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Patterns of Faunal Diversity and Species Abundance of Non-Volant Small Mammals on Negros Island, Philippines

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... Wild populations living far from all man-made habitats also exist, but are rather an exception. Such wild populations may be found in the mossy forests in the Philippines (Heideman et al., 1987) or on Seram (Kitchener, Schmitt & Maharadatunkamsi, 1994), in rain forests of Western Bengal (Ghose, 1976) or in steppic habitat in Rajasthan (Rana & Prakash, 1979). Because of its commensal habits, S. murinus has been passively transported by humans and is now spread over the entire Old World tropics, from India to the Malay Archipelago and Japan (Yosida, 1982) and, more recently, to the east coast of Africa and Madagascar (Heim de Balsac & Lamotte, 1957;Hutterer & Tranier, 1990). ...
... Samples of several Asiatic populations of shrews identified as Suncus murinus were collected in the following localities (see also (Heideman et al., 1987). Commensal animals sampled in Japan and Nepal and kept in captivity for several months were obtained by courtesy of Dr A. Nagel. ...
... Although almost exclusively linked to human activities, few Southeast Asian populations of S. murinus secondarily live in primary forests. This is the case on Negros Island in the Philippines (Heideman et al., 1987). These wild S. murinus do not constitute a separate lineage relative to the commensal forms, but are part of a single clade which invaded tropical Asia. ...
Protein electrophoresis was used to assess the phylogenetic relationships of populations of the phenotypically variable Asian house shrewSuncus murinus. These populations represent a sample of both commensal and wild forms. They were compared to another taxon,S. montanus, which was formerly considered conspecific withS. murinus. Suncus dayiwas used as an outgroup in all phylogenetic reconstructions. Within theS. murinuslineage, the allozyme data show very low levels of genetic differentiation among both wild and commensal Southeast Asian and Japanese samples when compared to the Indian populations. This pattern is consistent with the classical hypothesis of a recent introduction by man in Eastern Asia. The higher genetic diversity found withinS. murinusfrom India, as well as previous mitochondrial and karyological results suggest that this area is the probable centre of origin for the species. Although the lack of gene flow betweenS. murinusandS. montanusis clearly established in an area of sympatry in Southern India, one Asian house shrew sampled in Nepal was more closely related toS. montanus. This could either reflect the retention of an ancestral polymorphism, or result from a hybridization episode betweenS. murinusandS. montanus. Similar conclusions were also suggested in mitochondrial DNA studies dealing with animals sampled in the Northern parts of the Indian subcontinent. Clearly, further data onSuncusfrom this area are needed in order to assess these hypotheses.
... During the harvesting operations, this cat is often captured by the natives, who form a ring around the last patch of standing cane." Subsequent studies have provided some support for this observation (Alcala & Brown 1969;Heideman et al. 1987;Lorica & Oliver 2006;Fernandez & de Guia 2011), though all were limited by focusing on captive animals, dealing with small samples, or relying on problematic methodologies. These islands have few or no native, non-volant small mammals (only two are known from Negros, three from Panay, and none are known from Cebu or Masbate), and two civets (family Viverridae) occur on all four islands (Steppan et al. 2003;Heaney et al. 2010). ...
... Scats from palm civets are about the same size as those of leopard cats, but contain more fruits than small vertebrates (Rabinowitz 1991;Joshi et al. 1995;Shore et al. 2005;Nakashima et al. 2010). Upon analysis of each scat, samples containing seeds and plant matter other than grass were discarded, as they could belong to either species of omnivorous viverrid in the area (Rabor 1977;Heideman et al. 1987). Finally, DNA analysis to confirm the origin of some scats was performed by the laboratory of Prof. Worawidh Wajjwalku at Kasetsart University. ...
... Lorica & Heaney to high-elevation forest (Heideman et al. 1987;Steppan et al. 2003;Heaney et al. 2010), and neither was present in our samples. Suncus murinus, the Asian house shrew, occurs in human habitation and agricultural areas on Negros, but it also was not found in this study. ...
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Concerns about vulnerability of mammalian carnivores to extinction, especially on small islands, appear to conflict with prior reports of endemic populations of leopard cat Prionailurus bengalensis (Kerr, 1792) surviving in agricultural landscapes on oceanic islands. We investigated the persistence of the Visayan leopard cat (P. b. rabori) in the sugarcane fields on Negros, an oceanic island in central Philippines. A population remained throughout the year at our study site on a sugarcane farm, and reproduction was noted. Non-native rodents form the bulk of the cat’s diet, followed by reptiles, birds, amphibians, and insects. Prey species identified from the samples commonly occur in agricultural areas in the Philippines. Prey composition did not vary significantly with respect to wet and dry season, or sugarcane harvest cycle. This study provides evidence that an intensively managed agricultural landscape on this oceanic island supports a native obligate carnivore that subsists primarily on exotic rats. This study supports a prior prediction that leopard cats will show flexibility in prey selection on islands with few or no native small mammal prey species, but in this case they do so not by switching to other vertebrates and invertebrates, but rather to exotic pest species of rodents.
... murinus (Linnaeus, 1766) The Asian house-shrew occurs widely in Asia and Indo-Australia; it now occurs throughout the Philippines, though it is not native to the country. It is abundant in urban and agricultural areas; on islands with low mammal species richness such as Negros, it is sometimes abundant in disturbed forest and occasionally in primary forest (Heideman et al., 1987;), but on islands of average species richness, it is usually rare or absent from forest (Heaney et al., 1989, unpubl. data; Rickart et al., 1993). ...
... Records from the 1960s also show it to be present in mossy forest at ca. 1500 m. This distribution is similar to that found on Negros, where it occurs in high-density agricultural areas and in mossy forest at an elevation of 1500– 1650 m (Heideman et al., 1987; Heaney et al., 1989). Both Camiguin and Negros have very few native rodents, and this may influence the ability of nonnative species to invade the forest since the nonnatives are absent or very rare in mature forest on species-rich islands such as Leyte, Luzon, and Mindanao (Heaney et al., 1989Heaney et al., , 1999, unpubl. ...
... Temminck, 1844 Previously known in the Philippines as Rattus rattus and Rattus mindanensis, the Oriental house rat is a widespread nonnative rodent in the Philippines; it occurs from Afghanistan to Indomalaya, New Guinea, and Micronesia (Musser & Carleton, 1993). This 140–200-g rat is most often found as a pest in urban and agricultural areas but can be common in disturbed forest up to 1800 m (Heideman et al., 1987; Heaney et al., 1989 Heaney et al., , 1991 Rickart et al., 1993). On Camiguin, we found R. tanezumi to be very abundant in heavily disturbed agricultural land at 150 m (Site 3;Table 1). ...
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Biodiversity surveys in the 1960s and 1990s on Camiguin Island, a geologically young, volcanically active oceanic island surrounded by deep water, have demonstrated the presence of 24 species of land mammals. Three species (one insectivore and two rodents) are not native to the Philippines, but all others (one insectivore, 12 bats, one monkey, four rodents, two small carnivores, and one ungulate) are indigenous. Among those captured in the 1990s were two previously unknown species of murid rodents in the genera Apomys and Bullimus that are endemic to Camiguin. The discovery of two new species on such a small island (ca. 265 km2) is remarkable; Camiguin is currently the smallest island in the Philippines known to have unique species of mammals. Total species richness of nonvolant mammals on Camiguin is low, but relative to islands that were once part of Pleistocene Greater Mindanao, Camiguin is not depauperate. However, its fauna is not ecologically balanced in the same way as the faunas of the islands that were part of Greater Mindanao: ground-living shrews (Crocidura) and rodents (Apomys, Bullimus, Crunomys, and Rattus) from lowland forest, along with some large mammals (Macaca, Paradoxurus, and Sus) are well represented on Camiguin, but all the arboreal small mammals that characterize lowland forest on Mindanao (Sundasciurus, Exilisciurus, Cynocephalus, and Tarsius), ground-living small mammals from montane habitats (Urogale, Podogymnura, Batomys, Limnomys, and Tarsomys), and one large mammal (Cervus) are absent. Additionally, at least two genera of fruit bats (Haplonycteris and Megaerops) that are fairly common in lowland rain forests on Mindanao are absent on Camiguin. The presence of some nonvolant mammals demonstrates that dispersal across the deep but narrow intervening channel takes place, but the presence of two species endemic to Camiguin and the absence of other species that are present on nearby Mindanao implies that dispersal probably is rare. The Asian house shrew (Suncus murinus) was remarkably abundant in primary forest at high elevation; this species has also been found to be abundant in montane primary forest on Negros Island, which also has low total species richness. Species richness of small nonflying mammals was greatest at fairly high elevation.
... The Binukid farmers on Kitanglad are very familiar with this animal since it often frequents human habitations near forested areas and is believed to occasionally raid their poultry. In Philippine forest, it is omnivorous, feeding on fruits and small mammals (Alcala & Brown, 1969;Heideman et al., 1987;Heaney et al., 1999). ...
... Malindang (Musser, 1994;Musser & Heaney, 1992). The Kitanglad mammal fauna exceeds the total number known from the entire island of Negros, where 49 native mammals have been documented through extensive surveys Heideman et al., 1987;Utzurrum, 1995). ...
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Field surveys within and adjacent to the Mt. Kitanglad Nature Park in the Kitanglad Range of Bukidnon Province, north-central Mindanao, from 1992 to 1999, along with examination of previously existing specimens, have allowed us to document the local presence of 58 species of mammals, 53 native and five non-native. These include one gymnure (Erinaceidae), two shrews (Soricidae), one tree shrew (Tupaiidae), one flying lemur (Cynocephalidae), 14 fruit bats (Pteropodidae), eight roundleaf and horseshoe bats (Rhinolophidae), nine evening bats (Vespertilionidae), one mastiff bat (Molossidae), two primates (Tarsiidae and Cercopithecidae), three squirrels (Sciuridae), 14 mice and rats (Muridae), two civets (Viverridae), one pig (Suidae), and one deer (Cervidae). Mt. Kitanglad Nature Park has one of the most diverse mammal faunas in the Philippines, exceeding that of the more widely known Mt. Apo. Three species, a bat (Alionycteris paucidentata) and two native mice (Crunomys suncoides and Limnomys bryophilus), are currently known only from high elevations in the Kitanglad Range. Species richness of bats declined with increasing elevation, but richness of non-volant small mammals increased five-fold from lowlands to a peak at ca. 2250 m, and then declined with further increases in elevation. We found distinctive mammal communities in lowland rainforest (up to about 1200 m elevation), montane rainforest (ca. 1200 m to 1900 m), and mossy rainforest (2000 m to the peak at 2950 m). We conclude that all three rainforest types, at all elevations, are important to the success of the park as a biological reserve. Over-hunting of large mammals and illegal logging both pose serious problems. Lowland rainforest has been removed on much of Mindanao, including the vicinity of the park, and thus is the habitat type that is currently most threatened. Habitat destruction, especially of lowland rainforest, threatens the mammals in the Kitanglad Range as well as the economic and social stability of the human population of northern Mindanao.
... Maxomys whiteheadi, L. sabanus, R. exulans, T. tana and S. muelleri were generally found all over the elevation (Heideman et al., 1987;Nor, 2001). Somehow during this study, M. rajah and S. lowii were only found at the lowland at KNP. ...
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The study of non-volant small mammals was conducted at Kubah National Park (KNP), Sarawak for eight days between November 2018 until February 2019. The objective of this study was to determine the species diversity of non-volant small mammals at the highland (>750 – 805 m a.s.l.) and lowland (<200 m a.s.l.) at KNP. The elevation of Gunung Serapi is 911 m a.s.l. but the highest accessible area is at 805 m a.s.l. The distance between two sampling sites is approximately 4.5 km. Humidity and temperature measurement were also recorded at both sites. A total of 50 cage traps were set up at the highland and 50 cage traps at the lowland. The baits that were used in this study were oil palm and banana. A total of 26 individuals from 11 species, eight genera and four families were captured. However, there is no significant difference in species diversity between low and high elevations because the elevation of KNP was not high enough to distinguish species that are highland or lowland specialist.
... This hypothesis is based on a pattern documented on six mountains of the Philippines: Mount Isarog (Heaney et al., 1999), Mount Tapulao (Balete et al., 2009), and Mt. Bali-it, Balbalasang , on Luzon; Mount Guinsayawan on Negros Island (Heideman et al., 1987;Heaney et al., 1991); Mount Timpoong on Camiguin Island (Heaney et al., 2006a); and Mount Pangasugan on Leyte Island (Rickart et al., 1993). ...
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In 2005 and 2007, we conducted surveys of mammals along an elevational transect on Mount Palali (peak 1707 m) in the Caraballo Mountains, a poorly known mountain range on Luzon Island, Philippines. The surveys covered eight localities representing habitats from lowland agroforest and regenerating disturbed lowland rainforest at 780 m to mossy forest near the peak. We recorded 24 species, including one native shrew, one non-native shrew, five fruit bats, seven insectivorous bats, one monkey, six native rodents, two civets, and one pig. One species of Apomys is the newly described A. sierrae, and two species of Chrotomys are potentially undescribed species. Elevational patterns varied among mammals: bats were most diverse in the lowlands, native nonvolant small mammals had almost equal richness along the entire elevational transect, and most species of large mammals were present at all elevations. Bait attractiveness and diel activity pattern differed among native nonvolant small mammals: Apomys microdon and Rattus everetti were nocturnal and attracted to coconut baits, Chrotomys sp. 1 and 2 were mostly nocturnal and favored earthworms, Crocidura grayi showed no bait preference and was active during day and night, and A. sierrae showed no bait preference and was most active at night. The non-native Suncus murinus was restricted to highly disturbed areas at 780 m, whereas all native nonvolant small mammals were present in both degraded and undisturbed forest, supporting the hypothesis that non-native small mammals are not successful in invading native habitats on oceanic islands when the native community of small mammals is diverse. We conclude that all three rainforest types at all elevations, including newly regenerating forest, provide important habitat for mammals on Mount Palali.
... Knowing the distribution of organisms along an elevational gradient is critical to understanding the evolution and ecology of montane biotic systems, and to designing conservation strategies to maintain them. These reasons have motivated elevational surveys of small mammals in various areas of the world including Chile [1], Costa Rica [2], Malaysia [3], Philippines [4], [5], [6], Taiwan [7], and Tanzania [8]. Goodman, Ganzhorn & Rakotondravony [9] summarize some of the important biotic inventories along elevational gradients in Madagascar. ...
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Mt Kilimanjaro is Africa's highest mountain, and an icon for a country famous for its mammalian fauna. The distribution and abundance of small mammals on the mountain are poorly known. Here we document the distribution of shrews and rodents along an elevational gradient on the southeastern versant of Kilimanjaro. Five sites were sampled with elevational center points of 2000, 2500, 3000, 3500 and 4000 m, using a systematic methodology of standard traps and pitfall lines, to inventory the shrews and rodents of the slope. Sixteen species of mammal were recorded, including 6 shrew and 10 rodent species, and the greatest diversity of both was found at 3000 m, the elevational midpoint of the transect. No species previously unrecorded on Kilimanjaro were observed. Two genera of rodents that occur in nearby mountains (Hylomyscus and Beamys) were not recorded. Myosorex zinki, the only mammal endemic to Mt. Kilimanjaro, which previously was known by only a few specimens collected in the ericaceous or moorland habitat, was found in all but one (the lowest) of the sites sampled, and was one of the most widespread species of small mammal along the gradient. Two shrews (Crocidura allex and Sylvisorex granti) and one rodent (Dendromus insignis) were found throughout the entire transect, with Dendromus being observed at our highest trap point (4240 m). As in similar faunal surveys on other mountains of Tanzania, rainfall influenced the sample success of shrews, but not rodents. Trap success for rodents at 3500 m was notably low. This study contributes further justification for the conservation of the forest habitat of Mt. Kilimanjaro.
... Small mammals were trapped using Sherman live traps (2x2.5x6.5"), and mediumsized mammals were trapped using cage traps. Traps were set up in 'trap lines' following the approaches of Heideman et al. (1987) and Heaney et al. (1989). Traps were stationed in lines of 10 Sherman and 3 cages along a 60m transect with at least 5m spacing between each trap. ...
The distribution of mammals by elevation was studied in 1994-1995 in highland regions of Ba Vi and in 1993, 1996-1997 of Phansipang (northern Vietnam). Mammals were collected by trapping along transects with 25 m step or at sites of definite elevation. The maximum abundance of animals and the highest species diversity were determined in the middle range of Ba Vi at elevation from 400 to 800 m. At an elevation above 800 m these parameters decreased. In the high mountain range Phansipang, the maximum abundance of animals and the highest species diversity were registered at elevation from 1800 to 2000 m, in the places where polydominant rain tropical forests are substituted for moist tropical and subtropical forests. Relations between a specific biotope and elevational distribution of mammals are discussed. The role of human activities forcing various mammal species to occupy habitats at the elevations uncharacteristic of them is discussed also.
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1 It is widely accepted that tropical lowland rain forest holds the greatest diversity of organisms, and it is often implied that this general pattern is also true for virtually all individual higher-level taxa. Standardized elevational transect surveys of non-flying small mammals (Insectivora and Rodentia) on geologically old, species-rich islands in the Philippines consistently show maximum diversity and relative abundance in upper montane/lower mossy forest at 1500–2200 m, often exceeding lowland species richness and relative abundance by a factor of three or more.
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