlands (Whitten and Whitten 1992). The monsoon forests consist of several forest subtypes, notably moist deciduous forest, dry deciduous forest, dry thorn forest, and dry evergreen forest. Moist deciduous forests also occur as a band of lowland forest at the base of the hills and as gallery forests along streams, especially on Komodo Island. Dominant trees include Tamarindus indica and Sterculia foetida (Monk et al. 1997). The dry deciduous forest at altitudes below 200 m is J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 22 dominated by Protium javanicum, Schleichera oleosa, and Schoutenia ovata, whereas at medium altitudes, from 200 to 800 m, the dominant tree is Tabernaemontana floribunda. At these altitudes, lianas and climbers become common, especially the white-flowered liana Bauhinia. Above 1,000 m, Euphorbiaceae tend to become common and well represented (Monk et al. 1997). Dry thorn forest is another type of monsoon forest in this ecoregion, although little is left because it has been cleared by setting fires. This forest formation still exists along the southeast coast of Lombok and the southwest coast of Sumbawa but is being cleared in the latter region for road building, mine development, and a transmigration site (Monk et al. 1997). Dry evergreen forest occurs above dry deciduous forest and below the true evergreen montane forest, at 1,000 m above sea level on Mt. Batulante in northwest Sumbawa. Below 1,200 m on the north slopes, Albizia chinensis is a characteristic species. Other common species include Chionanthus, Prunus, and two Cryptocarya species. On many islands, drier areas in steep-sided valleys contain gallery forest. On Sumbawa, for instance, gallery forest is found from sea level to 2,000 m above sea level and is also present in lower montane forests (Monk et al. 1997). By contrast, the southern hill slopes along the southern coasts are kept moist during the dry season by the southeast trade winds, and dipterocarp rain forest occurs on the southwest hills of both Lombok and Sumbawa. Lombok also contains one of the few remaining patches of tropical semievergreen rain forest, at volcanic Mt. Rinjani, which acts as the major water catchment area for the whole island (Monk et al. 1997). Twenty-meter-high mixed montane forests of Podocarpus and Engelhardia are found from about 1,200 to 2,100 m, with lianas, epiphytes, and orchids such as Corybas, Corymborkis, and Malaxis very much in evidence. At higher elevations of up to 2,700 m, Casuarina junghuhniana forests occur. Toward the summit, from 3,300 to 3,400 m, the rocky ridges were once covered with lichens, mosses, grasses, herbs, and some ferns but are now being eroded. On Sumbawa, the south slopes of Mt. Batulante above 1,000 m are covered with a Cryptocarya-Meliaceae montane forest, although species composition varies with moisture. This forest is dominated by two species of Cryptocarya, one in the drier and usually lower forest (from 1,000 to 1,500 m above sea level) and the other at higher or moister sites. Drier, stonier slopes in poorer forest are the only places where lianas are common. Further east, from the eastern part of Flores to Alor, the forests are dominated by Pterocarpus indicus (Monk et al. 1997). There are also two types of savanna in this ecoregion: a Borassus flabellifer savanna that occurs from sea level to 400 m on Komodo, Rinca, and the north and south coasts of Flores; and the Ziziphus mauritiana savanna, which occurs on more sandy clay alluvial, and sometimes water-logged, soil. The dominant grasses are Eulalia leschenaultiana, spear grass (Heteropogon contortus, Themeda frondosa), and Themeda triandra (Monk et al. 1997). Biodiversity Features J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 23 This area, part of the Wallacean sub-region, includes a mix of Asian and Australian fauna, and because of the long years of isolation from the mainland it harbors many endemic mammals and birds. Most of the endemic mammals occur on Komodo and Flores eastward, rather than Lombok and Sumbawa. One of the important and better-known endemic species in this ecoregion is the Komodo dragon (Varanus komodoensis), the largest lizard in the world. The mammal faunal in this ecoregion consists of fifty species, including five ecoregional endemics, including the critically endangered Flores shrew (Suncus mertensi) and the vulnerable Komodo rat (Komodomys rintjanus) (table 1) (IUCN 2000). With the exception of the New Caledonia dry forests, with six endemic mammals, the five endemic mammals in this ecoregion are more than are found in any other dry forest ecoregion in the Indo-Pacific. Table 1. Endemic and Near-Endemic Mammal Species. Family Species Soricidae Suncus mertensi* Pteropodidae Pteropus lombocensis* Vespertilionidae Nyctophilus heran* Muridae Bunomys naso* Muridae Komodomys rintjanus* An asterisk signifies that the species' range is limited to this ecoregion. This ecoregion also harbors about 273 bird species, of which 29 are endemic or near endemic (table 2). The ecoregion is consistent with the Northern Nusa Tenggara EBA (Stattersfield et al. 1998). Of the twenty-nine restricted-range species in the EBA, seventeen are found nowhere else in the world. Three are endemic and threatened, including the endangered Flores monarch (Monarcha sacerdotum) and the vulnerable Wallace's hanging-parrot (Loriculus flosculus) and Flores crow (Corvus florensis). In addition, the white-rumped kingfisher (Caridonax fulgidus) is the sole representative of an endemic monotypic genus. Table 2. Endemic and Near-Endemic Bird Species. Family Common Name Species Columbidae Dusky cuckoo-dove Macropygia magna J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 24 Columbidae Flores green-pigeon Treron floris* Columbidae Pink-headed imperial-pigeon Ducula rosacea Columbidae Dark-backed imperial-pigeon Ducula lacernulata Psittacidae Wallace's hanging-parrot Loriculus flosculus* Loriidae Olive-headed lorikeet Trichoglossus euteles Strigidae Flores scops-owl Otus alfredi* Strigidae Wallace's scops-owl Otus silvicola* Alcedinidae Cinnamon-backed kingfisher Todirhamphus australasia Alcedinidae White-rumped kingfisher Caridonax fulgidus* Meliphagidae Sunda honeyeater Lichmera lombokia* Pachycephalida Bare-throated whistler Pachycephala nudigula* Rhipiduridae Brown-capped fantail Rhipidura diluta* Monarchidae Flores monarch Monarcha sacerdotum* Corvidae Flores crow Corvus florensis* Campephagidae Sumba cuckoo-shrike Coracina dohertyi Campephagidae Flores minivet Pericrocotus lansbergei* Turdidae Chestnut-backed thrush Zoothera dohertyi Muscicapidae Flores jungle-flycatcher Rhinomyias oscillans Zosteropidae Yellow-spectacled white-eye Zosterops wallacei Zosteropidae White-browed white-eye Lophozosterops superciliaris* Zosteropidae Dark-crowned white-eye Lophozosterops dohertyi* Zosteropidae Flores white-eye Heleia crassirostris* Sylviidae Russet-capped tesia Tesia everetti* J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 25 Sylviidae Timor leaf-warbler Phylloscopus presbytes Dicaeidae Golden-rumped flowerpecker Dicaeum annae* Dicaeidae Black-fronted flowerpecker Dicaeum igniferum* Dicaeidae Red-chested flowerpecker Dicaeum maugei Nectariniidae Flame-breasted sunbird Nectarinia solaris An asterisk signifies that the species' range is limited to this ecoregion. The Komodo dragon deserves special mention. It is the largest lizard species in the world. Varanus komodoensis occupies five islands: Komodo, Padar, Rinca, Gili Motang, and Flores. These animals range from sea level to approximately 450 m in elevation, mainly in tropical deciduous monsoon forest, tropical savanna, and grassland. They feed on a wide variety of animal food, including insects, lizards, snakes, birds, deer, wild boar, monkeys, and bird eggs; they also feed on carrion. Adults may have a foraging range of 500 ha. There are approximately 4,000 protected individuals in Komodo National Park (Monk et al. 1997). Current Status During World War II, logging and cultivation destroyed much of the forest cover of Lombok, east Flores, and the small islands of Adonara, Solor, Lomblen, Pantar, and Alor (Monk et al. 1997). The Lombok dipterocarp forest is almost depleted by commercial logging, and the forest of Sumbawa is partially covered by a mining concession (Monk et al. 1997). More than half of this ecoregion's natural habitat has been cleared, mainly for agriculture. Except for the island of Sumbawa, which still contains a large block of intact forest, most of the islands in this group have only fragments of natural habitat remaining. The twentyeight protected areas include about 10 percent of the ecoregion area, but most of the protected areas are small, with the average size being only 144 km2 (table 3). Komodo National Park, the most famous wild area in the Lesser Sundas, a World Heritage Site, and an important tourist destination, is only one of two reserves that is greater than 500 km2, but most of this park is marine. The park harbors the Komodo dragon. Lowlands are underrepresented in the protected areas system even though this habitat supports most of the ecoregion's species; for instance, most of Sumbawa's endemic birds are associated with lowland monsoon forest (Monk et al. 1997). Table 3. WCMC (1997) Protected Areas That Overlap with the Ecoregion. Protected Area J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea Area (km2) IUCN Category 26 Pulau Sangiang 150 PRO Pulau Moyo 160-222 VI Tambora Utara GR 480 PRO Tambora Selatan 150 VI Gunung Rinjani 830 + 760 ext. II Pulau Panjang 200 PRO Hutan Dompu Complex 110 PRO Gunung Olet Sangenges NR 280-350 PRO Suranadi 5 V Pulau Rakit 20 PRO Batu Gendang Forest 150 PRO Pantai Palolowaru 5 PRO Selahu Legini Complex 320-500 VI Kurung Baya/Varanus 40 PRO East Timor 210 PRO Tanjung Kerita Mese 300 PRO Danau Rana Mese 2 PRO Danau Kelimutu 20 DE Danau Sano 8 PRO Gunung Ambu Lombo 40 PRO Tuti 60 V Tanjung Watupayung 90 PRO Hadekawa-Labelakang 250 PRO Gunung Muna 100-150 PRO J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 27 Adonara NR 20 PRO Pulau Rusa 10 PRO Egon-Iliwuli 30 PRO Lewotobi 8 VI Total 4,048+ Ecoregion numbers of protected areas that overlap with additional ecoregions are listed in brackets. Increasing population pressure has also resulted in high rates of deforestation (WWFIndonesia n.d.). In the dry season, fires often are set to clear the understory and to encourage new growth as forage for domestic animals. This has been done since prehistoric times-not for domestic animals, but for attracting game (introduced) into areas of new grass growth. This practice has resulted in the proliferation of fire-resistant trees such as Casuarina junghuhniana and formation of grassland over an extensive area of these islands. Most of the remaining forest is confined to the steepest slopes and the tops of mountains (Whitten and Whitten 1992). Poorly managed tourism, especially in the Komodo National Park and on Lombok, has also caused environmental degradation (WWF-Indonesia n.d.). Types and Severity of Threats The future threats will continue to be deforestation, an increasing population and their demands on the environment, intentionally set fires, and agricultural land development. Justification of Ecoregion Delineation The drier forests in Nusa Tenggara were placed in three ecoregions that corresponded to the biogeographic units identified in Monk et al (1997). These are Lesser Sundas Deciduous Forest, which includes the chain of islands extending from Lombok, Sumbawa, Komodo, Flores, and the smaller satellite islands corresponding to the Flores biogeographic unit; Timor and Wetar Deciduous Forests, corresponding to the Timor biogeographic unit; and the Sumba Deciduous Forests, corresponding to the Sumba biogeographic unit. All three ecoregions belong to the tropical dry forests biome. References References for this ecoregion are currently consolidated in one document for the entire Indo-Pacific realm. Indo-Pacific Reference List J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 28 5. Seram rain forests The Seram Rain Forests are part of the region known as Wallacea, which contains a very distinctive fauna representing a mix of Asian and Australasian species. This small island ecoregion contains sixteen bird species, including a monotypic bird genus, that are found nowhere else on Earth. Nearly a fifth of the original forest cover has been cleared, mostly along the northern coast. However, large areas of contiguous, intact forest still exist, and the conservation status of this ecoregion is relatively stable. Location and General Description Seram Island, Indonesia Photograph by Anasia-Cruise Southeastern Asia: Island of Seram in Indonesia 7,500 square miles (19,400 square kilometers) -about the size of New Jersey Tropical and Relatively Subtropical Moist Stable/Intact Broadleaf Forests This ecoregion represents the semievergreen and moist forests of Seram and associated islands in the easternmost section of the Indonesian Archipelago. Based on the Köppen climate zone system, this ecoregion falls in the tropical wet climate zone (National Geographic Society 1999). Seram is part remnant crustal fragment, probably from the Australian continent, and part of the volcanic Inner Banda Arc. Consequently, the surface geology of Seram is complex, consisting of older metamorphic schists and gneiss, younger volcanics, and recent alluvium (Monk et al. 1997). The interior of the island is mountainous, with several ranges reaching more than 1,000 m. The highest point on the island is the 3,027-m Merkele ridge (Stattersfield et al. 1998). The natural vegetation of Seram is tropical lowland evergreen, semi-evergreen, and montane rain forest (Stattersfield et al. 1998). Semi-evergreen rain forest with trees that reach 30 m or more is a predominant forest type in this ecoregion. Rattans that exceed 100 m can be found in mature forests. The middle and lower layers include representatives of the Amaryllidaceae, sedges, and large ferns Angiopteris and Marattia, as well as climbers such as Freycinetia, Gnetum, Mucuna, Bauhinia, Piper, and Smilax. Most of the remaining dipterocarp forests are dominated by the endemic Shorea selanica, which can represent about 30 percent of the individual trees and 76 percent of the basal area in the forest. Also common are Anisoptera thurifera, Hopea gregaria, H. iriana, H. novoguineensis, Shorea assamica, S. montigena, S. selanica, and Vatica rassak (Monk et J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 29 al. 1997). This ecoregion also contains patches of ultrabasic rocks. The forests on these soils generally are poor in species, low, and shrubby. Tertiary limestone outcrops occur in the lowlands and on many mountains such as the Murkele Ridge and the top ridge of the central Mt. Binaiya, Seram's highest mountain (Monk et al. 1997). In Seram's montane forests, the Fagaceae are represented by only two species. Castanopsis buruana dominates between 400 and 1,400 m above sea level, where individuals tend to clump together, and Lithocarpus celebicus is found along ridges. Above 2,400 m on Mt. Binaiya, a low, open scrubby woodland contains Dacrydium spp., Myrica spp., Rapanea spp., Rhamnus spp., Rhododendron spp., and Vaccinium spp. Tree ferns are also important and include Cyathea binayana and C. pukuana, which form distinctive groves that support many epiphytic ferns. Pockets of this tree-fern savanna extend to the summit along with low Vaccinium woodland. At the highest points, from 2,700 to 3,000 m above sea level, grassland dominates and is characterized by several endemic herbs such as Viola binayensis, Pterostylis papuanum, and Euphrasia ceramensis (Monk et al. 1997). Biodiversity Features The overall richness and endemism of this ecoregion are low to moderate when compared with those of other ecoregions in Indo-Malaya. The islands are part of Wallacea, a unique region that supports a mixture of Asian and Australian fauna. The montane area of Seram supports the greatest number of endemic mammals of any island in the region (Flannery 1995). The ecoregion harbors thirty-eight mammal species and includes nine species that are endemic or near endemic (table 1), several of which are limited to montane habitats (Flannery 1995). The Seram flying-fox (Pteropus ocularis) and spiny Seram rat (Melomys feliceus) are considered vulnerable (IUCN 2000). The mammals found on Seram include Asian species (Murid rodents) as well as Australasian marsupials. Table 1. Endemic and Near-Endemic Mammal Species. Family Species Perorictidae Rhyncholemes prattorum* Pteropodidae Pteropus chrysoproctus Pteropodidae Pteropus ocularis Pteropodidae Pteropus argenatatus* Muridae Rattus feliceus* J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 30 Muridae Melomys fulgens Muridae Melomys aerosus* Muridae Melomys fraterculus* Muridae Stenomys ceramicus* An asterisk signifies that the species' range is limited to this ecoregion. The ecoregion harbors more than 213 bird species (Wikramanyake et al. 2001), of which 33 are endemic or near endemic (table 2). The ecoregion corresponds to the Seram EBA. The EBA contains thirty restricted-range species, including fourteen that are found nowhere else on Earth. Five species are threatened. The vulnerable Moluccan scrubfowl (Megapodius wallacei) is also found on Buru and Halmahera. The remaining four species are found nowhere else: the endangered black-chinned monarch (Monarcha boanensis) and vulnerable salmon-crested cockatoo (Cacatua moluccensis), purple-naped lory (Lorius domicella), and lazuli kingfisher (Todirhamphus lazuli). The bicoloured whiteeye (Tephrozosterops stalkeri), the sole member of its genus, is also found only on Seram. The fourteen endemic restricted-range birds can be divided into three groups: five species found generally in lowland forests (below 1,000 m), three species found in montane forests above 1,000 m, and six species found in both lowland and montane habitats (Stattersfield et al. 1998). The ecoregion also harbors the largest bird in the Moluccas, the two-wattled cassowary (Casuarius casuarius) (Whitten and Whitten 1992). Table 2. Endemic and Near-Endemic Bird Species. Family Common Name Species Accipitridae Rufous-necked sparrowhawk Accipiter erythrauchen Megapodiidae Forsten's scrubfowl Megapodius forstenii Megapodiidae Moluccan scrubfowl Megapodius wallacei Columbidae White-eyed imperial-pigeon Ducula perspicillata Columbidae Long-tailed mountain-pigeon Gymnophaps mada Cacatuidae Salmon-crested cockatoo Cacatua moluccensis* Loriidae Red lory Eos bornea Loriidae Blue-eared lory Eos semilarvata* J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 31 Loriidae Purple-naped lory Lorius domicella* Cuculidae Pied bronze-cuckoo Chrysococcyx crassirostris Strigidae Moluccan hawk-owl Ninox squamipila Tytonidae Lesser masked-owl Tyto sororcula Alcedinidae Lazuli kingfisher Todirhamphus lazuli* Meliphagidae Olive honeyeater Lichmera argentauris Meliphagidae Seram honeyeater Lichmera monticola* Meliphagidae Seram myzomela Myzomela blasii* Meliphagidae Wakolo myzomela Myzomela wakoloensis Meliphagidae Seram friarbird Philemon subcorniculatus* Pachycephalida Drab whistler Pachycephala griseonota Rhipiduridae Streaky-breasted fantail Rhipidura dedemi* Monarchidae Black-chinned monarch Monarcha boanensis* Monarchidae Moluccan flycatcher Myiagra galeata Oriolidae Seram oriole Oriolus forsteni* Campephagidae Moluccan cuckoo-shrike Coracina atriceps Campephagidae Pale-grey cuckoo-shrike Coracina ceramensis Turdidae Moluccan thrush Zoothera dumasi Sturnidae Long-crested myna Basilornis corythaix* Muscicapidae Cinnamon-chested flycatcher Ficedula buruensis Zosteropidae Ambon white-eye Zosterops kuehni* Zosteropidae Bicoloured white-eye Tephrozosterops stalkeri* Zosteropidae Grey-hooded white-eye Lophozosterops pinaiae* Sylviidae Chestnut-backed bush-warbler Bradypterus castaneus J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 32 Dicaeidae Ashy flowerpecker Dicaeum vulneratum An asterisk signifies that the species' range is limited to this ecoregion. Current Status Nearly a fifth of the original forest of this ecoregion has been cleared, mostly along the northern coast. However, large areas of contiguous, intact forest still exist. Therefore, the conservation status of this ecoregion is relatively stable. Seven protected areas cover 3,121 km2 (16 percent) of the ecoregion area, and one-Manusela National Park-is more than 2,000 km2 (table 3). This last reserve, with a wide range of forest types, conserves the cassowary (Casuarius casuarius). However, wildlife trade has been strong since historical times, and it may threaten some bird species such as the salmon-crested cockatoo (Cacatua moluccensis). The Trans-Seram Highway also threatens forest habitat by illegal logging, land clearance, and soil erosion. Table 3. WCMC (1997) Protected Areas That Overlap with the Ecoregion. Protected Area Area (km2) IUCN Category Sabuda Tataruga 10 IV Manusela 2,340 II Wae Bula 600 PRO Gunung Sahuai 120 PRO Pulau Kassa 1 IV Pulau Pombo 20 I Laut Banda 30 I Pulau Manuk 1 ? Total 3,122 Ecoregion numbers of protected areas that overlap with additional ecoregions are listed in brackets. Seram's moist lowland forests are being exploited by logging companies, primarily for their valuable damar trees (Agathis) (Whitten and Whitten 1992). The best dipterocarp stands were depleted by commercial loggers before the 1950s, and many other species J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 33 were overexploited by intensive logging in the 1970s (Monk et al. 1997). With no airport and only rudimentary ground transport, Seram is remote. Although this promotes conservation in many ways, it also prevents conservation employees from guarding boundaries, enlisting the support of local people, and conducting biological surveys (Whitten and Whitten 1992). Types and Severity of Threats The north Seram dipterocarp forests are still dominated by the endemic Shorea selanica and therefore are especially vulnerable to logging (Monk et al. 1997). The commercial wildlife trade is another significant threat. Parrots are captured and exported for the pet trade, with many casualties (Whitten and Whitten 1992). Justification of Ecoregion Delineation The Sula Islands were included within the Sulawesi Lowland Rain Forests and the Aru Islands in the Vogelkop-Aru Lowland Rain Forests. Buru Island, identified as a distinct subunit (13c) by MacKinnon (1997) and as an EBA (Stattersfield et al. 1998), was delineated as a distinct ecoregion, the Buru Rain Forests. Seram, the larger island to the east of Buru, was also delineated as an ecoregion: Seram Rain Forests. The larger Halmahera Rain Forests ecoregion includes Obi Island, which MacKinnon (1997) recognized as a separate subunit (13b) from Halmahera Island (subunit 13a). We created the Banda Sea Islands Moist Deciduous Forests by combining the islands in the Kai and Tanimbar archipelagos, which were distinguished as a biogeographic unit by Monk et al. (1997). The primary vegetation on the islands in both these archipelagos is moist deciduous forests and semi-evergreen forests, whereas the vegetation in the other, nearby large islands (Seram and Aru) is evergreen rain forests (Monk et al. 1997). References References for this ecoregion are currently consolidated in one document for the entire Indo-Pacific realm. Indo-Pacific Reference List J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 34 6. Sulawesi lowland rain forests Dumoga National Park, Sulawesi, Indonesia Photograph by © WWF-Canon/William F. RODENBURG Indonesia: Island 44,900 square miles (116,300 of Sulawesi square kilometers) -- about the size of Tropical and Pennsylvania Subtropical Moist Broadleaf Forests Relatively Stable/Intact The Sulawesi Lowland Rain Forests harbor some of the most unique animals on Earth. The islands are located in the region known as Wallacea, which contains a distinctive fauna representing a mix of Asian and Australasian species. A fruit-eating pig with huge tusks, a dwarf buffalo, endemic macaques, and cuscuses exemplify a truly unique mammal community. Sulawesi, like the hub of a wheel, is surrounded by a variety of exotic ocean basins, including the Flores Sea, the Banda Sea, the Molucca Sea, the Java Sea, and the Straits of Makassar, as well as the diverse islands of Borneo, Java, Flores, Halmahera, and the Philippines. More than half of the original forest has been cleared, and most of the remaining forests have been reduced to fragments. Location and General Description This ecoregion represents the lowland forests (less than 1,000 m) on Sulawesi and the surrounding islands of Banggai and Sula to the east and Talaud and Sangihe to the north. Sulawesi is almost completely mountainous. There are no extensive lowlands on Sulawesi, with large areas above 1,000 m and the highest elevation at 3,455 m on Mt. Rantemario. Sangihe is mountainous, reaching an elevation of 1,784 m, whereas Talaud is low-lying. The physiography of the Sula Islands is hilly, with mountains over 800 m only on the island of Taliabu (Stattersfield et al. 1998). The upland areas (more than 1,000 m) of Sulawesi form a separate ecoregion, the Sulawesi montane rain forests. Based on the Köppen climate zone system, this ecoregion falls in the tropical wet climate zone (National Geographic Society 1999). Sulawesi has a complex geologic history and is composed of three geologic provinces based on that history. West and East Sulawesi form two of the geologic provinces, separated by the Palu-Koro fault, which runs from the town of Palu to the Gulf of Bone. The third geologic province consists of the Tokala region on the northeast peninsula, the Banggai Islands, Butung Island, and the Sula Islands. East and West Sulawesi collided approximately 13-19 million years ago, and ultrabasic rocks were exposed as East Sulawesi overrode the western portion. The forces that caused the collision are still at work, and Sulawesi is being torn apart today. The surface geology of Sulawesi is a diverse patchwork of ophiolites, Mesozoic sedimentary rocks, Tertiary sedimentary and igneous rocks, and Quaternary volcanics and sediments. Active volcanoes are located on J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 35 the northern arm of Sulawesi (Whitten et al. 1987). The lowland forest is predominantly tropical lowland evergreen and semi-evergreen rain forest, with some monsoon forests at the tip of the southeast peninsula and small areas of freshwater and peat swamp forest (Whitten et al. 1987; Stattersfield et al. 1998). Distinctive forest types on limestone are distributed around southern Sulawesi and on ultrabasic soils in scattered locations all around the island. The lowland and hill forests contain the most tree species, although these forests are not dominated by any one tree family; only seven dipterocarp species are found in Sulawesi (compared with 267 and 106 in Borneo and Sumatra, respectively). The dipterocarps include Anisoptera costata, Hopea celebica, H. gregaria, Shorea assamica, Vatica rassak, and V. flavovirens. Distinctive ebonies (Diospyros spp.) were common in dense clumps in the lowland forests. Palms are common in the lowland forest, including Oncosperma horridum, Liculala celebensis, Pinanga, Areca, Caryota, and Livistona rotundifolia (Whitten et al. 1987). The isolated Sula Islands, just off Sulawesi's east coast, receive rain from both the northwest and the southeast and have volcanic soils that create excellent growth conditions (Monk et al. 1997). Aopa Swamp, 100 km west of Kendari, is a major area of peat swamp that varies seasonally in extent from about 150 to 314 km2. The dominant tree species in this forest include Casuarina spp., Eugenia spp., Geunsia paloensis, Premna foetida, Metroxylon sagu, Pholidocarpus spp., Licuala spp., Arenga spp., Oncosperma spp., and Corypha spp. Sedges such as Scleria spp. also occur along with 5-m tall Pandanus spp., at least two species of climbing rattan, and epiphytic Lecanopteris ant-ferns (Whitten et al. 1987). Freshwater swamp forest is characterized by grassy areas near open water, with palms and pandans on firmer ground and ubiquitous pitcher plants (Nepenthes). Riverine forest dominated by tall Eucalyptus deglupta is found in the Sopu Valley northeast of Lake Lindu and Mt. Nokilalaki (Whitten et al. 1987). This ecoregion also includes karst (limestone) areas that have a relative paucity of trees and tree species because of their shallow soils and steep slopes, resulting from the high solubility of limestone rocks. High calcium levels in the soil give rise to distinctive tolerant plant communities but support certain snail species limited to limestone forest as well as the large swallowtail butterfly (Graphium androcles) (Whitten et al. 1987). Infertile ultrabasic substrates, with serpentine and peridotite rocks, contain unique forests with a high degree of plant endemism. Common species include ironwood (Metrosideros), Agathis, Calophyllum, Burseraceae, Sapotaceae, and dipterocarps (Vatica and Hopea celebica). Myrtaceae (Eugenia, Kjellbergiodendron, and Metrosideros) are dominant in the low and regular canopy. There is little marketable timber in such forests (Whitten et al. 1987). J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 36 Biodiversity Features Wallace's Line, running from between Bali and Lombok and between Sulawesia and Borneo, marks the location of a deep oceanic trench and the point over which land animals and plants could not cross easily. Similarly, Lydekker's Line, running from between Timor and the Australian shelf to between Halmahera, Seram, and New Guinea, marks the point where Australasian flora and fauna could not easily pass. Sulawesi lies between these two lines. Sulawesi's location, geologic history, and long geographic isolation have created Sulawesi's distinctive fauna. There is variability, different among various animal and plant groups, in the amount of interchange between other biogeographic areas in the region, which led to the evolution of a large number of species endemic to the island. Although not species-rich relative to Borneo or Java, Sulawesi is high in endemicity because of its long isolation from Asia and Australia in Wallacea. This ecoregion exhibits high plant endemism, and the several distinct forest types provide habitat for the highest number of endemic mammals in Asia and several endemic birds (Whitten et al. 1987). Of the 104 mammal species in the ecoregion, 29 are endemic or near endemic (table 1). Whereas the two cuscuses have Australasian affinities (i.e., the Peleng cuscus [Phalanger pelengensis] and dwarf cuscus [Strigocuscus celebensis]), the remainder of Sulawesi's mammals have Asian origins, including the crested macaque (Macaca nigra), moor macaque (M. maura), booted macaque (M. ochreata), lowland anoa (Bubalus depressicornis), spectral tarsier (Tarsius spectrum), and babirusa (Babyrousa babyrussa) (Flannery 1995). The crested macaque, moor macacque, and lowland anoa are considered endangered (IUCN 2000). Table 1. Endemic and Near-Endemic Mammal Species. Family Species Phalangeridae Phalanger pelengensis* Sorcidae Crocidura elongata Sorcidae Crocidura lea Sorcidae Crocidura levicula Pteropodidae Acerodon humilis* Pteropodidae Neopteryx frosti* Pteropodidae Nyctimene minutus Rhinolophidae Hipposideros inexpectatu J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 37 Vespertilionidae Pipistrellus minahassae Vespertilionidae Hesperoptenus gaskelli Tarsiidae Tarsius pelengensis Cercopithecidae Macaca maura* Cercopithecidae Macaca ochreata* Cercopithecidae Macaca nigra Suidae Babyrousa babyrussa Bovidae Bubalus depressicornis* Muridae Rattus koopmani* Muridae Rattus xanthurus Muridae Rattus bontanus* Muridae Rattus elaphinus* Muridae Maxomys hellwaldii* Muridae Haeromys minahassae* Muridae Margaretamys beccarii* Muridae Taeromys celebensis* Muridae Taeromys punicans* Muridae Taeromys taerae* Muridae Echiothrix leucura* Muridae Melomys fulgens Muridae Melomys caurinus* An asterisk signifies that the species' range is limited to this ecoregion. Sulawesi contains a depauperate bird fauna but with high levels of endemicity (Stattersfield et al. 1998). The origin of Sulawesi's birds is predominantly Asian (Whitten et al. 1987). The bird fauna consists of about 337 species, of which 70 are endemic or J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 38 near-endemic species (table 2). The ecoregion also overlaps the lowland portions of the Sulawesi EBA and completely overlaps both the Sangihe and Talaud and Banggai and Sula Islands EBAs and the Salayar and Bonerate Islands Secondary Area (Stattersfield et al. 1998). Of the seventy restricted-range birds in these three EBAs, thirty-two bird species are found nowhere else in the world but this lowland ecoregion. Thirteen additional species are found only in this ecoregion and the adjacent montane ecoregion (and nineteen more species are found only in the uplands of Sulawesi) (Stattersfield et al. 1998). One species found on Sangihe, the cerulean paradise flycatcher (Eutrichomyias rowleyi), is critically endangered, and the red-and-blue lory (Eos histro), Sangihe hanging-parrot (Loriculus catamene), and elegant sunbird (Aethopyga duyvenbodei), all found on Sangihe and Talaud, are endangered (Stattersfield et al. 1998). Table 2. Endemic and Near-Endemic Bird Species. Family Common Name Species Accipitridae Small sparrowhawk Accipiter nanus Megapodiidae Sula scrubfowl Megapodius bernsteinii* Megapodiidae Maleo Macrocephalon maleo Rallidae Platen's rail Aramidopsis plateni Rallidae Bare-faced rail Gymnocrex rosenbergii Rallidae Isabelline waterhen Amaurornis isabellinus Columbidae Dusky cuckoo-dove Macropygia magna Columbidae Sulawesi ground-dove Gallicolumba tristigmata Columbidae Maroon-chinned fruit-dove Ptilinopus subgularis* Columbidae White-bellied imperial-pigeon Ducula forsteni Columbidae Pink-headed imperial pigeon Ducula rosacae Columbidae Grey-headed imperial-pigeon Ducula radiata Columbidae Grey imperial-pigeon Ducula pickeringii Columbidae Silver-tipped imperial-pigeon Ducula luctuosa* Psittacidae Yellowish-breasted racquet-tail Prioniturus flavicans* Psittacidae Moluccan hanging-parrot Loriculus amabilis J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 39 Psittacidae Sangihe hanging-parrot Loriculus catamene* Psittacidae Pygmy hanging-parrot Loriculus exilis* Loriidae Red-and-blue lory Eos histrio* Loriidae Yellow-and-green lorikeet Trichoglossus flavoviridis Cuculidae Sulawesi hawk-cuckoo Cuculus crassirostris Cuculidae Bay coucal Centropus celebensis* Strigidae Ochre-bellied hawk-owl Ninox ochracea Strigidae Speckled hawk-owl Ninox punctulata Tytonidae Minahassa owl Tyto inexspectata Tytonidae Taliabu owl Tyto nigrobrunnea* Tytonidae Sulawesi owl Tyto rosenbergii Caprimulgidae Diabolical nightjar Eurostopodus diabolicus Caprimulgidae Sulawesi nightjar Caprimulgus celebensis* Alcedinidae Sulawesi kingfisher Ceyx fallax* Alcedinidae Lilac kingfisher Cittura cyanotis* Alcedinidae Black-billed kingfisher Pelargopsis melanorhyncha* Alcedinidae Talaud kingfisher Todirhamphus enigma* Alcedinidae Green-backed kingfisher Actenoides monachus* Alcedinidae Scaly kingfisher Actenoides princeps Meropidae Purple-bearded bee-eater Meropogon forsteni Coraciidae Purple-winged roller Coracias temminckii Bucconidae Sulawesi hornbill Penelopides exarhatus* Bucconidae Knobbed hornbill Aceros cassidix J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 40 Acanthizidae Rufous-sided gerygone Gerygone dorsalis Pachycephalida Sulphur-bellied whistler Pachycephala sulfuriventer Pachycephalida Drab whistler Pachycephala griseonota Rhipiduridae Rusty-flanked fantail Rhipidura teysmanni Monarchidae Cerulean paradise-flycatcher Eutrichomyias rowleyi* Monarchidae White tipped monarch Monarcha everetti* Dicruridae Sulawesi drongo Dicrurus montanus Corvidae Banggai crow Corvus unicolor* Campephagidae Cerulean cuckoo-shrike Coracina temminckii Campephagidae Pied cuckoo-shrike Coracina bicolor* Campephagidae White-rumped cuckoo-shrike Coracina leucopygia Campephagidae Sula cuckoo-shrike Coracina sula* Campephagidae Slaty cuckoo-shrike Coracina schistacea* Campephagidae White-rumped triller Lalage leucopygialis* Turdidae Rusty-backed thrush Zoothera erythronota* Sturnidae Pale-bellied myna Acridotheres cinereus Sturnidae Sulawesi myna Basilornis celebensis Sturnidae Helmeted myna Basilornis galeatus* Sturnidae White-necked myna Streptocitta albicollis Sturnidae Bare-eyed myna Streptocitta albertinae* Sturnidae Fiery-browed myna Enodes erythrophris Sturnidae Finch-billed myna Scissirostrum dubium* Muscicapidae Henna-tailed jungle-flycatcher Rhinomyias colonus* Muscicapidae Rufous-throated flycatcher Ficedula rufigula* J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 41 Zosteropidae Sulawesi white-eye Zosterops consobrinorum* Zosteropidae Black-ringed white-eye Zosterops anomalus Sylviidae Sulawesi leaf-warbler Phylloscopus sarasinorum Dicaeidae Crimson-crowned flowerpecker Dicaeum nehrkorni Dicaeidae Red-chested flowerpecker Dicaeum maugei Dicaeidae Grey-sided flowerpecker Dicaeum celebicum* Nectariniidae Elegant sunbird Aethopyga duyvenbodei* An asterisk signifies that the species' range is limited to this ecoregion. Four of Sulawesi's amphibians have Sundaland affinities, and two have Australasian roots. Thirty-eight of Sulawesi's sixty-three snake species are found on both sides of Wallace's Line. There are large reptiles of conservation significance: the sailfin lizard (Hydrosaurus amboinensis), saltwater crocodile (Crocodylus porosus), and reticulated python (Python reticulatus) (Whitten et al. 1987). Sulawesi's flora is most closely related to the floras of dry areas in the Philippines, Moluccas, Lesser Sundas, and Java. The lowland forests have affinities to New Guinea, whereas the upland areas are more related to Borneo (Whitten et al. 1987). Three Centres of Plant Diversity are located in lowland Sulawesi: Dumoga-Bone National Park, Limestone Flora of Sulawesi, and Ultramafic Flora of Sulawesi (Davis et al. 1995). Current Status More than half of the original forest has been cleared, and the remaining forests have been reduced to fragments except for a few fairly large blocks that are still intact. There are thirty-eight protected areas that cover 9,460 km2 (8 percent) of the ecoregion area (table 3). Seven of these reserves are more than 500 km2, but none are more than 1,100 km2. Table 3. WCMC (1997) Protected Areas That Overlap with the Ecoregion. Protected Area Area (km2) IUCN Category Karakelang Utara 260 VI Karakelang Selatan 80 VI J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 42 Gunung Sahendaruman 100 PRO Dua Saudara 120 I Pulau Taliabu 700 PRO Pulau Seho 20 I Pati-Pati 20 IV Lombuyan I, II 1,070 IV Gunung Lokon 20 VI Gunung Manembo-Nembo 50 IV Dumoga [AA0124] 940 ? Mas Popaya Raja 40 I Tanggale 10 PRO Panua 410 I Marisa 30 PRO Pulau Una-una 70 PRO Tanjung Api 50 I Bangkiriang 390 PRO Morowali [AA0124] 870 I Mamuja/Tapalang 150 PRO Danau Towuti 560 V Lamiko-miko 280 PRO Lasolo-Sampara 320 PRO Lamedae 30 I Rawa Aopa Watumohai 1,030 II Tanjung Batikolo 30 IV J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 43 Lampoko Mampie 30 IV Danau Tempe 310 PRO Bontobahari 50 IV Tanjung Peropa 300 IV Polewai 100 PRO Tanjung Amelango 10 IV Kaya Kuku 40 PRO Buton Utara 700 IV Napabalano 10 I Kokinawe 30 PRO Lambu Sango NR 200 PRO Tirta Rimba 30 V Total 9,460 Ecoregion numbers of protected areas that overlap with additional ecoregions are listed in brackets. Sulawesi still supports some lowland moist forests on steep slopes that are unsuitable for agriculture. However, large areas in the south and some parts of the center and north of the island have been cleared for permanent and shifting cultivation (IUCN 1991). The lowland peneplain dry forest is completely gone because of large-scale agricultural plantations, transmigration, logging, and local clearance. The riverine forest in the Dumoga Valley is now the site of a major irrigation scheme, and some of the limestone vegetation has been destroyed by quarrying to supply the Tonasa cement factories. During the dry season, cattle farmers set fires to encourage the growth of young grass, and repeated burnings have resulted in a persistent grassland vegetation in some areas and a savanna with fire-resistant trees in others. Uncontrolled exploitation for the oil in its heartwood has depleted stands of the sandalwood tree Santalum album, even in protected areas such as Paboya Reserve (Whitten et al. 1987; Whitten, pers. comm., 2000). Sangihe and Talaud were largely deforested by 1920, and there is minimal natural forest remaining on these islands. A survey has been proposed to determine appropriate locations for additional protected areas around remaining forest (Stattersfield et al. 1998). J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 44 Most of Taliabu, the largest island in the Sula Islands, is still forested, but there has been large-scale logging in the lowlands. The other main Sula Islands, Sanana and Mangole, have been heavily degraded. Extensive lowland forest still remains on the Banggai Islands (Stattersfield et al. 1998). Types and Severity of Threats Uncontrolled and illegal logging will continue to be the biggest threat to the integrity of the remaining forests. This situation has been and will be exacerbated by lack of authority and implementation of existing environmental laws. Justification of Ecoregion Delineation There have been several attempts to divide the bioregion into biogeographic units (MacKinnon 1997; Stattersfield et al. 1998; van Balgooy 1971, cited in Monk et al. 1997; MacKinnon and Artha 1981; MacKinnon and MacKinnon 1986; MacKinnon et al. 1982; van Steenis 1950; Udvardy 1975). Because many of the islands have distinct natural faunal communities and a high degree of endemism (Monk et al. 1997), the more recent attempts have used faunal dissimilarities-especially birdsto identify distinct biogeographic units (MacKinnon 1997; Stattersfield et al. 1998; MacKinnon and Artha 1981; MacKinnon and MacKinnon 1986). Because detailed floral data are largely unavailable across most of the bioregion, we followed these authors in delineating ecoregions based on distribution of biomes and vertebrate communities. On Sulawesi Island we delineated two ecoregions: the Sulawesi Lowland Rain Forests and Sulawesi Montane Rain Forests. These represent the tropical lowland and montane tropical moist forests, respectively. The small patch of monsoon forests on the southwest peninsula of Sulawesi and on Butung Island (Whitmore 1984) were included in the Sulawesi Lowland Rain Forests but should be considered a distinct habitat type in an ecoregion-based conservation assessment to ensure representation. References References for this ecoregion are currently consolidated in one document for the entire Indo-Pacific realm. Indo-Pacific Reference List J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 45 7. Sulawesi montane rain forests Sulawesi, Indonesia Photograph by David Olson The Sulawesi Montane Rain Forest harbor some of the most unique animals on Earth. The islands are located in the region known as Wallacea, which contains a distinctive fauna representing a mix of Asian and Australasian species. A fruit-eating pig with large curly tusks, a dwarf buffalo, four monkey species, and cuscuses exemplify a truly unique mammal community. Like the hub of a wheel, Sulawesi is surrounded by a variety of exotic ocean basins, including the Flores Sea, the Banda Sea, the Moluccas Sea, the Java Sea, and the Straits of Makassar, as well as the diverse islands of Borneo, Java, Flores, Halmahera, and the Philippines. Although more than half of the original forest has been cleared, Sulawesi still supports tracts of montane moist forests in areas of steep slopes that are unsuitable for agriculture. Location and General Description This ecoregion represents these montane forests above 1,000 m, whereas the lowlands constitute a separate ecoregion. Most of Sulawesi lies above 500 m, and about 20 percent of the total land areaTropical and mostly the central region-is above 1,000 m Subtropical Moist Broadleaf Forests (Whitten et al. 1987). Based on the Köppen Relatively climate zone system, this ecoregion falls in Stable/Intact the tropical wet climate zone (National Geographic Society 1999). As might be surmised from its shape, Sulawesi has a complex geologic history and is composed of three geologic provinces based on that history. West and East Sulawesi form two of the geologic provinces, separated by the Palu-Koro fault, which runs from the town of Palu to the Gulf of Bone. The third geologic province consists of the Tokala region on the northeast peninsula, the Banggai Islands, Butung Island, and the Sula Islands. East and West Sulawesi collided approximately 13-19 million years ago, and ultrabasic rocks were exposed as East Sulawesi overrode the western portion. The forces that caused the collision are still at work, and Sulawesi is being torn apart today. The surface geology of Sulawesi is a diverse patchwork of ophiolites, Mesozoic sedimentary rocks, Tertiary sedimentary and igneous rocks, and Quaternary volcanics and sediments. Active volcanoes are located on the northern arm of Sulawesi (Whitten et al. 1987). Southeastern Asia: Island of Sulawesi in Indonesia 29,300 square miles (75,800 square kilometers) -- about the size of West Virginia and Connecticut combined J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 46 Above 1,000 m, forest trees become shorter and less massive, and epiphytes such as orchids become more common. Whereas the forests of Sulawesi's lowlands are not dominated by any particular tree family, the forests in the lower montane region are dominated by oaks (four species of Lithocarpus) and chestnut (two species of Castanopsis). An example association includes Phyllocladus, Agathis dammara, and Eugenia dominated by Castanopsis. Upper montane forest contains conifers (pines and related Gymnosperms such as Podocarpus spp., Dacrycarpus spp., Dacrydium spp., Phyllocladus spp.) and the magnificent and commercially important Agathis spp. (Whitten et al. 1987). The highest peaks have sub-alpine forests with yet smaller trees whose branches bear epiphytic lichens and a ground cover of shrubs, colorful herbs, and grasses (Whitten et al. 1987). Biodiversity Features Wallace's Line, running from between Bali and Lombok and between Sulawesia and Borneo, marks the location of a deep oceanic trench and the point over which land animals and plants could not cross easily. Similarly, Lydekker's Line, running from between Timor and the Australian shelf to between Halmahera, Seram, and New Guinea, marks the point where Australasian flora and fauna could not easily pass. Sulawesi lies between these two lines. Sulawesi's location, geologic history, and long geographic isolation have created Sulawesi's distinctive fauna. There is variability, different among various animal and plant groups, in the amount of interchange between other biogeographic areas in the region, which led to the evolution of a large number of species endemic to the island. Although not species-rich relative to Borneo or Java, Sulawesi is high in endemicity because of its long isolation from Asia and Australia in Wallacea. This ecoregion exhibits high plant endemism, and the several distinct forest types provide habitat for the highest number of endemic mammals in Asia and several endemic birds (Whitten et al. 1987). The ecoregion harbors 102 mammal species, of which 33 species are endemic or near endemic (table 1). Together with the lowland forests, the montane forests of Sulawesi have the highest recorded number of endemic mammals among the Indo-Pacific ecoregions. These endemic species include the endangered mountain anoa (Bubalus quarlesi) and crested macaque (Macaca nigra) and the vulnerable babirusa (Babyrousa babyrussa) and Sulawesi montane long-nosed squirrel (Hyosciurus heinrichi) (Flannery 1995; IUCN 2000). Table 1. Endemic and Near-Endemic Mammal Species. Family Species Sorcidae Crocidura elongata Sorcidae Crocidura lea J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 47 Sorcidae Crocidura levicula Rhinolophidae Hipposideros inexpectatu Vespertilionidae Pipistrellus minahassae Vespertilionidae Hesperoptenus gaskelli Tarsiidae Tarsius pumilus* Tarsiidae Tarsius dianae* Cercopithecidae Macaca nigra Suidae Babyrousa babyrussa Bovidae Bubalus quarlesi* Sciuridae Hyosciurus heinrichi* Sciuridae Prosciurillus weberi* Sciuridae Prosciurillus abstrusus* Muridae Rattus mollicomulus* Muridae Rattus xanthurus Muridae Rattus marmosurus* Muridae Maxomys dollmani* Muridae Maxomys wattsi* Muridae Crunomys celebensis* Muridae Bunomys coelestis* Muridae Bunomys prolatus* Muridae Bunomys fratrorum* Muridae Bunomys heinrichi* Muridae Bunomys penitus* Muridae Eropeplus canus* J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 48 Muridae Margaretamys elegans* Muridae Margaretamys parvus* Muridae Taeromys hamatus* Muridae Taeromys arcuatus* Muridae Melasmothrix naso* Muridae Melasmothrix rhinogradoi* Muridae Melasmothrix macrocercus* An asterisk signifies that the species' range is limited to this ecoregion. There are approximately 168 bird species listed as resident in the ecoregion, of which 44 species are endemic or near endemic (table 2). The ecoregion also overlaps the montane portions of the Sulawesi EBA (Stattersfield et al. 1998). Of the fifty-four restricted-range bird species found in the EBA, fourteen species are found in both lowland and montane Sulawesi, and twenty-two species are only found in the uplands of Sulawesi. Nineteen of these montane species are found nowhere else on Earth. Two montane bird species are classified as threatened: the endangered Lompobattang flycatcher (Ficedula bonthaina) and the vulnerable Matinan flycatcher (Cyornis sanfordi) (Stattersfield et al. 1998). Table 2. Endemic and Near-Endemic Bird Species. Family Common Name Species Accipitridae Small sparrowhawk Accipiter nanus Megapodiidae Maleo Macrocephalon maleo Rallidae Platen's rail Aramidopsis plateni Rallidae Bare-faced rail Gymnocrex rosenbergii Rallidae Isabelline waterhen Amaurornis isabellinus Scolopacidae Sulawesi woodcock Scolopax celebensis* Columbidae Sulawesi ground-dove Gallicolumba tristigmata Columbidae Red-eared fruit-dove Ptilinopus fischeri* Columbidae White-bellied imperial-pigeon Ducula forsteni J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 49 Columbidae Grey-headed imperial-pigeon Ducula radiata Columbidae Sombre pigeon Cryptophaps poecilorrhoa* Loriidae Yellow-and-green lorikeet Trichoglossus flavoviridis Cuculidae Sulawesi hawk-cuckoo Cuculus crassirostris Strigidae Ochre-bellied hawk-owl Ninox ochracea Tytonidae Minahassa owl Tyto inexspectata Caprimulgidae Diabolical nightjar Eurostopodus diabolicus Alcedinidae Scaly kingfisher Actenoides princeps Meropidae Purple-bearded bee-eater Meropogon forsteni Coraciidae Purple-winged roller Coracias temminckii Meliphagidae Dark-eared honeyeater Myza celebensis* Meliphagidae Greater streaked honeyeater Myza sarasinorum* Pachycephalida Olive-flanked whistler Hylocitrea bonensis* Pachycephalida Maroon-backed whistler Coracornis raveni* Pachycephalida Sulphur-bellied whistler Pachycephala sulfuriventer Rhipiduridae Rusty-flanked fantail Rhipidura teysmanni Dicruridae Sulawesi drongo Dicrurus montanus Campephagidae Cerulean cuckoo-shrike Coracina temminckii Campephagidae Pygmy cuckoo-shrike Coracina abbotti* Turdidae Geomalia Geomalia heinrichi* Turdidae Sulawesi thrush Cataponera turdoides* Turdidae Great shortwing Heinrichia calligyna* Sturnidae Pale-bellied myna Acridotheres cinereus J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 50 Sturnidae Sulawesi myna Basilornis celebensis Sturnidae Fiery-browed myna Enodes erythrophris Muscicapidae Lompobattang flycatcher Ficedula bonthaina* Muscicapidae Matinan flycatcher Cyornis sanfordi* Muscicapidae Blue-fronted flycatcher Cyornis hoevelli* Zosteropidae Black-ringed white-eye Zosterops anomalus Zosteropidae Streak-headed white-eye Lophozosterops squamiceps* Sylviidae Chestnut-backed bush-warbler Bradypterus castaneus Sylviidae Sulawesi leaf-warbler Phylloscopus sarasinorum Timaliidae Malia Malia grata* Dicaeidae Crimson-crowned flowerpecker Dicaeum nehrkorni Fringillidae Mountain serin Serinus estherae An asterisk signifies that the species' range is limited to this ecoregion. Two Centres of Plant Diversity are found in the uplands of Sulwesi: Dumoga-Bone National Park and Pegunungan Latimojong. The montane forests of Dumoga-Bone National Park contain a rich gene pool of timber trees and rattans and are dominated by Eugenia, Shorea, and Agathis, with an abundance of rattans in the understory. The lower montane forests of Pegunungan Latimojong and contain Lithocarpus, Phyllocladus hypophyllus, Podocarpus steupi, and Taxus sumatrana, whereas the upper montane areas contain Vaccinium and Rhododendron vanvuurenii, Hypericum leschenaultii, and Drimys piperata. The area extends to 3,455 m and contains extensive sub-alpine vegetation above 3,200 m (Davis et al. 1995). Current Status This ecoregion is still largely intact, with about three-quarters of the original habitat remaining. Most of the habitat destruction has occurred in the southwestern portion, and large blocks of forest remain in the northern and eastern montane areas of the island. The twenty-nine protected areas cover 23 percent of the ecoregion (table 3). The average size of a protected area in this ecoregion is 602 km2, and there are five protected areas that exceed 1,000 km2. Table 3. WCMC (1997) Protected Areas That Overlap with the Ecoregion. J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 51 Protected Area Area (km2) IUCN Category Gunung Kelabat 30 DE Gunung Soputan 120 VI Gunung Simbalang 270 PRO Dumoga 1,890 ? Dolongan 1 IV Pinjan/Tanjung Matop 6 IV Kelompok Hutan Buol Toli-toli 3,920 PRO Kelompok extension 630 PRO Gunung Sojol 690 PRO Palu Mountains 3,190 PRO Wera 4 V Lore Lindu 2,220 II Palu Mountains 320 PRO Palu Mountains 130 PRO Morowali [AA0123] 1,150 I Rompi 170 PRO Rangkong 310 PRO Lamiko-miko 280 PRO Mambuliling 110 PRO Pegunungan Latimojong 510 VI Lampoko Mampie 20 IV Peg. Feruhumpenai 860 I J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 52 Danau Matano 290 V Bulu Saraung 50 I Sungai Camba 10 IV Karaenta 4 I Bantimurung 5 I Gunung Lompobatang 180 PRO Tirta Rimba 90 V Total 17,460 Ecoregion numbers of protected areas that overlap with additional ecoregions are listed in brackets. Types and Severity of Threats The steep slopes and the relative lack of commercially valuable tree species help to discourage logging activity. However, the logging that has occurred has had devastating effects on the landscape and the ecosystems; for instance, extensive erosion on surrounding deforested slopes has clogged the irrigation systems of the once fertile rice fields of Palu Valley (Whitten et al. 1987). Hunting and anthropogenic fires are also serious threats to the wildlife assemblages and habitat. Hunters set fires to facilitate hunting of anoa, creating montane meadows. Upper montane and sub-alpine forests are subject to periods of drought, during which the oil-rich leaves of Rhododendron, Vaccinium, and Gaultheria easily catch fire. With repeated burning, alang-alang grass (Imperata cylindrica) may become dominant. Other threats include transmigration and local clearance (Whitten et al. 1987). Justification of Ecoregion Delineation There have been several attempts to divide the bioregion into biogeographic units (MacKinnon 1997; Stattersfield et al. 1998; van Balgooy 1971, cited in Monk et al. 1997; MacKinnon and Artha 1981; MacKinnon and MacKinnon 1986; MacKinnon et al. 1982; van Steenis 1950; Udvardy 1975). Because many of the islands have distinct natural faunal communities and a high degree of endemism (Monk et al. 1997), the more recent attempts have used faunal dissimilarities-especially birdsto identify distinct biogeographic units (MacKinnon 1997; Stattersfield et al. 1998; MacKinnon and Artha 1981; MacKinnon and MacKinnon 1986). Because detailed floral data are largely unavailable across most of the bioregion, we followed these authors in delineating ecoregions based on distribution of biomes and vertebrate communities. J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 53 On Sulawesi island we delineated two ecoregions: the Sulawesi Lowland Rain Forests and Sulawesi Montane Rain Forests. These represent the tropical lowland and montane tropical moist forests, respectively. The small patches of monsoon forests on the southwest peninsula of Sulawesi and on Butung Island (Whitmore 1984) were included in the Sulawesi Lowland Rain Forests but should be considered a distinct habitat type in an ecoregion-based conservation assessment to ensure representation. References References for this ecoregion are currently consolidated in one document for the entire Indo-Pacific realm. Indo-Pacific Reference List J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 54 8. Sumba deciduous forests Sumba Island, Indonesia Photograph by © WWF-Canon/Edward PARKER The Sumba Deciduous Forests are found on the single island of Sumba and are part of the region known as Wallacea, which contains a distinctive fauna representing a mix of Asian and Australasian species. Although vertebrate diversity is low, the ecoregion contains seven bird species found nowhere else in the world and several other birds with very limited ranges. As a result of forest clearance and repeated burning for grazing and agriculture, the forested area of Sumba has declined significantly over the last century. Location and General Description This ecoregion represents the semievergreen forests on the island of Sumba, in the eastern Indonesian Archipelago. The surface geology of Sumba is composed primarily of sandstone and mudstone, with Tropical and Critical/Endangered some igneous intrusions overlain by recent Subtropical Dry Broadleaf limestone (Whitten and Whitten 1992). Forests Sumba is believed to be a fragment of the Australian continental crust that was separated some 20 million years ago, well before the neighboring outer arc island of Timor (Monk et al. 1997). The island is quite rugged, consisting of deeply dissected plateaus. There is very little area above 1,000 m, and the highest point on the island is 1,225 m (Stattersfield et al. 1998). Precipitation in Sumba is seasonal, and based on the Köppen climate zone system, this ecoregion falls in the tropical dry climate zone (National Geographic Society 1999). Southeastern Asia: Island of Sumba in Indonesia 4,200 square miles (10,800 square kilometers) -about twice the size of Delaware The naturally dominant vegetation of the island was deciduous monsoon forest (Stattersfield et al. 1998). However, the southern hill slopes along the southern coasts, which remain moist during the dry season, are covered with lowland evergreen rain forest. The most extensive and important of these rain forest areas is the Mt. Wanggameti-Laiwanga forest complex in East Sumba, a major water catchment. In East Sumba there are extensive gallery forests in ravines and along rivers that form riparian corridors across open grasslands or savannas. The savanna understory includes an endemic insectivorous sundew (Drosera indica) (Monk et al. 1997). Biodiversity Features The ecoregion harbors seventeen mammal species, but none are considered to be endemic or even near endemic. J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 55 The avifauna of this ecoregion is highly distinctive, with both Asian and Australian influences, although the total diversity is low. There are approximately 180 bird species on the island, and 12 of these species are endemic or near endemic (table 1). The ecoregion corresponds to the Sumba EBA. The Sumba EBA contains twelve restrictedrange bird species, seven of which are found nowhere else on Earth. Four of these species are considered vulnerable: Sumba buttonquail (Turnix everetti), red-naped fruit-dove (Ptilinopus dohertyi), Sumba boobook (Ninox rudolfi), and Sumba hornbill (Aceros everetti). These threatened species have specific habitat needs that make them susceptible to forest clearance (Stattersfield et al. 1998). Table 1. Endemic and Near-Endemic Bird Species. Family Common Name Species Turnicidae Sumba buttonquail Turnix everetti* Columbidae Sumba green-pigeon Treron teysmannii* Columbidae Red-naped fruit-dove Ptilinopus dohertyi* Strigidae Sumba boobook Ninox rudolfi* Alcedinidae Cinnamon-backed kingfisher Todirhamphus australasia Bucconidae Sumba hornbill Aceros everetti* Campephagidae Sumba cuckoo-shrike Coracina dohertyi Turdidae Chestnut-backed thrush Zoothera dohertyi Muscicapidae Flores jungle-flycatcher Rhinomyias oscillans Muscicapidae Sumba flycatcher Ficedula harterti* Zosteropidae Yellow-spectacled white-eye Zosterops wallacei Nectariniidae Apricot-breasted sunbird Nectarinia buettikoferi* An asterisk signifies that the species' range is limited to this ecoregion. Current Status Almost three quarters of the ecoregion area has been burnt for hunting or cleared, mostly for agriculture or firewood extraction. A few small, intact patches exist but are scattered in isolated fragments. Most of the original monsoon forests have been replaced by savanna and grassland (Monk et al. 1997). The four small (average size 83 km2) protected areas include about 3 percent (330 km2) of the J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 56 ecoregion area (table 2). Table 2. WCMC (1997) Protected Areas That Overlap with the Ecoregion. Protected Area Area (km2) IUCN Category Watu Manggota 20 VI Manupeu 180 VI Luku Meloto 60 PRO Laiwangi-Wanggameti NP ? ? Gunung Wanggameti 70 DE Manupea-Tanadaru NP ? ? Total 330 Ecoregion numbers of protected areas that overlap with additional ecoregions are listed in brackets. Pressures from the rapidly increasing, poor population are intense in this ecoregion (WWF-Indonesia n.d.), and nearly three-quarters of this ecoregion has been deforested, with only isolated fragments of natural habitat remaining. Types and Severity of Threats Threats include deforestation, burning of grasslands to establish agricultural fields, livestock grazing, and poaching (WWF-Indonesia n.d.). Much of the forest has already been replaced by fire-resistant casuarinas or eucalypts and extensive deciduous scrub. For instance, the ecoregion's dry thorny forest, which is especially vulnerable to clearance by fire, has almost completely disappeared (Monk et al. 1997). Justification of Ecoregion Delineation The drier forests in Nusa Tenggara were placed in three ecoregions that corresponded to the biogeographic units identified in Monk et al (1997): Lesser Sundas Deciduous Forests, which includes the chain of islands extending from Lombok, Sumbawa, Komodo, Flores, and the smaller satellite islands corresponding to the Flores biogeographic unit; Timor and Wetar Deciduous Forests, corresponding to the Timor biogeographic unit; and the Sumba Deciduous Forests, corresponding to the Sumba biogeographic unit. All three ecoregions belong to the tropical dry forests biome. J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 57 References References for this ecoregion are currently consolidated in one document for the entire Indo-Pacific realm. Indo-Pacific Reference List J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 58 9. Timor and Wetar deciduous forests Wetar Island, Indonesia Photograph by Anasia-Cruise Southeastern Asia: Islands of Timor and Wetar in Indonesia 12,900 square miles (33,500 square kilometers) -- about twice the size of Hawaii The Timor and Wetar Deciduous Forests are found on both inner and outer island arcs at the collision point of the Eurasian and Australian tectonic plates. The seasonally dry forests found in this dynamic geologic setting are part of the region known as Wallacea, which contains a very distinctive fauna representing a mix of Asian and Australasian species. Nearly twothirds of the original extent of forest has been cleared, and the ecoregion contains only fragments of natural habitat, which are themselves threatened. Location and General Description This ecoregion represents the semievergreen dry forests of Timor, Wetar, and some smaller islands in the provinces of Nusa Tenggara and Maluku in the eastern Indonesian Archipelago. This ecoregion has a dry climate, with the most xeric being the mountains of Timor. Moa, in the Leti Islands, receives an average of 1,329 mm rainfall spread over just sixty-six days of the year. Based on the Köppen climate zone system, this ecoregion falls in the tropical dry climate zone (National Geographic Society 1999). The geology of the islands is a combination of inner and outer volcanic island arcs. Wetar, Romang, Damar, and the Banda Islands are part of the inner arc, and Timor, the Leti Islands, Sermata, and Babar are part of the outer arc. The inner arc islands are a result of the subduction and partial melting of the Australian tectonic plate below the Eurasian plate. With the exception of Wetar, the inner arc islands represent young volcanoes that have coalesced with lava and sediment. The basement rock of the outer islands, on the other hand, is composed of actual continental margin from the Australian plate that has not been subducted. These outer islands are less than 4 million years old. The resulting surface geology consists of complex sedimentary and metamorphic rocks: uplifted coral reefs over complex basement rocks (Monk et al. 1997). Tropical and Subtropical Dry Broadleaf Forests Critical/Endangered The forest types in the ecoregion are dry deciduous, dry evergreen, and thorn forests. Below 1,000 m the common tree species include Sterculia foetida and Calophyllum teysmannii (both of which produce oil-bearing seeds) and Aleurites moluccana. The lowland monsoon forests are dominated by Pterocarpus indicus, especially in the lowland monsoon forest remnants of West Timor and in the well-drained, dry soils north of Oebelo on the Bena coastal plain in south Timor (Monk et al. 1997). Semi-evergreen rain forest is found on southern hill slopes at Buraen, which are kept moist by southeast trade winds, and on the Damar Islands (Monk et al. 1997). East Timor's few remaining forest patches contain the last natural stands of Eucalyptus urophylla (now widely used in J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 59 plantations) and Santalum album, the sandalwood tree (Whitten and Whitten 1992). The shrub layer in these forests includes Verbenaceae, Rubiaceae, and Euphorbiaceae, and the herbs include Acanthaceae, Tacca palmata, the root parasite Balanophora fungosa, and ground orchids such as Corymborkis (Monk et al. 1997). Four types of savanna are found here, each characterized by palm, Eucalyptus, Acacia spp., and Casuarina spp. On Timor's larger coastal plains, the vegetation ranges from grassland to open stands of deciduous trees, with increasing forest cover toward the moister southern mountains. Biodiversity Features This ecoregion has the greatest number of bird species of any tropical dry forest ecoregion in the Indo-Pacific region. Because of the long isolation with the mainland communities, there are several endemic species from several taxonomic groups. The ecoregion has thirty-eight mammal species, five of which are endemic or near endemic (table 1). Both Asian species and an Australasian cuscus (Phalanger orientalis timorensis) are found on the islands. Crocidura tenuis (Soricidae), possibly introduced by man, and the Flores giant rat (Papagomys armandvillei) are considered vulnerable (IUCN 2000). Table 1. Endemic and Near-Endemic Mammal Species. Family Species Sorcidae Crocidura tenuis* Pteropodidae Pteropus chrysoproctus Rhinolophidae Rhinolophus canuti Muridae Papagomys armandvillei* Muridae Rattus timorensis* An asterisk signifies that the species' range is limited to this ecoregion. The bird fauna consists of about 229 species. The bird fauna also represents a mix of mostly Asian species with some Australasian birds. Endemism is extremely high for these islands, with thirty-five species that are endemic or near endemic (table 2). The ecoregion encompasses with the Timor and Wetar EBA (Stattersfield et al. 1998). Thirtyfive restricted-range bird species are found in the Timor and Wetar EBA, twenty-three of which are found nowhere else on Earth. Five of these species are considered vulnerable: black cuckoo-dove (Turacoena modesta), Wetar ground-dove (Gallicolumba hoedtii), Timor green-pigeon (Treron psittacea), Timor imperial-pigeon (Ducula cineracea), and iris lorikeet (Psitteuteles iris). J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 60 Table 2. Endemic and Near-Endemic Bird Species. Family Common Name Species Columbidae Dusky cuckoo-dove Macropygia magna Columbidae Black cuckoo-dove Turacoena modesta* Columbidae Wetar ground-dove Gallicolumba hoedtii* Columbidae Timor green-pigeon Treron psittacea* Columbidae Pink-headed imperial-pigeon Ducula rosacea Columbidae Timor imperial-pigeon Ducula cineracea* Psittacidae Olive-shouldered parrot Aprosmictus jonquillaceus* Loriidae Olive-headed lorikeet Trichoglossus euteles Loriidae Iris lorikeet Psitteuteles iris* Alcedinidae Cinnamon-backed kingfisher Todirhamphus australasia Acanthizidae Plain gerygone Gerygone inornata* Meliphagidae White-tufted honeyeater Lichmera squamata Meliphagidae Yellow-eared honeyeater Lichmera flavicans* Meliphagidae Black-chested honeyeater Lichmera notabilis* Meliphagidae Crimson-hooded myzomela Myzomela kuehni* Meliphagidae Black-breasted myzomela Myzomela vulnerata* Meliphagidae Streak-breasted honeyeater Meliphaga reticulata* Meliphagidae Timor friarbird Philemon inornatus* Pachycephalida Fawn-breasted whistler Pachycephala orpheus* Oriolidae Timor oriole Oriolus melanotis* Oriolidae Timor figbird Sphecotheres viridis* Oriolidae Wetar figbird Sphecotheres hypoleucus* J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 61 Turdidae Chestnut-backed thrush Zoothera dohertyi Turdidae Orange-banded thrush Zoothera peronii Muscicapidae Black-banded flycatcher Ficedula timorensis* Muscicapidae Timor blue-flycatcher Cyornis hyacinthinus* Muscicapidae Timor bushchat Saxicola gutturalis* Zosteropidae Timor white-eye Heleia muelleri* Sylviidae Timor stubtail Urosphena subulata Sylviidae Timor leaf-warbler Phylloscopus presbytes Sylviidae Buff-banded bushbird Buettikoferella bivittata* Estrildidae Tricolored parrotfinch Erythrura tricolor Estrildidae Timor sparrow Padda fuscata* Dicaeidae Red-chested flowerpecker Dicaeum maugei Nectariniidae Flame-breasted sunbird Nectarinia solaris An asterisk signifies that the species' range is limited to this ecoregion. Timor also harbors the endemic and rare Timor python (Python timoriensis) (Whitten and Whitten 1992). Current Status Other than one remaining large block of forest near the center of Timor Island, this ecoregion contains only fragments of natural habitat. Nearly two-thirds of the original extent of forest has been cleared, mostly for agriculture. Most of the original monsoon forest on these islands has been replaced by savanna and grassland. On East Timor, the south escarpment of the Fuiloro limestone plateau originally was covered by primary rain forest, but in the 1950s this area was degraded to secondary forest. Wetar is threatened by poorly managed gold mines that have been passed from company to company, causing major environmental damage. There are twenty-four protected areas that include roughly 10 percent (3,661 km2) of the ecoregion area, but all are small, with the average size being only 152 km2 (Monk et al. 1997) (table 3). Table 3. WCMC (1997) Protected Areas That Overlap with the Ecoregion. J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 62 Protected Area Area (km2) IUCN Category Gunung Api 1 I Pulau Damar 200 PRO Pulau Babar 620 PRO Gunung Arnau 420 PRO Pulau Kambing 20 PRO Danau Ira Lalora-Pulau Yaco 120 PRO Lore 110 ? Gunung Futumasin 30 PRO Gunung Diatuto 40 PRO Gunung Talamailu 200 ? Sungai Clere GR 300 ? Tilomar 160 PRO Gunung Mutis 330 PRO Gunung Timau 340 PRO Maubesi 80 I Keluk Kupang 730 I Baun Forest 80 PRO Dataran Bena 100 VI Manipo 50 V Teluk Pelikan 30 PRO Watu Panggota/Bondokapu 30 PRO Bakau Perhatu 20 PRO J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 63 Tanjung Pukuwatu 60 PRO Pulau Dana 10 PRO Total 4,081 Ecoregion numbers of protected areas that overlap with additional ecoregions are listed in brackets. Types and Severity of Threats Deforestation is occurring very rapidly as people burn the forests for hunting, shifting cultivation, and fodder production (Whitten and Whitten 1992; Monk et al. 1997; WWF-Indonesia n.d.). Logging has also grown in importance; for instance, Damar Island was densely forested until the late 1980s, when logging began on a large scale to supply timber to the outer arc islands, where the forests had already been more heavily exploited. As a result, fire-resistant Casuarina junghuhniana grows in pure stands in cleared areas, and Mt. Mutis, on West Timor, is covered almost exclusively by Eucalyptus urophylla (Monk et al. 1997). This problem is worsening as the human populations expand. Savanna areas are especially prone to erosion. This ecoregion is highly threatened. In previous centuries, many forest resources such as sandalwood were depleted through uncontrolled exploitation (Monk et al. 1997). Justification of Ecoregion Delineation The drier forests in Nusa Tenggara were placed in three ecoregions that corresponded to the biogeographic units identified in Monk et al (1997). These are Lesser Sundas Deciduous Forests, which includes the chain of islands extending from Lombok, Sumbawa, Komodo, Flores, and the smaller satellite islands corresponding to the Flores biogeographic unit; Timor and Wetar Deciduous Forests, corresponding to the Timor biogeographic unit; and the Sumba Deciduous Forests, corresponding to the Sumba biogeographic unit. All three ecoregions belong to the tropical dry forests biome. References References for this ecoregion are currently consolidated in one document for the entire Indo-Pacific realm. Indo-Pacific Reference List J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 64 10. Mindanao-Eastern Visayas rain forests This ecoregion features lowland and hill forests on a number of large Philippine islands that, though currently disconnected, were part of one island during the height of the last ice age. These islands harbor Philippine warty pigs, Philippine deer, the Philippine tarsier, flying lemurs, and some of the last strongholds of the charismatic Philippine eagle. Most lowland forest has been cleared from the islands, but some large patches of hill and montane forest remain (the montane forest is a separate ecoregion). Tiny Camiguin Island, with two strictly endemic mammals of its own, is a unique feature of this ecoregion. Mt. Konduko, Biliran, Phillippines Photograph by L. Heaney Location and General Description This ecoregion includes the lowland (less than 1,000 m elevation) on the main islands of Mindanao, Samar, Leyte, Bohol, and 40,600 square Southeastern numerous smaller satellite islands, Asia: Philippines miles (105,100 square kilometers) including Biliran and Basilan. The climate -- about the size of Tropical and of the ecoregion is tropical wet (National Ohio Subtropical Moist Geographic Society 1999). The northern Broadleaf Critical/Endangered Visayas (northern portions of Samar and Forests Leyte) are in the main typhoon track that so strongly influences the more northerly Philippine islands. These typhoons typically occur from July to November: As much as one-third of an island's total annual precipitation may be collected during typhoon events. Mindanao is south of the main typhoon track (Dickinson et al. 1991). Mindanao and the Visayas were transported across the western Pacific to their present location during the last 25 million years. Most of these islands have been uplifted above water only in the last 15 million years or less (Hall and Holloway 1998). During the Pleistocene, Mindanao, Samar, Leyte, and Bohol were all one island-Greater Mindanaoand their faunal affinities to each other persist to this day (Heaney 1986; Heaney and Regalado 1998). Vegetation types on Mindanao and in the Eastern Visayas originally included beach forest, mangroves, lowland rain forest, and more open forest at higher elevations up to 1,000 m (Stattersfield et al. 1998). The stunted beach forest contains Casuarina and Barringtonia mixed with other lowland species. Palms, vines, bamboo, and Pterocarpus indicus are present only in rare backJ. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 65 beach swamps. This habitat type is extremely rare because of coastal habitation (Heaney and Regalado 1998). The dominant forest type in the Mindanao lowlands and the rest of the Philippines was dipterocarp forest. This group of trees is known as Philippine mahogany in the timber trade. This forest type occurred from sea level to elevations of 400 m or higher. Individual dipterocarps occur to 1,500 m. Philippine dipterocarp forest is quite tall (45-65 m) and dense, with three canopy layers. Lianas and bamboo are rare in mature forest but common in poorly developed evergreen forest. Ferns, orchids, and other epiphytic plants are found on the larger trees. At higher elevations there are only two canopy layers, tree stature is lower, and there are more epiphytes. Upper hill dipterocarp forest is found at elevations of 650 to about 1,500 m and contains dominant Shorea polysperma and oaks, chestnuts, and elaeocarps (Heaney and Regalado 1998). Biodiversity Features Mindanao and its neighbor, Basilan, situated adjacent to the Sulu Archipelago, have been influenced by immigration from Borneo, although in recent millennia movement has been primarily in the other direction (Dickinson et al. 1991). During the most recent ice ages, the Mindanao faunal region has developed its own unique fauna, with a large number of endemic vertebrates. Tiny Camiguin Island (ca. 265 km2) contains two strictly endemic and as yet undescribed mammal species: a small forest mouse (Apomys sp.) and a large moss-mouse (Bullimus sp.) (Heaney and Tabaranza 1995), in addition to an endemic frog. Several taxa found on Mindanao, only a short distance away, are absent from Camiguin, including squirrels, some murid rodents, flying lemurs, tarsiers, and deer. Camiguin is the smallest island in the Philippines known to have unique mammal species. Consisting of a series of active volcanic cones reaching a maximum elevation of 1,713 m, the island is surrounded by deep water. Fortunately, the island still has good forest cover (Heaney et al. 1998). There is also variation within the island of Mindanao. Thirty-one bird species are polytypic on the island. Sixteen of these variations are based on differences between isolated mountain ranges, and seven species have races associated with the Zamboanga Peninsula and Basilan Island. There are three species that vary between the uplands and lowlands (Dickinson et al. 1991). Approximately 80 percent of Greater Mindanao's nonvolant mammal species are found nowhere else in the world. Whereas flying lemurs, tree shrews, tree squirrels, and tarsiers are found on the islands of Greater Mindanao, they are not found on the other large Philippine island, Luzon, just 25 km from the northern tip of Samar (Heaney and Regalado 1998). More than 30 percent of nonvolant mammals in the ecoregion are endemic to Mindanao only, but the other islands in the ecoregion generally share their species with Mindanao. However, tiny Dinagat island, located just north of Mindanao, contains three of its own endemic mammals (Heaney 1986), including the endangered Dinagat Island cloud-rat (Crateromys australis). There are sixteen endemic or nearendemic mammal species in the ecoregion (table 1). J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 66 Table 1. Endemic and Near-Endemic Mammal Species. Family Species Erinaceidae Podogymnura aureospinula* Soricidae Crocidura beatus Tupaiidae Urogale everetti Cynocephalidae Cynocephalus volans Pteropodidae Ptenochirus minor Rhinolophidae Hipposideros coronatus* Sciuridae Sundasciurus philippinensis Muridae Bullimus bagobus Muridae Batomys salomonseni Muridae Batomys russatus* (Dinagat only) Muridae Crateromys australis*(Dinagat only) Muridae Crunomys melanius* Muridae Apomys sp. D*(Camiguin only) Muridae Bullimus sp. A*(Camiguin only) Muridae Tarsomys echinatus* Sciuridae Exilisciurus concinnus An asterisk signifies that the species' range is limited to this ecoregion. An endemic subspecies of Philippine deer (Cervus mariannus nigricans) is limited to Mindanao. Philippine deer are widespread (though patchily distributed) in the Philippines, being found on Luzon, Mindoro, Samar, Leyte, Mindanao, and the Basilan Islands. The subspecies is threatened by habitat loss and hunting (Wemmer 1998). It has been reported that the endangered Visayan or Philippine spotted deer (Cervus alfredi) was potentially found on Bohol Island, but it seems likely that these reports refer to Cervus mariannus. Cervus alfredi is not found on Bohol (Oliver et al. 1991; Wemmer 1998). J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 67 The kagwang (Cyanocephalus volans), or Philippine flying lemur, is also endemic to Greater Mindanao; the only other species of this unique order of mammals is found in Malaysia and Indonesia. These small nocturnal mammals glide between trees for distances up to 135 m. Fortunately, the kagwang actually prefers second-growth forests to old growth, so they are more secure than other Philippine mammals (Heaney and Regalado 1998). However, the species is still considered vulnerable (IUCN 2000). The ecoregion also supports a population of the Philippine warty pig (Sus philippensis), which the IUCN considers rare and declining. The Philippine warty pig is widely but patchily distributed in the still-forested areas of Luzon, Mindoro, Samar, Leyte, Mindanao, and some of the smaller satellite islands. Many of these forested areas are found in existing national parks. The Philippine warty pig is closely related to Sus barbatus of the Greater Sundas and was once thought to be a subspecies, analogous to the Palawan bearded pig (Sus barbatus ahoenobarbus). This species is still threatened by hunting and habitat loss (Oliver 1993). Greater Mindanao is also home to an endemic primate, the Philippine tarsier (Tarsius sychrita), which is found on Samar, Leyte, Dinagat, Siargao, Bohol, Mindanao, and Basilan. Although they are also found in primary forests and mangroves, these highly charismatic small mammals seem to prefer second-growth forests, and they are not considered threatened by the IUCN (Nowak 1999a). The Philippine tree shrew (Urogale everetti), in the order Scandentia, which is found on Mindanao, Dinagat, and Siargao Islands, represents an endemic, monotypic genus. Worldwide there are sixteen species of tree shrew, a diurnal animal that resembles a squirrel but whose dentition, circulatory system, and large braincase are more like those of primates (Nowak 1999a). This species is considered vulnerable (IUCN 2000). Greater Mindanao also supports an endemic genus of Erinaceidae, Podogymura. There are two moonrat species in this genus, both of which are found in Greater Mindanao. One species is found in the adjacent montane ecoregion of Mindanao (P. truei), and the other (P. aureospinula) is found in the lowland forest of Dinagat, Siargao, and the Bucas Grande Islands (Heaney et al. 1998). Lowland Greater Mindanao is home to endangered mammals also found in other parts of the Philippines, including the golden-capped fruit bat (Acerodon jubatus) and the mottlewinged flying-fox (Pteropus leucopterus) (found on Luzon and Dinagat) (Heaney et al. 1998; IUCN 2000). This ecoregion overlaps with the Mindanao and Eastern Visayas EBA, with the exception of the montane areas above 1,000 m, which have been given their own ecoregion. The EBA contains fifty-one restricted-range birds, twenty-four (or possibly twenty-five) of which are lowland and hill forest specialists and are thus resident in this ecoregion. All the restricted-range birds are forest species. The ecoregion contains thirty-six endemic or near-endemic bird species (Kennedy et al. 2000; table 2). Nine of these species are threatened, including the endangered Mindanao bleeding-heart (Gallicolumba criniger). J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 68 The remainder of the threatened species are considered vulnerable. This situation should be contrasted with the adjacent upland Mindanao montane rain forests ecoregion. Although the upland ecoregion contains more restricted-range species, only one of these is considered threatened (Stattersfield et al. 1998; Collar et al. 1999). Table 2. Endemic and Near-Endemic Bird Species. Family Common Name Species Rallidae Brown-banded rail Lewinia mirificus Columbidae Mindanao bleeding-heart Gallicolumba criniger* Columbidae Mindanao brown-dove Phapitreron brunneiceps Columbidae Grey imperial-pigeon Ducula pickeringii Cuculidae Black-faced coucal Centropus melanops* Strigidae Mindanao eagle-owl Mimizuku gurneyi Apodidae Philippine needletail Mearnsia picina Alcedinidae Silvery kingfisher Alcedo argentata* Alcedinidae Blue-capped kingfisher Actenoides hombroni Bucconidae Mindanao hornbill Penelopides affinis Bucconidae Samar hornbill Penelopides samarensis* Bucconidae Writhed hornbill Aceros leucocephalus Pittidae Azure-breasted pitta Pitta steerii* Eurylaimidae Wattled broadbill Eurylaimus steerii* Eurylaimidae Visayan wattled broadbill Eurylaimus samarensis* Rhipiduridae Blue fantail Rhipidura superciliaris* Monarchidae Short-crested monarch Hypothymis helenae Monarchidae Celestial monarch Hypothymis coelestis Muscicapidae Little slaty flycatcher Ficedula basilanica* J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 69 Muscicapidae Cryptic flycatcher Ficedula crypta Pycnonotidae Zamboanga bulbul Ixos rufigularis* Pycnonotidae Yellowish bulbul Ixos everetti Sylviidae Long-tailed bush-warbler Bradypterus caudatus Sylviidae Rufous-headed tailorbird Orthotomus heterolaemus Sylviidae Yellow-breasted tailorbird Orthotomus samarensis* Sylviidae White-browed tailorbird Orthotomus nigriceps* Sylviidae White-eared tailorbird Orthotomus cinereiceps* Timaliidae Striated wren-babbler Ptilocichla mindanensis* Timaliidae Pygmy babbler Stachyris plateni Timaliidae Rusty-crowned babbler Stachyris capitalis Timaliidae Brown tit-babbler Macronous striaticeps Timaliidae Miniature tit-babbler Micromacronus leytensis Paridae White-fronted tit Parus semilarvatus Dicaeidae Whiskered flowerpecker Dicaeum proprium Dicaeidae Olive-capped flowerpecker Dicaeum nigrilore Dicaeidae Flame-crowned flowerpecker Dicaeum anthonyi An asterisk signifies that the species' range is limited to this ecoregion. In addition to the restricted-range species, several widespread threatened species are found in the ecoregion, including the critically endangered Philippine eagle (Pithecophaga jeffreyi) and Philippine cockatoo (Cacatua haematuropygia). Four additional widespread but vulnerable species are also found in the ecoregion (Stattersfield et al. 1998; Collar et al. 1999). The critically endangered Philippine crocodile (Crocodylus mindorensis) was historically found on Jolo, Luzon, Mindoro, Masbate, Samar, Negros, Busuanga, and Mindanao, but the only remaining populations are found on Mindoro, Negros, Mindanao, and Busuanga. The current wild population may be approximately 100 nonhatchlings (Ross 1998). J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 70 Mt. Apo, on Mindanao, is considered a Centre of Plant Diversity (Davis et al. 1995). This spectacular mountain in the southern portion of the Central Cordillera contains primary lowland forest and lower montane forests as well as montane forests found in the Mindanao Montane Rain Forests [IM0128] ecoregion. Much of the lowland forest below 1,000 m has been cleared, but dipterocarp forest is found from 1,000 to 1,600 m. Current Status All the islands in the ecoregion were once completely forested, but there is little forest left on most islands, and especially little lowland forest left. The dire situation in the lowlands of Mindanao and Eastern Visayas is highlighted by the contrast in conservation status between the lowland ecoregion and the adjacent upland Mindanao Montane Rain Forests [IM0128] ecoregion. Although the upland ecoregion contains more restricted-range species, only one of these is considered threatened. In fact, the Mindanao and Eastern Visayas EBA contains more threatened birds than any other EBA in the southeast Asian island region, and all but one of these are found in the lowlands (Stattersfield et al. 1998; Collar et al. 1999). Bohol is heavily deforested, and almost all of the island's natural forest is to be found in Rajah Sikatuna National Park (RSNP). The conditions in this 9,023-ha area are good, however, and the Philippine Department of Natural Resources is actively reforesting the edges of the park. Both of the Eastern Visayan endemic birds and all four of Bohol's endemic bird subspecies can be found in RSNP. Problems of firewood and rattan collection, hunting and trapping, and slash-and-burn agriculture are effectively limited to the eastern portions of the park (Brooks et al. 1995). Samar and Leyte each have two areas of closed-canopy forest remaining. The largest blocks are found on Samar. Three of these patches are found in areas of suspended timber license agreements and the remaining forest block, on Leyte, is found in the Philippine National Oil Company Tungonan Forest Reserve (Development Alternatives 1992). By 1988, approximately 29 percent of Mindanao's forest remained, including both primary and secondary forests (Stattersfield et al. 1998). There is much less today. The Zamboanga Peninsula on southwest Mindanao contains a number of isolated fragments, the largest of which is found in the watershed of Zamboanga City. The remaining patches are scattered in hill and montane areas around the peninsula. These patches contained evidence of recent logging in 1992. In southern Mindanao, some large areas of forest remain in hill and montane areas. Political instability, lack of access, and poor commercial values have helped protect some of these areas. Ironically, some of the areas, which had been under now-suspended timber license agreements, are threatened by encroaching agriculture and fire. There are other large blocks of forest in the rest of Mindanao, but they are similarly limited to hill and montane areas; there is very little lowland dipterocarp forest remaining on Mindanao (Development Alternatives 1992). Southern Mindanao is faced with political instability that poses a challenge for active conservation. J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 71 Aerial surveys of Basilan in 1992 revealed less than 2 percent natural forest remaining. Unfortunately, Basilan is also subject to political insurgency that makes active conservation efforts quite difficult (Stattersfield et al. 1998). Both the Philippine warty pig and Philippine deer suffer from intense hunting pressure and fragmentation of their remaining habitats. The pigs are in an especially poor situation because they tend to raid crops and are regarded as pests; consequently, no protections are in place for them (Oliver 1993). Table 3 details the existing protected areas on the island. Table 3. WCMC (1997) Protected Areas That Overlap with the Ecoregion. Protected Area Area (km2) IUCN Category Mado Hot Spring 20 III Sohoton Natural Bridge 40 III Imelda Lake 40 II Mahagnao Volcano 30 II Lake Danao 5 IV Rajah Sikatuna 110 II Rizal 10 III Initao 10 V Mt. Malindang [IM0302] 160 II Mt. Apo [IM0302] 130 II Lake Butig 6 V Liguasan March GRBS 410 IV Lake Buluan 80 IV Agusan Marsh 810 PRO Basilan 90 II Total 1,951 J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 72 Ecoregion numbers of protected areas that overlap with additional ecoregions are listed in brackets. Types and Severity of Threats Many of the factors that have contributed to the loss of habitat in the past still present threats to the future of these forests. They include firewood and rattan collection, hunting and trapping, slash-and-burn agriculture, and commercial forestry (Brooks et al. 1995). Justification of Ecoregion Delineation MacKinnon (1997) identified seven subunits in the Philippines, and the Philippine Biodiversity Action Plan (Philippine BAP 1997) demarcated fifteen biogeographic units. Udvardy (1975) identified the Philippines as a single biogeographic province. We delineated nine ecoregions in the Philippine islands, including Palawan. We deviated from Udvardy (1975), MacKinnon (1997), Stattersfield (1998), and the Philippine BAP (1997) in varying degrees and based our delineation of the Philippine ecoregions on Heaney (1993), with the exception of Camiguin, which Heaney separated. The islands of Leyte, Samar, Dinagat, and Bohol were combined with the lowland rain forests of Mindanao island to form the Mindanao-Eastern Visayas Rain Forests. We also included the Basilan Islands off the southwest peninsula of Mindanao in this ecoregion, based on Heaney (1993). In Mindanao we used the 1,000-m contour from the DEM (USGS 1996) to delineate the montane forests from the lowland forests. The montane forests of Mindanao were placed into their own ecoregion, the Mindanao Montane Rain Forests. In our delineation of the Mindanao-Eastern Visayas Rain Forests and Mindanao Montane Rain Forests ecoregions, we deviated from MacKinnon (1997). MacKinnon placed both of Mindanao's lowland and montane forests in a single subunit (26c). The Basilan Islands were part of subunit 26d, and the islands of Leyte and Samar made up subunit 26e. References References for this ecoregion are currently consolidated in one document for the entire Indo-Pacific realm. Indo-Pacific Reference List J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 73 11. Mindanao montane rain forests This ecoregion features the montane forests on the island of Mindanao. This disjunct ecoregion harbors Philippine warty pigs, Philippine deer, and some of the last strongholds of the charismatic Philippine eagle. Because most of the lowland forest on Mindanao has been cleared, the remaining montane forests are some of the last vestiges of wild Mindanao. Location and General Description Mt. Kataglad, Mindanao, Philippines Photograph by Tom Brooks 7,000 square miles (18,200 square kilometers) -Tropical and about the size of Subtropical Moist Delaware and Broadleaf Rhode Island Forests combined Indo-Malay The climate of the ecoregion is tropical wet (National Geographic Society 1999), with temperature and rainfall modified by the elevation, which reaches up to 2,700 m. There are extensive, disjunct areas of the island above 1,000 m. Mindanao generally is south of the typhoon track of the storms that usually hit the northern Philippines from July to November (Dickinson et al. 1991). Critical/Endangered Mindanao and the Visayas have been transported across the western Pacific to their present location during the last 25 million years. Most of these islands have been uplifted above water only in the last 15 million years (Hall and Holloway 1998). During much of the Pleistocene, Mindanao and the Eastern Visayas (Samar, Leyte, and Bohol) were all one island-Greater Mindanao (Heaney 1986)-but the higher elevations of this larger island generally were limited to what is now Mindanao. Vegetation types in the montane forests of Mindanao consist of hill dipterocarp forests, lower and upper montane forest, elfin woodland (mossy forest), and summit grasslands (Davis et al. 1995). The dominant forest type in Mindanao and the rest of the Philippines was dipterocarp forest. Whereas upper hill dipterocarp forest is found at elevations of 650-1,000, individual dipterocarps occur to 1,500 m, and on Mt. Apo, primary dipterocarp forest occurs from 1,000 to 1,600 m. Upper hill dipterocarp forest on Mt. Apo is dominated by the dipterocarps Hopea plagata, Shorea guiso, and Dipterocarpus grandiflorus and species of Cinnamomum, Lithocarpus, Homalanthus, and Musa. There are many epiphytes, mostly ferns and orchids. Tree ferns (Cyathea) and palms (Areca) are also found in the understory (Davis et al. 1995; Heaney and Regalado 1998). The transition zone between dipterocarp forest and montane forest includes increasing numbers of tree ferns, pandans, rattans, and Angiopteris; the dipterocarps Shorea almon, S. polysperma, and Lithocarpus spp.; and Agathis philippensis (Davis et al. 1995). J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 74 On Mt. Apo, montane forest occurs above approximately 2,000 m. Dominant genera include Lithocarpus, Cinnamomum, Melastoma, Caryota, Calamus, Ficus, Agathis, and numerous Lauraceae (Davis et al. 1995). The mossy upper montane forest generally is found at elevations from 1,200 m to 1,500 m (Davis et al. 1995; Lewis 1988), where humidity is constantly high. This stunted, single-story, moss- and epiphyte-covered forest contains tree ferns up to 10 m high (Dickinson, Kennedy, and Parkes 1991). All surfaces are covered or draped with lichens, bryophytes, begonias, orchids, aroids, Selaginella, and Nephrolepis ferns (Davis et al. 1995). Biodiversity Features Mindanao and its neighbor, Basilan, situated adjacent to the Sulu Archipelago, have been influenced by animal dispersal from Borneo, although in recent millennia movement has been primarily in the other direction (Dickinson et al. 1991). Over the course of the most recent ice ages, the Mindanao faunal region has developed its own unique fauna, with a number of endemic vertebrates. There is also variation within the island of Mindanao. Thirty-one bird species are polytypic on the island. Sixteen of these variations are based on differences between isolated mountain ranges, and seven species have races associated with the Zamboanga Peninsula and Basilan Island. There are three species that vary between the uplands and lowlands (Dickinson et al. 1991). Approximately 80 percent of Greater Mindanao's nonvolant mammal species are found nowhere else in the world. Although flying lemurs, tree shrews, tree squirrels, and tarsiers are found on the islands of Greater Mindanao, they are not found on the other large Philippine island of Luzon, just 25 km away from the northern tip of Samar (Heaney and Regalado 1998). More than 30 percent of nonvolant mammals in the ecoregion are endemic to Mindanao only, but the other islands share their species with Mindanao. However, tiny Camiguin and Dinagat islands, located north of Mindanao, contain two and three, respectively, of their own endemic mammals (Heaney 1986; Heaney et al. 1995). Fourteen mammal species are endemic or near endemic to the ecoregion (table 1) Table 1. Endemic and Near-Endemic Mammal Species. Family Species Erinaceidae Podogymnura truei* Soricidae Crocidura beatus Soricidae Crocidura grandis* Tupaiidae Urogale everetti J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 75 Pteropodidae Alionycteris paucidentata* Sciuridae Petinomys crinitus* Sciuridae Sundasciurus philippinensis Sciuridae Exilisciurus concinnus Muridae Bullimus bagobus Muridae Limnomys sibuanus* Muridae Tarsomys apoensis* Muridae Batomys salomonseni Muridae Crunomys suncoides* Muridae Limnomys sp. B* An asterisk signifies that the species' range is limited to this ecoregion. An endemic subspecies of Philippine deer (Cervus mariannus nigricans) is limited to Mindanao. Philippine deer are widespread (though distributed patchily) in the Philippines, being found on Luzon, Mindoro, Samar, Leyte, Mindanao, and the Basilan Islands. The subspecies (and species) is threatened by habitat loss and hunting (Wemmer 1998). The Philippine tree shrew (Urogale everetti), which is found on Mindanao, Dinagat, and Siargao islands, represents an endemic, monotypic genus. There are sixteen species of tree shrews, a diurnal animal that resembles a squirrel but whose dentition, circulatory system, and large braincase are more like those of primates (Nowak 1999a). This species is considered vulnerable (IUCN 2000). The ecoregion also supports a population of the Philippine warty pig (Sus philippensis), which the IUCN considers rare and declining. The Philippine warty pig is widely distributed in the still-forested areas of Luzon, Mindoro, Samar, Leyte, Mindanao, and some of the smaller satellite islands. Many of these forested areas are found in existing national parks. The Philippine warty pig is closely related to Sus barbatus of the Greater Sundas and was once thought to be a subspecies, analogous to the Palawan bearded pig (Sus barbatus ahoenobarbus). The Philippine warty pig is still threatened by hunting and habitat loss (Oliver 1993). Greater Mindanao also supports an endemic genus of Erinaceidae, Podogymura. There are two moonrat species in this genus, both of which are found in Greater Mindanao and one of which is found in the montane regions of Mindanao. The exclusively montane J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 76 species P. truei, or Mindanao moonrat, is common in montane and mossy forest at elevations from 1,400 m to 2,800 m. Contrary to the IUCN listing, it is not threatened (Heaney et al. 1998; Heaney, pers. comm., 2000). Montane Mindanao is also home to the endangered Greater Mindanao shrew (Crocidura grandis) and the widespread (within the Philippines) but endangered golden-crowned fruit bat (Acerodon jubatus) (Heaney et al. 1998). This ecoregion overlaps with the Mindanao and the Eastern Visayas EBA, but only the montane portions of it. The EBA contains fifty-one restricted-range birds, twenty-six (or possibly twenty-seven) of which are montane and mossy forest specialists and are thus resident in this ecoregion. All of the restricted-range birds are forest species. There are thirty-four endemic or near-endemic bird species in the ecoregion (Kennedy et al. 2000; table 2). Only one of these species is considered threatened: the vulnerable blue-capped kingfisher (Actenoides hombroni). This situation should be contrasted with the adjacent lowland Mindanao and Eastern Visayas rain forests. Although it supports fewer restricted range species, the lowland ecoregion contains nine threatened species (Stattersfield et al. 1998; Collar et al. 1999). Table 2. Endemic and Near-Endemic Bird Species. Family Common Name Species Columbidae Mindanao brown-dove Phapitreron brunneiceps Psittacidae Mindanao racquet-tail Prioniturus waterstradti* Loriidae Mindanao lorikeet Trichoglossus johnstoniae* Strigidae Mindanao scops-owl Otus mirus* Strigidae Mindanao eagle-owl Mimizuku gurneyi Apodidae Whitehead's swiftlet Aerodramus whiteheadi Apodidae Philippine needletail Mearnsia picina Alcedinidae Blue-capped kingfisher Actenoides hombroni Bucconidae Mindanao hornbill Penelopides affinis Bucconidae Writhed hornbill Aceros leucocephalus Rhipiduridae Black-and-cinnamon fantail Rhipidura nigrocinnamomea* J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 77 Campephagidae McGregor's cuckoo-shrike Coracina mcgregori* Sturnidae Apo myna Basilornis miranda* Muscicapidae Mindanao jungle-flycatcher Rhinomyias goodfellowi* Muscicapidae Cryptic flycatcher Ficedula crypta Laniidae Mountain shrike Lanius valdirostris Zosteropidae Mindanao white-eye Lophozosterops goodfellowi* Zosteropidae Cinnamon white-eye Hypocryptadius cinnamomeus* Sylviidae Long-tailed bush-warbler Bradypterus caudatus Sylviidae Rufous-headed tailorbird Orthotomus heterolaemus Timaliidae Bagobo babbler Trichastoma woodi* Timaliidae Pygmy babbler Stachyris plateni Timaliidae Rusty-crowned babbler Stachyris capitalis Timaliidae Brown tit-babbler Macronous striaticeps Timaliidae Miniature tit-babbler Micromacronus leytensis Estrildidae Red-eared parrotfinch Erythrura coloria* Dicaeidae Whiskered flowerpecker Dicaeum proprium Dicaeidae Olive-capped flowerpecker Dicaeum nigrilore Dicaeidae Flame-crowned flowerpecker Dicaeum anthonyi Nectariniidae Grey-hooded sunbird Aethopyga primigenius* Nectariniidae Mt. Apo sunbird Aethopyga boltoni* Nectariniidae Linas sunbird Aethopyga linaraborae* Fringillidae Mountain serin Serinus estherae Fringillidae White-cheeked bullfinch Pyrrhula leucogenis J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 78 An asterisk signifies that the species' range is limited to this ecoregion. In addition to the restricted-range species, the critically endangered Philippine eagle (Pithecophaga jeffreyi) and vulnerable spotted imperial-pigeon (Ducula carola) are found in the ecoregion (Stattersfield et al. 1998; Collar et al. 1999). Mt. Apo on Mindanao is considered a Centre of Plant Diversity (Davis et al. 1995). This spectacular mountain in the southern portion of the Central Cordillera contains primary lowland forest and lower montane forests as well as montane forests found in the Mindanao Montane Rain Forests [IM0128] ecoregion. Much of the lowland forest below 1,000 m has been cleared, but dipterocarp forest is found from 1,000 to 1,600 m. Current Status Although the remaining forest is found in isolated patches, most forest remaining on the island of Mindanao is contained in this upland ecoregion. This is in contrast to the largely deforested lowlands of Mindanao and the Eastern Visayas. The conservation status of restricted-range birds is illustrative in this respect: although the upland ecoregion contains more restricted-range species than the lowlands, only one of the upland species is considered threatened, compared with nine threatened species in the lowlands (Stattersfield et al. 1998; Collar et al. 1999). By 1988, approximately 29 percent of Mindanao's forest, including both primary and secondary forests, remained (Stattersfield et al. 1998). There is less today (Development Alternatives 1992). The Zamboanga Peninsula on southwest Mindanao contains a number of isolated fragments, the largest of which is found in the watershed of Zamboanga City. The remaining patches are scattered in hill and montane areas around the peninsula. These patches contained evidence of recent logging in 1992. In southern Mindanao, some large areas of forest remain in hill and montane areas. Political instability, lack of access, and poor commercial values have helped protect some of these areas. Ironically, some of the areas, which had been under now-suspended timber license agreements, are threatened by encroaching agriculture and fire. There are other large blocks of forest in the rest of Mindanao, but they are similarly limited to hill and montane areas (Development Alternatives 1992). Varying levels of protection have been accorded to the protected areas in the ecoregion; by 1988, approximately 50 percent of Mt. Apo National Park had been deforested. In addition to the classic sequence of logging, invasion by kaingineros, and associated hunting and burning, the park is administered by more than one Bureau of Forest Development office and is sometimes occupied by rebel groups (Lewis 1988). Table 3 details the existing protected areas in the ecoregion. Table 3. WCMC (1997) Protected Areas That Overlap with the Ecoregion. J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 79 Protected Area Area (km2) IUCN Category Sacred Mountain 10 III Rungkunan 10 V Pantuwaraya Lake 8 V Salikata 10 V Lake Dapao 20 V Mt. Kitanglad 250 II Mt. Apo [IM0156] 420 II Mainit Hot Spring 30 PRO Mt. Malindang [IM0156] 300 II Total 1,058 Ecoregion numbers of protected areas that overlap with additional ecoregions are listed in brackets. Types and Severity of Threats Habitat destruction is the main threat to biodiversity in the Philippines. Logging and shifting cultivation (kaingin) are cited as the primary forces of habitat conversion. Logging takes many forms, from industrial scale to smaller-scale operations that use water buffalo to haul logs out of the forest (Davis et al. 1995). Both the Philippine warty pig and Philippine deer suffer from intense hunting pressure and fragmentation of their remaining habitats. The pigs are in an especially poor situation because they tend to raid crops and are regarded as pests; consequently, there are no effective protections in place for them (Oliver 1993). Justification of Ecoregion Delineation MacKinnon (1997) identified seven subunits in the Philippines, and the Philippine Biodiversity Action Plan (Philippine BAP 1997) demarcated fifteen biogeographic units. Udvardy (1975) identified the Philippines as a single biogeographic province. We delineated nine ecoregions in the Philippine islands, including Palawan. We deviated from Udvardy (1975), MacKinnon (1997), Stattersfield et al. (1998), and the Philippine BAP (1997) to varying degrees and based our delineation of the Philippine ecoregions on Heaney (1993). J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 80 The islands of Leyte, Samar, Dinagat, and Bohol were combined with the lowland rain forests of Mindanao Island to form the Mindanao-Eastern Visayas Rain Forests. We also included the Basilan Islands off the southwest peninsula of Mindanao in this ecoregion, based on Heaney (1993). In Mindanao we used the 1,000-m contour from the DEM (USGS 1996) to delineate the montane forests from the lowland forests. The montane forests of Mindanao were placed into their own ecoregion, the Mindanao Montane Rain Forests. In our delineation of the Mindanao-Eastern Visayas Rain Forests and Mindanao Montane Rain Forests ecoregions, we deviated from MacKinnon (1997). MacKinnon placed both of Mindanao's lowland and montane forests in a single subunit (26c). The Basilan Islands were part of subunit 26d, and the islands of Leyte and Samar made up subunit 26e. References References for this ecoregion are currently consolidated in one document for the entire Indo-Pacific realm. Indo-Pacific Reference List J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 81 12. Mindoro rain forests Called the dark island by outsiders because of a virulent strain of malaria, Mindoro is located between the large islands of Luzon and the Sundaaffiliated Palawan, and it shares faunal attributes of both islands. However, Mindoro was isolated from Luzon and Palawan throughout the Pleistocene and retains its own unique character, including an endemic water buffalo species (Heaney 1986). Unfortunately, Mindoro is one of the most severely deforested islands in the country (Heaney and Mittermeir 1997). Only the most rugged portions of the island's central spine has been spared from commercial logging, and the forest is still under pressure. Mindoro, Philippines Location and General Description Photograph by Tom Brooks This ecoregion includes the island of Mindoro and the Semirara Islands. The climate of the ecoregion is tropical wet (National 3,900 square miles Geographic Society 1999). The western coast Philippines: (10,100 square Island of of Mindoro experiences a wet season during kilometers) -Mindoro the southwest monsoon of June to October about half the size and a dry season during the November to of Massachusetts Tropical and February northeast monsoon because of the Subtropical Moist Critical/Endangered central mountains (High Rolling Mountains) Broadleaf Forests (Collins et al. 1991). The High Rolling Mountains dominate the central portions of the island and rise to a maximum elevation of approximately 2,500 m at Mt. Halcon and Mt. Baco. Mindoro (along with Palawan and the Calamianes) was rifted (below water) from the Asian mainland approximately 32 million years ago, transported through seafloor spreading across the growing South China Sea, added to the growing Philippine Archipelago approximately 17 million years ago, and uplifted above water approximately 5-10 million years ago (Hall and Holloway 1998; Dickinson, Kennedy, and Parkes 1991). Mindoro is separated from Palawan to the south and Luzon to the north by deepwater channels and has not been connected to those islands during the recent past (Pleistocene) (Heaney 1986). Vegetation types on Mindoro include lowland evergreen rain forest to approximately 400 m or higher, open forest from about 650 to 1,000 m, and mossy forest above. Only small patches remain of the lowland evergreen dipterocarp rain forest that would have dominated the lowland eastern portions of the island. Semideciduous forest would have predominated on the western half of the island. Limited stands of Mindoro pine (Pinus merkusii) are J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 82 found at elevations of 600 m or less in the northern portions of the island (Stattersfield et al. 1998; Development Alternatives 1992). Biodiversity Features Of the forty-two indigenous mammal species found on Mindoro, close to 20 percent endemic or near endemic (table 1). The nonendemic mammals are also found on Luzon. An endemic rat (Rattus mindorenis) is closely related to Rattus tiomanicus, and the endemic genus Anonomomys is most closely related to the genus Haeromys, from Palawan and some of its satellite islands. Thus colonization of Mindoro has occurred from both Luzon and Palawan (Heaney 1986). Table 1. Endemic and Near-Endemic Mammal Species. Family Species Sorcidae Crocidura mindorus Bovidae Bubalus mindorensis* Muridae Rattus mindorensis* Muridae Anonymomys mindorensis* Muridae Crateromys paulus* Muridae Apomys gracilirostris* Muridae Apomys sp. E* Pteropodidae Pteropus sp. A* An asterisk signifies that the species' range is limited to this ecoregion. The most unique animal feature of Mindoro must be the tamaraw (Bubalus mindorensis), or dwarf water buffalo. There were perhaps 10,000 living at all elevations on the island at the turn of the century. The tamaraw, like the anoas of Sulawesi (Anoa spp.), are wary forest animals, just over 1 m tall at the shoulder (Heaney and Regalado 1998; Nowak 1999a). Tamaraws sometimes are placed in the same genus as anoas. They should not be confused with the carabao of the Philippines, which is a small variety of domesticated Asian water buffalo (B. bubalus). The tamaraw needs both dense vegetation for resting and open grazing land. It is unclear whether tamaraws need wallows. Unfortunately, they are confined to areas of grassland that have taken the place of the native forest. Adult bulls are largely solitary and aggressive toward each other. Young males (up to five years) form bachelor groups. Females are found alone, with a bull, or with up to three young of differing ages. Young are born during Mindoro's June-November rainy season and stay with their mother J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 83 until the age of 1.5 to 4.5 years (Nowak 1999a). Mindoro also supports a population of the Philippine warty pig (Sus philippensis), which the IUCN considers rare and declining (IUCN 2000). The Philippine warty pig is widely distributed in the still-forested areas of Luzon, Mindoro, Samar, Leyte, Mindanao, and some of the smaller satellite islands. Many of these forested areas are found in existing national parks. The Philippine warty pig is closely related to Sus barbatus of the Greater Sundas and was once thought to be a subspecies, analogous to the Palawan bearded pig (Sus barbatus ahoenobarbus). This species is still threatened by hunting and habitat loss (Oliver 1993). An endemic subspecies of the Philippine deer (Cervus mariannus barandanus) is found on Mindoro. Although Philippine deer are native to Luzon, Mindoro, Samar, Leyte, Mindanao, and the Basilan Islands, C. m. barandanus is found only on Mindoro. The population of this subspecies is considered to be at risk over its limited range on the island (Wemmer 1998). Greater Mindoro is home to the critically endangered Illin hairy-tailed cloud rat (Crateromys paulus), the endangered Mindoro shrew (Crocidura mindorus), and the more widespread (within the Philippines) but endangered golden-crowned fruit bat (Acerodon jubatus) (IUCN 2000). This ecoregion corresponds exactly with the Mindoro EBA. The Mindoro EBA contains ten restricted-range birds, six of which are threatened. The Mindoro ecoregion contains eleven endemic or near-endemic bird species (Kennedy et al. 2000; table 2). Two bird species, the Mindoro bleeding-heart (Gallicolumba platenae) and the black-hooded coucal (Centropus steerii), are considered critically endangered, and four species are considered vulnerable: Mindoro imperial-pigeon (Ducula mindorensis), ashy thrush (Zoothera cinerea), Luzon water-redstart (Rhyacornis albiventris), and scarlet-collared flowerpecker (Dicaeum retrocinctum). Three of these species, the Mindoro bleeding-heart, the Mindoro imperial pigeon, and the black-hooded coucal, are strict island endemics (Collar et al. 1999; Stattersfield et al. 1998). Mindoro's endemic birds can be split into montane and lowland species. Although both are in urgent need of conservation, the situation for the lowland species is particularly dire because the lowland forests are almost entirely gone (Dutson et al. 1992). Mindoro is also an important wintering and staging area for ducks and other waterbirds (Bagarinao 1998). Table 2. Endemic and Near-Endemic Bird Species. Family Common Name Species Columbidae Mindoro bleeding-heart Gallicolumba platenae* Columbidae Mindoro imperial-pigeon Ducula mindorensis* Cuculidae Black-hooded coucal Centropus steerii* J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 84 Strigidae Mindoro scops-owl Otus mindorensis* Strigidae Mantanani scops-owl Otus mantananensis Bucconidae Mindoro hornbill Penelopides mindorensis* Pachycephalida Green-backed whistler Pachycephala albiventris Laniidae Mountain shrike Lanius validrostris* Turdidae Ashy thrush Zoothera cinerea Muscicapidae Luzon redstart Rhyacornis bicolor Dicaeidae Scarlet-collared flowerpecker Dicaeum retrocinctum An asterisk signifies that the species' range is limited to this ecoregion. The type specimen of the critically endangered Philippine crocodile (Crocodylus mindorensis) was collected in Mindoro's Naujan Lake (Bagarinao 1998). They were historically found on the islands of Luzon, Mindoro, Masbate, Samar, Jolo, Negros, Busuanga, and Mindanao, but the only remaining populations are found on Mindoro, Negros, Mindanao, and Busuanga. The only protected population of Philippine crocodiles is in Lake Naujan National Park on Mindoro. Whereas the decline of the species initially was driven by overexploitation, habitat loss and human persecution are now the principal threats to the Philippine crocodile. Surveys in 1980-1982 revealed a total wild population of approximately 500-1,000 individuals, but current wild populations may be approximately 100 nonhatchlings. Captive breeding efforts are being led by the Crocodile Farming Institute, an entity of the Philippine government (Ross 1998; IUCN 2000). Lubang Island, near Mindoro and Luzon, is a poorly known island surrounded by deep water channels; it may well represent a small but distinct center of endemism (L. Heaney, pers. comm., 2000). Current Status The only remaining intact forests in Mindoro are found along the top of the mountain ridge that divides the island. On the eastern side of the ridge commercial logging ended long enough ago that the remaining intact forests are buffered by secondary forests that have reestablished a closed condition, yet these same forests are again under threat from poaching and kaingin (slash-and-burn) agriculture. On the western side of the ridges, however, perennial fires in adjacent grasslands used for pasture are eating into the forest (Development Alternatives 1992). Only 8.5 percent of Mindoro was forested in 1988 (SSC 1988). Several reserves have been established in Mindoro, beginning in 1936, but these have J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 85 proved to be less than effective (Heaney and Regalado 1998). Table 3 details the existing protected areas on the island. Table 3. WCMC (1997) Protected Areas That Overlap with the Ecoregion. Protected Area Area (km2) IUCN Category Lake Naujan 90 IV Mts. Iglit-Baco 790 II F.B. Harrison 1,800 IV Total 2,680 Ecoregion numbers of protected areas that overlap with additional ecoregions are listed in brackets. The largest protected area on Mindoro is Mounts Iglit-Baco National Park, which is one of two ASEAN Natural Heritage Sites in the Philippines (the other is Mount Apo National Park on Mindanao). The park covers the east-west divide and includes several physiographic regions and an important tamaraw population. Small patches of dipterocarp and mossy forest can be found in the park. The park is inhabited by the Mangyan tribal people, and much of the reserve consists of fire-maintained grassland with Imperata cylindrica and Sacchareum spontaneum. The combination of burning to maintain pasture for domestic cattle, ranching, and uncontrolled hunting activities leaves this protected area substantially altered, and the park is an insecure refuge for the endangered tamaraw (Collins et al. 1991). Lake Naujan National Park is the only protected area in the Philippines that protects the critically endangered Philippine crocodile (Ross 1998). The tamaraw numbered approximately 10,000 animals at the turn of the century and approximately 1,000 by 1949, and today estimates range from 100 to 200 animals (Collins et al. 1991; Heaney and Regalado 1998). A well-funded conservation program aimed at captive breeding of tamaraws has been an expensive failure (Heaney and Regalado 1998). Types and Severity of Threats Hunting by local people is a threat to all large mammals in the ecoregion, including the tamaraw, Philippine deer, and Philippine warty pig (Hedges, in press). Forestry activities and kaingin (slash-and-burn) agriculture continue to fragment and destroy the remaining habitat. J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 86 Although it is smaller and not as rich as some of the larger Philippine islands, Mindoro faces high levels of faunal endangerment because a larger proportion of its fauna is endangered; this level of endangerment is well-correlated with the degree of deforestation on the respective islands (Heaney 1993). Justification of Ecoregion Delineation MacKinnon (1997) identified seven subunits in the Philippines, and the Philippine Biodiversity Action Plan (Philippine BAP 1997) demarcated fifteen biogeographic units. Udvardy (1975) identified the Philippines as a single biogeographic province. We delineated nine ecoregions in the Philippine islands, including Palawan. We deviated from Udvardy (1975), MacKinnon (1997), Stattersfield (1998), and the Philippine BAP (1997) in varying degrees and based our delineation of the Philippine ecoregions on Heaney (1993). MacKinnon (1997) designates Mindoro island as subunit 26f and includes the Lubang Islands. We delineated the island of Mindoro as the Mindoro Rain Forests. References References for this ecoregion are currently consolidated in one document for the entire Indo-Pacific realm. Indo-Pacific Reference List J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 87 13. Luzon montane rain forests The ecoregion has suffered from human exploitation but still contains some of the most extensive forests left in the Philippines. Therefore, the montane forests are extremely valuable for the wide range of endemic species they support and for their role in preventing soil erosion and protecting water quality. It is one of the biologically least known ecoregions in the Philippines. Location and General Description Sierra Madre Mountains, Philippines Photograph by Jorgen Thomsen/ Conservation International 3,200 square miles (8,300 square kilometers) -Tropical and about half the size Subtropical Moist of Hawaii Broadleaf Forests Critical/Endangered Philippines Luzon is located in the western Pacific Ocean. It is the largest island in the Philippines and lies at the northern end of the island group. The Luzon Montane Rain Forests ecoregion comprises the high elevations of several mountain ranges including the Northern and Southern Sierra Madre, which parallels the northeastern coastline of Luzon. Also included in this ecoregion are Mt. Sapocoy, Mt. Magnas, and Mt. Agnamala in the northern Central Cordillera and the Zambales Mountains in the west. The geologic history of the Philippines is very complex and has had tremendous influence on the biota found there. Luzon has developed many unique species of plants and animals as a result of its long-standing isolation from other landmasses. Parts of the Luzon highlands were established as a result of volcanic activity and the friction of the Australian and Asian plates at least 15 million years ago. The highlands began to take their current form over the next 10 million years. Luzon is therefore oceanic in character, having never been connected to mainland Asia. Even during the Pleistocene, as world sea levels fell 120 m, Luzon expanded to become a larger island including the modern islands of Polillo, Marinduque, and Catanduanes but never connected to other regions of the Philippines or to mainland Asia (Heaney and Regalado 1998). Annual rainfall in the ecoregion can be as high as 10,000 mm in some areas, or about quadruple what the Luzon Rain Forests [IM0123] receive. When the rain falls, varies with the mountain ranges. The Sierra Madres are only mildly seasonal, with a dry period occurring from December to April. The mountains of the northern Central Cordillera and the Zambales Mountains are more strongly seasonal, receiving a bit less rainfall and having a longer dry period. These forests are also affected by typhoons that sweep across the South China Sea, hitting the western side of Luzon every few years. These typhoons are a major element of disturbance. J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 88 The montane forests of this ecoregion begin at about 1,000 m and are characterized by the appearance of oak and laurel species. These trees gradually replace the dipterocarp trees that dominate at lower elevations. The oaks and laurels do not have the wide buttresses of lower-elevation trees. Montane forests in general are shorter in stature than lowland forests and have less undergrowth. Epiphytes and vines (particularly pandans of the genus Freycinetia) and moss-covered branches are very common in the montane forests. The decreased temperature that accompanies increasing elevation slows the decomposition of debris (Heaney and Regalado 1998). This makes the forest floor thick with humus. The highest elevations of the montane forests sometimes are called upper montane forest or elfin forest. These forests are not treated as a separate ecoregion; rather, they are considered montane forests in extreme. Trees branches appear to be many times thicker than they actually are because the moss covering the branch is so thick. Tree height may be only a few meters, and plants that are not typically epiphytes become aerial because of the thick moisture and abundant organic material on and around trees here. Many of the endemic animal species of the Philippines are found as burrowers in the matty soil of this high-elevation forest (Heaney and Regalado 1998). Biodiversity Features The Luzon Montane Rain Forests [IM0122] ecoregion has eight species of nearendemic mammals and one species that is strictly endemic (table 1). The strictly endemic Palanan shrew-mouse, Archboldomys musseri, is known only from two specimens taken at about 1,650 m from Mt. Cetaceo in the northern Sierra Madres (Danielsen et al. 1994; Heaney et al. 1998). Species listed as threatened (VU or above) include three near-endemic species: Luzon pygmy fruit bat (Otopteropus cartilagonodus), Luzon short-nosed rat (Tryphomys adustus), and long-nosed Luzon forest mouse (Apomys sacobianus). Table 1. Endemic and Near-Endemic Mammal Species. Family Species Pteropodidae Otopteropus cartilagonodus Muridae Apomys abrae Muridae Apomys datae Muridae Apomys microdon Muridae Apomys sacobianus Muridae Archboldomys musseri* J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 89 Muridae Bullimus luzonicus Muridae Phloeomys pallidus Muridae Tryphomys adustus An asterisk signifies that the species' range is limited to this ecoregion. Five relatively large mammals inhabit the ecoregion: long-tailed macaque (Macaca fascicularis), Philippine warty pig (Sus philippensis), Philippine brown deer (Cervus mariannus), Malay civet (Viverra tangalunga), and common palm civet (Paradoxurus hermaphroditus). All are fairly widespread, and none are listed as threatened by IUCN (2000). However, habitat destruction affects all these species, and hunting affects all but the civets (Heaney et al. 1998). Additionally, the Philippine brown deer is said to be declining and is listed as data deficient by IUCN, although the species is not uncommon in appropriate habitat (Heaney et al. 1998; Wemmer 1998). The ecoregion contains twenty-eight near-endemic bird species and no strict endemics (table 2). At least eleven threatened bird species (IUCN categories VU and above) occur in the ecoregion, and two others may be present, but their distributions are poorly known (brown-banded rail [Lewinia mirificus] and Luzon buttonquail [Turnix worcesteri]) (Collar et al. 1999). Whiskered pitta (Pitta kochi) is a species typical of mossy montane forests (Dickinson et al. 1991) and is listed as vulnerable (Collar et al. 1999). Whiskered pittas usually are found above 1,000 m in the Sierra Madre and Central Cordillera among oaks, 5-12 m high, with a fern and rhododendron understory. They hunt for invertebrate prey on the ground and are known to regularly forage where Philippine warty pigs have rooted over the soil and exposed prey (Poulsen 1995). The bird is sometimes described as uncommon to rare (Dickinson et al. 1991; Kennedy et al. 2000), although in appropriate habitat it may actually be quite common (Poulsen 1995). This discrepancy probably results from undersampling (and general lack of knowledge) of habitat throughout the pitta's range. Table 2. Endemic and Near-Endemic Bird Species. Family Common Name Species Turnicidae Spotted buttonquail Turnix ocellata Turnicidae Luzon buttonquail Turnix worcesteri Rallidae Brown-banded rail Lewina mirificus Columbidae Luzon bleeding-heart Gallicolumba luzonica Columbidae Flame-breasted fruit-dove Ptilinopus marchei J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 90 Columbidae Cream-breasted fruit-dove Ptilinopus merrilli Psittacidae Luzon racquet-tail Prioniturus montanus Cuculidae Scale-feathered malkoha Phaenicophaeus cumingi Cuculidae Rufous coucal Centropus unirufus Strigidae Luzon scops-owl Otus longicornis Bucconidae Luzon hornbill Penelopides manilloe Pittidae Whiskered pitta Pitta kochi Laniidae Grey-capped shrike Lanius validirostris Turdidae Ashy thrush Zoothera cinerea Muscicapidae Luzon redstart Rhyacornis bicolor Timaliidae Golden-crowned babbler Stachyris dennistouni Timaliidae Chestnut-faced babbler Stachyris whiteheadi Sylviidae Philippine bush-warbler Cettia seebohmi Sylviidae Long-tailed bush-warbler Bradypterus caudatus Muscicapidae Rusty-flanked jungle-flycatcher Rhinomyias insignis Muscicapidae Ash-breasted flycatcher Muscicapa randi Muscicapidae Blue-breasted flycatcher Cyornis herioti Pachycephalida Green-backed whistler Pachycephala albiventris Rhabdornithidae Long-billed rhabdornis Rhabdornis grandis Dicaeidae Flame-crowned flowerpecker Dicaeum anthonyi Fringillidae White-cheeked bullfinch Pyrrhula leucogenis Estrildidae Green-faced parrotfinch Erythrura viridifacies Oriolidae White-lored oriole Oriolus albiloris An asterisk signifies that the species' range is limited to this ecoregion. J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 91 Another vulnerable species, typical of montane forests, is the flame-breasted fruit-dove (Ptilinopus marchei), a beautiful bird with a crimson orange breast and matching crown. It is the largest fruit-dove in the Philippines, and although it has probably always been uncommon and local, it appears to be particularly sensitive to the threats facing much of the ecoregion's biodiversity. The fruit-dove is not found in areas highly susceptible to habitat destruction, found in neither logged nor selectively logged areas (Poulsen 1995). The fruit-dove is also hunted for food and the pet trade (Collar et al. 1999). Current Status There are no recent estimates of primary forest cover for the Philippines. If one adds the total forest cover for the provinces containing the bulk of the ecoregion, about 482,000 ha of forest remained in 1992 (Development Alternatives 1992). A small portion of the ecoregion's montane forest occurs outside these provinces, yet the overall figure is still too high because much of the forest is below 1,000 m. More importantly, the figure is too high because the amount of forest has declined a great deal since 1992. We know this because forest cover for the same provinces totaled more than a million hectares in the early to mid-1980s, for a decline of 55 percent since 1992, and deforestation has continued, with some slowdown since 1994 (figures based on 1981 and 1984 data, depending on the particular province) (Development Alternatives 1992). The two largest remaining forested areas in the ecoregion are in the montane portions of the northern Sierra Madres, which have remained inaccessible, and the northern Central Cordillera. The Northern Sierra Madre Natural Park (also known as the Palanan complex or wilderness) is the main focus of conservation in the region, although the park covers only the lowland portions of the forest and should be extended to incorporate the higher elevations as well (Mallari and Jensen 1993; Poulsen 1995) (table 3). Other sites in the northern Sierra Madres that have been identified as important areas for biodiversity include Mt. Cetaceo and Mt. Los Dos Cuernos. Neither of these sites receives any formal protection, and both are being cleared (Collar et al. 1999). Recent field work in the northern Central Cordillera has documented extensive forest managed by the traditional cultural groups. Nearly unknown biologically until recently, it is now suspected of being one of the biologically richest and most important areas in the country (Heaney and Mallari in press). Table 3. WCMC (1997) Protected Areas That Overlap with the Ecoregion. Protected Area Area (km2) IUCN Category PNOC 1636 [IM0123] 200 ? Ecoregion numbers of protected areas that overlap with additional ecoregions are listed in brackets. Types and Severity of Threats J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 92 Habitat conversion is the primary threat to the ecoregion. Commercial logging (both legal and illegal) continues to have a devastating effect on biodiversity. Conversion of highland areas to "large-scale plantations is currently expanding, causing both displacement of subsistence farmers (who then move further upslope) and increased erosion, which is already a serious problem" (Heaney et al. 1999: 314). New roads and mining projects directly threaten forests but also make forests more susceptible exploitation as a result of increased accessibility. Subsistence hunting and capture of species for a growing wildlife pet trade adversely affect many of the ecoregion's species. Justification of Ecoregion Delineation MacKinnon (1997) identified seven subunits in the Philippines, and the Philippine Biodiversity Action Plan (Philippine BAP 1997) demarcated fifteen biogeographic units. Udvardy (1975) identified the Philippines as a single biogeographic province. We delineated nine ecoregions in the Philippine islands, including Palawan. We deviated from Udvardy (1975), MacKinnon (1997), Stattersfield et al. (1998), and the Philippine BAP (1997) to varying degrees and based our delineation of the Philippine ecoregions primarily on Heaney (1993). In Luzon we delineated three ecoregions, which correspond to MacKinnon's subunit 26a. First, we used the 1,000-m contour from the DEM (USGS 1996) to delineate the montane forests from the lowland forests. The Luzon Montane Rain Forests are made up primarily of the montane moist evergreen forests along the Sierra Madre, northern Central Cordillera, and Zambales mountain ranges. MacKinnon (1997) showed an area of freshwater swamp forests as part of the original vegetation of Luzon Island, which we combined with the remaining lowland forest of Luzon to form the Luzon Rain Forests. These freshwater swamps, in the valley to the east of the Zambales Mountain Range and in the Cagayan River plains, have been converted to rice fields (D. Madulid, pers. comm., 1999). Following Stattersfield et al. (1998) and Dickinson et al. (1991), we placed the Lubang Islands with the Luzon Rain Forests. The Banguet pine Pinus insularis (also known as P. kesiya)-dominated conifer forests in the Central Cordillera were designated as the Luzon Tropical Pine Forests. References References for this ecoregion are currently consolidated in one document for the entire Indo-Pacific realm. Indo-Pacific Reference List J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 93 14. Luzon tropical pine forests Bontoc, Mountain Province, Philippines Photograph by Larry Heaney The mountainous, often cloudy Luzon Pine Forests are a unique habitat in the Philippines. Regular fires have led to a parkland structure of grass with widely dispersed trees and prevented broadleaf trees from establishing over large areas. However, more typical montane forest covers so much of the ecoregion that some experts would lump the pine forests with the Luzon Montane Rain Forest ecoregion. The ecoregion has been exploited for its trees and mineral resources in the past; shifting agriculture and mining are the greatest current threats. Location and General Description Southeastern Asia: Island of Luzon in the Philippines 2,700 square miles (7,100 square kilometers) -about the size of Delaware and Rhode Island combined Luzon is located in the western Pacific Ocean. It is the largest island in the Philippines and lies at the northern end of the island group. The Luzon Pine Forests [IM0302] ecoregion is entirely within the Tropical and Subtropical Central Cordillera Mountain Range of Coniferous northwestern Luzon and includes all Critical/Endangered Forests regions above 1,000 m, except a large tract of montane forest at the northern end (see ecoregion 102). Included in this ecoregion are some of Luzon's highest mountain peaks: Mt. Puguis, Mt. Polis, Mt. Data, and Mt. Pulog. The geologic history of the Philippines is very complex and has had tremendous influence on the biota currently found there. Luzon has developed many unique species of plants and animals as a result of its long-standing isolation from other landmasses. Parts of the Luzon highlands were established as a result of volcanic activity and the friction of the Australian and Asian plates at least 15 million years ago. The highlands began to take their current form over the next 10 million years. Luzon is therefore oceanic in character, having never been connected to mainland Asia. Even during the Pleistocene, as world sea levels fell 120 m, Luzon expanded to become a larger island including the modern islands of Polillo, Marinduque, and Catanduanes but never connecting to other regions of the Philippines or to mainland Asia (Heaney and Regalado 1998). The ecoregion receives about 2,500 mm of rain a year (even more than 4,000 mm on Mt. Pulog in some years), but this rain is highly seasonal. Often the pine forests of Luzon are categorized as monsoon forests because of the pronounced dry period (November-April) between rain (May-August, with July and August being the wettest). The temperature of J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 94 the pine forests averages about 20°C and rarely exceeds 26°C. The forests are affected by typhoons that sweep across the South China Sea, hitting the western side of Luzon every few years. These typhoons are a major element of disturbance. Most of the ecoregion's area is covered by grassland that is interspersed with trees. The pronounced dry periods and periodic fires favor the Benguet pine or saleng (Pinus insularis, also known as P. kesiya). This species ranges in elevation between 1,000 and 2,500 m and is also present in mainland Asia. However, montane forest interdigitates with the pine forests and covers much of the ecoregion's surface area. This is particularly true in Balbalasang-Balbalan protected area, which is nearly all montane forest. Species lists for the ecoregion therefore include many animals that are limited to montane forests within the ecoregion and are not even found in pine forests. Biodiversity Features The ecoregion is inhabited by nine near-endemic mammal species and seven strictly endemic mammals (table 1). All but one of these species belongs to the mouse and rat family, Muridae, which has undergone extensive adaptive radiation on Luzon. Cloud rats (genera Crateromys and Phloeomys) occur only in the northern Philippines and are perhaps the most unique and characteristic of mammals here. These large, bushy-tailed creatures look more like squirrels than typical rats (Heaney and Regalado 1998). All are subject to heavy hunting pressures (Heaney et al. 1998). Three relatively large mammals occur in the ecoregion: long-tailed macaque (Macaca fascicularis), Philippine warty pig (Sus philippensis), and Malay civet Viverra tangalunga). All are fairly widespread, and none are listed as threatened by IUCN (IUCN 2000). However, habitat destruction affects all three species, and hunting affects all but the Malay civet. Table 1. Endemic and Near-Endemic Mammal Species. Family Species Pteropodidae Otopteropus cartilagonodus Muridae Abditomys latidens Muridae Apomys abrae Muridae Apomys datae Muridae Apomys sacobianus Muridae Batomys dentatus* Muridae Batomys granti J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 95 Muridae Bullimus luzonicus Muridae Carpomys melanurus* Muridae Carpomys phaeurus* Muridae Celaenomys silaceus* Muridae Chrotomys whiteheadi* Muridae Crateromys schadenbergi* Muridae Phloeomys pallidus Muridae Rhynchomys soricoides* Muridae Tryphomys adustus An asterisk signifies that the species' range is limited to this ecoregion. The Luzon Pine Forests [IM0302] are home to twenty-three near-endemic birds, none of which are strictly limited to the ecoregion (table 2). Perhaps the bird species most characteristic of the ecoregion is also one of its most widespread worldwide. Red crossbill (Loxia curvirostra), known primarily from high-latitude coniferous forests, reaches its southernmost extent in the Old World in the mountains of the Central Cordillera (the populations in Central America, notably Nicaragua, are further south). Dickinson et al. (1991) listed eleven other bird species that typify the pine forests, including one tit (Parus elegans), a nuthatch (Sitta frontalis), and thrush (Turdus poliocephalus). These groups of birds are familiar to many people of northern latitudes and are indicative of the unique habitat this ecoregion represents within the Philippines. Table 2. Endemic and Near-Endemic Bird Species. Family Common Name Species Turnicidae Luzon buttonquail Turnix worcesteri Rallidae Brown-banded rail Lewina mirificus Columbidae Luzon bleeding-heart Gallicolumba luzonica Columbidae Flame-breasted fruit-dove Ptilinopus marchei Psittacidae Luzon racquet-tail Prioniturus montanus Cuculidae Scale-feathered malkoha Phaenicophaeus cumingi J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 96 Strigidae Luzon scops-owl Otus longicornis Apodidae Whitehead's swiftlet Aerodramus whiteheadi Pittidae Whiskered pitta Pitta kochi Laniidae Grey-capped shrike Lanius validirostris Turdidae Ashy thrush Zoothera cinerea Muscicapidae Luzon redstart Rhyacornis bicolor Timaliidae Golden-crowned babbler Stachyris dennistouni Timaliidae Chestnut-faced babbler Stachyris whiteheadi Sylviidae Philippine bush-warbler Cettia seebohmi Sylviidae Long-tailed bush-warbler Bradypterus caudatus Muscicapidae Rusty-flanked jungle-flycatcher Rhinomyias insignis Muscicapidae Ash-breasted flycatcher Muscicapa randi Pachycephalida Green-backed whistler Pachycephala albiventris Rhabdornithidae Long-billed rhabdornis Rhabdornis grandis Dicaeidae Flame-crowned flowerpecker Dicaeum anthonyi Fringillidae White-cheeked bullfinch Pyrrhula leucogenis Oriolidae White-lored oriole Oriolus albiloris An asterisk signifies that the species' range is limited to this ecoregion. Current Status The status of the ecoregion is hard to assess from published sources. Pines from the ecoregion have been exploited for a long time. Resin from the pines provided an important commercial source of turpentine during the Spanish colonial period (Heaney and Regalado 1998). One of the protected areas in the ecoregion, Mt. Pulog, is the highest peak in Luzon (table 3). Mt. Pulog is important globally as a center of plant diversity with many local endemics (Davis et al. 1995) and as a key site for threatened birds, with six (Collar et al. 1999). The mountain, like much of the ecoregion, is valuable for water quality in Luzon. J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 97 The size of the park (11,500 ha) may be adequate, although there was initial difficulty in demarcating the park boundary when it was established in 1987 (Davis et al. 1995). The park continues to have problems with agricultural encroachment and wildlife exploitation (Collar et al. 1999). Mt. Polis is also listed as a key site for threatened birds, with six species having been recorded. Unfortunately, little forest remains on Mt. Polis, and it receives no legal protection (Collar et al. 1999). Table 3. WCMC (1997) Protected Areas That Overlap with the Ecoregion. Protected Area Area (km2) IUCN Category Balbalasang-Balbalan 160 VI Mt. Data 300 V Bessang Pass 10 III Mts. Pulog, Nueva Vizcaya, Ifugao, Benguet 80 II Total 550 Ecoregion numbers of protected areas that overlap with additional ecoregions are listed in brackets. Types and Severity of Threats The population growth of the Philippines and the extreme poverty of many has forced people to cultivate land at increasingly high altitudes. Population growth and rural poverty are definitely threats to the ecoregion. More typical threats to the biodiversity of the ecoregion exist but are perhaps symptoms of these overarching problems. Fire and habitat conversion are the greatest threats, and they are in some ways the same problem. Fire is a natural process in the ecoregion, but human-induced fire is being used to clear extensive areas for growing vegetables and cut flowers. The increased number of fires limits regeneration of some forest plants; it also makes the region susceptible to invasive grasses (Davis et al. 1995). Logging (both legal and illegal) is a threat everywhere in the Philippines and is another means of habitat destruction. Mining is an ever-present threat. The Central Cordillera is famous for its mineral wealth, including copper and gold. In fact, most of the mountain range is included within mining application and exploration areas. Wildlife exploitation is also a serious problem in many parts with species being sought for trade and consumption. Justification of Ecoregion Delineation J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 98 MacKinnon (1997) identified seven subunits in the Philippines, and the Philippine BAP (1997) demarcated fifteen biogeographic units. Udvardy (1975) identified the Philippines as a single biogeographic province. We delineated nine ecoregions in the Philippine islands, including Palawan. We deviated from Udvardy (1975), MacKinnon (1997), Stattersfield et al. (1998), and the Philippine BAP (1997) to varying degrees and based our delineation of the Philippine ecoregions primarily on Heaney (1993). In Luzon we delineated three ecoregions, which correspond to MacKinnon's subunit 26a. First, we used the 1,000-m contour from the DEM (USGS 1996) to delineate the montane forests from the lowland forests. The Luzon Montane Rain Forests are made up primarily of the montane moist evergreen forests along the Sierra Madre, northern Central Cordillera, and Zambales mountain ranges. MacKinnon (1997) shows an area of freshwater swamp forests as part of the original vegetation of Luzon Island, which we combined with the remaining lowland forest of Luzon to form the Luzon Rain Forests. These freshwater swamps, in the valley to the east of the Zambales Mountain Range and in the Cagayan River plains, have been converted to rice fields (D. Madulid, pers. comm., 1999). Following Stattersfield et al. (1998) and Dickinson et al. (1991), we placed the Lubang Islands with the Luzon Rain Forests. The conifer forests in the Central Cordillera dominated by Banguet pine (Pinus insularis, also known as P. kesiya) were designated as the Luzon Tropical Pine Forests. References References for this ecoregion are currently consolidated in one document for the entire Indo-Pacific realm. Indo-Pacific Reference List J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 99 15. Luzon rain forests The Luzon Rain Forests ecoregion is rich in endemic species and also contains one of the largest populations of the Philippine eagle Pithecophaga jefferyi. A high proportion of the ecoregion was originally forested, but now very little of this forest remains. However, the ecoregion has managed to retain one of the largest remaining tracts of primary forest in the Philippines. Location and General Description Near Sierra Madre Mountains, Philippines Luzon is located in the western Pacific Ocean. It is the largest island in the Photograph by Jorgen Thomsen/ Philippines and lies at the northern end of Conservation International the island group. The lowland rain forests ecoregion comprises all the areas below 1,000 m on Luzon and a few isolated 36,900 square Southeastern volcanic mountains in the south of the miles (95,500 Asia: Luzon island that exceed 1,000 m: Mt. Maquiling, square kilometers) Island in the Mt. Banashaw, Mt. Isarog, Mayon Volcano, -- about the size of Philippines Indiana and Rhode and Bulusan Volcano. The broad Cagayan Island combined River valley to north is sheltered from Tropical and Subtropical Moist typhoons lying between the two north-south Critical/Endangered Broadleaf mountain ranges: the Cordillera Central in Forests the west and the Sierra Madre to the east. The fertile soil of the Cagayan Valley is the biggest rice-growing region in the country. Southern Luzon is also agricultural but is subject to typhoons and comprises less area as Luzon narrows southward. Several neighboring island groups are also part of the ecoregion, including the Batanes and Babuyan Islands to the north (rather isolated but placed here for convenience), Polillo and Catanduanes to the east, and Marinduque to southwest. The geologic history of the Philippines is very complex and has had tremendous influence on the biota currently found there. Luzon has developed many unique plant and animal species as a result of its long-standing isolation from other landmasses. Parts of the Luzon highlands were established as a result of volcanic activity and the friction of the Australian and Asian plates at least 15 million years ago. The highlands began to take their current form over the next 10 million years. Luzon therefore is oceanic in character, having never been connected to mainland Asia. Even during the Pleistocene, as world sea levels fell 120 m, Luzon expanded to become a larger island including the modern islands of Polillo, Marinduque, and Catanduanes but never connecting to other regions of the Philippines or to mainland Asia (Heaney and Regalado 1998). Temperatures in Luzon vary greatly with elevation, but within the lowlands temperatures J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 100 are fairly uniform at about 25-28°C. Rainfall in the lowlands is seasonally variable, with four distinct types. Southwestern Luzon and Marinduque Island receive rain uniformly throughout the year. The Cagayan valley and the eastern portion of the Bataan Peninsula do not have pronounced seasons but are dry from November to April. The southeastern portions of Luzon and Polillo and Catanduanes Islands have no dry season but do have a period of increased rainfall from May to January. Northwestern Luzon has two distinct seasons, being wet from May to October and dry November to April. Lowland vegetation of Luzon is dominated by dipterocarp trees with wide buttresses at the base. These massive trees are 1-2 m in diameter and up to 60 m high. The canopy height of mature lowland forests tends to be uneven. In areas of disturbance, rattans and lianas receive the light they need to flourish in the understory. There tends to be an abundant herbaceous undergrowth, and ferns and orchids are prevalent on large branches of tall trees. Other natural habitats in the ecoregion include mangrove forests and beach forests (consisting of Casuarinas and Barrintonia) near the coasts. There also were natural grasslands in valley bottoms and on plateaus, as evidenced by the presence of several endemic buttonquail taxa needing grasslands (Collar et al. 1999). Biodiversity Features In terms of mammalian endemism, perhaps the most significant area of endemism in the ecoregion is Mt. Isarog, but this has only recently become apparent. Mt. Isarog is an extinct volcano and the second highest peak in southern Luzon at 1,966 m (Mt. Mayon is higher at 2,462 m). The unique character and geographic isolation of Isarog make it difficult to lump with the other two montane ecoregions of Luzon [IM0122], [IM0302], so it is considered here as part of the Luzon rain forests. The Luzon rain forest ecoregion as a whole has ten species of near-endemic mammals and five strictly endemic species (table 1). Three of the five strict endemics are found only on Mt. Isarog, and none was been described before 1981: Isarog shrew-mouse (Archboldomys luzonensis), Isarog striped shrew-rat (Chrotomys gonzalesi), and Isarog shrew-rat (Rhynchomys isarogensis). All three consume earthworms, and the latter two are strongly vermivorous. The southern Luzon giant cloud rat (Phloemys cumingi) is another of the ecoregion's strict endemics found on Isarog, but it is also found at other locations. The fifth strict endemic is the northern Luzon shrew-mouse (Crunomys fallax), known from a single specimen collected at about 300 m in the northern Sierra Madres. Table 1. Endemic and Near-Endemic Mammal Species. Family Species Pteropodidae Otopteropus cartilagonodus Muridae Abditomys latidens J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 101 Muridae Apomys abrae Muridae Apomys datae Muridae Apomys microdon Muridae Apomys sacobianus Muridae Archboldomys luzonensis* Muridae Batomys granti Muridae Bullimus luzonicus Muridae Chrotomys gonzalesi* Muridae Crunomys fallax* Muridae Phloemys cumingi* Muridae Phloemys pallidus Muridae Rhynchomys isarogensis* Muridae Tryphomys adustus An asterisk signifies that the species' range is limited to this ecoregion. The ecoregion has thirteen mammal species that are listed by IUCN as threatened (categories VU and above) (IUCN 2000). One of these species is the golden-crowned flying-fox (Acerodon jubatus). It is probably the largest bat in the world (at more than 1.2 kg, perhaps reaching 1.5 kg) and is widespread in the Philippines but has undergone a precipitous decline because of heavy hunting and habitat destruction (Heaney and Regalado 1998). Five large mammals inhabit the ecoregion: long-tailed macaque (Macaca fascicularis), Philippine warty pig (Sus philippensis), Philippine brown deer (Cervus mariannus), Malay civet (Viverra tangalunga), and common palm civet (Paradoxurus hermaphroditus). All are fairly widespread, and none are listed as threatened by IUCN (IUCN 2000). However, habitat destruction affects all these species, and hunting affects all but the civets. Additionally, the Philippine brown deer is said to be declining and is listed as data deficient by IUCN, although the species is not uncommon in appropriate habitat (Heaney et al. 1998; Wemmer 1998). The lowland forests of Luzon contain thirty-four near-endemic bird species and six strict endemics (table 2). Only two of the strict endemics are threatened (IUCN categories VU J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 102 and above): green racquet-tail Prioniturus luconensis and Isabela oriole Oriolus isabellae (Collar et al. 1999). Researchers in Luzon feared that Isabela oriole was extinct (Mallari and Jensen 1993; Poulsen 1995) until two recent reports of its existence in northern Luzon (Gamauf and Tebbich 1995; van der Linde 1995). Although these observations are encouraging, some doubt has been raised about the level of scrutiny the records were subject to (Collar et al. 1999). The green racquet-tail's decline is thought to be similar to that of many other parrots of the Philippines: it was common several decades ago but has become very rare recently as a result of deforestation and collection for the pet trade (Poulsen 1995; Snyder et al. 2000). However, Kennedy et al. (2000) stated that the decline of green racquet-tail may be less straightforward because it appears not to be heavily subject to the pet trade or to deforestation. The parrot pet trade in the Philippines has had a tremendous negative impact on certain species. The most notable example is the Philippine cockatoo (Cacatua haematuropygia), which at one time was widespread throughout the Philippines but is now scarce (if not extinct) in Luzon, an enormous decline caused almost entirely by the pet trade (Snyder et al. 2000). Table 2. Endemic and Near-Endemic Bird Species. Family Common Name Species Turnicidae Spotted buttonquail Turnix ocellata Turnicidae Luzon buttonquail Turnix worcesteri Rallidae Brown-banded rail Lewina mirificus Columbidae Luzon bleeding-heart Gallicolumba luzonica Columbidae Whistling green-pigeon Treron formosae Columbidae Flame-breasted fruit-dove Ptilinopus marchei Columbidae Cream-breasted fruit-dove Ptilinopus merrilli Psittacidae Luzon racquet-tail Prioniturus montanus Psittacidae Green racquet-tail Prioniturus luconensis* Cuculidae Red-crested malkoha Phaenicophaeus superciliosus* Cuculidae Scale-feathered malkoha Phaenicophaeus cumingi Cuculidae Rufous coucal Centropus unirufus Strigidae Luzon scops-owl Otus longicornis J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 103 Strigidae Ryukyu scops-owl Otus elegans Bucconidae Luzon hornbill Penelopides manilloe Pittidae Whiskered pitta Pitta kochi Campephagidae Blackish cuckoo-shrike Coracina coerulescens Turdidae Ashy thrush Zoothera cinerea Timaliidae Luzon wren-babbler Napothera rabori* Timaliidae Golden-crowned babbler Stachyris dennistouni Timaliidae Chestnut-faced babbler Stachyris whiteheadi Timaliidae Luzon striped-babbler Stachyris striata* Sylviidae Philippine bush-warbler Cettia seebohmi Sylviidae Long-tailed bush-warbler Bradypterus caudatus Sylviidae Grey-backed tailorbird Orthotomus derbianus Muscicapidae Rusty-flanked jungleflycatcher Rhinomyias insignis Muscicapidae Ash-breasted flycatcher Muscicapa randi Muscicapidae Furtive flycatcher Ficedula disposita* Muscicapidae Blue-breasted flycatcher Cyornis herioti Muscicapidae Luzon redstart Rhyacornis bicolor Monarchidae Short-crested monarch Hypothymis helenae Monarchidae Celestial monarch Hypothymis coelestis Pachycephalidae Green-backed whistler Pachycephala albiventris Paridae White-fronted tit Parus semilarvatus Rhabdornithidae Long-billed rhabdornis Rhabdornis grandis Dicaeidae Flame-crowned flowerpecker Dicaeum anthonyi J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 104 Zosteropidae Lowland white-eye Zosterops meyeni Estrildidae Green-faced parrotfinch Erythrura viridifacies Oriolidae White-lored oriole Oriolus albiloris Oriolidae Isabela oriole Oriolus isabellae* An asterisk signifies that the species' range is limited to this ecoregion. The Philippine eagle is the national bird of the Philippines and the most famous animal in the country. Unfortunately, the eagle is critically endangered, existing in primary lowland forests of Samar, Leyte, Mindanao, and Luzon. The large area needs of the eagle coupled with the bird's low reproductive rate have made it highly susceptible to deforestation. The two largest remaining populations are in Luzon and Mindanao, although precise numbers of individuals are still speculative. Population numbers are estimated mainly by assumptions of remaining forest cover, range size, percentage occupancy, and the number of immature birds that territories include. Luzon is thought to have between 52 and 104 eagles, but probably around 78 (Collar et al. 1999). The survival of the Philippine eagle is being watched as a benchmark of the health of the Philippines environment as a whole. The survival of the species is largely tied to the protection of the few remaining large tracts of forest. Such protection would benefit many other forest-dwelling species. Current Status The largest remaining forested area in the ecoregion is in the lowlands of the northern Sierra Madres, which have remained inaccessible. The Northern Sierra Madre Natural Park (also know as the Palanan complex or wilderness) has recorded most of the ecoregion's endemic bird species and is a stronghold for the long-tailed macaque, Philippine warty pig, and Philippine brown deer. Luzon's population of Philippine eagles is joined by thirteen other threatened bird species. The park receives funding and attention from the Department of Environment and Natural Resources (DENR), several conservation organizations, the Global Environment Facility, and the European Commission. However, the park is threatened by plans to construct roads that would transect the park and is subject to high levels of encroachment (several towns are within the park boundary). Other sites, which include lowlands in the northern Sierra Madres and have been identified as important areas for biodiversity, include Mt. Cetaceo and Mt. Los Dos Cuernos. Neither of these sites receives any formal protection, and both are being cleared (Collar et al. 1999). As noted earlier, Mt. Isarog National Park (table 3) is of major importance for endemic mammals. Mt. Isarog also contains four threatened bird species. The national park status of the Mt. Isarog has not effectively protected it thus far. Encroachment on the park has led to a population of several hundred people who live in the park. Also, deforestation continues at the hands of "well-financed commercial ventures" (Collar et al. 1999, p. 57; J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 105 see also Heaney et al. 1999 and Heaney and Regalado 1998). However, the Haribon Foundation does have an active conservation program at Mt. Isarog that is again gaining strength. Table 3. WCMC (1997) Protected Areas That Overlap with the Ecoregion. Protected Area Area (km2) IUCN Category Paoay Lake 7 III Cassamata Hill 40 V Northern Luzon Heroes Hill 110 III Lake Malimanga 6 IV Subic-Bataan Extension 90 PRO Bataan 310 II Roosevelt 20 V Biak-na-Bato 60 V Hinulugang Taktak 2 III Quezon National Park 30 III Mts. Palay-Palay-Mataas Na Gulod 40 II Fugo Island 140 V Palaui 90 V Magapit 90 IV Penablanca 120 V Callao Cave 8 V Northern Sierra Madre 260 II Fuyot Spring 30 III PNOC 1636 [IM0122] 1,060 ? J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 106 Mts. Banahaw San Cristobal 30 II Taal Volcano 20 III Bicol 30 II Quezon National Park 10 III Libmanan Cave 20 III Catanduanes 270 PRO Caramoan 30 III Bulusan Volcano 50 II Manlelung Spring 5 III Minalungao 1 V Capas Death March Monument 1 III Mt. Arayat 80 V Aurora Memorial 120 V Mt. Isarog 150 II Mayon Volcano 80 III Total 3,410 Ecoregion numbers of protected areas that overlap with additional ecoregions are listed in brackets. The smaller islands in the ecoregion have not fared well. Marinduque contained only 317 ha of primary forest in 1992, compared with 813 ha in 1984, for a loss of 61 percent (Development Alternatives 1992). The current amount of primary forest on the island is unknown. Early this century, Polillo was so forested that McGregor wrote that "he had never seen an island with 'so large a proportion of the area covered with trees,'" but today no forest remains (McGregor 1910 quoted in Collar et al. 1999, p. 55). Forest cover maps from 1992 showed that Catanduanes contained some forested areas. Central Catanduanes is listed as an important area for biodiversity, but it currently receives no protection (although it is a proposed watershed reserve) (Collar et al. 1999). Types and Severity of Threats There are no recent estimates of primary forest cover for the Philippines. J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 107 Originally 95 percent of the Philippines was covered by rain forest (Heaney and Regalado 1998). Intact lowland forest made up less than 6 percent of the total land area of the country about a decade ago, with another 12 percent being classified as degraded (Collins et al. 1991). From these figures, the state of the forest in the Philippines could be the poorest in all of tropical Asia (Poulsen 1995). In the intervening decade, more forest has been cleared as a result of unsustainable shifting agricultural practices, legal logging, and illegal logging. These threats are still present and are the main future threats to biodiversity, followed by unsustainable hunting and collection for trade. The Luzon Rain Forests [IM0123] ecoregion has been greatly modified by human activities. This is probably unavoidable given the high rate of population increase and the size of the population as a whole. Three of the country's largest cities are in this ecoregion: Manila, Quezon City, and Caloocan City. Many of the lowland areas were converted into agriculture long ago, but recent forest clearing is a tremendous problem. Still, Luzon's estimated 24 percent total forest cover was and probably is better than that of the Philippines as a whole. This is probably because Luzon has smaller trees than the rest of the country because of increasingly frequent typhoons in the northern Philippines (Collar et al. 1999), rugged mountains, and independent ethnic communities. Justification of Ecoregion Delineation MacKinnon (1997) identified seven subunits in the Philippines, and the Philippine Biodiversity Action Plan (Philippine BAP 1997) demarcated fifteen biogeographic units. Udvardy (1975) identified the Philippines as a single biogeographic province. We delineated nine ecoregions in the Philippine islands, including Palawan. We deviated from Udvardy (1975), MacKinnon (1997), Stattersfield et al. (1998), and the Philippine BAP (1997) to varying degrees and based our delineation of the Philippine ecoregions primarily on Heaney (1993). In Luzon we delineated three ecoregions, which correspond to MacKinnon's subunit 26a. First, we used the 1,000-m contour from the DEM (USGS 1996) to delineate the montane forests from the lowland forests. The Luzon Montane Rain Forests are made up primarily of the montane moist evergreen forests along the Sierra Madre, northern Central Cordillera, and Zambales mountain ranges. MacKinnon (1997) shows an area of freshwater swamp forests as part of the original vegetation of Luzon Island, which we combined with the remaining lowland forest of Luzon to form the Luzon Rain Forests. These freshwater swamps, in the valley to the east of the Zambales Mountain Range and in the Cagayan river plains, have been converted to rice fields (D. Madulid, pers. comm., 1999). Following Stattersfield et al. (1998) and Dickinson et al. (1991), we placed the Lubang Islands and Batanes and Babuyan groups with the Luzon Rain Forests. The Banguet pine (Pinus insularis, also known as P. kesiya)-dominated conifer forests in the Central Cordillera were designated as the Luzon Tropical Pine Forests. References References for this ecoregion are currently consolidated in one document for the J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 108 entire Indo-Pacific realm. Indo-Pacific Reference List 16. Palawan rain forests Palawan represents a bridge between the Sunda Shelf and Philippine bioregions and contains faunal elements from both, as well as it own unique elements. This ecoregion, though more intact than any other region in the Philippines, is under great pressure from logging interests. Coron Island, Philippines Photograph by Sterling Zumbrunn/Conservation International Philippines: Islands of Palawan, Balabac, Ursula, and the Calamain Group 5,500 square miles (14,300 square kilometers) -about the size of Connecticut and Rhode Island combined Tropical and Critical/Endangered Subtropical Moist Broadleaf Forests Location and General Description This ecoregion includes the island Palawan plus Balabac, Ursula Island, and the Calamian Group. Palawan itself is the sixth largest of the Philippine Islands. The climate of the ecoregion is tropical wet (National Geographic Society 1999). In northwest Palawan, a dry season lasts from November to May while the wet season lasts from June to October; the rest of the island experiences a short, one- to three-month dry season. The east coast becomes progressively drier than the west coast from north to south (Davis et al. 1995). Palawan (along with the Calamianes and the island of Mindoro) was rifted (below water) from the Asian mainland approximately 32 million years ago, transported through seafloor spreading across the growing South China Sea, added to the growing Philippine Archipelago approximately 17 million years ago, and uplifted above water approximately 5-10 million years ago (Hall and Holloway 1998; Dickinson, Kennedy, and Parkes 1991). Metamorphic rocks are found in the northern portion of the island north of Mt. St. Paul. Volcanic rocks are found in the vicinity of Cleopatra's Needle, just south of Mt. St. Paul. Mt. St. Paul itself and the El Nido Cliffs are karst landscapes. The southern third of the island, south of the Quezon-Aboabo Gap, is dominated by ultramafics mixed with volcanic rocks and Tertiary limestone. Tertiary sandstones and shales occur along the southwest coast (Davis et al. 1995). The channel between Palawan and Borneo is about 145 m deep. During the middle Pleistocene, sea levels were 160 m lower than today, and the islands were connected. During the last ice age (late Pleistocene), sea level was approximately 120 m below current levels, and Palawan was separated from ice age Borneo by a narrow channel. Palawan has always remained separated from the rest of the Philippines. Palawan is long J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 109 and narrow, consisting of a steep mountain range whose highest point is 2,085 m (Mt. Mantalingajan). More than 45 percent of Palawan consists of mountains with slopes greater than 30 percent (Davis et al. 1995). Vegetation types on Palawan are diverse and include beach forest, tropical lowland evergreen dipterocarp rain forest, lowland semi-deciduous forest, montane forest, and ultramafic and limestone forest. Beach forest merges with other forest types away from the coast and includes Calophyllum inophyllum, Canarium asperum var. asperum, Pometia pinnata, Palaquium dubardii, and Ficus spp. (Davis et al. 1995). The lowland evergreen dipterocarp rain forest, which naturally occupies 31 percent of the island, is dominated by Agalai spp., Dipterocarpus gracilis, D. grandiflorus, Ficus spp., Tristania spp., Exocarpus latifolius, and Swintonia foxworthyi. Sygium spp., Dracontomelon dao, and Pongamia pinnata are emergent. Lianas and cycads are common. In southern Palawan, a Casuarina sp. dominates in the lowland forests (Davis et al. 1995). The eastern half of the island is in a rain shadow and contains moist semi-deciduous forests. Soils are thin on the steeper slopes and support medium-sized trees (up to 15 m tall), which shed their leaves during the March-May dry season. The rainy season is JuneJuly. Common tree species include Pterocymbium tinctorium, Pterospermum diversifolium, Hymenodictyon spp., and Garuga floribunda (Davis et al. 1995). Montane forests, found between 800 and 1,500 m, are dominated by Tristania spp., Casuarina spp., Swietenia foxworthyi, and Litsea spp. in the lower elevations. Upper montane forest trees include Agathis philippinensis, Dacrydium pectinatum, Podocarpus polystachyus, Gnetum latifolium, Cycas wadei, Cinnamomum rupestre, Nepenthes philippinensis, and Angiopteris spp. (Davis et al. 1995). Limestone forests are found on the islets surrounding Palawan and over large areas in the southern portions of the island. Represented are Euphorbia trigona, Aglaia argentea, and Antidesma, Drypetes, Gomphandra, Sterculia, Pleomele, and Begonia spp. (Davis et al. 1995). Victoria Peak, in south-central Palawan, contains the largest region of ultramafic forest on the island. Although many of the ultramafic tree species are shared with semideciduous forest, several species, including Scaevola micrantha, Brackenridgea palustris var. foxworthi, Exocarpus latifolius, and Phyllanthus lamprophyllus are believed to be heavy metal indicators (Davis et al. 1995). Biodiversity Features Relative to the size of Palawan, the ecoregion contains a rich fauna, including several groups that are not found in the rest of the Philippines (carnivores, pangolins, porcupines, and some insectivores) (Heaney 1986). J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 110 There are many endemic mammals in Palawan, but nearly all the genera (96 percent) are also found in Borneo. Of twenty-five indigenous nonvolant mammal species, eleven (44 percent) are endemic to Palawan, and the remainder are shared with Borneo. Therefore, the greater Palawan region is rightly considered part of the Sunda Shelf bioregion rather than that of the Philippines. The large number of endemic species but few endemic genera of Palawan are consistent with a separation of Borneo and Palawan of approximately 160,000 (since the middle Pleistocene) (Heaney 1986). There are fifteen endemic or near-endemic mammals in greater Palawan (table 1). Table 1. Endemic and Near-Endemic Mammal Species. Family Species Pteropodidae Acerodon leucotis* Cervidae Axis calamianensis* Sciuridae Sundasciurus steerii* Sciuridae Sundasciurus moellendorfi* Sciuridae Sundasciurus rabori* Sciuridae Hylopetes nigripes* Muridae Chiropodomys calamianensis* Muridae Maxomys panglima* Muridae Palawanomys furvus* Hystricidae Hystrix pumila* Sorcidae Crocidura palawanensis* Muridae Haeromys sp. A* Sciuridae Sundasciurus hoogstraali* Sciuridae Sundasciurus juvencus* Tupaiidae Tupaia palawanensis* An asterisk signifies that the species' range is limited to this ecoregion. The Calamian deer (Axis calamianensis) is found only in the Calamian Islands, where it survives in low densities on Busuanga, Calauit, and Culion Islands. The only protected J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 111 area for this species was established to protect free-ranging African ungulates on Calauit Island (Wemmer 1998). Balabac, Palawan, and the Calamian Islands also provide habitat for an endemic subspecies of the bearded pig (Sus barbatus ahoenobarbus), another subspecies of which is widely distributed in the Greater Sundas. The IUCN considers this species to be rare and declining. This species naturally inhabits tropical evergreen rain forest but is able to use a wide variety of habitats within forests. They are quite dependent on fruit supplies but consume a wide variety of foods. Directional large-scale population movements in scattered or condensed herds lasting days, weeks, or even months are reported for other subspecies in Borneo and Sumatra; this is generally associated with the mast fruiting of dipterocarps. Such movements have not been reported from the Philippines (Oliver 1993). Several of Palawan's endemic mammals are considered threatened. Three endemic mammal species are considered endangered, including the Calamian deer, a Sunda tree squirrel (Sundasciurus juvencus) (recommended for delisting; Heaney et al. 1998), and the Palawan rat (Palawanomys furvus), which was collected only four times in 1962. A subspecies of mouse deer, the Balabac chevrotain (Tragulus napu nigricans), which is confined to Balabac Island, is also considered endangered. Five endemic mammal species are considered vulnerable, including Acerodon leucotis, the Palawan treeshrew (Tupaia palawanensis), the Palawan stink badger (Mydaus marchei), the Palawan binturong (Arctictis binturong whitei), and a Sunda tree squirrel (Sundasciurus rabori) (IUCN 2000). As with mammals, Philippine birds in general show a strong Bornean affinity, and it is clear that the main pathway of Asian immigration to the Philippines was through Palawan; of 395 Philippine breeding species, 137 (35 percent) also breed in Borneo. Palawan birds exhibit strong differentiation at the subspecific level when compared with its nearest Philippine neighbor, Mindoro. This is in contrast to the other partial land bridge between Borneo and the Philippines, the Sulu Islands, which have not differentiated significantly from Mindanao. Borneo and Palawan share twenty-three bird species that are not found in the rest of the Philippines. The Asian genera Polyplectron, Malacocincla, Malacopteron, Dinopium, Aegithina, Criniger, Seicercus, and Gracula are found only in Palawan within the Philippines (Dickinson et al. 1991). The island forms an important bird migration route between Borneo and the rest of the Philippines for southern migrants (Davis et al. 1995). This ecoregion corresponds exactly with the Palawan EBA (Stattersfield et al. 1998). The EBA contains twenty restricted-range birds, seventeen of which are found nowhere else on Earth and five of which (Palawan peacock-pheasant [Polyplectron emphanum], grey imperial-pigeon [Ducula pickeringii], blue-headed racquet-tail [Prioniturus platenae], falcated wren-babbler [Ptilocichla falcata], and Palawan flycatcher [Ficedula platenae]) are considered vulnerable (Collar 1999). All these vulnerable birds are dependent on lowland and hill forest (Collar et al. 1999; Stattersfield et al. 1998). There are twenty endemic or near-endemic bird species in the Palawan ecoregion (Kennedy et al. 2000; J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 112 table 2). Table 2. Endemic and Near-Endemic Bird Species. Family Common Name Species Phasianidae Palawan peacock-pheasant Polyplectron emphanum* Columbidae Grey imperial-pigeon Ducula pickeringii Psittacidae Blue-headed racquet-tail Prioniturus platenae* Strigidae Mantanani scops-owl Otus mantananensis Strigidae Palawan scops-owl Otus fuliginosus* Apodidae Palawan swiftlet Aerodramus palawanensis* Bucconidae Palawan hornbill Anthracoceros marchei* Monarchidae Blue paradise-flycatcher Terpsiphone cyanescens* Irenidae Yellow-throated leafbird Chloropsis palawanensis* Muscicapidae Palawan flycatcher Ficedula platenae* Muscicapidae Palawan blue-flycatcher Cyornis lemprieri* Muscicapidae White-vented shama Copsychus niger* Pycnonotidae Sulphur-bellied bulbul Ixos palawanensis* Timaliidae Ashy-headed babbler Malacocincla cinereiceps* Timaliidae Palawan babbler Malacopteron palawanense* Timaliidae Falcated wren-babbler Ptilocichla falcata* Timaliidae Palawan striped-babbler Stachyris hypogrammica* Paridae Palawan tit Parus amabilis* Paridae White-fronted tit Parus semilarvatus Dicaeidae Palawan flowerpecker Prionochilus plateni* An asterisk signifies that the species' range is limited to this ecoregion. J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 113 The critically endangered Philippine crocodile (Crocodylus mindorensis) was historically found on the islands of Luzon, Mindoro, Masbate, Samar, Jolo, Negros, Busuanga, and Mindanao. Busuanga contains one of the only remaining populations (others are found on Mindoro, Negros, and Mindanao). Whereas the decline of the species was initially driven by overexploitation, habitat loss and human persecution are now the principal threats to the Philippine crocodile. Surveys in 1980-1982 revealed a total wild population of approximately 500-1,000 individuals, but current wild populations may be approximately 100 nonhatchlings. Captive breeding efforts are being led by the Crocodile Farming Institute, an entity of the Philippine government (Ross 1998). A total of 1,522 (Davis et al. 1995) to 1,672 (Quinnell and Balmford 1988) vascular plants have been identified on Palawan, and it is estimated that more than 2,000 species are present on the island. As detailed earlier, Palawan has an extremely diverse range of vegetation types for the Philippines. A small number of dipterocarps, an important timber tree group, are present on the island, as well as a variety of medicinal plants used by ethnic tribes and plants used in ceremony and as ornamentals (Davis et al. 1995). Current Status Almost all of the Philippines was once completely forested (Dickinson et al. 1991). As of 1988, Palawan contained 7,410 km2 (54 percent) of total forest remaining (SSC 1988). At the time this was the highest percentage of any of the Philippines' large islands. Later aerial surveys (Development Alternatives 1992) indicated that significant reductions in closed-canopy forest cover had occurred since 1988 as a result of recent logging. As seen from the air, the lowlands and hillsides consist of slash-and-burn agriculture up to the edges of natural forest in the highlands. Closed-canopy forest caps only the highest areas on the island. Palawan's forests are of low commercial value because of the small number of dipterocarps, and until the last twenty years Palawan's forests were ignored in favor of the more valuable forests of Luzon and Mindanao. Government logging regulations setting guidelines for minimum diameter, minimum rotation length, and replanting have been largely ignored (Quinnell and Balmford 1988). Because of a generally high population density in other parts of the Philippines, large numbers of shifting cultivators (kaingineros) are attracted to Palawan to eke out a living on the hillsides of the island, and their cumulative impact is enormous (Quinnell and Balmford 1988). All of Palawan was declared a Fauna and Flora Watershed Reserve, and this includes a variety of protected areas, including national parks, wilderness areas, experimental forests, forest research reserves, game refuges, wildlife sanctuaries, museum reservations and research sites, tourist zones, and marine reserves. J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 114 Recent reports in the international press indicate (and have been confirmed, L. Heaney, pers. comm., 2000) that the situation in Palawan has stabilized, that large-scale logging has been halted, and that a balance is being achieved between economic development and conservation; future monitoring will determine whether this is remains true. Types and Severity of Threats Habitat destruction is the main threat to biodiversity in the Philippines, and Palawan, though currently in better condition, is no different. Logging and shifting cultivation (kaingin) are cited as the primary forces of habitat conversion. Logging takes many forms, from industrial scale to smaller-scale operations that use water buffalo to haul logs out of the forest. Mangroves are used locally for firewood, dyes, and tannins (Davis et al. 1995), and they are sometimes removed to make way for fishponds (Quinnell and Balmford 1988). Hunting and the wild pet trade are also significant threats in Palawan. Leopard cats have been hunted for their pelts and are sold when kittens as pets (Heaney and Regalado 1998). The Palawan binturong is hunted for meat and as pets, and the pangolin is hunted for its hide (Quinnell and Balmford 1988). The Palawan peacock-pheasant (Dickinson et al. 1991; Collar et al. 1999), blue-headed racquet-tail (Collar et al. 1999), Philippine cockatoos (Cacatua haematuropygia), and blue-naped parrots (Tanygnathus lucionensis) (Quinnell and Balmford 1988) apparently are suffering greatly from the pet trade. The final destination for these birds often is the United States (Quinnell and Balmford 1988). Ornamental plant collecting, especially for the orchids (Phalaenopsis amabilis and Paphiopedilum argus), pitcher plants (Nepenthes spp.), palms (Veitchia merrillii), and aroids (Amorphophallus spp. and Alocasia spp.) threatens some plant populations (Davis et al. 1995). A valuable resin, known as Manila copal, is collected from Agathis dammara trees. This collection weakens the trees, and slackening production and disease combined with overexploitation are threatening the species (Davis et al. 1995; Quinnell and Balmford 1988). Currently, Palawan's mineral wealth (chromite, copper, iron, manganese, mercury, and nickel) has not been extensively exploited, but the possible future extraction of these minerals represents a potential threat (Quinnell and Balmford 1988). Justification of Ecoregion Delineation MacKinnon (1997) identified seven subunits in the Philippines, and the Philippine Biodiversity Action Plan (Philippine BAP 1997) demarcated fifteen biogeographic units. Udvardy (1975) identified the Philippines as a single biogeographic province. We delineated nine ecoregions in the Philippine islands, including Palawan. We deviated from Udvardy (1975), MacKinnon (1997), Stattersfield et al. (1998), and the Philippine BAP (1997) to varying degrees and based our delineation of the Philippine ecoregions on Heaney (1993). J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 115 We placed Palawan, Calamian Islands, and Cuyo Islands into a single ecoregion, the Palawan Rain Forests. Palawan has closer zoogeographic affinities to Borneo. References References for this ecoregion are currently consolidated in one document for the entire Indo-Pacific realm. J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 116 17. Sulu Archipelago rain forests Although these islands represent transitional stepping stones from the island of Borneo to Mindanao in the Philippines, they have evolved their own distinctive faunas. Almost no forest remains on Sulu, and only the eastern portion of Tawitawi is forested. The islands are extremely politically unstable, which exacerbates a difficult conservation situation. Location and General Description Tawitawi Island, Sulu archipelago, Philippines Photograph by Arvin Diesinos 900 square miles (2,300 square kilometers) -Tropical and about the size of Subtropical Moist Rhode Island Broadleaf Forests Critical/Endangered Indo-Malay This ecoregion includes the main islands of Jolo (Sulu) and Tawitawi and the surrounding smaller islands from Sibutu up to but not including Basilan Island. The climate of the ecoregion is tropical wet (National Geographic Society 1999). There are apparently short (two-week) dry seasons in January and May on Tawitawi (Allen 1998). The Sulus are located south of the main typhoon track that so strongly influences the more northerly Philippine islands (Dickinson et al. 1991). The Philippines are essentially accreted terrains, an accretion of previously isolated island archipelagos that were brought together during the collision and partial subduction of large oceanic tectonic plates. The precursors of the Sulu Islands were an arc of submarine volcanoes that have existed for at least 25 million years. However, the Sulus were not clearly above-water islands until within the last 15 million years (Hall and Holloway 1998). The islands are low-lying and coralline (limestone). Bongao Peak, on Bongao, reaches 300 m, and Mt. Sibangkok, the highest point on the central ridge that divides Tawitawi, reaches 532 m (Allen 1998). During the Pleistocene, the majority of the present Sulu Archipelago was one island, separated from Basilan-Mindanao to the north and greater Sibutu (and Borneo) to the south by deepwater channels of 205 m and 290 m depths, respectively. The distances between these ice age islands were not great, however (Heaney 1986). Vegetation types in the Sulu Archipelago originally included beach forest, lowland rain forest, scrub forest, and mangroves (Stattersfield et al. 1998). Beach forest is composed of Barringtonia, Caesalpinia, and Terminalia. A small patch of this forest type may be found on Simunul, but this is generally an endangered habitat because of coastal development (human habitation and cultivation, coconut plantations). Formerly the most prevalent forest type on the islands (as with the rest of the Philippines), lowland rain J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 117 forest, or dipterocarp forest, is now mostly cleared. Represented dipterocarp genera include Anisoptera, Dipterocarpus, Hopea, and Shorea. Little information is available about the native scrub forest, which has been extensively cleared as well. Mangroves are found on the coasts throughout the Archipelago but are especially extensive on Tawitawi; mangroves around Bongao have been cleared. The principal mangrove genera include Rhizophora, Ceriops, Brugueira, Sonneratia, Avicennia, and Nypa (palms) (Allen 1998). Biodiversity Features Unlike that of Palawan, which is located between Borneo and the Philippines, the Sulu Archipelago's fauna is not Sundaic (Allen 1998) and, though rather small, is poorly known biologically (L. Heaney, pers. comm., 2000). Palawan was the main pathway for immigrants from Borneo to the Philippines, and the Sulus have many taxa that are identical to or derived from taxa in Mindanao. Even Sibutu, close to Borneo and separated from the rest of the Sulus by the Sibutu Passage, contains an avifauna more closely related to the Sulus than to Borneo (Dickinson et al. 1991). Although there are some Sulu birds with Sundaic distributions, the avifauna of the Archipelago is essentially Philippine (Dutson et al. 1992). The Sulu hornbill (Anthracoceros montani) is one example of an animal whose likely closest relative, the black hornbill (Anthracoceros malayanus), is from Borneo. There is a cline of relatedness to Borneo as one moves north among the islands. Sibutu contains birds of Bornean origin that are not found on Tawitawi (Allen 1998). The Sulus (Sangasanga, Bongao, Simunul, Tawitawi) also support a population of slow loris (Nycticebus coucang), a Sundaic primate that is not found in the remainder of the Philippines (Heaney 1986). There is one endemic mammal in the ecoregion (table 1). The Tawitawi Island rat (Rattus taitawiensis) is considered vulnerable (IUCN 2000). Table 1. Endemic and Near-Endemic Mammal Species. Family Species Muridae Rattus tawitawiensis* An asterisk signifies that the species' range is limited to this ecoregion. A new pig species, Sus spp. nov., is being described from the Sulus on the basis of MtDNA and skull measurements from a dead specimen (Rose and Grubb, in prep.). The same subspecies of bearded pig found on Borneo (Sus barbatus barbatus) is also found in the southwestern Sulus (Sibutu and Tawitawi), and this species can still be observed crossing open water to reach these islands from Borneo. It is unknown whether these over-water migrations are related to periodic eruptions on the Bornean mainland (Oliver 1993). This ecoregion overlaps exactly with the Sulu Archipelago EBA. The EBA contains nine restricted-range birds, four of which are limited to the Sulus. All the restricted-range J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 118 birds are forest species. Ten bird species qualify as endemic or near endemic to this ecoregion (Kennedy et al. 2000; table 2). Included in the ecoregion are the critically endangered Sulu bleeding-heart (Gallicolumba menagei), Tawitawi brown-dove (Phapitreron cinereiceps), and Sulu hornbill (Anthracoceros montani) and the endangered blue-winged racquet-tail (Prionoturus verticalis). Several endemic bird subspecies may warrant elevation to species status upon detailed review (Stattersfield et al. 1998; Collar et al. 1999). Table 2. Endemic and Near-Endemic Bird Species. Family Common Name Species Columbidae Sulu bleeding-heart Gallicolumba menagei* Columbidae Dark-eared dove Phapitreron cinereiceps* Columbidae Grey imperial-pigeon Ducula pickeringii Psittacidae Blue-winged racquet-tail Prioniturus verticalis* Strigidae Mantanani scops-owl Otus mantananensis Apodidae Philippine needletail Mearnsia picina Bucconidae Sulu hornbill Anthracoceros montani* Monarchidae Celestial monarch Hypothymis coelestis Pycnonotidae Yellowish bulbul Ixos everetti Timaliidae Brown tit-babbler Macronous striaticeps An asterisk signifies that the species' range is limited to this ecoregion. Several widespread but threatened species also occur on the islands, including the critically endangered Philippine cockatoo (Cacatua haematuropygia) and vulnerable rufous-lored kingfisher (Todirhamphus winchelli) (Collar et al. 1999; Stattersfield et al. 1998). The critically endangered Philippine crocodile (Crocodylus mindorensis) was historically found on Jolo (as well as Luzon, Mindoro, Masbate, Samar, Negros, Busuanga, and Mindanao), but the only remaining populations are found on Mindoro, Negros, Mindanao, and Busuanga. The current wild population may be approximately 100 nonhatchlings (Ross 1998). Current Status J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 119 There is almost no forest remaining on Jolo (Sulu) Island, and only the eastern and north-central portions of Tawitawi are forested (Stattersfield et al. 1998; Allen 1998). The majority of Tawitawi was selectively logged in the 1960s and early 1970s (Allen 1998). Apparently, there were plans to replace the remaining forests of Tawitawi with oil palm plantations. The situation on the smaller islands is mixed. Sibutu and Simunul have been largely cleared (Stattersfield et al. 1998). Simunul has some patches of forest remaining that support populations of Philippine cockatoo (Cacatua haematuropygia), blue-naped parrot (Tanygnathus lucionensis), and blue-backed parrot (T. sumatranus) (Dutson et al. 1992). Sibutu has considerable secondary forest, and the island supports numerous Sulu subspecies. The last forests of Sangasanga were cleared in 19921993. The island of Bongao still supports forests; an unidentified jungle flycatcher collected in 1973 has not been observed since (Dutson et al. 1992). Small islands in the Tandubas group (which also includes Tawitawi and Sangasanga) still have small forest tracts that reportedly maintain populations of the endemic Sulu bleeding-heart and Sulu hornbill (Stattersfield et al. 1998). The main population center is on Bongao, where a busy port exists. Tawitawi is not heavily populated, but future economic development on the island is a concern (Allen 1998; Stattersfield et al. 1998). Table 3 details the existing protected area on the islands. Table 3. WCMC (1997) Protected Areas That Overlap with the Ecoregion. Protected Area Area (km2) IUCN Category Mt. Dajo 40 IV Ecoregion numbers of protected areas that overlap with additional ecoregions are listed in brackets. Types and Severity of Threats In general, habitat loss is the main threat to wildlife, but hunting is also a problem. Small-scale logging continues to destroy the remaining habitat (Stattersfield et al. 1998). Justification of Ecoregion Delineation MacKinnon (1997) identified seven subunits in the Philippines, and the Philippine BAP (Philippine DENR & UNEP 1997) demarcated fifteen biogeographic units. Udvardy (1975) identified the Philippines as a single biogeographic province. We delineated nine ecoregions in the Philippine islands, including Palawan. We deviated from Udvardy (1975), MacKinnon (1997), Stattersfield et al. (1998), and the Philippine BAP (Philippine DENR & UNEP 1997) to varying degrees and based our delineation of the Philippine ecoregions J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 120 on Heaney (1993). The islands of the Sulu Archipelago were delineated as a separate ecoregion, the Sulu Archipelago Rain Forests. This ecoregion includes the Tawitawi Group, Tapul Group, Jolo Group, and Samales Group of islands. These islands, with a lowland moist or semievergreen moist forest vegetation (Whitmore 1984), are also an EBA (Stattersfield et al. 1998) and have been identified as a distinct biounit by MacKinnon (1997) and a biogeographic zone by the Philippine BAP (Philippine DENR & UNEP 1997) References References for this ecoregion are currently consolidated in one document for the entire Indo-Pacific realm. J. S. Tasirin: Suplemen Kuliah Ekologi Hutan Wallacea 121