Abstract

     The mammalian fauna of the Philippine Islands is now known to include 201 species: 21 are marine mammals, 172 are native terrestrial mammals, and 7 are introduced, non-native species that occur in the wild. With 111 (64%) of the terrestrial native species as endemics, the archipelago has one of the highest per-area levels of endemism in the world, on the basis of both absolute numbers and percentage. Since the last checklist was published, in 1987, sixteen new species have been discovered —currently one of the highest rates of discovery in the world (Morell, 1986); fourteen of the species listed here are not yet formally described. For each species, we provide citation to the original description (except when the species has yet to be formally described), the English common name, the documented distribution, a summary of habitat data, and an assessment of conservation status. At least 52 native species are threatened (many seriously endangered) as a result of forest habitats, over-hunting, and destruction of cave and marine ecosystems. Further basic research and implementation of effective habitat protection programs, especially for the forests, are essential to protect this remarkably diverse and endangered fauna.

---

Click below to jump down to the:
Introduction
Content of Synopsis
Distributional Patterns
Fossil Mammals
Compiling Procedures
Common Name
Distribution
Museum Abbreviation
Habitat
Status
Authorship Note

---

Introduction
     The mammalian fauna of the Philippine Islands is remarkably diverse and species-rich. As documented in this study, the terrestrial fauna is now known to include 172 native species (plus seven introduced species, most of which are widespread in the country), which is one of the highest levels diversity on a per-area basis in the world. An additional 22 species of marine mammals have also been reported. Moreover, most of the species are found nowhere else: of the 172 terrestrial species, 111 (64%) are endemic. As noted by Cole et al. (1994) in their review of global patterns of mammalian diversity, only Madagascar has a higher percentage of unique species. However, only about 80 of the 100 mammal species on Madasgascar are endemic, and Madagascar has twice the land area of the Philippines (Heaney, 1993).  Although no comprehensive analysis has yet been done, we believe that the Philippines have the highest level of endemic mammalian species richness of any country on on a per-unit-area basis.
     These species include some of the most spectacular radiations of mammals. The murid rodent fauna of Luzon has long been considered to be one of the most remarkable assemblages of mammals (Thomas, 1898). To name only a few, there exist giant arboreal animals with long, lush coats of shiny black fur (Oliver et al., 1993); exceptionally long-snouted, hopping rodents that feed principally on earthworms in mossy forest (Rickart et al., 1991); tiny brown mice that dig through leaf-litter in their search for insects (Rickart et al., 1991); and beautiful mice with large dark eyes, pure-white bellies and auburn heads and backs that scamper along forest floor and through tree-tops (Balete and Heaney, in press). It is not surprising that these creatures have generated great interest over the century since biologists first learned of their existence from the native peoples of the Central Cordillera of Luzon.
     The Philippines are exceptional in second sense. As we report here, recent field work has produced 16 new species of mammals in the last 10 years. Since the last checklist was published in 1987 (Heaney et al., 1987). This rate of discovery is matched in only a few other countries (Brazil and Peru), and those are more than 10 times as large as the Philippines (Morell, 1996; Wilson & Reeder, 1993). Because all of the newly discovered species are endemic, the estimate of the rate of endemism for the Philippines has risen, and likely will continue to rise.
     Finally, and most important, the mammalian fauna of the Philippines is exceptional in a third sense. The most recent International Union for the Conservation of Nature (IUCN) Red Data Lists (Baillie &
Groombridge, 1996) includes 49 threatened mammals from the Philippines, which is seventh in total number for any country but first when the sizes of the countries are considered. The recognition that the Philippines have the most extensively endangered mammalian fauna in the world is supported by the even more recent Philippine Red Data List (Wildlife Conservation Society of the Philippines, 1997), which lists 52 species as endangered in the country. One of the primary purposes of this synopsis is to provide consistent information of the conservation status of all mammalian species in the country, with an indication of the limitations of current information, so as to serve as a guide both to current conservation efforts and to research that will provide even better information in the future.


Return to top

Content of Synopsis
    We have compiled an updated checklist of all mammalian species known to occur in the Philippines, including 14 as yet undescribed species. We include, for the first time, a complete listing of the marine mammals known or likely to occur in the Philippines, based on a large amount of recent field work by M. L. Dolar and her colleagues. This checklist contains updated and more detailed information on the distribution of each species than has been available previously, and for the first time we have added brief synopses of the habitat associations of every species.
     Under species entries, we have cited relevant publications that have appeared since the last checklist (Heaney et al., 1987), along with critical earlier references. For additional earlier publications, readers should consult that publication (Heaney et al., 1987) and the annotated bibliography of Balete et al. (1992).

Return to top


       Distributional Patterns of Philippine mammals: In describing the distributions of species, we often refer to the zoogeographic regions of the country; these are shown here (from Heaney, 1986). The map is based on the extent of islands in the Philippines during the late Pleistocene period, when the development of immense continental glaciers had, in effect, removed water from the world's oceans, so that sea level was 120m lower than it is today (Heaney, 1991a). Each of these Pleistocene islands defines the limit of a distinct fauna (Alcala, 1976; Dickerson, 1928; Everett, 1889; Heaney, 1985b, 1986). The three largest faunal regions thus defined are the Luzon Faunal Region (including Luzon, Catanduanes, Marinduque, Polillo, and several small islands), the Mindanao Faunal Region (including Mindanao, Basilan, Bohol, Leyte, Samar, and adjacent small islands), and the Palawan Faunal Region (including Palawan, Balabac, Busuanga, Culion, Cuyo, and adjacent small islands). Some other islands coalesced into islands larger than those of today (e.g., there was a single island from Tawi-tawi to Jolo, and an island including Negros, Panay, Cebu, and Masbate), where as other islands remained isolated (e.g., Camiguin, Sibuyan, Siquijor, and Mindoro).
      The recognition of these faunal regions is an important aid in understanding the present distribution of Philippine mammals. However, our knowledge of distribution is far from complete. Many small islands have never been surveyed, and many moderately large islands are poorly known. New distributional records are discovered by virtually every regional survey, particularly among the more poorly known taxa (especially bats and rodents). Further field work is essential to determine the distributions and habitat requirements of many species, to document the faunas of the smaller island groups, and to carry out the alpha-taxonomic studies that are necessary to determine species limits and to discover new species.
     The distribution of marine mammals in this synopsis is based on surveys, published records of occurrence, strandings, fishery by-catch, and skeletons found on beaches. Not all of the marine habitats have been surveyed, and thus the absence of records of a given species from any given area may result from lack of surveys rather than real absence. Most surveys have been focused on the waters of the Visayan region such as the eastern Sulu Sea, Tanon Strait, Bohol Sea, Panay Gulf, and parts of the Visayan Sea. Other areas recently surveyed include the southern Sulu Sea, Davao Gulf, and parts of the Celebes Sea in the southern Philippines. No thorough survey has been done in northern Philippine waters, although reports of cetacean sightings and strandings from this region have been included in this checklist.

Return to top

Fossil Mammals
      As noted in the last checklist of Philippine mammals (Heaney et al., 1987), very little is known about the fossil mammals of the Philippines. All known species are large-bodied (elephants, rhinoceros, deer, pig, and buffalo; Fox & Peralta, 1974; Groves, 1984, 1985); their large size contributes to the likelihood of both preservation and discovery. Smaller species are currently entirely unknown, although they undoubtedly were present. The discovery of new material is certain to add an exciting new dimension to our understanding of the history of the fauna, and may provide clues to the long-term interactions between humans and the native fauna of the country as well as provide the means to test many of the current models of speciation, colonization, and extinction of the Philippine mammals (e.g., Heaney, 1986, 1991b).

Return to top

Procedures Used in Compiling This Synopsis:
     As with our prior checklist (Heaney et al., 1987), this paper is intended as a working list of the mammals of the Philippines. We have not provided full taxonomic synonymies; earlier papers (Alcasid, 1970; Taylor, 1934) should be consulted for this information. We have used Wilson and Reeder (1993) as our starting point for both species names, taxonomic authorities, and for higher categories.
       We have not included mention of subspecies designations in this list for two reasons. First, the use of subspecies names implies detailed knowledge of geographic variation, and such information is rarely available for Philippine mammals. Virtually all subspecies names now in use need to be critically reevaluated. Second, we believe that use of subspecies would distract the reader from more crucial issues of general distribution and conservation status.

Return to top

      Common Name: Because common names often assist in increasing public interest in animals, we have chosen in this paper to provide English common names for Philippine mammals. In one respect, this has been a relatively easy matter: because English is a foreign language, there were virtually no English common names in use until about 100 years ago. English names have thus been coined almost entirely by biologists. With a few exceptions, we have chosen not to include names in the many languages of the Philippines because many local names are in use, with different names often for the same species in different linguistic areas, and sometimes a single name is in use for different species in different linguistics areas. Most of the few that we have included are coming into use as English names (i.e, kagwang and tamaraw). A complete compilation of native names would be a worthwhile and instructive effort, but is beyond the scope of this paper.
     In choosing English common names to include here, we have retained most names that are well-established; these are for species that are widespread in Asia, marine mammals, or large, conspicuous species. Some small species that occur outside of the country have established English common names, but often several have been used. In such cases, we have retained the names where we felt they were suitable, but in some instances we have coined new names that are more informative and more consistent with the biology of the animals. For the many species for which no English names are in use, we have coined names. In doing so, we have tried to make the names informative about the ecology, distribution, or appearance of the species. In some instances this has simply involved translating the Latin species name where that is suitably descriptive. We have avoided long names and patronyms, and have attempted to make names interesting to the public, for whom these names are primarily intended. Where two roughly equivalent common names are available, we list both, with our preferred name listed first.

Return to top

      Distribution: The distribution of many species of Philippine mammals is poorly known. This is true primarily because little effort has been made to survey many islands, and even less in the many mountainous areas to which many species are restricted (e.g., Mindanao; Sanguila & Tabaranza, 1979; Tabaranza, 1979). This problem is being addressed by ourselves and others, and is critically important to forming a detailed and accurate picture of mammalian distribution in this geographically and ecologically complex archipelago. In this synopsis we restrict our statement of distribution to a summary of the islands on which each species has been found, with the exception of Luzon and Mindanao; on those two large islands we report each province in which a species has been found. The provinces that are shown are those that existed during the late 1980's and early 1990's when we conducted our studies; readers should be aware that provinces in the Philippines are split as the human population grows, and so their boundaries are unstable. For example, South Cotabato Province was split in two in late 1995, with the northern portion retaining the same name and the southern being named Sarangani Province.
      The term "faunal region" refers to the centers of endemism that are defined by the extent of Pleistocene islands. The six largest faunal regions are Greater Luzon, Greater Mindanao, Greater Palawan, Greater Negros-Panay, Greater Mindoro, and Greater Sulu, but others (e.g., Sibuyan and Camingiun) often have endemic species as well.
     In addition to the problems posed by insufficient surveys, many specimens collections in the United States and the Philippines, including some that formed the basis for past publications, were mistakenly identified. This is not surprising, given the difficulties inherent in identifying many species, the lack of opportunity for comparison with type specimens, and the poor quality of some early descriptions. However, it is a serious problem that requires very careful work in the future and skeptical review of earlier publications. We have taken a conservative approach to this problem in this synopsis: with few exceptions, we accept identifications only of those specimens that have been examined and identified unequivocally by the authors. Virtually all have been seen by the first author. The few exceptions involve species that are very poorly known, for which the published record forms virtually the only information available, or for some equivalent crucial reason. For each Philippine island or province listed in a species entry, we have indicated a single museum containing at least one specimen whose identification we have verified. Additional records from the same islands or provinces often are available in additional museums, but we cite only one.

Return to top


Museums are indicated by the following abbreviations:

AMNH - American Museum of Natural History, New York.
ANM - Australian National Museum, Sydney.
BMNH - Natural History, London.
CUCV - Cornell University Vertebrate Collections, Ithaca.
DMNH - Delaware Museum of Natural History, Greenville.
FMNH - Field Museum of Natural History, Chicago.
FSM - Florida State Museum of Natural History, Gainesville.
MCZ - Museum of Comparative Zoology, Harvard University, Cambridge.
MMNH - Bell Museum of Natural History, University of Minnesota, St. Paul.
MSU-IIT - Mindanao State University-Iligan Institute of Technology, Iligan City.
PAWB- Protected Areas and Wildlife Bureau, Quezon City.
PNM - Philippine National Museum, Manila.
ROM - Royal Ontario Museum, Toronto.
SMF - Senckenberg Museum, Frankfurt.
SU - Museum of Natural History, Silliman University, Dumaguete.
SUML- Marine Laboratory, Silliman University, Dumaguete.
UIMNH - University of Illinois Museum of Natural History, Urbana.
UMMZ - University of Michigan Museum of Zoology, Ann Arbor.
UPD- Museum of Birds and Mammals, University of the Philippines at Diliman, Quezon City.
UPLB - Museum of Natural History, University of the Philippines at Los Banos, College.
USNM - United States National Museum of Natural History, Washington, D.C.

Return to top

      Habitat:  This section of each species account is a brief summary of the primary habitats utilized by the species. In each case, we indicate the types of forest in which the species has been documented to occur, the elevational range, and the relative abundance in each of the major habitats, to the extent that these are known. Whenever possible, published sources are cited for the information, but we also relied heavily on our personal observations where these significantly supplemented the published sources. These comments are based entirely on specimens for which we believe to be correctly identified.
     The principal natural terrestrial habitat in the Philippines occur along elevational gradients, with lowland forest, montane forest, and mossy forest as the primary elements. The elevational range of the habitats varies among the mountains based on peak elevation, topography, and annual rainfall. The habitats occurs lower on mountains, near the coast and in areas with high rainfall and higher on mountains that have high peaks, are far away from the shore, and are relatively dry. Lowland forest is dominated by species of the dipterocarp family; trees often have large buttresses, and often exceed 40 m or more in height. Lowland forest ranges from sea level up to 700 m on small, low-lying islands and to 1500 m on large, mountainous islands. Montane forest is not dominated by a single family of trees. It has trees that typically lack buttresses and reach 12 m -30 m in height; it occurs from 700 m to 1000 m on small, low-lying islands and from 1500 to as much as about 2300 m on large, mountainous islands. Mossy forest is characterized by short, gnarly trees (usually 3 - 8 m)m by having moss covering most tree trunks and branches, and by having most ground surfaces covered by moss over a thick layer of humic soil. Conifers with low height but large girth (up to 2 m dbh) are found in some mossy forests. "Heath" vegetation, dominated by woody shrubs, often occurs on exposed ridges within mossy forest. Mossy forest occurs from as low as 1000 m (rarely 800 m) on small islands (at the peaks) and from 2100 m and above on the highest mountains. For a given mountain range, the elevation of these types of forest may overlap, and there is always fairly gradual transition between them. Note that for some species of mammals we know only the elevational range, not the actual habitat in which the species occurred.
      We use "primary forest" to refer to the forest that has not been disturbed significantly by major human activities such as logging. "Secondary forests" have been disturbed, often heavily, usually by logging or burning; they range from well-regenerated second growth of natural forests, to planted forests of exotic species, to scrubby areas with scattered trees. We use "agricultural areas" to refer to a range of habitats, from coconut palm plantations to mixed pasture, fields, and orchards, and varying densities of human habitation.

Return to top

     Status: This section provides a brief statement of the conservation status of each species based on published information and our personal observations. For each species, we indicate our best estimate of the general stability of the species' population(s), abundance, and degree of vulnerability to known threats (especially habitat destruction and hunting). Where the information is available, we indicate how a species' present status differs from that in the past. (Gonzales & Alcala, 1969; Rabor, 1966, 1968).
     We have used the following terms to describe relative abundance, in order of increasing abundance: extinct, rare, uncommon, moderately common, common, and abundant. "Endemic" refers to a species restricted to a defined area; thus, a species might be said to be endemic to Camiguin Island, which means that it is found nowhere else. It should be noted that such a species could also be said to be endemic to the Philippines. "Native" species occur naturally in the Philippines; all endemic species are native, but not all native species are endemic. For example, the palm civet (Paradoxurus hermaphroditus) is native to the Philippines, but it is also present in much of South and Southeast Asia. "Non-native" also called "exotic" species are those that do not occur naturally in the Philippines; rather, they have been brought into the Philippines by humans, often accidentally, such as the rats and mice that arrived on ships. "Commensal" species are often, but not always, closely with human settlements. These commensal species often live in human houses and other buildings; except for a few species of bats, all such species in the Philippines are non-native. All marine mammals listed here are native to the Philippines, but none are endemic.

  Official/Legal Status: Many of the species listed herein are included in various official listings of species of conservation concern and/or protected status under national or international agreement. These include CITES (Convention on International Trade in Endangered Species), IUCN (International Union for the Conservation of Nature), and U.S. ESA (United States Endangered Species Act). We include reference to all of these listings in this synopsis, current through December 1996; as more is learned about the conservation status of the mammals of the Philippines, many more species will be added to these lists.
      Users of this synopsis should be aware that the intent and coverage of each of these lists varies greatly. Coverage by the US ESA is especially limited with respect to Philippine species, because the US ESA deals primarily (but not exclusively) with species in the United States and its territories. Species are listed by CITES only if it is believed that these species are threatened by international trade. It should be noted that CITES lists some species that are moderately common in the wild, but are believed to be highly vulnerable if trade were to be unrestricted, such as African Elephants. On the other hand CITES does not list many species that are seriously endangered because the are not believe to be threatened by international trade. Even species that are threatened by local trade are not included by CITES if there is no international trade. Additionally, CITES lists species that are common if there is reason to believe that an endangered species might be misidentified as that species (thus increasing the likelihood of trade); for example, all species of Pteropus are listed by CITES because several species are endangered as a result of international trade and most species are quite similar in appearance. In short, CITES listing is really not a threatened species status category at all, but a regulatory mechanism for international trade. U.S. ESA listing are also potentially misleading, but for different reasons. Both CITES and U.S. ESA listings are often inconsistent with ICUN listings, and they can be positively misleading if misunderstood. The IUCN Red List (Bailie & Groonbridge, 1996) is the most comprehensive and consistent of these listings; it is intended to cover all parts of the world and all groups of mammals (as well as other taxa) equally. However, it is too limited. In past, attention was often focused ("charismatic") animal because these were often the only species for which adequate information was available; only in the 1996 list are smaller species given equal attention. However, even the 1996 list has limitations. First, data are often lacking on the conservation status (and ecology) of small mammals in the tropics. Second, incorrect information in past lists has sometimes been carried over such as the listings of Podogymnura truei, which is probably not threatened, as discussed below). Third, the IUCN does not includes species that have been discovered but have not yet been formally described. In the case of the Philippines, where 14 species are currently in the process of being described (as listed below), this constitutes a major portion of the fauna.
     Finally, it should be noted that the Filipino is one of the first countries in the tropics to have its own Red Data Book (Wildlife Conservation Society of the Filipino, 1997). The mammal section in that book was largely taken from anearlier draft of this paper and represents an effort to present a comprehensive statement of current knowledge about the conservation status of the various species. However, as noted in that volume and in this paper, information on many species is very incomplete, and neither publication should be taken as more than one step in the process of acquiring the needed data.

Return to top

      A note on authorship: The bulk of this paper was written by the first two authors, with the exception of the sections on marine mammals, which were written principally by M. L. Dolar. Other authors reviewed the manuscript, made available unpublished notes, and/or made available specimens in collections they care for. All authors reviewed the manuscript and worked to develop a consensus on the many issues that are summarized here.

©2002 Heaney et al., Mammals of the Philippines
The Field Museum, All Rights Reserved
1400 S. Lake Shore Dr., Chicago, IL 60605-2496
312.922.9410

Technical Support
webmaster@fieldmuseum.org

Copyright Information | Linking Policy