{"paper_id":"093177cd-40e3-4f3f-8720-9a4f3138865e","body_text":"1 \n \nResearch Article 1 \nSyafiq, Badli-Sham, Fariduddin, Izham, Aqmal-Naser, Farah, Gukaaneswaran, Nadhirah, 2 \nFarhana, Shahril-Ridhwan, Joehan, Kei Li, Shu Xin, Fahmi-Ahmad, Rizal, and Ahmad  |  3 \nHerpetofauna of Bukit Maras, Terengganu, Peninsular Malaysia 4 \nHerpetofauna Diversity of The Disturbed and Isolated Bukit Maras in Terengganu, Peninsular 5 \nMalaysia 6 \nMuhamad F. Syafiq1, Baizul Hafsyam Badli-Sham2, Syed Ahmad Fariduddin3, Mohd Izham 7 \nMohd A Wahid2, Mohamad Aqmal-Naser1, Farah Hazirah Amir2, Gukaaneswaran Kaliyappan2, 8 \nNur Nadhirah Sapri2, Nurziattul Farhana Nordin2, Muhammad Shahril-Ridhwan2, Joehan 9 \nAzzimin2, Tng Kei Li2, Ng Shu Xin2, Muhammad Fahmi-Ahmad2, Syed Ahmad Rizal2 and 10 \nAmirrudin B. Ahmad1,2,* 11 \nAffiliations.  12 \n1 Institute of Tropical Biodiversity and Sustainable Development, Universiti Malaysia 13 \nTerengganu, 21030 Kuala Nerus, Terengganu, Malaysia  14 \n2 Faculty of Science and Marine Environment, Universiti Malaysia Terengganu, 21030 Kuala 15 \nNerus, Terengganu, Malaysia 16 \n3 501-A, Taman Seri Kolam, Jalan Sultan Sulaiman, 20000 Kuala Terengganu, Terengganu, 17 \nMalaysia 18 \n 19 \n*Corresponding author: Amirrudin B. Ahmad, email: amirrudin@umt.edu.my 20 \n  21 \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n2 \n \nAbstract 22 \nWe present the first checklist of herpetofauna in Bukit Maras based on surveys conducted from 23 \n2019 to 2023. Visual Encounter Surveys (VES) and drift-fenced pitfall traps were employed as 24 \ncollection methods. Our study documented a total of 55 herpetofauna species, comprising 23 25 \namphibians and 32 reptiles. Among these, the critically endangered species, Manouria emys 26 \n(according to the IUCN Red List) is a species of high conservation concern. The non-asymptotic 27 \nnature of the Species Accumulation Curve (SAC) suggests that further sampling efforts could 28 \nreveal additional species. Species-habitat network analysis revealed variations in species 29 \ncomposition across different habitat types. Notably, secondary forest exhibited higher 30 \nherpetofauna diversity compared to agricultural areas. Therefore, the conservation of remaining 31 \nsecondary forest in Bukit Maras is crucial for preserving its herpetofauna and mitigating 32 \nanthropogenic impacts on this disturbed and isolated ecosystem. 33 \nKey words 34 \nAgricultural area, amphibians, anthropogenic disturbance, habitat island, reptiles, secondary 35 \nforest, Southeast Asia 36 \nAcademic editor: [academic editor to complete] | publication data [copyeditor/layout editor to 37 \ncomplete] 38 \nCitation: [copyeditor/layout editor to complete] 39 \n  40 \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n3 \n \nIntroduction 41 \nPeninsular Malaysia boasts remarkable biodiversity (Myers et al. 2000). However, rampant 42 \ndeforestation threatens this richness (Sodhi et al. 2004). Driven by factors like urban sprawl, 43 \nlogging, and agriculture, primary forest degradation has fragmented these once-continuous 44 \nlandscapes (Hadad et al. 2015; Magintan et al. 2017). This has resulted in isolated hills 45 \nsurrounded by human-modified areas. Research suggests these isolated hills can serve as refugia 46 \nfor displaced amphibians and reptiles, even harboring new species discoveries (Quah et al. 2013; 47 \nGrismer et al. 2014a, 2016a). Documenting herpetofauna diversity in these areas is crucial before 48 \nlocal populations disappear. 49 \nIn the Terengganu state, over 308,000 hectares of native forest have been cleared for agricultural 50 \nactivities, primarily for oil palm, rubber plantation, and crop cultivation (Alam et al. 2012). 51 \nWhile the existing body of researches showed that many herpetofauna studies have been 52 \nconducted in the primary forest of this state (Grismer et al. 2013a, 2013b, 2014b, 2014c, 2015, 53 \n2016b, 2018; Chan et al. 2014; Sumarli et al. 2015, 2016; Nur Amalina et al. 2017; Shahirah-54 \nIbrahim et al. 2018; Quah et al. 2021; Badli-Sham et al. 2023; Syafiq et al. 2023, 2024), only a 55 \nfew have focused on the disturbed forested areas (Badli-Sham et al. 2019; Fatihah-Syafiq et al. 56 \n2020; Komaruddin et al. 2020). Hence, there is a significant gap in knowledge on the 57 \nherpetofauna of these habitats, which necessitates further study. 58 \nBukit Maras (BM) is an isolated hill range surrounded by the human settlement and agricultural 59 \nmatrix. Approximately 40% of the forested area at BM have been converted to orchards. The 60 \nsecondary forest in these hills regenerated after the land abandonment (about 15 years ago), but 61 \nthis area also faces the threat of deforestation due to current expansion of orchard areas. Given 62 \nthe alarming rate of habitat loss, these disturbances could severely impact herpetofauna species 63 \nthat may rely on these habitats as refugia.  64 \nRegrettably, information on the herpetofauna in this hill is virtually unknown. To our knowledge, 65 \nthere is only a single inventory study in Bukit Maras but which focused on the chiropteran 66 \nspecies (Sulaiman and Lian 2011). The study recorded a total of 14 bat species in BM, with one 67 \nof the species categorized as \"Near Threatened\" by the IUCN Redlist. Therefore, this study aims 68 \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n4 \n \nto produce the first report of herpetofauna in BM. Bukit Maras is a non-protected area thus, this 69 \ninvestigation could provide valuable information on what species inhabit BM before we lose 70 \nthem due to the deforestation.   71 \nMethods 72 \nStudy area 73 \nBukit Maras is situated in the Kuala Nerus district, Terengganu, surrounded by human settlement 74 \nand agricultural areas (Fig. 1). It has a total of approximately 2200 hectares of land comprised of 75 \nagricultural areas, secondary forest, and primary forest (underexplored).  76 \nThere are three main sampling sites: 77 \n• Site 1 (5°25'20.98\"N 103°1'18.18\"E) – This hill has a large soil road from the foothill to 78 \nthe summit as a hiking trekking trail. The landscape is characterized by an agricultural 79 \nmatrix and an open area for car parking at the foothill. Crop plantation areas also can be 80 \nfound along the trekking trails up to 200 meters. Examples of the planted crops are durian 81 \ntrees (Durio spp.), petai trees (Parkia speciosa), and banana plants (Musa spp.). Two 82 \ndisturbed small streams can be found in the agricultural areas, which originated from the 83 \nintact secondary and primary forest. 84 \n• Site 2 (5°24'33.77\"N 103°1'27.63\"E) – This hill also has a large soil road from the 85 \nfoothill to the summit as a hiking trekking trail. Oil palm and rubber plantations are the 86 \nprimary agricultural plantations here. Secondary forest vegetation situated mainly at the 87 \npeak. A disturbed small stream flow through the agricultural area.  88 \n• Site 3 (5°24'35.4\"N 103°00'47.4\"E) – This site is a secondary forest characterized by a 89 \nsmall stream covered by a few dipterocarp tree species left undisturbed from 90 \ndeforestation in the previous years. This site has the minimal disturbance compared to 91 \nother two sites but the threat of deforestation for orchard expansion is imminent.  92 \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n5 \n \n93 \nFigure 1. A. Location of Bukit Maras in Terengganu, Peninsular Malaysia. B–C. QGIS 94 \ngenerated map showing the area of BM surrounded by the agricultural, rural, and urban areas. 95 \nSample collection and preservation 96 \nThe surveys were conducted opportunistically from July 2019–October 2019, 4 July 2020, 12 97 \nJuly 2020, 26–27 July 2020, 17 October 2020, 12–14 November 2020, 1 November 2020, 26 98 \nNovember 2020, 28 November 2020,15 January 2021 and between 31 December 2022 to 11 99 \nMarch 2023. Initially, pilot surveys were done opportunistically in 2019, and a systematic survey 100 \nwas in the planning for the subsequent years but due to the Covid 19 situation during the former 101 \nyears (2020–2021), opportunistic surveys approach was implemented at this site to standardize 102 \nthe sampling effort. Surveyed areas spanned the disturbed and undisturbed small streams, 103 \nsecondary forests, and agricultural areas. The primary forest was remained untapped due to its 104 \ninaccessibility. We sampled 13 sampling points grouped into four categories: agricultural area 105 \n(AA); secondary forest (SF); agricultural area stream (STA); secondary forest stream (STF) 106 \n(Table 1). We divided the habitat types into these four categories as we expect these four habitat 107 \ntypes will consist of different species composition and demonstrate the importance of each 108 \nhabitat for this species.  109 \nSouth \nChina \nSea \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n6 \n \nTable 1. Habitat categories studied in Bukit Maras 110 \nHabitat Characteristics Sampling \nSite \nSampling \nPoint \nAgricultural \narea \nMainly composed of durian (Durio sp.), banana \n(Musa sp.), petai (Parkia speciosa), oil palm (Elaeis \nguineensis) and rubber (Hevea brasiliensis) \nplantation. Scatterly located from the foothill up to \nelevation of 200 metres a.s.l. \nSite 1, \nSite 2 \n4 \nSecondary \nforest \nForest remnants composed mainly of native species, \nunmanaged environment, with formation of \nunderstory and canopy. \nAll sites 4 \nAgricultural \narea stream \nSmall stream in the agricultural area with no canopy \nformation \nSite 1, \nSite 2 \n3 \nSecondary \nforest stream \nSmall stream in the agricultural area with canopy \nformation \nSite 1, \nSite 3 \n2 \n 111 \n  \nA \n B \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n7 \n \n  \n  \nC \n D \nE\n F\nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n8 \n \n  \nFigure 2. Habitat types in Bukit Maras A–D. agricultural area E. secondary forest F. stream at 112 \nsecondary forest G–H. stream at agricultural area. 113 \nTo maximize the sampling effort, three types of collecting methods were employed: Visual 114 \nEncounter Survey (VES), drift-fenced pitfall traps, and acoustic sampling. The samplings were 115 \nexecuted during the day (10:00 to 13:00 h) and at night (20:00 to 23:00 h) to record both diurnal 116 \nand nocturnal species. The VES activities were conducted with search parties consisting of three 117 \nto four persons. This method was executed during the day and night to sample diurnal and 118 \nnocturnal species in the area. A set of pitfall traps consisted of three 18L buckets and aluminum 119 \nzinc as the fence. The buckets were buried two meters apart from each other and were arranged 120 \nin approximately straight lines. Two sets of pitfall traps were set up randomly at different 121 \nlocations. The first one was deployed at the foothill near the agricultural area, and the second set 122 \nwas deployed at the hilly area in the secondary forest. The vocalization method also was used to 123 \nrecord the species based on the frog calls. The recorded sound can be used for species-specific 124 \nsound characteristic description in future study (Chan et al., 2020; Quah et al., 2021). A total of 125 \n816 man-hours (204 hours/person) were spent for the herpetofauna surveys.  126 \nCaptured specimens were identified based on their morphological characteristic, following Berry 127 \n(1975) for amphibians, Grismer (2011) for lizards, Auliya (2007) for freshwater turtles and 128 \ntortoises, and Das (2012) for snakes. Photographs of live specimens were taken with the Canon 129 \n3000D and Sony A6000 digital cameras. Amphibian nomenclature follows the Amphibian 130 \nSpecies of the World database (Frost 2024), while the reptile nomenclature follows The Reptile 131 \nDatabase (Uetz et al. 2023). Only a few samples were taken for voucher specimens to minimize 132 \nG\n H\nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n9 \n \nthe extirpation of the current population. Euthanized voucher specimens were then preserved 133 \nwith 10% formalin before being transferred into 70% ethanol for long-term storage and 134 \ndeposited at the General Biology Lab, Universiti Malaysia Terengganu, and catalogued under 135 \nUMT Zoological Collection (UMTZC). 136 \nData analysis 137 \nThe individual-based and coverage-based rarefaction and extrapolation analysis was constructed 138 \nby using the “iNEXT” R package (Hsieh et al. 2016) to determine the adequacy of our sampling 139 \neffort at BM and for both amphibian and reptile groups. The sampling achieves completeness 140 \nwhen the curve is plateauing for the former and achieve completeness value which is 1.0 for the 141 \nlatter. This analysis permits comparison of diversity with Hill numbers of order q for unequal 142 \nsampling effort between two or more sites or groups. There are three types of order q for Hill 143 \nnumbers namely, species richness (q=0), Shannon’s diversity (q=1) and Simpson’s diversity 144 \n(q=3). We only utilized diversity order of species richness (q=0) for this study. We also used this 145 \nanalysis to compare the herpetofauna diversity between the four habitat types. For the 146 \ncomparison of habitat types, we combined both amphibians and reptiles data as herpetofauna 147 \ndata to utilize in the comparison analysis. Non-overlap curves strongly indicate significant 148 \ndifference and vice versa for both individual-based and coverage-based rarefaction and 149 \nextrapolation curves.  150 \nFor species-habitat network, we built two networks using both abundance and incidence data 151 \nacting as links while habitat types and herpetofauna species acting as nodes (Marini et al. 2019). 152 \nThe analysis was performed using the “bipartite” R package (Dormann et al. 2009). All analyses 153 \nwere executed in RStudio software (RStudio Team 2023). 154 \nResults 155 \nA total of 55 herpetofauna species from six amphibian families (Bufonidae – 2 spp., 156 \nDicroglossidae – 5 spp., Megophryidae – 3 spp., Microhylidae – 4 spp., Ranidae – 5 spp., 157 \nRhacophoridae – 4 spp.) and nine reptile families (Agamidae – 7 spp., Gekkonidae – 7 spp., 158 \nScincidae – 4 spp., Varanidae – 1 spp., Colubridae – 9 spp., Pythonidae – 1 spp., Viperidae – 1 159 \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n10 \n \nspp., Trionychidae – 1 spp., Testudinidae – 1 spp.) were recorded in this study (Table 2). Based 160 \non IUCN Redlist (2024), there is only single species (Manouria emys) listed under \"Critically 161 \nEndangered\" status in this study, whereas the rest of the species are categorized as \"Least 162 \nConcern\". 163 \nTable 2. A species checklist of amphibians and reptiles recorded in Bukit Maras and their IUCN 164 \nstatus. 165 \nNo Taxa IUCN \n AMPHIBIANS \n \n Bufonidae  \n1 Duttaphrynus melanostictus (Schneider, 1799) LC \n2 Ingerophrynus parvus (Boulenger, 1887) LC \n Dicroglossidae  \n3 Fejervarya limnocharis (Gravenhorst, 1829) LC \n4 Limnonectes blythii (Boulenger, 1920) LC \n5 Limnonectes deinodon Dehling, 2014 LC \n6 Limnonectes hascheanus (Stoliczka, 1870) LC \n7 Limnonectes malesianus (Kiew, 1984) LC \n Megophryidae  \n8 Leptobrachella sola (Matsui, 2006) LC \n9 Leptobrachium hendricksoni Taylor, 1962 LC \n10 Pelobatrachus nasuta (Schlegel, 1858) LC \n Microhylidae  \n11 Kaloula pulchra Gray, 1831 LC \n12 Microhyla berdmorei (Blyth, 1856) LC \n13 Microhyla heymonsi Vogt, 1911 LC \n14 Microhyla mantheyi Das, Yaakob & Sukumaran, 2007 LC \n Ranidae  \n15 Humerana miopus (Boulenger, 1918) LC \n16 Hylarana glandulosa (Boulenger, 1882) LC \n17 Hylarana labialis (Boulenger, 1887) LC \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n11 \n \n18 Hylarana laterimaculata (Barbour & Noble, 1916) LC \n19 Hylarana nicobariensis (Stoliczka, 1870) LC \n Rhacophoridae \n \n20 Nyctixalus pictus (Peters, 1871) LC \n21 Polypedates discantus Rujirawan, Stuart & Aowphol, 2013 LC \n22 Polypedates leucomystax (Gravenhorst, 1829) LC \n23 Theloderma licin McLeod & Ahmad, 2007 LC \n REPTILES  \n LIZARDS \n \n Agamidae  \n24 Acanthosaura armata (Gray, 1827) LC \n25 Bronchocela cristatella (Kuhl, 1820) LC \n26 Calotes versicolor (Daudin, 1802) LC \n27 Draco sumatranus Schlegel, 1844 LC \n28 Gonocephalus grandis (Gray, 1845) LC \n29 Gonocephalus liogaster (Günther, 1872) LC \n30 Leiolepis belliana (Hardwicke & Gray, 1827) LC \n Gekkonidae  \n31 Cyrtodactylus consobrinus (Peters, 1871) LC \n32 Cyrtodactylus quadrivirgatus Taylor, 1962 LC \n33 Gehyra mutilata (Wiegmann, 1834) LC \n34 Gekko monarchus (Schlegel, 1836) LC \n35 Hemidactylus frenatus Duméril & Bibron, 1836 LC \n36 Hemidactylus platyurus (Schneider, 1797) LC \n37 Hemiphyllodactylus typus Bleeker, 1860 LC \n Scincidae  \n38 Dasia olivacea Gray, 1839 LC \n39 Eutropis multifasciata (Kuhl, 1820) LC \n40 Lipinia vittigera (Boulenger, 1894) LC \n41 Lygosoma siamensis Siler, Heitz, Davis, \nFreitas, Aowphol, Termprayoon & Grismer, 2018 \nLC \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n12 \n \n Varanidae \n \n42 Varanus salvator (Laurenti, 1768) LC \n SNAKES \n \n Colubridae  \n43 Ahaetulla prasina (Boie, 1827) LC \n44 Boiga cynodon (Boie, 1827) LC \n45 Dendrelaphis caudolineatus (Gray, 1834) LC \n46 Dendrelaphis cyanochloris (Wall, 1921) LC \n47 Dendrelaphis pictus (Gmelin, 1789) LC \n48 Dendrelaphis striatus (Cohn, 1905) LC \n49 Gonyosoma oxycephalum (Boie, 1827) LC \n50 Lycodon subcinctus Boie, 1827 LC \n51 Pseudorhabdion longiceps (Cantor, 1847) LC \n Pythonidae  \n52 Malayopython reticulatus (Schneider, 1801) LC \n Viperidae \n \n53 Tropidolaemus wagleri (Boie, 1827) LC \n FRESHWATER TURTLE AND TORTOISE  \n Testudinidae  \n54 Manouria emys (Schlegel & Müller, 1844) CR \n Trionychidae  \n55 Dogania subplana (Geoffroy Saint-Hilaire, 1809) LC \nNotes: IUCN STATUS: LC = Least Concerned; CR = Critically Endangered.  166 \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n13 \n \n  \n  \n   \nA \nC \n D\nE\n F\nB \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n14 \n \n  \n  \n  \nFigure 3. Amphibians from Bukit Maras A. Duttaphrynus melanostictus B. Ingerophrynus 167 \nparvus C. Fejevarya limnocharis D. Limnonectes deinodon E. Leptobrachium hendricksonii F. 168 \nG\n H\nI\nK\n L\nJ \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n15 \n \nMegophrys nasuta G. Microhyla heymonsi H. Microhyla mantheyi I. Humerana miopus J. 169 \nHylarana labialis K. Polypedates leucomystax L. Theloderma licin 170 \n    \n   \n  \nA \nC \n D\nE\n F\nB \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n16 \n \n \n  \n  \n  \nFigure 4. Lizards from Bukit Maras A. Acanthosaura armata B. Bronchocela cristatella C. 171 \nCalotes versicolor D. Gonocephalus grandis E. Gonocephalus liogaster F. Leiolepis belliana G. 172 \nCyrtodactylus consobrinus H. Cyrtodactylus quadrivirvagtus I. Gekko monarchus J. 173 \nHemiphyllodactylus typus K. Lygosoma siamensis L. Lipinia vittigera 174 \nG\nH\nI\nK\n L\nJ \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n17 \n \n     \n \n \n \n \n \n   \nA \nC \n D\nE\n F\nB \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n18 \n \n    \n    \n    \nFigure 5. Freshwater tortoise and turtle and snakes from Bukit Maras A. Ahaetulla prasina B. 175 \nBoiga cynodon C. Dendrelaphis caudolineatus D. Dendrelaphis cyanochloris E. Dendrelaphis 176 \npictus F. Dendrelaphis striatus G. Gonyosoma oxycephalum H. Lycodon subcinctus I. 177 \nMalayopython reticulatus J. Tropidolaemus wagleri K. Manouria emys L. Dogania subplana 178 \nG\n H\nI\nK\n L\nJ \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n19 \n \nThe individual-based rarefaction and extrapolation curves for the total herpetofauna and each of 179 \namphibian and reptilian assemblages implied that additional species can be detected in BM when 180 \nmore sampling efforts are executed as the extrapolated curves for all three showed no sign of 181 \nplateauing just yet. At the extrapolated curves, it is estimated that 60 herpetofauna species can be 182 \ndiscover in BM, which to be specific a potential discovery of an additional two species of 183 \namphibians and three species of reptiles with additional efforts. Coverage-based rarefaction and 184 \nextrapolation curves are in accord with the individual-based rarefaction and extrapolation curves 185 \nas the curves approaching the sampling completeness value. All curves are non-overlapped 186 \nwhich indicated that the species richness (q=0) for all curves are significantly different. 187 \n 188 \nFigure 6. A. Individual-based rarefaction (solid line segment) and extrapolation (dotted line 189 \nsegment) sampling curves with 95% confidence interval (shaded areas) (left panel) and B. 190 \ncoverage-based rarefaction (solid line segment) and extrapolation (dotted line segment) sampling 191 \ncurves with 95% confidence interval (shaded areas) (right panel) for herpetofauna sampled in 192 \nBukit Maras, Terengganu, Peninsular Malaysia.  193 \nIn general, the Calotes versicolor had the highest abundance compared to other species in Bukit 194 \nMaras. It also contributed the highest number of individuals in agricultural area (AA). The AA is 195 \nalso the site with the highest number of individuals occupied by the herpetofauna. Species 196 \nrichness-wise, the seconday forest (SF) had the highest number of species among the habitats. 197 \nThe Fejevarya limnocharis had the highest number of frequency as it can be found across all 198 \nfour habitats. Each of the habitat type composed of different set of species composition (Fig. 7). 199 \nIn specific, there are 36 species of herpetofauna in SF and the highest number of individuals is 200 \nEutropis multifasciata (12 individuals), followed by Microhyla heymonsi (11 individuals) and 201 \nA\n B\nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n20 \n \nLimnonectes hascheanus (7 individuals). Meanwhile, AA had only 17 species and dominated by 202 \nC. versicolor (68 individuals), followed by Hemidactylus frenatus (17 individuals) and 203 \nDuttaphrynus melanostictus (15 individuals). For riparian areas, stream of secondary forest 204 \n(STF) had 16 species, and dominated by Limnonectes deinodon (36 individuals), followed by 205 \nHylarana labialis (25 individuals) and Leptobrachium hendricksoni (16 individuals). On the 206 \nother hand, stream of agricultural area (STA) had only nine species, dominated by Polypedates 207 \nleucomystax (23 individuals), followed by Fejevarya limnocharis (19 individuals) and H. labialis 208 \n(17 individuals).  209 \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n21 \n \n  \nFigure 7. Species-habitat network in Bukit Maras. A. Abundance-based species-habitat network 210 \nB. Incidence-based species-habitat network. The left panel represents nodes for habitat types 211 \nwhile the right panel represents nodes for each species. The width of the links represents the 212 \nA \n B \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n22 \n \nnumber of individuals (abundance-based) and frequency of occurrence (incidence-based). The 213 \nnodes are arranged from the highest to lowest (abundance/frequency). 214 \nThe diversity order of species richness (q=0) for both curves for secondary forest (SF) and 215 \nsecondary forest’s stream (STF) are not yet approaching asymptote even at the extrapolated 216 \ncurves indicating that more species can be discovered in both habitats. On the other hand, both 217 \ncurves for agricultural area (AA) and agricultural area’s stream (STA) showed a sign of levelling 218 \noff at the extrapolated curves indicating that the sampling in these areas are almost complete.  219 \n 220 \nFigure 8. A. Individual-based rarefaction (solid line segment) and extrapolation (dotted line 221 \nsegment) sampling curves with 95% confidence interval (shaded areas) and B. coverage-based 222 \nrarefaction (solid line segment) and extrapolation (dotted line segment) sampling curves with 223 \n95% confidence interval (shaded areas) for the herpetofauna data of four habitat types: green 224 \n(secondary forest); red (agricultural area). The solid dots/triangles represent the reference 225 \nsamples. Hill numbers of order (q = 0) or species richness was measured for both curves. 226 \nDiscussion 227 \nThe forested area at the human-induced landscape is disappearing at an unprecedented rate, 228 \nleaving \"islands\" of forest remnants (Sodhi et al. 2010). This habitat may become refuge to the 229 \nperturbed herpetofauna and our study demonstrated that by documenting a total of 55 species of 230 \nherpetofauna in Bukit Maras. This record is a preliminary checklist and additional species record 231 \nin this baseline data is promising, as indicated by the non-asymptotic individual rarefaction and 232 \nextrapolation curves. The extrapolated curves indicated that an additional five herpetofauna 233 \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n23 \n \nspecies can be found with additional efforts. Some species of herpetofauna particularly snakes 234 \ncan be elusive and previous study on temporal snake diversity in Terengganu suggested that 235 \nsurveys should be intensified especially during the raining season (Syafiq et al. 2023). This 236 \nwould not only increase the chance to encounter elusive snake species but as well as amphibian 237 \nspecies (Badli-Sham et al. 2023).  238 \nBukit Maras is a disturbed landscape dominated by secondary forest and agricultural areas. 239 \nHerpetofauna are sensitive to land-use changes (Sodhi et al. 2008). The ongoing orchard 240 \nexpansion threatens the discovery of new species and the persistence of elusive ones. Our 241 \nfindings, aligning with previous research, show that secondary forests have higher herpetofauna 242 \ndiversity compared to agricultural areas (Fig. 8). This difference is likely due to species loss in 243 \nagricultural areas, where suitable habitat is limited (Fig. 7). Forest-dwelling herpetofauna, known 244 \nfor their high site fidelity (Vitt and Caldwell 2001; Hillers et al. 2008), are particularly 245 \nvulnerable to local extirpation in such modified habitats. Studies in oil palm plantations 246 \n(Gillespie et al. 2012; Faruk et al. 2013) demonstrate a similar pattern, with generalist species 247 \ndominating disturbed areas. Our results in the agricultural areas of Bukit Maras reflect this trend. 248 \nWithout action to curb orchard expansion, biotic homogenization, where species diversity 249 \nreduced and dominated by only a few common species, is a looming threat. 250 \nThe presence of the critically endangered species, Manouria emys tortoise in the secondary forest 251 \nhighlights the high conservation value of Bukit Maras' remaining forests. This discovery 252 \nunderscores the urgency for immediate action by policymakers, local authorities, and the public 253 \nto protect these vital habitats. 254 \nGiven the widespread occurrence of secondary forests around human settlements, practical 255 \nconservation efforts should focus on preserving these areas and their riparian zones (Chazdon et 256 \nal. 2009; Pirnat and Hladnik 2016). While not a perfect substitute for primary forests, secondary 257 \nforests can support a significant diversity of herpetofauna (Thompson and Donnelly 2018). Their 258 \nmix of vegetation and microhabitats from both primary and disturbed forests creates a more 259 \nfavourable environment for herpetofauna compared to other human-modified landscapes (Luja et 260 \nal. 2008). Protecting these secondary forests can serve as a buffer zone, mitigating anthropogenic 261 \ndisturbances and safeguarding the remaining herpetofauna in Bukit Maras. 262 \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806\n\n24 \n \nConclusion 263 \nOur study helps fill a critical knowledge gap by investigating the herpetofauna of isolated hills 264 \nwithin Terengganu's human-modified landscape. This research focused on Bukit Maras, but 265 \nsimilar isolated hills in Terengganu, such as Besar Hill, Chendering Hill, and Jong Hill, warrant 266 \nfurther investigation using similar methods. Further surveys in Bukit Maras' northern region 267 \ncould reveal additional species. This study provides valuable baseline data for future monitoring 268 \nefforts to assess the impacts of environmental changes on herpetofauna in human-modified 269 \nlandscapes. 270 \nAcknowledgements 271 \nWe thank Universiti Malaysia Terengganu for the research equipment used during this study. 272 \nThe first author (MFS) is deeply grateful for the Tuanku Canselor Scholarship, generously 273 \nfunded by Universiti Malaysia Terengganu, which greatly supported first author throughout his 274 \nstudy. We also thank to all undergraduate students for their assistance in the field. The 275 \nDepartment of Wildlife and National Parks is dully acknowledged for permission to conduct this 276 \nstudy (Permit no.: T-00563-16-17). We thank the anonymous reviewer for their helpful 277 \ncomments. 278 \nAuthors’ Contributions 279 \nMFS collected the data, served as authority on species identification, provide photograph of the 280 \nspecimens, analyzed the data, wrote, and revised the manuscript. BHBS collected the data, 281 \nserved as authority on species identification, provide photograph of the specimens and 282 \nconstructed the map figure. SAF, MIMAW, MAN, FHA, GK, NNS, NFN, MS-R, JA, TKL, 283 \nNSX, MFA, SAR collected the data. 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Accessed on: 2024–03–11 447 \nVitt LJ, Caldwell JP (2001) The effects of logging on reptiles and amphibians of tropical forests. 448 \nIn: Fimbel, R.A., Grajal, A., Robinson, J. (Eds.), The Cutting Edge: Conserving Wildlife in 449 \nLogged Tropical Forests. Columbia University Press, New York, 239–259. 450 \nAppendix 451 \nTable A. List of voucher specimens from Bukit Maras catalogued with the UMTZC voucher 452 \ncode numbers. 453 \nVoucher No. (UMTZC) Species Name \n1701 Gonocephalus liogaster \n1705 Cyrtodactylus quadrivirgatus \n1706 Phrynoides aspera \n1772 Hylarana nicobarensis \n1773 Calotes versicolor \n1774 Eutropis multifasciata \n1882 Cyrtodactylus consobrinus \n1823 Hemiphyllodactylus typus \n1825 Limnonectes deinodon \n1828 Hylarana labialis \n1831 Pseudorhabdion longiceps \n 454 \nAuthor-formatted, not peer-reviewed document posted on 14/03/2024. DOI:  https://doi.org/10.3897/arphapreprints.e122806","source_license":"CC-BY-4.0","license_restricted":false}