Diversity, Encounter Rate and Detection of Non-Volant Nocturnal Mammals on Two Malaysian Islands
Main Article Content
Abstract
Nocturnal mammals constitute a crucial component of tropical faunal diversity, but not much is known about the effects of anthropogenic disturbance on the habitat use and detectability of these species. We investigated which habitat and environmental variables impact the detectability of non-volant nocturnal arboreal mammals across varying habitat types at two tropical islands with different levels of anthropogenic development in Malaysia. We conducted night transect line and point count surveys following pre-existing paths in Penang Island and Langkawi Island between 2019 and 2020. We used a head torch with red filter and a thermal imaging device (FLIR) to enhance animal detection success. We calculated the encounter rates (individual km–1) for each species as a proxy for abundance. Overall, we detected 17 species, but did not find higher species diversity in intact forested environments compared to disturbed areas. Encounter rates of the most observed species were influenced by ‘time after sunset’ on the highly developed island of Penang, whereas on the rural island of Langkawi, detection was higher in sites with better canopy connectivity. Different species of non-volant nocturnal arboreal mammals use their respective habitats differently and thus, are differently impacted by varying levels of anthropogenic activities. Our results provided baseline data on the diversity, encounter rate, and detectability of these highly elusive species, which can also help to further improve methodologies for the detection of nocturnal wildlife.
Article Details
This work is licensed under a Creative Commons Attribution 4.0 International License.
References
Abernethy K A, Coad L, Taylor G, Lee M E and Maisels F. (2013). Extent and ecological consequences of hunting in Central African rainforests in the twenty-first century. Philosophical Transactions of the Royal Society B: Biological Sciences 368(1625): 20120303. https://doi.org/10.1098/rstb.2012.0303
Achard F. (2002). Determination of deforestation rates of the world’s humid tropical forests. Science 297(5583): 999–1002. https://doi.org/10.1126/science.1070656
Akbar Z and Ariffin M K E. (1997). A comparision of small mammal abundance between a primary and disturbed lowland rain forest in peninsular Malaysia. Malayan Nature Journal 50: 201–206.
Ando M and Shiraishi S. (1993). Gliding flight in the Japanese giant flying squirrel Petaurista leucogenys. Journal of the Mammalogical Society of Japan 18(June): 19–32. https://doi.org/10.11238/jmammsocjapan.18.19
Aplin K. (2017). Trichys fasciculata. The IUCN Red List of Threatened Species 2017: e.T22132A22232753. https://doi.org/10.2305/IUCN.UK.2017-2.RLTS.T22132A22 232753.en
Aplin K and Lunde D. (2016). Iomys horsfieldii (errata version published in 2017). The IUCN Red List of Threatened Species 2016: e.T10845A115099730. https://doi.org/10.2305/IUCN.UK.2016-3.RLTS.T10845A22249234.en
Azlan J M and Sharma D S K. (2006). The diversity and activity patterns of wild felids in a secondary forest in Peninsular Malaysia. Oryx 40(1): 36–41. https://doi.org/10.1017/S0030605306000147
Azlan J M. (2006). Mammal diversity and conservation in a secondary forest in Peninsular Malaysia. Biodiversity and Conservation 15(3): 1013–1025. https://doi.org/10.1007/s10531-004-3953-0
Bailey L L, Simons T R and Pollock K H. (2004). Estimating site occupancy and species detection probability parameters for terrestrial salamanders. Ecological Applications 14(3): 692–702. https://doi.org/10.1890/03-5012
Barret E. (1985). The ecology of some nocturnal, arboreal mammals in the rainforest of Peninsular Malaysia. PhD diss., University of Cambridge.
Bashir A Z. (2014). Wildlife diversity near natural saltlicks in the Ulu Muda Forest Reserve. Petaling Jaya. http://awsassets.wwf.org.my/downloads/ulu_muda_biodiversity_ report_2014.pdf (accessed on 19 March 2022).
Bates D, Mächler M, Bolker B and Walker S. (2015). Fitting linear mixed-effects models using lme4. Journal of Statistical Software 67(1): 1–48. https://doi.org/10.18637/jss.v067.i01
Beier P. (2006). Effects of artificial night lighting on terrestrial mammals. In C. Rich and T. Longcore (eds.), Ecological consequences of artificial night lighting. Washington, DC: Island Press, 19–42.
Berglund L G. (1998). Comfort and humidity. ASHRAE Journal 40(8): 35.
Bista D, Baxter G S, Hudson N J, Lama S T and Murray P J. (2022). Effect of disturbances and habitat fragmentation on an arboreal habitat specialist mammal using GPS telemetry: A case of the red panda. Landscape Ecology 37(3): 795–809. https://doi.org/10.1007/s10980-021-01357-w
Boeadi and Steinmetz R. (2008). Galeopterus variegatus. The IUCN Red List of Threatened Species 2008: e.T41502A10479343. https://doi.org/10.2305/IUCN.UK.2008.RLTS.T41502A10479343.en
Bowmaker J K. (2008). Evolution of vertebrate visual pigments. Vision Research 48(20): 2022–2041. https://doi.org/10.1016/j.visres.2008.03.025
Buckland S, Rexstad E, Marques T and Oedekoven C. (2015). The basic method. In Distance sampling: Methods and applications. Methods in Statistical Ecology. Cham, Springer: 3–13. https://doi.org/10.1007/978-3-319-19219-2_1
Buckland S T, Anderson D R, Burnham K P, Laake J L, Borchers D L and Thomas L. (2004). Advanced distance sampling. Oxford: Oxford University Press. https://doi.org/10.1093/oso/9780198507833.001.0001
Carthew S and Goldingay R L. (1997). Non-flying mammals as pollinators. Trends in Ecology and Evolution 12(3): 104–108. https://doi.org/10.1016/S0169-5347(96)10067-7
Catling P C, Burt R J and Kooyman R. (1997). A comparison of techniques used in a survey of the ground-dwelling and arboreal mammals in forests in north-eastern New South Wales. Wildlife Research 24(4): 417–432. https://doi.org/10.1071/WR96073
Connell J H. (1978). Diversity in tropical rain forests and coral reefs. Science 199(4335): 1302–1310. https://doi.org/10.1126/science.199.4335.1302
Crane M J, Lindenmayer D B and Cunningham R B. (2014). The value of countryside elements in the conservation of a threatened arboreal marsupial Petaurus norfolcensis in agricultural landscapes of south-eastern Australia: The disproportional value of scattered trees. PLoS ONE 9(9): e107178. https://doi.org/10.1371/journal.pone.0107178
Cudney‐Valenzuela S J, Arroyo‐Rodríguez V, Morante‐Filho J C, Toledo‐Aceves T and Andresen E. (2022). Tropical forest loss impoverishes arboreal mammal assemblages by increasing tree canopy openness. Ecological Applications 33(1): e2744. https://doi.org/10.1002/eap.2744
da Rosa C A, Secco H, Carvalho N, Maia A C and Bager A. (2018). Edge effects on small mammals: Differences between arboreal and ground-dwelling species living near roads in Brazilian fragmented landscapes. Australian Ecology 43(1): 117–126. https://doi.org/10.1111/aec.12549
de Thoisy B, Brosse S and Dubois M A. (2008). Assessment of large-vertebrate species richness and relative abundance in Neotropical forest using line-transect censuses: What is the minimal effort required? Biodiversity and Conservation 17(11): 2627– 2644. https://doi.org/10.1007/s10531-008-9337-0
Department of Statistics Malaysia. (2010). Taburan penduduk dan ciri-ciri asas demografi 2010. Putrajaya: Department of Statistics Malaysia. https://www.mycensus.gov.my/index.php/ms/produk-banci/penerbitan/banci-2010/664-taburan-penduduk-dan-ciri-ciri-asas-demografi-2010
Deuskar C, Baker J and Mason D. (2015). East Asia’s changing urban landscape: Measuring a decade of spatial growth. Washington, DC: World Bank Publications.
Dirzo R, Young H S, Galetti M, Ceballos G, Isaac N J B and Collen B. (2014). Defaunation in the Anthropocene. Science 345(6195): 401–406. https://doi.org/10.1126/science.1251817
Dornelas M. (2010). Disturbance and change in biodiversity. Philosophical Transactions of the Royal Society B: Biological Sciences 365(1558): 3719–3727. https://doi.org/10.1098/rstb.2010.0295
Duckworth J W, Timmins R, Choudhury A, Chutipong W, Willcox D, Mudappa D, Rahman H, Widmann P, Wilting A and Xu W. (2016). Paradoxurus hermaphroditus. The IUCN Red List of Threatened Species 2016: e.T41693A45217835. https://doi.org/10.2305/IUCN.UK.2016-1.RLTS.T41693A45217835.en
Duckworth J W. (1998). The difficulty of estimating population densities of nocturnal forest mammals from transect counts of animals. Journal of Zoology 246(4): 466–468. https://doi.org/10.1111/j.1469-7998.1998.tb00183.x
Duckworth J W. (2016). Petaurista petaurista (errata version published in 2017). https://doi.org/10.2305/IUCN.UK.2016-3.RLTS.T16723A22272173.en
Dzulhelmi N and Abdullah M T. (2010). Distribution of the Sunda colugo (Galeopterus variegatus) in Malaysia. Tropical Life Sciences Research 21(2): 69–83.
Einoder L D, Southwell D M, Gillespie G R, Fisher A, Lahoz-Monfort J J and Wintle B A. (2018). Optimising broad-scale monitoring for trend detection: review and re-design of a long-term program in northern Australia. In S Legge, D B Lindenmayer, N M Robinson, B C Scheele, D M Southwell, B A Wintle (eds.), Monitoring threatened species and ecological communities. Australia: CSIRO Publishing, 165–178.
Emmons L H and Gentry A H. (1983). Tropical forest structure and the distribution of gliding and prehensile-tailed vertebrates. The American Naturalist 121(4): 513–524. https://doi.org/10.1086/284079
Ferrari M C, Sih A and Chivers D P. (2009) The paradox of risk allocation: A review and prospectus. Animal Behaviour 78: 579–585. https://doi.org/10.1016/j.anbehav.2009.05.034
Focardi S, Marinis A M D, Rizzotto M and Pucci A. (2001). Comparative evaluation of thermal infrared imaging and spotlighting to survey wildlife. Wildlife Society Bulletin 29(1): 133–139. https://www.jstor.org/stable/3783989
Fragoso J M V, Levi T, Oliveira L F B, Luzar J , Overman, H, Read J M and Silvius K M. (2016). Line transect surveys underdetect terrestrial mammals: Implications for the sustainability of subsistence hunting. PLoS ONE 11(4): 1–18. https://doi.org/10.1371/journal.pone.0152659
Francis C. (2019). Field guide to the mammals of South-east Asia. London, UK: Bloomsbury Publishing.
Garvey S M, Templer P H, Pierce E A, Reinmann A B and Hutyra L R. (2022). Diverging patterns at the forest edge: Soil respiration dynamics of fragmented forests in urban and rural areas. Global Change Biology 28(9): 3094–3109. https://doi.org/10.1111/gcb.16099
Gaynor K M, Hojnowski C E, Carter N H and Brashares J S. (2018). The influence of human disturbance on wildlife nocturnality. Science 360(6394): 1232–1235. https://doi.org/10.1126/science.aar7121
Gibson L, Lynam A J, Bradshaw C J, He F, Bickford D P, Woodruff D S, Bumrungsri S and Laurance W F. (2013). Near-complete extinction of native small mammal fauna 25 years after forest fragmentation. Science 341(6153): 1508–1510. https://doi.org/10.1126/science.1240495
Glen A S, Cockburn S, Nichols M, Ekanayake J and Warburton B. (2013). Optimising camera traps for monitoring small mammals. PLoS ONE 8(6): e67940. https://doi.org/10.1371/journal.pone.0067940
Green N S, Wildhaber M L, Albers J L, Pettit T W and Hooper M J. (2020). Efficient mammal biodiversity surveys for ecological restoration monitoring. Integrated Environmental Assessment and Management (Special Series) 00(0), 1–13. https://doi.org/10.1002/ieam.4324
Guillera‐Arroita G. (2017). Modelling of species distributions, range dynamics and communities under imperfect detection: advances, challenges and opportunities. Ecography 40(2): 281–295. https://doi.org/10.1111/ecog.02445
Halle S and Stenseth N C. (2012). Activity patterns in small mammals: An ecological approach. Berlin Heidelberg: Springer.
Hart D. (2007). Predation on primates: A biogeographical analysis. In S L Gursky and K A I Nekaris (eds.), Primate anti-predator strategies. Boston, MA: Springer, 27–59. https://doi.org/10.1007/978-0-387-34810-0_2
Hart A G, Dawson M, Fourie R, MacTavish L and Goodenough A E. (2022). Comparing the effectiveness of camera trapping, driven transects and ad hoc records for surveying nocturnal mammals against a known species assemblage. Community Ecology 23(1): 27–39. https://doi.org/10.1007/s42974-021-00070-7
Henrich M, Niederlechner S, Kröschel M, Thoma S, Dormann C F, Hartig F and Heurich M. (2020). The influence of camera trap flash type on the behavioural reactions and trapping rates of red deer and roe deer. Remote Sensing in Ecology and Conservation 6(3): 399–410. https://doi.org/10.1002/rse2.150
Hodgkison R, Balding S T, Zubaid A and Kunz T H. (2003). Fruit bats (Chiroptera: Pteropodidae) as seed dispersers and pollinators in a lowland Malaysian rain forest. Biotropica 35(4): 491–502. https://doi.org/10.1111/j.1744-7429.2003.tb00606.x
Huang J C C, Abdullah N I, Nurul-Ain E, Mohd Sah S A and Lim L S. (2019). The first record of Coelops frithii (Blyth, 1848) to Penang, Malaysia, with a note of the potential acoustic plasticity of the species. Journal of Bat Research and Conservation 12(1): 21–26. https://doi.org/10.14709/Barbs.12.1.2019.03
Jacobson A, Dhanota J, Godfrey J, Jacobson H, Rossman Z, Stanish A, Walker H and Riggio J. (2015). A novel approach to mapping land conversion using Google Earth with an application to East Africa. Environmental Modelling and Software 72: 1–9. https://doi.org/10.1016/j.envsoft.2015.06.011
Joshi A R, David Smith J L and Cuthbert F J. (1995). Influence of food distribution and predation pressure on spacing behavior in palm civets. Journal of Mammalogy 76(4): 1205–1212. https://doi.org/10.2307/1382613
Kawanishi K. (1999). Preliminary analysis on abundance of large mammals at Sungai Relau, Taman Negara. Journal of Wildlife and National Parks Malaysia 17: 62–82. https://ci.nii.ac.jp/naid/10013281218/
Laurance W F. (1991). Ecological correlates of extinction proneness in Australian tropical rain forest mammals. Conservation Biology 5(1): 79–89. https://doi.org/10.1111/j.1523-1739.1991.tb00390.x
Lee P. (1986). Ecological studies on two sympatric Petaurista species in Taiwan. Bulletin of the Institute of Zoology, Academia Sinica 25: 113–124.
Leman M S, Komoo I, Mohamed K R, Ali C A and Unjah T. (2007). Geopark as an answer to geoheritage conservation in Malaysia: The Langkawi Geopark case study. Bulletin of the Geological Society of Malaysia 53(June): 95–102. https://doi.org/10.7186/bgsm53200715
Leman M S, Komoo I, Mohamed K R, Ali C A, Unjah T, Othman K and Yasin M H M. (2008). Geology and geoheritage conservation within Langkawi Geopark, Malaysia. Unesco Globle Geoparks Network. http://www.globalgeopark.org/Articles/6337.htm
Lim L S, Struebig M J, Zalipah M N, Mohd-Adnan A, Senawi J, Zubaid A, Mohd Sah S A and Rossiter S J. (2019). Bats from the understorey of lowland tropical rainforests across Peninsular Malaysia. Journal of Bat Research and Conservation 12(1): 68–82. https://doi.org/10.14709/BarbJ.12.1.2019.10
Lim N. (2007). Colugo: The flying lemur of South-East Asia. Singapore: Draco Publishing and Distribution.
Lim N T L, Giam X, Byrnes G and Clements G R. (2013). Occurrence of the Sunda colugo (Galeopterus variegatus) in the tropical forests of Singapore: A Bayesian approach. Mammalian Biology 78(1): 63–67. https://doi.org/10.1016/j.mambio.2012.06.008
Lim N T and Ng P K. (2010). Population assessment methods for the Sunda colugo Galeopterus variegatus (Mammalia: Dermoptera) in tropical forests and their viability in Singapore. Raffles Bulletin of Zoology 58(1): 157–164.
Lima S L and Dill L M. (1990). Behavioral decisions made under the risk of predation: A review and prospectus. Canadian Journal of Zoology 68: 619–640. https://doi.org/10.1139/z90-092
Linley G D, Pauligk Y, Marneweck C and Ritchie E G. (2020). Moon phase and nocturnal activity of native Australian mammals. Australian Mammalogy 43(2): 190–195. https://doi.org/10.1071/AM19070
Lit E, Idris M I, Marni W, Mortada M J and Abdullah M. (2011). Species diversity of small mammals in Lubuk Sembilang Recreational Park, Langkawi Island, Kedah. RIMBA2: Regional Sustainable Development in Malaysia and Australia January: 124–135.
Lo S, Maseri M, Kamarul H and Amir A. (2018). A preliminary camera traps assessment of terrestrial vertebrates at different elevation gradients in Gunong Stong State Park, Kelantan, Malaysia. Malayan Nature Journal 70: 3–11.
Lopes K G and Bicca-Marques J C. (2017). Ambient temperature and humidity modulate the behavioural thermoregulation of a small arboreal mammal (Callicebus bernhardi). Journal of Thermal Biology 69: 104–109. https://doi.org/10.1016/j.jtherbio.2017.06.010
Lowrey C and Longshore K M. (2017). Tolerance to disturbance regulated by attractiveness of resources: A case study of desert bighorn sheep within the river mountains. Nevada Western North American Naturalist 77(1): 82–98. https://doi.org/10.3398/064.077.0109
Luna-Jorquera G, Fernández C E and Rivadeneira M M. (2012). Determinants of the diversity of plants, birds and mammals of coastal islands of the Humboldt current systems: Implications for conservation. Biodiversity and Conservation 21(1): 13– 32. https://doi.org/10.1007/s10531-011-0157-2
Main D C J, Stokes J E, Reader J D and Whay H R. (2012). Detecting hoof lesions in dairy cattle using a hand-held thermometer. Veterinary Record 171: 504–506. https://doi.org/10.1136/vr.100533
Malayan Nature Society. (1999). Nature trails of Penang Island. Kuala Lumpur: Malayan Nature Society.
Marsden S J, Loqueh E, Takuo J M, Hart J A, Abani R, Ahon D B, Annorbah N N, Johnson R and Valle S. (2016). Using encounter rates as surrogates for density estimates makes monitoring of heavily-traded grey parrots achievable across Africa. Oryx 50(4): 617–625. https://doi.org/10.1017/S0030605315000484
Marshall A R, Lovett J C and White P C L. (2008). Selection of line-transect methods for estimating the density of group-living animals: Lessons from the primates. American Journal of Primatology 70(5): 452–462. https://doi.org/10.1002/ajp.20516
Masum K M, Mansor A, Sah S A M, Lim H S and Hossain M K. (2017). Effect of differential forest management on biodiversity in a tropical hill forest of Malaysia and implications for conservation. Biodiversity and Conservation 26(7): 1569–1586. https://doi.org/10.1007/s10531-017-1318-8
McComb B, Zuckerberg B, Vesely D and Jordan C. (2010). Monitoring animal populations and their habitats: A practitioner’s guide. Boca Raton, FL: CRC Press. https://doi.org/10.1201/9781420070583
McCullagh P and Nelder J A. (2019). Generalized linear models, 2nd ed. New York: Routledge. https://doi.org/10.1201/9780203753736
McShea W J, Stewart C, Peterson L, Erb P, Stuebing R and Giman B. (2009). The importance of secondary forest blocks for terrestrial mammals within an Acacia/ secondary forest matrix in Sarawak, Malaysia. Biological Conservation 142(12): 3108–3119. https://doi.org/10.1016/j.biocon.2009.08.009
Medway L. (1969). The wild mammals of Malaya and offshore islands including Singapore. Oxford: Oxford University Press. https://doi.org/10.2307/1378329
Meijaard E. (2003). Mammals of south-east Asian islands and their late Pleistocene environments. Journal of Biogeography 30(8): 1245–1257. https://doi.org/10.1046/j.1365-2699.2003.00890.x
Miard P. (2020). Distribution, methodological validation and ecology of nocturnal island mammals in Peninsular Malaysia. PhD diss, Universiti Sains Malaysia.
Miller D L, Rexstad E, Thomas L, Marshall L and Laake J L. (2019). Distance sampling in R. Journal of Statistical Software 89(1): 1–28. https://doi.org/10.18637/jss.v089.i01
Miller M W. (2016). Distance sampling: Methods and applications. Bird Study 63(1): 152–153. https://doi.org/10.1080/00063657.2016.1148352
Mohd Sah S A, Nor Z, Nurul-Ain E, Ganesan M and Mark R. (2006a). Mammals and other vertebrate fauna in Teluk Bahang and Bukit Panchor Recreational Forests. In M Masshor and Z Mohd Yunus (Eds.), Ecological survey on natural forests: A case study of Teluk Bahang and Bukit Panchor. Penang: Penerbit Universiti Sains Malaysia, 64–101.
Mohd Sah S A, Nurul-Ain E, Nor Z, Mark Z and Ganesan M. (2006b). Mammals and other vertebrates survey in Balik Pulau and Pantai Acheh. In M Masshor and Z Mohd Yunus (Eds.), Ecological survey on mangrove forests: A case study of Balik Pulau and Pantai Acheh. Penang: Penerbit Universiti Sains Malaysia, 33–72.
Monterroso P, Alves P C and Ferreras P. (2013). Catch me if you can: Diel activity patterns of mammalian prey and predators. Ethology 119: 1044–1056. https://doi.org/10.1111/eth.12156
Morris E K, Caruso T, Buscot F, Fischer M, Hancock C, Maier T S, Meiners T, Müller C, Obermaier E, Prati D, Socher S A, Sonnemann I, Wäschke N, Wubet T, Wurst S and Rillig M C. (2014). Choosing and using diversity indices: Insights for ecological applications from the German biodiversity exploratories. Ecology and Evolution 4(18): 3514–3524. https://doi.org/10.1002/ece3.1155
Murcia C. (1995). Edge effects in fragmented forests: Implications for conservation. Trends in Ecology & Evolution 10(2): 58–62. https://doi.org/10.1016/S0169-5347(00)88977-6
Nakashima Y, Inoue E, Inoue-Murayama M and Sukor J A. (2010). High potential of a disturbance-tolerant frugivore, the common palm civet Paradoxurus hermaphroditus (Viverridae), as a seed disperser for large-seeded plants. Mammal Study 35(3): 209–215. https://doi.org/10.3106/041.035.0307
Nasir D. (2013). Natural history of the Colugo. Bangi, Selangor: Penerbit Universiti Kebangsaan Malaysia.
Nekaris K A I. (2001). Activity budget and positional behavior of the Mysore slender loris (Loris tardigradus lydekkerianus): Implications for slow climbing locomotion. Folia Primatologica 72(4): 228–241. https://doi.org/10.1159/000049942
Nekaris K A I, Blackham G V and Nijman V. (2008). Conservation implications of low encounter rates of five nocturnal primate species (Nycticebus spp.). Asian Biodiversity Conservation 17: 733–747. https://doi.org/10.1007/s10531-007-9308-x
Nekaris K A I, Pambudi J A A, Susanto D, Ahmad R D and Nijman V. (2014). Densities, distribution and detectability of a small nocturnal primate (Javan slow loris Nycticebus javanicus) in a montane rainforest. Endangered Species Research 24(2): 95–103. https://doi.org/10.3354/esr00585
Nekaris K A I, Poindexter S and Streicher U. (2020). Nycticebus coucang. The IUCN Red List of Threatened Species 2020: e.T163017685A17970966. https://doi.org/10.2305/IUCN.UK.2020-2.RLTS.T163017685A17970966.en
Nor S M, Mohd Sah S A, Nurul-Ain E, Nor Zalipah M, Ahmad N, Ahmad M Y, Marj R D, Ganesan M and Ariffin A S. (2007). Small mammal surveys at several localities on Langkawi Island, Kedah, Malaysia. Malaysian Forester 70(2): 123–131.
Oksanen J, Blanchet F G, Friendly M, Kindt R, Legendre P, McGlinn D, Minchin P R, O’Hara R B, Simpson G L, Solymos P, Stevens M H H, Szoecs E and Wagner H. (2019). Vegan: Community ecology package. R package version 2.5-2. Cran R [Software].
Oliver K, Ngoprasert D and Savini T. (2019). Slow loris density in a fragmented, disturbed dry forest, north‐east Thailand. American Journal of Primatology 81(3): e22957. https://doi.org/10.1002/ajp.22957
Othman S. (Ed.). (2000). Special issue: Vertebrate fauna of Krau Wildlife Reserve. Journal of Wildlife and Parks 18: 1–106. http://www.wildlife.gov.my/images/document/penerbitan/jurnal/Jil182000.pdf
Palmeirim A F, Peres C A and Vieira M V. (2020). Optimizing small mammal surveys in Neotropical fragmented landscapes while accounting for potential sampling bias. Mammalian Biology 100(1): 81–90. https://doi.org/10.1007/s42991-020-00012-2
Paterson J D. (1981). Postural-positional thermoregulatory behaviour and ecological factors in primates. Canadian Review of Physical Anthropology 3: 3–11.
Pereira B C, Ferreguetti Á C and Bergallo H G. (2017). Factors affecting mammalian encounter rates in transect surveys: A case study in Ilha Grande State Park, State of Rio de Janeiro, Brazil. Oecologia Australis 21(4): 422–430. https://doi.org/10.4257/oeco.2017.2104.06
PERHILITAN. (2017). Red list of mammals for Peninsular Malaysia Version 2.0. Kuala Lumpur: Department of Wildlife and National Parks (PERHILITAN) Peninsular Malaysia.
Phillips H R P, Newbold T and Purvis A. (2017). Land-use effects on local biodiversity in tropical forests vary between continents. Biodiversity and Conservation 26(9): 2251–2270. https://doi.org/10.1007/s10531-017-1356-2
Plumptre A J, Sterling E J and Buckland S T. (2013). Primate census and survey techniques. In E Sterling, N Bynum and M Blair (Eds.), Primate ecology and conservation. Oxford: Oxford University Press, 10–26. https://doi.org/10.1093/acprof:oso/9780199659449.003.0002
Rinehart K A, Elbroch L M and Wittmer H U. (2014). Common biases in density estimation based on home range overlap with reference to pumas in Patagonia. Wildlife Biology 20(1): 19–26. https://doi.org/10.2981/wlb.12100
Rocha E C, de Lima C A, Batista N L and da Silva Castro A L. (2021). Encounter rates and density of medium and large-sized mammals with nocturnal habits in southern Amazon, Brazil. Current Science 121(8): 1113. https://doi.org/10.18520/cs/v121/i8/1113-1119
Rode-Margono E J, Nijman V, Wirdateti and Nekaris K A I. (2014). Ethology of the critically endangered Javan slow loris Nycticebus javanicus É. Geoffroy Saint-Hilaire in West Java. Asian Primates Journal 4(2): 27–38.
Rode-Margono E J and Nekaris K A I. (2014). Impact of climate and moonlight on a venomous mammal, the Javan slow loris (Nycticebus javanicus Geoffroy, 1812). Contributions to Zoology 83(4): 217–225. https://doi.org/10.1163/18759866-08304001
Ross J, Brodie J, Cheyne S, Chutipong W, Hedges L, Hearn A, Linkie M, Loken B, Mathai J, McCarthy J, Ngoprasert D, Tantipisanuh N, Wilting A and Haidir I A. (2015). Hemigalus derbyanus. The IUCN Red List of Threatened Species 2015: e.T41689A45216918.
Ruppert N B, Asyraf M and Shahrul Anuar M S. (2015). Diversity and biomass of terrestrial small mammals at a Malaysian primary rainforest (Segari Melintang forest reserve, Peninsular Malaysia). Journal of Tropical Life Sciences 5(1): 3–34. https://doi.org/10.11594/jtls.05.01.07
Sala O E, Chapin F S, Armesto J J, Berlow E, Bloomfield J, Dirzo R, Huber-Sanwald E, Huenneke L F., Jackson R B, Kinzig A and Leemans R. (2000). Global biodiversity scenarios for the year 2100. Science 287: 1770–1774.
https://doi.org/10.1126/science.287.5459.1770
Santini L, González‐Suárez M, Russo D, Gonzalez‐Voyer A, von Hardenberg A and Ancillotto L. (2019). One strategy does not fit all: Determinants of urban adaptation in mammals. Ecology Letters 22(2): 365–376. https://doi.org/10.1111/ele.13199
Shafie N. (2016). Ecological aspects of fruit bats (Megachiroptera) in Penang Island, Malaysia. PhD diss., Universiti Sains Malaysia. https://doi.org/10.1017/ CBO9781107415324.004
Silveira L, Jácomo A T A and Diniz-Filho J A F. (2003). Camera trap, line transect census and track surveys: A comparative evaluation. Biological Conservation 114(3): 351–355. https://doi.org/10.1016/S0006-3207(03)00063-6
Sompud J, Lu S, Pamin D, Sompud C B and Mojiol A R. (2016). Nocturnal terrestrial mammals population in Sabah Forest Industries (SFI) forest plantation, Sipitang. Transactions on Science and Technology 3(1–2): 123–129. https://doi.org/10.1039/b008095k
Southern H N, Watson J S and Chitty D. (1946). Watching nocturnal animals by infra-red radiation. The Journal of Animal Ecology 15(2): 198. https://doi.org/10.2307/1558
Starr C, Nekaris K A I, Streicher U and Leung L K P. (2011). Field surveys of the vulnerable pygmy slow loris Nycticebus pygmaeus using local knowledge in Mondulkiri Province, Cambodia. Oryx 45(1): 135–142. https://doi.org/10.1017/S0030605310001316
Sutherland W J. (2002). Mammals. In W J Sutherland (ed.), Ecological census techniques. Cambridge, UK: Cambridge University Press, 260–278.
Tahir Z M, Mohidin F S M and Rosely N F N. (2020). Visual tree analysis of rain trees (Samanea saman) in Universiti Sains Malaysia, Main Campus. IOP Conference Series: Earth and Environmental Science 549(1): 012032. https://doi.org/10.1088/1755-1315/549/1/012032
Thompson W I. (Ed.). (2004). Sampling rare or elusive species: Concepts, designs, and techniques for estimating population parameters. Washington, DC: Island Press.
Timmins R and Duckworth J. (2015). Tragulus kanchil. The IUCN Red List of Threatened Species 2015: e.T136297A61978576. https://doi.org/10.2305/IUCN.UK.2015-2.RLTS.T136297A61978576.en
Umapathy G and Kumar A. (2000). The occurrence of arboreal mammals in the rain forest fragments in the Anamalai Hills, south India. Biological Conservation 92(3): 311– 319. https://doi.org/10.1016/S0006-3207(99)00097-X
Underwood A H, Derhè M A and Jacups S. (2022). Thermal imaging outshines spotlighting for detecting cryptic, nocturnal mammals in tropical rainforests. Wildlife Research 49(6): 491–499. https://doi.org/10.1071/WR21130
Vernes K. (2001). Gliding performance of the northern flying squirrel (Glaucomys sabrinus) in mature mixed forest of eastern Canada. Journal of Mammalogy 82(4): 1026–1033. https://doi.org/10.1644/1545-1542(2001)082%3C1026:GPOTNF%3E2.0.CO;2
Vestal B M and Hill J L. (1972). Pattern vision of deermice (Peromyscus) under red light. Journal of Mammalogy 53(2): 374–376. https://doi.org/10.2307/1379179
Vitousek P M. (1997). Human domination of earth’s ecosystems. Science 277(5325): 494–499. https://doi.org/10.1126/science.277.5325.494
Voskamp A, Rode E J, Coudrat C N Z, Wirdateti, Abinawanto, Wilson R J and Nekaris K A I. (2014). Modelling the habitat use and distribution of the threatened Javan slow loris Nycticebus javanicus. Endangered Species Research 23(3): 277–286. https://doi.org/10.3354/esr00574
Wells K, Corlett R T, Lakim M B, Kalko E K V and Pfeiffer M. (2009). Seed consumption by small mammals from Borneo. Journal of Tropical Ecology 25(5): 555–558. https://doi.org/10.1017/S0266467409990058
Wells K, Kalko E K V, Lakim M B and Pfeiffer M. (2007). Effects of rain forest logging on species richness and assemblage composition of small mammals in Southeast Asia. Journal of Biogeography 34(6): 1087–1099. https://doi.org/10.1111/j.1365-2699.2006.01677.x
Wells K, Lakim M B and O’Hara R B. (2014). Shifts from native to invasive small mammals across gradients from tropical forest to urban habitat in Borneo. Biodiversity and Conservation 23(9): 2289–2303. https://doi.org/10.1007/s10531-014-0723-5
Wells K, Pfeiffer M, Lakim M B and Linsenmair K E. (2004). Use of arboreal and terrestrial space by a small mammal community in a tropical rain forest in Borneo, Malaysia. Journal of Biogeography 31(4): 641–652. https://doi.org/10.1046/j.1365-2699.2003.01032.x
Wearn O R, Carbone C, Rowcliffe JM, Pfeifer M, Bernard H and Ewers R M. (2018). Land-use change alters the mechanisms assembling rainforest mammal communities in Borneo. Journal of Animal Ecology 88(1), 125–137. https://doi.org/10.1111/1365-2656.12903
Weng Chan N. (1998). Environmental hazards associated with hill land development in Penang Island, Malaysia: Some recommendations on effective management. Disaster Prevention and Management: An International Journal 7(4): 305–318. https://doi.org/10.1108/09653569810230148
Wilson E O. (2002). The future of life. London: Little Brown.
Whittaker R H. (1977). Evolution of species diversity in land communities. Evolutionary Biology 10: 1–67. https://doi.org/10.1007/978-1-4615-6953-4_1
Wiens F and Zitzmann A. (1999). Predation on a wild slow loris (Nycticebus coucang) by a reticulated python (Python reticulatus). Folia Primatologica 70(6): 362–364. https://doi.org/10.1159/000021719
Wiens F. (2002). Behavior and ecology of wild slow lorises (Nycticebus coucang): Social organization, infant care system, and diet. PhD diss., Universität Bayreuth, Germany.
Winter Y and von Helversen O. (2001). Bats as pollinators: Foraging energetics and floral adaptations. In L Chittka and J D Thomson (eds.), Cognitive ecology of pollination. Cambridge University Press, 148–170. https://doi.org/10.1017/CBO9780511542268.009
Yasuda M, Miura S, Ishii N, Okuda T and Hussein N A. (2009). Fallen fruits and terrestrial vertebrate frugivores: A case study in a lowland tropical rainforest in Peninsular Malaysia. In P-M Forget, P Hulme, J E Lambert and S B Vander Wall (eds.), Seed fate: Predation, Dispersal and seedling establishment. Wallingford: CABI, 151– 174. https://doi.org/10.1079/9780851998060.0151
Zuur A F, Leno E N and Elphick C S. (2010). A protocol for data exploration to avoid common statistical problems. Methods Ecology and Evolution 1: 3–14. https://doi.org/10.1111/j.2041-210X.2009.00001.x
Zuur A F, Leno E N, Walker N J, Saveliev A A and Smith G M. (2009). Mixed effects models and extensions in ecology with R (Vol. 574). New York: Springer. https://doi.org/10.1007/978-0-387-87458-6