archaeological, linguistic, or human biological evidence. However, it is certain that later Polynesian voyaging canoes had the technical capability to reach the Americas, and to return. That the Polynesians did indeed reach South America and return is strongly suggested by preserved remains of the sweet potato (Ipomoea batatas), a South American domesticate, recovered from several prehistoric Polynesian sites (such as the Tangatatau rockshelter on Mangaia, Cook Islands). Further evidence of contact has recently been provided by the bones of chickens (Gallus gallus, a Polynesian domesticate) from a pre-Columbian archeological site in Chile. More controversial has been the claim that Polynesians may have introduced the technology of plank-built canoes to the Chumash of the Channel Islands, off the coast of California. SEE ALSO THE FOLLOWING ARTICLES Archaeology / Cook Islands / Easter Island / Exploration and Discovery / Human Impacts, Pre-European / Kon-Tiki / Polynesian Voyaging / Tonga FURTHER READING Bellwood, P. 1985. Prehistory of the Indo-Malaysian archipelago, revised ed. Honolulu: University of Hawaii Press. Irwin, G. 1992. The prehistoric exploration and colonisation of the Pacific. Cambridge: Cambridge University Press. Kirch, P. V. 2000. On the road of the winds: an archaeological history of the Pacific Islands before European contact. Berkeley: University of California Press. Kirch, P. V., and R. C. Green. 2001. Hawaiki, Ancestral Polynesia: an essay in historical anthropology. Cambridge: Cambridge University Press. Lilley, I., ed. 2006. Archaeology of Oceania: Australia and the Pacific Islands. Oxford: Blackwell Publishing. Spriggs, M. 1997. The island Melanesians. Oxford: Blackwell Publishing. PHILIPPINES, BIOLOGY RAFE M. BROWN University of Kansas, Lawrence ARVIN C. DIESMOS National Museum of the Philippines, Manila The Philippines (Fig. 1) is one of the Earth’s most spectacular island archipelagoes. The country spans the Asian– Australian faunal zone interface at the sharpest biotic demarcation (Wallace’s Line) on the planet. Although collectively comprising a land mass approximately the size of the U.S. state of Arizona, the Philippines is a complex archipelago with more than 7100 distinct islands. Geographically situated on the edge of multiple colliding tectonic plates, the Philippines has an ancient and complex geological history that has only recently come to light. Ancient land mass movements, environmental gradients along steep volcanic slopes, and sea level–induced alterations of connectivity between neighboring islands have all presumably fueled the in situ evolutionary process of diversification on a magnitude seen in few other island systems. The result is a spectacular array of biodiversity, unparalleled levels of vertebrate endemism, and some of the planet’s most spectacular examples of life on islands. The Philippines is a global superpower of biodiversity that may possess one of the highest concentrations of vertebrate life on Earth. FIGURE 1 The position of the Philippines (darkly shaded islands) in relation to surrounding Southeast Asian and Australasian land masses. The positions of Wallace’s and Huxley’s lines are indicated for reference. Wallace’s line marks the edge of the Sunda Shelf and the transition between Asian and Australian faunal regions. Huxley’s line separates Palawan and associated land bridge islands from the truly oceanic portions of the Philippines. PHILIPPINES, BIOLOGY Gillespie08_P.indd 723 723 4/30/09 10:57:15 AM FIGURE 2 Species exemplars from the extraordinarily diverse endemic Philippine rodent radiation: (A) Luzon hairy-tailed rat, Batomys granti; (B) Cordillera striped earth-mouse, Chrotomys whiteheadi; (C) Luzon needle-nosed shrew-rat, Rynchomys cf soricoides; (D) Kalinga shrewmouse, Archboldomys kalinga; (E) silver earth-mouse, Crotomys silaceus; and (F) common Philippine forest rat, Rattus everetti. Photographs by Rafe M. Brown. GEOGRAPHICAL AND GEOLOGICAL SETTING The Philippines is situated south of Taiwan and north of Borneo and Sulawesi (between the equator and the Tropic of Cancer), separated from the Asian mainland by the South China Sea. Collectively the archipelago covers 2 million km2 with a total land mass of approximately 300,000 km2. The western part of the archipelago (Palawan, Balabac, Busuanga, Coron, and smaller associated islands) has had a stable geologic configuration and has been intermittently connected (or nearly connected) to northern Borneo via a narrow land bridge during maximum exposure of the world’s largest continental shelf (the Sunda Shelf ). The remaining portions of the Philippines are classified as oceanic islands and are believed to have risen directly from the ocean floor as a result of volcanic activity associated with subduction of the Philippine Plate at the western boundary of the Pacific Ring of Fire. BIODIVERSITY AND ENDEMISM The Philippines is recognized internationally as a global stronghold of biodiversity. The shallow, warm seas sur724 Gillespie08_P.indd 724 rounding the country support the richest coral reef communities on the planet and are literally the epicenter of Southeast Asian and southwestern Pacific marine diversity. Terrestrial ecosystems in the Philippines are similarly diverse, supporting a wealth of natural resources, habitat heterogeneity, and a rich array of species diversity. High levels of alpha-diversity (species richness) and beta-diversity (variation among adjacent regions) place the Philippines among the world’s most biodiversity-rich countries. Some of this diversity is truly astonishing. Highlights include dwarf water buffalos or “Tamaraw” (Bubalus mindorensis), the largest extant forest eagle in the world (Pithecophaga jefferyi), a radiation of the world’s largest flowers (genus Rafflesia), the rarest batagurid turtle in the world (Siebenrockiella leytensis), an extensive and exceptionally diverse rodent radiation (including such novelties as raccoon-sized “cloud rats” and needle-nosed shrew-rats that feed entirely on worms; Fig. 2), four endemic genera of snakes, five endemic species of spectacularly maned wild pigs, and such “living fossils” as the primitive and relictual flat-headed frog (Barbourula busuangenis). The fossil record shows that many extraordinary, large-bodied animals have gone extinct with the arrival of modern humans, including dwarf buffalos, elephants, and giant land tortoises. Terrestrial ecosystems are fantastically diverse. Recent summaries of birds recognized 593 species (32% endemic); mammal diversity currently is estimated at 175 native terrestrial mammals (65% endemic). Recent reviews of the classification of Philippine amphibians and reptiles recognized 105 species of amphibians (79% endemic) and 264 reptiles (68% endemic). These estimates emphasize conspicuous terrestrial vertebrates, but total country estimates (Table 1) are awe inspiring, with as many as 15,000 plants (and their relatives) and 38,000+ animals (vertebrates and invertebrates), for a startling total of 53,500 species. These numbers should be viewed as conservative approximations; numerous recent studies have shown that terrestrial biodiversity of the Philippines is substantially underestimated, in some cases grossly so. In poorly studied groups such as earthworms, more than a hundred TABLE 1 A Summary of Recent Estimates of Total Countrywide Philippine Species Diversity Taxonomic Group Philippine Total Vertebrates Invertebrates Plants Others (algae, lichens, fungi, etc.) Total 3,308–3,325 34,940–35,000 14,000–15,310 6,100+ 53,500+ PHILIPPINES, BIOLOGY 4/22/09 3:33:49 PM FIGURE 3 Estimated Cenozoic movements of Philippine land masses (modified from work of Hall 1998) 10, 20, and 30 million years ago. new species have recently been discovered. Molecular phylogenetic studies have confirmed the exceptional species diversity of numerous Philippine radiations; many have uncovered suites of previously unrecognized, cryptic species. Several of these key studies of species boundaries have produced startling results and increased species diversity in selected groups by 30–50%. Virtually every molecular phylogenetic study that has been published in the last 10 years has included the discovery of new hidden species, including many groups of frogs, lizards, insects, worms, forest mice, bats, rats, and birds. One example is the case of the Philippine forest frogs of the genus Platymantis. Past studies were based solely on morphology; species numbers grew slowly from 7 to 12 named species between 1950 and 1990. It was at this point that a group of herpetologists began to focus on bioacoustic characters (analysis of the mating calls of male frogs) and applied molecular techniques (DNA sequence data) to the problem of species boundaries in this group. By emphasizing different aspects of the phenotype, these workers provided fine-scale taxonomic partitioning of FIGURE 4 Spectacular Philippine endemic species: (A) the endemic Philippine freshwater crocodile, Crocodylus mindorensis (photograph by Rafe M. Brown); (B) the newly discovered Calayan Island flightless rail, Gallirallus calayanensis (photograph by Marge Babon/CEAE); (C) the giant flower Rafflesia manillana (photograph by Arvin Diesmos); (D) and one of the world’s only fruit-eating monitor lizards, Varanus olivaceus (photograph by Charles Linkem). PHILIPPINES, BIOLOGY Gillespie08_P.indd 725 725 4/30/09 10:38:46 AM forest frogs and opened the way for a flood of discoveries and new species descriptions. It is clear that Platymantis, currently estimated at 27 described species, represents one of the Earth’s major island radiations of frogs. Several dozen species remain to be described. When the same analytical tools are applied to other frog groups throughout the archipelago, it becomes clear that the species diversity of Philippine Amphibia has been underestimated by as much as 30–40%. Despite the lack of a complete knowledge of the biodiversity of the Philippines, today’s taxonomic estimates allow researchers to calculate the number of species per unit area. When this calculation is carried out for terrestrial species, given the available land mass (300,000 km2), the end result is the highest concentration of biodiversity on Earth. BIOGEOGRAPHY AND PROCESSES OF DIVERSIFICATION FIGURE 5 Spectacular Philippine endemic species: (A) the recently discovered Mindoro stripe-faced flying fox, Styloctenium mindorensis (photograph by Jake Esselstyn); (B) the rarest batagurid turtle in the world, the Philippine forest turtle, Siebenrockiella leytensis (photograph by Rafe M. Brown); (C) the endemic Philippine dwarf water buffalo or Tamaraw, Bubalus mindorensis (photograph courtesy of Department of the Environment and Natural Resources–Protected Areas and Wildlife Bureau Tamaraw Conservation Program); (D) the brightly colored Igorot frog, Rana igorota (photograph by Rafe M. Brown). 726 Gillespie08_P.indd 726 Several generations of biologists have converged on a startling consensus: The extraordinary biodiversity of the Philippines has likely been produced by three major processes of the geographic template (i.e., geology + climate). We consider these process to be the major “generators” of diversity in the Philippines. The first involves the complex geological history of the archipelago. Briefly, the formation of the Philippines involves more than 50 million years of collisions of multiple separate plates of the earth’s crust, resulting in a history characterized by constantly changing configurations of islands. The Philippines of 10, 20, or 30 million years ago looked almost nothing like the country today (Fig. 3). The inferred result of these ancient movements of land masses is that the distant ancestors of many of today’s Philippine endemic species may have first invaded and become isolated on Philippine paleo-islands several to many tens of millions of years before present. These events appear to have given rise to deep phylogenetic diversity and the presence of numerous “old endemics,” or highly divergent, taxonomically distinctive taxa (Figs. 4 and 5). The second generator of biodiversity in the Philippines is the process of evolutionary differentiation along dramatic elevational, atmospheric, and environmental gradients from sea level to 2000+ meters (Fig. 6). The oceanic portions of the Philippines are islands produced primarily by volcanism. Many of the smaller islands (and many isolated peaks within larger islands) were formed as active volcanoes arose from the ocean floor over the last 100 million years. Steep elevational gradients (replicated PHILIPPINES, BIOLOGY 4/30/09 10:38:49 AM numerous times along dozens of isolated volcanic peaks) have given rise to atmospheric variation, microclimate variability, and forest community heterogeneity—resulting in biodiversity gradients along these sheer mountain slopes. As a result, the study of Philippine biodiversity is largely the study of species succession along environmental gradients. The last 50 years of biodiversity research in the Philippines have documented the enormous impact that elevation-associated atmospheric and habitat gradients that have played a prominent role in the evolutionary processes of diversification. The third generator of Philippine biodiversity (and the one that is, for better or for worse, most often invoked) is the repeated “species pump” action of rising and falling sea levels between the mid- to late Pleistocene (350,000–12,000 years ago). Repeated cooling episodes during this recent time period resulted in the capture of the Earth’s water at the expanding polar ice caps and a concomitant lowering of sea levels throughout the globe. In the Philippines the result of these sea level oscillations was repeated episodes of connectivity and isolation between islands separated by channels of 120–180 meters (Fig. 7). Biogeographers now recognize at least eight faunal provinces that correspond to these Pleistocene Aggregate Island Complexes (PAICs), which represent maximum exposure of expanded islands. Each PAIC is a major center of biodiversity and endemism, with distinctive flora and fauna. The tracing of submarine bathymetric contours to reveal Pleistocene exposure of land provides biologists with an estimate of former paleoisland connection and serves as a basis for predictions of taxonomic affinity (in the absence of phylogenetic data) for the species inhabiting those islands. This exercise has also identified several minor subcenters of biological diversity in the form of small islands. Finally, it is clear that both in situ evolutionary diversification and repeated colonization events have contributed to the accumulation of diversity in the archipelago. The development of molecular phylogenetic methods has allowed for unprecedented study of both biodiversity in general and the historical and temporal framework for FIGURE 6 Striking elevational gradients and volcanic landscapes of the Philippines: (A) Pagudpud area, northern Luzon Island; (B) Lake Danao, a high-elevation volcanic crater lake in northern Leyte Island; (C) the saw-toothed peaks of Mt. Guiting-guiting, Sibuyan Island; and (D) terraced fields along the slopes of the northern Cordillera Mountain range (between Bontok and Banaue). Photographs by Rafe M. Brown. PHILIPPINES, BIOLOGY Gillespie08_P.indd 727 727 4/30/09 10:38:50 AM FPO FIGURE 7 A detailed map of the Philippines (light green shaded islands) with Pleistocene Aggregate Island Complexes (PAICs) indicated by tracing of 120 m underwater bathymetric contour. Areas shaded purple indicate shallow seas that may have been exposed as many as ten times during the middle- to late Pleistocene. These land-positive connections (land bridges) may have allowed for past floral and faunal exchange between islands that today are isolated by marine channels. the generation of Philippine biodiversity. Several modern phylogenetic studies have produced surprises that contradict earlier hypotheses and threaten to topple the prevailing view of diversification in the Philippines. The result is an emerging consensus suggesting that Philippine biodiversity is far more complex than previously thought. Recent studies of Philippine flying lizards, forest geckos, spotted stream frogs, and forest mice suggest 728 Gillespie08_P.indd 728 that for some taxa, periodic sea level oscillations have had a profound impact on the production and distribution of species diversity. These studies have all demonstrated a near complete adherence of species to PAIC boundaries and Pleistocene seashores. For much of the terrestrial vertebrate life in the Philippines, these historical events have played a predominant role in shaping species distributions—much more so, in fact, than the PHILIPPINES, BIOLOGY 4/22/09 3:33:56 PM traditional biogeographic variables such as island size and distance from mainland. This dominant paradigm of Philippine biogeography is so prevalent, in fact, that the shared presence of species across PAIC boundaries is now viewed as an extreme outlier. For the most part, this has been a healthy development for the understanding of Philippine biodiversity. By heightened attention to the unique evolutionary histories of PAIC endemics, biogeographers are now beginning to arrive at an appreciation of Philippine biodiversity that is concordant with evolutionary history—one that recognizes distinct and cohesive lineage segments as evolutionary species. The end result is that a truly evolutionary classification of Philippine vertebrate life is now perceived as an obtainable goal. Several recent studies have dramatically upset accepted scenarios concerning the predominant processes of evolutionary diversification in the Philippines. One study, a phylogenetic analysis of Asian “Fanged Frogs” conducted by Ben Evans and colleagues, revealed a pervasive and entirely unexpected zoogeographic link between Sulawesi and the Philippines. In that study, DNA sequence data revealed that multiple over-water dispersal events have allowed for successive waves of exchange between Sulawesi and the Philippines. As might be expected, the oldest invasions gave rise to proportionately more species and have spread across more PAICs than the younger arrivals. This Philippine–Sulawesi connection stands in opposition to all previous zoogeographic evidence provided by the last 25 years of studies of Philippine vertebrates, particularly mammals. Additional studies have upset the land-bridge (Palawan) versus oceanic (remaining Philippines) dichotomy. Phylogenetic studies of fanged frogs, spiny rats, shrews, litter frogs, spotted stream frogs, slender stream frogs, emerald tree skinks, flying lizards, geckos, fresh water fish, river shrimp, and numerous groups of insects all demonstrate that a large portion of Palawan’s endemics are most closely related to the truly oceanic portions of the Philippines, to the exclusion of the (expected) species from the islands of the Sunda Shelf. In fact, many Palawan endemics are nested within the truly Philippine radiation, suggesting recent dispersal to Palawan from the oceanic portions of the Philippines, and a lack of faunal exchange with Borneo. The simple depiction of Palawan as a faunal extension of northern Borneo (based largely on mammal and bird taxonomy) is a taxon-biased expectation that is not supported by the bulk of available phylogenetic evidence. In fact, Palawan is an exceptional amalgamation of greatly divergent ancient taxa, recent dispersal events from Borneo (e.g., mammals), and much older biogeographic elements that are nested within Philippine radiations and represent old dispersal events from the oceanic portions of the archipelago. In this sense, the slender island of Palawan represents a true biogeographic novelty, literally a crossroads spanning the most prominent biogeographic boundary in the world and one of the exceptions to expectations based on Wallace’s and Huxley’s Lines. An additional class of exceptions to the PAIC-centric tradition of biogeography in the Philippines is the increasingly prevalent pattern of micro-endemism on small land bridge islands. Because small land bridge (or even deep-water) islands next to the larger land masses in the Philippines were expected to possess a nested subset of species diversity observed on large neighboring islands, they have often gone unsurveyed and unscrutinized by biologists. Many of these tiny, seemingly irrelevant islands are now known to harbor endemics, some of which are spectacularly divergent evolutionary novelties. Examples include Lubang, Camiguin Sur, Dinagat, Siargao, Sibuyan, and the Gigante Islands. All demonstrate extensive levels of endemism, commensurate with their status as deepwater minor subcenters of diversity. Recent work suggests that our collective knowledge of these minor subcenters of Philippine biodiversity is far from complete. A final exception to the prevailing paradigm of PAIClevel partitioning of biodiversity of the Philippines is an emerging pattern of autochthonous speciation on the large islands. Numerous recent phylogenetic studies involving amphibians, reptiles, shrews, bats, rats, birds, plants, and insects have drawn attention to patterns of endemism within the large islands of Luzon, Palawan, Mindanao, Negros, Panay, and Samar-Leyte. These studies provide evidence of evolutionary and ecological processes (e.g., other than rising sea level vicariance) fueling diversification. Many of these are very interesting because they provide support for adaptive evolution, sympatric, and/or ecological speciation within larger islands. As such, these examples stand in contrast to the “passive” process of non-adaptive divergence following isolation inferred for the diversification of many other groups of species in the archipelago. HUMAN HISTORY AND IMPACT Colonizing from mainland Asia, humans first arrived in the Philippines many thousands of years ago. Before the arrival of the Spanish (sixteenth century), the Philippines was 90–95% forested, with only scattered settlements surrounding various coastal areas. Despite a growing population and a great demand for timber, the next 300 years probably resulted in the loss of only an additional 20% of forest cover. We know that at the turn of the twentieth century, Cebu Island (Ferdinand Magellan’s capitol) was PHILIPPINES, BIOLOGY Gillespie08_P.indd 729 729 4/22/09 3:33:59 PM almost completely deforested and that nearby islands of Negros, Guimaras, and Panay were covered by less than half of their original forest area. But other islands maintained vast expanses of intact forest, and perhaps as much as 70% of the country’s forest persisted. The industrial and agricultural revolution, periods of American and Japanese occupation, plus the period of economic expansion following World War II took an extremely heavy toll on Philippine forests. During this the last century, it is estimated that the Philippines lost between 50% and 70% of its original forest cover. During this post-war period, many factors weighed heavily on Philippine biodiversity, including high human population growth associated with the industrial revolution (and a concomitant application of low-level pressure on forests; Fig. 8), a long tradition of colonial control and exploitation, shortcomings in the country’s (U.S.-installed) education system, crippling poverty in many undeveloped portions of the country, and chronic government graft FIGURE 8 Primary causes of deforestation in the Philippines: (A) Large-scale commercial logging was introduced to the Philippines by the Americans. In this image, loggers use refurbished World War II weapons carriers to skid mature hardwood trunks from lowland forest of Mt. Busa, South Cotobato Province, southern Mindanao Island. (B) Small-scale timber poaching and clearing of land for agricultural purposes. Kaingineros like this man from Mt. Malinao, Albay Province, Luzon Island, routinely clear small patches of forest, burn debris, and then plant a single crop of rapidly-growing vegetables for sustenance and sale at local markets. Photographs by Rafe M. Brown. 730 Gillespie08_P.indd 730 and corruption that have pandered to a wealthy elite and foreign hegemony. However, by far the most destructive force has been unchecked environmental destruction associated with largescale logging and mining. Large-scale logging and mining were introduced to the country during the American occupation. Over the last century, the Philippines has been the target of wholesale exploitation of natural resources combined with wide-scale plantation-style agricultural efforts (principally hemp, bananas, copra, and sugar cane), which have converted many of the country’s most fertile natural ecosystems into monoculture and barren wasteland. The end result of this unchecked colonial rule and political instability has been systematic exploitation of Philippine natural resources, and catastrophic environmental destruction at rates exceeding those almost anywhere on the planet. Originally >90% forested, the Philippines now retain only 4–8% of its original old-growth forest. CONSERVATION PROSPECTS AND CHALLENGES FOR THE FUTURE The Philippines shares only with Madagascar the distinction of being both a megadiverse country and a Global Conservation Hotspot. Despite a greater understanding of Philippine biodiversity gained during the past 15 years, such knowledge is accumulating too slowly to conserve and stem the loss of the archipelago’s spectacular biodiversity. There can be no doubt: the Philippines is of the planet’s highest conservation priorities. Part of the roots of the global conservation crisis currently unfolding in the Philippines is related to the so-called Linnaean shortfall: the disparity between where taxonomists have focused their attention over the last 400 years (e.g., predominantly vertebrates, selected insect groups, and angiosperms) and where the greatest proportions of undescrbed biodiversity remain to be studied (invertebrates, other plants, parasites, and soil microbes, among many other groups). The Linnaean shortfall represents a failure of taxonomy and at the same time a tremendous opportunity for future generations of researchers who choose to focus their attention on these lesser-known groups. Nevertheless, because a lack of knowledge of biodiversity in part contributes to its wanton exploitation and neglect, the end result of this ignorance is biodiversity loss and extinction. Despite these bleak assessments, there is increasing cause for hope and renewed efforts towards fostering sustainable development, ecosystem restoration, and reliance on renewable natural resources (e.g., nontimber forest products such as rattan). The National Integrated Protected Areas System (NIPAS) Act has established a protocol for the establish- PHILIPPINES, BIOLOGY 4/30/09 10:42:08 AM ment and support of nationally protected areas, and establishment of new parks continues to this day at a growing rate. Grassroots community environmentalism has sprung up throughout the country, and a growing protectedarea system continues to spread coverage and jurisdiction over the remaining forested regions. The country’s National Commission on Indigenous Peoples has taken a strong leadership role in protecting natural resources through management of tribal ancestral homelands, and an expanding base of young, energetic conservation biologists have enthusiastically taken up the battle to protect the country’s natural heritage. There is great cause for hope in Philippine conservation, and unique Philippine flagship species (Fig. 9) play an increasingly important role in spreading environmental awareness to the Filipino public. An understanding of Philippine biodiversity will be greatly enhanced by renewed, vigorous attention to four primary avenues of conservation-related research. First and foremost, these include a need for large-scale, countrywide, and faunistically comprehensive surveys of the remaining natural habitats of the Philippines. Great progress has been made by a handful of researchers over the past 20 years, but this progress needs to be increased by an order of magnitude in both scope and urgency. The next several decades must see a massive resurgence in biodiversity studies if the Philippine conservation crisis (and expected catastrophic extinction event) is to be averted. Second, thorough taxonomic revisionary studies will be required to avert the Linnaean shortfall in the Philippines and promote biodiversity conservation as a global priority. This field of study is surprisingly thankless and increasingly threatened by changing academic landscapes and emphasis on “high-impact” publications in science. Third, the future is very bright for molecular phylogenetic studies of endemic Philippine radiations. Many of the truly spectacular Philippine radiations have gone unstudied. A much-needed synthesis of geological evidence and time-calibrated phylogenetic studies shows tremendous promise for exposing the temporal framework for evolutionary diversification of Philippine biodiversity. Finally, for a variety of reasons, it is clear that the future of biodiversity research and conservation in the Philippines is an effort that must be led by Filipinos. For this to occur, public and government perception of biologists, societal values, and educational emphasis must all shift to endorse the preservation of natural resources and environmental quality. Aside from their inherent value, biodiversity, forested ecosystems, and environmental health all provide societal services such as clean water, food, renewable resources, and buffering from inclement weather. Filipinos have a rich cultural and historical legacy that is tightly linked to their natural heritage through these natural resources. These must all be celebrated and preserved for future generations. FIGURE 9 Flagship species of Philippine biodiversity conservation and symbols of an emerging surge in environmentalism: (A) the poorly-known Philippine tarsier, Tarsius syrichta; (B) the largest eagle in the world, the Philippine “monkey-eating” eagle, Pithecophaga jefferyi. Photographs by Rafe M. Brown. SEE ALSO THE FOLLOWING ARTICLES Borneo / Frogs / Indonesia, Biology / Madagascar / Philippines, Geology / Sustainability / Wallace’s Line PHILIPPINES, BIOLOGY Gillespie08_P.indd 731 731 4/22/09 3:34:00 PM FURTHER READING Brown, R. M. 2007. Introduction, in Systematics and zoogeography of Philippine Amphibia. R. F. Inger. Kota Kinabalu, Malaysia: Natural History Publications, 1–17. Brown, R. M., A. C. Diesmos, and A. C. Alcala. 2002. The state of Philippine herpetology and the challenges for the next decade. The Silliman Journal 42: 18–87. Catibog-Sinha, C. S., and L. R. Heaney. 2006. Philippine biodiversity: principles and practice. Quezon City, Philippines: Haribon Foundation for Conservation of Natural Resources. Collar, N. J., N. A. D. Mallari, and B. Tabaranza, Jr. 1999. Threatened birds of the Philippines. Makati City, Philippines: Bookmark, Inc. Department of Environmental Resources and United Nations Environment Programme. 1997. Philippine biodiversity: as assessment and action plan. Makati City, Philippines: Bookmark, Inc. Diesmos, A. C., R. M. Brown, A. C. Alcala, R. V. Sison, L. E. Afuang, and G. V. A. Gee. 2002. Philippine amphibians and reptiles, in Philippine Biodiversity Conservation Priorities: a Second Iteration of the National Biodiversity Strategy and Action Plan. P. S. Ong, L. E. Afuang, and R. G. Rosell-Ambal, eds. Quezon City, Philippines: Department of the Environment and Natural Resources, 26–44. Environmental Center of the Philippines Foundation. 1998. Environment and natural resources atlas of the Philippines. Manila, Philippines: ECRF. Hall, R. 1998. The plate tectonics of Cenozoic SE Asia and the distribution of land and sea, in Biogeography and geological evolution of southeast Asia. R. Hall and J. D. Holloway, eds. Leiden: Brackhuys, 99–132. Mallari, N. A. D., B. R. Tabaranza, Jr., and M. J. Crosby. 2001. Key conservation sites in the Philippines. Makati City, Philippines: Bookmark, Inc. Posa, M. R. C., A. C. Diesmos, N. S. Sodhi, and T. M. Brooks. 2008. Hope for threatened tropical biodiversity: lessons from the Philippines. BioScience 58: 231–240. PHILIPPINES, GEOLOGY GRACIANO YUMUL, JR., AND CARLA DIMALANTA University of the Philippines, Quezon City KARLO QUEAÑO Department of Environment and Natural Resources, Quezon City, Philippines EDANJARLO MARQUEZ University of the Philippines, Manila Island arc systems such as the Philippines are produced through accretion brought about by collision of geologic blocks, resulting in volcanism and emplacement of crust and mantle fragments on land. The various igneous, sedimentary, and metamorphic rock types in island arcs reflect the complex processes involved in their generation and evolution. Island arc-related processes involve interactions of geological features (e.g., trenches, volcanoes, and faults) that result in specific tectonic evolution, geologic hazards, and mineral deposits. 732 Gillespie08_P.indd 732 GEOLOGIC AND TECTONIC SETTING OF AN ISLAND ARC SYSTEM The Philippines, an arc system lying off the Asian land mass, is trapped at the margins of the Eurasian–Sundaland and the Philippine Sea Plates. The northwest-southeast oblique convergence between these plates is currently being absorbed by two oppositely dipping subduction zones, namely (1) the Manila–Negros–Sulu–Cotobato trench system along which the marginal basins (i.e., South China Sea, Sulu Sea, and the Celebes Sea) on the eastern edge of the Eurasian–Sundaland Plate are being subducted west of the Philippine arc, and (2) the East Luzon Trough–Philippine Trench system, along which the West Philippine Basin of the Philippine Sea Plate is being subducted east of the arc (Fig. 1). The subduction zones extend approximately 1500 km, delineating an approximately 400-km-wide, seismically active deformed zone known as the Philippine Mobile Belt within the archipelago. Intense deformation also affects different parts of the Philippine Mobile Belt with the activity of the Philippine Fault Zone (Fig. 1). This major left-lateral strike-slip fault system transects the archipelago for more than 1200 km, from northwestern Luzon to eastern Mindanao. It has both transpressional and transtensional components and both horizontal and vertical displacements. Several major strike-slip faults branch out from the main trace of this major fault system. These faults and fault segments can be recognized on the basis of displacements in exposed rock sequences, fault scarps, sag ponds, and pressure ridges. Linear features on aerial photographs, satellite, and remote sensing images also serve to define these faults. The Philippine Fault Zone, which formed during the Middle Miocene, accommodates the lateral component corresponding to the excess stress resulting from the oblique convergence between the Philippine Sea Plate and the Philippine Mobile Belt through shear partitioning mechanism. Data obtained from the Global Positioning System networks within the Southeast Asian region have provided measurements of the convergence rate between the Sundaland–Eurasian margin and the Philippine Sea Plate. The Sunda block to the west of the Philippine archipelago is moving with respect to Eurasia at around 10 mm/year in the direction 78° east of south along its northern margin and ∼6 mm/year toward 61° east of south along its southern portion. On the eastern side, the Philippine Sea Plate is moving northwestward at approximately 7 cm/ year in the region northeast of Luzon and around 9 cm/ year southeast of Mindanao (see inset in Fig. 1). The Manila Trench on the western part of the archipelago continues on as collision zones in the central PHILIPPINES, GEOLOGY 4/22/09 3:34:02 PM