Effect of canal blocking on biodiversity of degraded peatlands: Insight from West Kalimantan

story species (herb and woody lianas), trees, and canals only at the edge of the forest, suggests that disturbances have occurred some time ago and there have been no recent massive disturbances in disturbed peat forests [102]. These indications were confirmed during conversations with local people during our surveys. They stated that the forest is now under the management of the local forestry agency, with the primary concerns being forest fires, illegal wildlife hunting, and selective logging of certain tree species that have reached a certain age (usually the sapling stage) or a diameter of less than 10 cm. During our surveys, we neither observed nor heard of any actions regarding the replanting of vegetation due to legal restrictions imposed on disturbed forest areas, suggesting that any vegetation growth occurs naturally. Protection is the NCS strategies also needed by the disturbed peat forest, as these forests have the potential to regen- erate naturally and only require support to boost recovery. Research has shown that a peat forest in Central Kalimantan that experienced a single burning event in 1997 demonstrated similar Shannon diversity and plant species counts to unburned forests after 13 years [99]. Similarly, research from Central Kalimantan also demonstrates that peat forests sub- jected to selective logging in the past have a strong natural regeneration potential, shown by their high resemblance to rel- atively undisturbed forests in terms of tree density, species richness, and a similar understory vegetation community [103].

Additionally, if illegal hunting is curtailed, the forest could benefit from the presence of mammals and birds, which would serve as seed dispersers and pollinators to aid in the regeneration process [104]. Despite instances of illegal activity such as burn-clearing, logging, and hunting, disturbed peat forests in Mempawah Landscape still hold promise as potential

PLOS One | https://doi.org/10.1371/journal.pone.0334014 October 8, 2025 16 / 23 habitats, and, if left to regenerate naturally with interventions focused on illegal activities protection, they can progress into more mature forests with high biodiversity [105]. In estate crops where peatland has already been converted to oil palm plantations, the NCS hierarchy to support biodiversity conservation and increase carbon sequestration includes protection and improved management. Since the vegetation complexity of estate crops cannot provide suitable habitats for a wide range of wildlife species, protection from hunting—which still occurs in these areas—is crucial to maintain existing biodiversity. Hunting, especially targeting rare and endangered species, needs to be minimized as much as possible, as some wildlife have large area coverage, which is a consequence of establishing a plantation in a previously forested area [106]. Moreover, according to our results, one factor that supports estate crops as wildlife habitats is the presence of understory vegetation. The plantation management protocols in oil palm plantation, such as undergrowth clearing for fruit harvesting, road development, pest and weed man- agement, or tillage, can potentially eliminate understory coverage, impacting the biodiversity of the estate crop [107–109]. Maintaining understory coverage in estate crops can be considered an improved management practice. The presence of understory plants does not compete with oil palms; instead, it provides multiple benefits, such as reducing CO₂ emis-

sions, increasing carbon stocks, enhancing biodiversity, improving water and soil nutrient balances, and reducing erosion [110,111]. Furthermore, replacing the homogeneous structure with heterogeneity in oil palm plantations can enhance habitat quality to sustain forest-like areas [112]. Oil palm plantations in peatland areas are well-known actors in the cascading effects of habitat quality degradation, one of which is biodiversity loss. In addition, plantations can harbour species, usu- ally common species, which are sometimes considered pests and potentially invasive and in long-term could increase the wild crop-raiding population thus affecting the forest [113]. The estate is also not being used by forest-specific species as habitat, especially those that are peat forest specialists [109]. Adopting the jangka benah strategy—a solution to address unauthorized oil palm plantations in forest areas by implementing oil palm agroforestry—could be a suitable practice for estate crops to diversify vegetation [114]. Rather than expanding land area, intensification within existing areas will likely increase yield and optimize biodiversity conservation [115]. Participation in oil palm certifications (roundtable on sustainable palm oil (RSPO) or Indonesian sustainable palm oil (ISPO) by companies or smallholders is another form of improved management that can help prevent biodiversity loss, with recent reports indicating positive impacts of these certifications on biodiversity [116].

The disturbed wet shrub areas in the Mempawah Landscape, which are dominated by open areas, require more inten- sive recovery actions. For disturbed wet shrublands already used for agriculture, improved management practices need to be applied. An agroforestry system that enriches the land with woody plants offers an alternative for farmers to increase yield potential while contributing to biodiversity conservation and climate change mitigation. In addition to agricultural use, some disturbed wet shrub areas are left open. For these areas, protection should be prioritized, as fire regimes pose one of the greatest threats to dry peatlands, especially since canals have disrupted their hydrological systems [15,117]. In disturbed wet shrub areas that experience recurring fires, the third level of NCS, which is restoration, is the only way to recover the area. Repeated burning proved to prevent regeneration [101]. Studies show that a single forest fire can transform an initially forested area into an open area, but regeneration can occur relatively quickly due to the fire’s facilitation of seed sprouting for some species [99,100]. However, recurring fires can deplete soil nutrients, reduce soil fertility, diminish canopy, and ground cover, eliminate vegetation roots, and ultimately halt the process of plant regrowth and seed sprouting, resetting the succession to level zero [101,118,119] This chain reaction can significantly decrease biodiversity levels and increase the

dominance of specific plants, such as herbaceous ferns, in an area. This condition aligns with the observation results drawn from disturbed wet shrub [120]. With the worsening effects of rising global temperatures and El Niño periods, there is a drive to improve attitudes and strengthen restoration projects like never before [15,121,122]. Repeated fires are one of the factors halting open peatland revegetation. It is crucial to consider this when conducting restoration in vast peat wet shrublands.

PLOS One | https://doi.org/10.1371/journal.pone.0334014 October 8, 2025 17 / 23 With environmental deterioration from many aspects, restoration projects need to be implemented hand in hand [105]. On the rewetting sites, vegetation planting needs to incorporate local communities into action. This makes sense not only in terms of effective funding but also by providing benefits for the community, such as allowing them to select local trees that potentially increase their well-being through non-timber forest products or adopting responsible sustainable logging practices [105,123–125]. Involving local champions in building local nurseries for peatland revegetation and decision- making in peatland restoration projects can also increase the chance of disturbed wet shrub recovery. After all, the pres- ence of tree vegetation in the area can reduce the severity level of fire impacts, thus preventing the global hazard caused by massive landscape fire [118,126]. Supporting information S1 Fig. Boxplot showing acoustic indices from four habitats. (TIF) S2 Table. Acoustic index statistics across habitat types. (XLSX) S3 Table. Acoustic index statistics across habitat types and months. (XLSX) S4 Table. Habitat statistics. (XLSX) S5 Table. Temperature and humidity statistics across habitat types. (XLSX) S6 Table. Generalized Linear Model (GLM) of wildlife survey. (DOCX) S7 Table. Result of analysis of deviance table (Type II tests) for the GLM model for wildlife survey. (DOCX) S8 Table. Post hoc test using Tukey HSD for wildlife survey. (DOCX)

S9 Table. Analysis of deviance table (Table II Wald chi-square tests). (DOCX) S10 Table. Pairwise differences of acoustic indices in each habitat type. (DOCX) S11 Table. Summary result of Linear Mixed Model from acoustic indices. (DOCX) S12 Table. Number of individual and species for every growth stage in each habitat. (DOCX) S13 Fig. Boxplot showing vegetation features from four habitats. (TIF) S14 File. Inclusivity in global research questionnaire. (DOCX)

PLOS One | https://doi.org/10.1371/journal.pone.0334014 October 8, 2025 18 / 23 Acknowledgments We sincerely thank Rio Syahrudin, Muhammad Akmal Ramadhan, Clarra Malia Salsabila, Ignasius Suriyanto, Moham- mad Jakaria, Simon Petrus, and Setia Budi Gunawan for their assistance in data collection. We also appreciate the contributions of the local community members who helped us obtain the data and the village governments for granting us permission to conduct our research in their administrative areas. Thanks to our colleagues from the Forest Resource Con- servation Department, Faculty of Forestry, Universitas Gadjah Mada, for supporting during bioacoustics data analyses. Special thanks to The Nature Conservancy (TNC) for initiating this research and to Yayasan Konservasi Alam Nusantara (YKAN) for providing valuable preliminary information about the sites. Author contributions Conceptualization: Muhammad Ali Imron. Data curation: Muhammad Ali Imron, Farah Dini Rachmawati, Tungga Dewi, Giot Marganti Ito Simanullang. Formal analysis: Farah Dini Rachmawati, Tungga Dewi, Giot Marganti Ito Simanullang. Funding acquisition: Muhammad Ali Imron, Dennis Albihad, Erin E. Poor. Investigation: Muhammad Ali Imron, Dennis Albihad. Methodology: Muhammad Ali Imron, Farah Dini Rachmawati, Tungga Dewi, Erin E. Poor. Project administration: Muhammad Ali Imron, Farah Dini Rachmawati, Dennis Albihad. Resources: Giot Marganti Ito Simanullang. Supervision: Muhammad Ali Imron. Visualization: Tungga Dewi. Writing – original draft: Muhammad Ali Imron, Farah Dini Rachmawati, Tungga Dewi.

Writing – review & editing: Muhammad Ali Imron, Dennis Albihad, Erin E. Poor. References 1. Apers S, De Lannoy GJM, Baird AJ, Cobb AR, Dargie GC, Del Aguila Pasquel J, et al. Tropical peatland hydrology simulated with a global land surface model. J Adv Model Earth Syst. 2022;14(3):e2021MS002784. https://doi.org/10.1029/2021MS002784 PMID: 35860446 2. Xu J, Morris PJ, Liu J, Holden J. PEATMAP: Refining estimates of global peatland distribution based on a meta-analysis. Catena. 2018;160:134– 40. https://doi.org/10.1016/j.catena.2017.09.010 3. Ribeiro K, Pacheco FS, Ferreira JW, de Sousa-Neto ER, Hastie A, Krieger Filho GC, et al. Tropical peatlands and their contribution to the global carbon cycle and climate change. Glob Chang Biol. 2021;27(3):489–505. https://doi.org/10.1111/gcb.15408 PMID: 33070397 4. Osaki M, Setiadi B, Takahashi H, Evri M. Peatland in Kalimantan. In: Osaki M, Tsuji N, eds. Tropical peatland ecosystems. Springer Japan; 2016. 91–112. https://doi.org/10.1007/978-4-431-55681-7_6 5. Osaki M, Nursyamsi D, Noor M, Wahyunto, Segah H. Peatland in Indonesia. In: Osaki M, Tsuji N, eds. Tropical peatland ecosystems. Springer Japan; 2016. 49–58. https://doi.org/10.1007/978-4-431-55681-7_3 6. Dohong A, Abdul Aziz A, Dargusch P. A review of techniques for effective tropical peatland restoration. Wetlands. 2018;38(2):275–92. https://doi. org/10.1007/s13157-018-1017-6 7. Widyastuti K, Imron MA, Pradopo ST, Suryatmojo H, Sopha BM, Spessa A, et al. PeatFire: an agent-based model to simulate fire ignition and

spreading in a tropical peatland ecosystem. Int J Wildland Fire. 2021;30(2):71. https://doi.org/10.1071/wf19213 8. Harrison ME, Wijedasa LS, Cole LES, Cheyne SM, Choiruzzad SAB, Chua L, et al. Tropical peatlands and their conservation are important in the context of COVID-19 and potential future (zoonotic) disease pandemics. PeerJ. 2020;8:e10283. https://doi.org/10.7717/peerj.10283 PMID: 33240628 9. Yule CM. Loss of biodiversity and ecosystem functioning in Indo-Malayan peat swamp forests. Biodivers Conserv. 2008;19(2):393–409. https://doi. org/10.1007/s10531-008-9510-5 10. Amalia A. Ecological restoration in protected area of peat swamp forest as an effort to build socio-ecological resilience in Sebangau National Park Central Kalimantan, Indonesia. bwp. 2018;1(1):1–12. https://doi.org/10.47266/bwp.v1i1.5

PLOS One | https://doi.org/10.1371/journal.pone.0334014 October 8, 2025 19 / 23 11. Locky DA, Bayley SE. Effects of logging in the southern boreal peatlands of Manitoba, Canada. Can J For Res. 2007;37(3):649–61. https://doi. org/10.1139/x06-249 12. Wösten JHM, Clymans E, Page SE, Rieley JO, Limin SH. Peat–water interrelationships in a tropical peatland ecosystem in Southeast Asia. Cat- ena. 2008;73(2):212–24. https://doi.org/10.1016/j.catena.2007.07.010 13. Warren M, Hergoualc’h K, Kauffman JB, Murdiyarso D, Kolka R. An appraisal of Indonesia’s immense peat carbon stock using national peatland maps: uncertainties and potential losses from conversion. Carbon Balance Manag. 2017;12(1):12. https://doi.org/10.1186/s13021-017-0080-2 PMID: 28527145 14. Novita N, Subarno, Lestari NS, Anshari GZ, Lugina M, Yeo S, et al. Natural climate solutions in Indonesia: wetlands are the key to achieve Indone- sia’s national climate commitment. Environ Res Lett. 2022;17(11):114045. https://doi.org/10.1088/1748-9326/ac9e0a 15. Imron MA, Widyastuti K, Al Bihad D, Satria RA, Prayoga W, Pradopo ST, et al. Beyond climatic variation: human disturbances alter the effective- ness of a protected area to reduce fires in a Tropical Peatland. Front For Glob Change. 2022;5. https://doi.org/10.3389/ffgc.2022.788023 16. Widyastuti K, Reuillon R, Chapron P, Abdussalam W, Nasir D, Harrison ME, et al. Assessing the impact of forest structure disturbances on the arboreal movement and energetics of orangutans—An agent-based modeling approach. Front Ecol Evol. 2022;10. https://doi.org/10.3389/

fevo.2022.983337 17. Fujita MS, Samejima H, Haryadi DS, Muhammad A, Irham M, Shiodera S. Low conservation value of converted habitat for avifauna in tropical peatland on Sumatra, Indonesia. Ecol Res. 2016;31(2):275–85. https://doi.org/10.1007/s11284-016-1334-2 18. Syaufina L, Abi Hamzah A. Changes of tree species diversity in peatland impacted by moderate fire severity at Teluk Meranti, Pelalawan, Riau Province, Indonesia. Biodiversitas. 2021;22(5). https://doi.org/10.13057/biodiv/d220555 19. Harrison ME, Capilla BR, Thornton SA, Cattau ME, Page SE. Impacts of the 2015 fire season on peat-swamp forest biodiversity in Indonesian Borneo. In: 15th International Peat Congress. 2016. 713–7. 20. Rahajoe JS, Alhamd L, Atikah TD, Pratama BA, Shiodera S, Kohyama TS. Floristic diversity in the peatland ecosystems of Central Kalimantan. Tropical peatland ecosystems. Springer Japan; 2016. 167–96. https://doi.org/10.1007/978-4-431-55681-7_11 21. Harrison ME, Rieley JO. Tropical peatland biodiversity and conservation in southeast Asia: foreword. Mires and Peat. 2018;22:1–7. https://doi. org/10.19189/MaP.2018.OMB.382 22. Tawaraya K, Turjaman M. Mycorrhizal fungi in Peatland. In: Tropical peatland ecosystems. Springer Japan; 2016. 237–44. https://doi. org/10.1007/978-4-431-55681-7_15 23. Gumiri S, Ardianor, Higashi S, Iwakuma T. Aquatic communities in peatland of Central Kalimantan. In: Tropical peatland ecosystems. Springer Japan; 2016. 227–36. https://doi.org/10.1007/978-4-431-55681-7_14

24. Posa MRC, Wijedasa LS, Corlett RT. Biodiversity and Conservation of tropical peat swamp forests. BioScience. 2011;61(1):49–57. https://doi. org/10.1525/bio.2011.61.1.10 25. Husson SJ, Limin SH, Adul, Boyd NS, Brousseau JJ, Collier S. Biodiversity of the Sebangau tropical peat swamp forest, Indonesian Borneo. Mires and Peat. 2018;22(SI). https://doi.org/10.19189/MaP.2018.OMB.352 26. Bonn A, Allott T, Evans M, Joosten H, Stoneman R. Peatland restoration and ecosystem services. Cambridge University Press; 2016. https://doi. org/10.1017/cbo9781139177788 27. Page SE, Baird AJ. Peatlands and global change: response and resilience. Annu Rev Environ Resour. 2016;41(1):35–57. https://doi.org/10.1146/ annurev-environ-110615-085520 28. Giesen W, Nirmala E. Tropical peatland restoration report: the Indonesian case. Kemitraan Kesejahteraan Hijau (Kehijauan Berbak); 2018. https:// doi.org/10.13140/RG.2.2.30049.40808 29. Wösten H, Budiman A. Rewetting of drained indonesian peatlands to mitigate carbon dioxide emission. In: 14th International peat congress; peat- lands in balance. International Peat Society. 1928;205(5):724. https://doi.org/10.1016/s0016-0032(28)91004-2 30. Herawati H, Ali Akbar A, Farastika D, Azmeri. Water table evaluation post the construction of canal blocks on peatland in West Kalimantan, Indone- sia. MATEC Web Conf. 2018;195:03016. https://doi.org/10.1051/matecconf/201819503016 31. Ota T, Kusin K, Kilonzi FM, Usup A, Moji K, Kobayashi S. Sustainable financing for payment for ecosystem services (PES) to conserve peat swamp

forest through enterprises based on swiftlets’ nests: an awareness survey in Central Kalimantan, Indonesia. Small-scale Forestry. 2020;19(4):521– 39. https://doi.org/10.1007/s11842-020-09452-7 32. Page S, Hosciło A, Wösten H, Jauhiainen J, Silvius M, Rieley J, et al. Restoration ecology of lowland tropical peatlands in southeast asia: current knowledge and future research directions. Ecosystems. 2008;12(6):888–905. https://doi.org/10.1007/s10021-008-9216-2 33. Yuwati TW, Rachmanadi D, Pratiwi, Turjaman M, Indrajaya Y, Nugroho HYSH, et al. Restoration of degraded tropical peatland in indonesia: a review. Land. 2021;10(11):1170. https://doi.org/10.3390/land10111170 34. Harrison ME, Ottay JB, D’Arcy LJ, Cheyne SM, Anggodo, Belcher C, et al. Tropical forest and peatland conservation in Indonesia: challenges and directions. People Nat. 2019;2(1):4–28. https://doi.org/10.1002/pan3.10060 35. BRG. Three years of peatland restoration in Indonesia. 2019. https://www.scribd.com/document/426381308/Buku-3-tahun-BRG-pdf 36. Harrison ME, Capilla BR, Thornton SA, Cattau ME, Page SE. Impacts of the 2015 fire season on peat-swamp forest biodiversity in Indonesian Borneo. In: 15th International Peat Congress. 2016. 713–7.

PLOS One | https://doi.org/10.1371/journal.pone.0334014 October 8, 2025 20 / 23 37. Thornton SA, Dudin, Page SE, Upton C, Harrison ME. Peatland fish of Sebangau, Borneo: diversity, monitoring and conservation. Mires Peat. 2018;22(Figure 1):1–25. https://doi.org/10.19189/MaP.2017.OMB.313 38. Erb WM, Barrow EJ, Hofner AN, Utami-Atmoko SS, Vogel ER. Wildfire smoke impacts activity and energetics of wild Bornean orangutans. Sci Rep. 2018;8(1):7606. https://doi.org/10.1038/s41598-018-25847-1 PMID: 29765067 39. Santosa Y, Yudea Utami C. Is it true that forest and land fires caused the extinction of biodiversity…?. IOP Conf Ser: Earth Environ Sci. 2020;528(1):012021. https://doi.org/10.1088/1755-1315/528/1/012021 40. Sanderfoot OV, Bassing SB, Brusa JL, Emmet RL, Gillman SJ, Swift K, et al. A review of the effects of wildfire smoke on the health and behavior of wildlife. Environ Res Lett. 2021;16(12):123003. https://doi.org/10.1088/1748-9326/ac30f6 41. Masy’ud B, Santosa Y, Aulia N, Gusnadi FD. The impact of fire in palm oil estate on the bird and butterfly species diversity: case study in RAJ Oil Palm Estate, South Sumatera. IOP Conf Ser: Earth Environ Sci. 2020;504(1):012002. https://doi.org/10.1088/1755-1315/504/1/012002 42. Slik JWF, Van Balen S. Bird community changes in response to single and repeated fires in a lowland tropical rainforest of eastern Borneo. Biodiv- ers Conserv. 2006;15(14):4425–51. https://doi.org/10.1007/s10531-005-4385-1

43. Printing organic electronics on flexible substrates. In: Conjugated polymers. CRC Press; 2006. 187–212. https://doi.org/10.1201/b10739-10 44. Santosa Y, Kwatrina RT. Effect of forest fire on mammals: comparisons of species diversity on different time-period and area. IOP Conf Ser: Earth Environ Sci. 2020;528(1):012023. https://doi.org/10.1088/1755-1315/528/1/012023 45. Meijaard E, Sheil D, Nasi R, Augeri D, Rosenbaum B, Iskandar D. Life after logging: reconciling wildlife conservation and production forestry in Indonesian Borneo. Bogor, Indonesia. 2005. 46. Blackham GV, Andri Thomas, Webb EL, Corlett RT. Seed rain into a degraded tropical peatland in Central Kalimantan, Indonesia. Biol Conserv. 2013;167:215–23. https://doi.org/10.1016/j.biocon.2013.08.015 47. Blackham GV, Webb EL, Corlett RT. Natural regeneration in a degraded tropical peatland, Central Kalimantan, Indonesia: Implications for forest restoration. For Ecol Manag. 2014;324:8–15. https://doi.org/10.1016/j.foreco.2014.03.041 48. Blackham GV, Corlett RT. Post-dispersal seed removal by ground-feeding rodents in tropical peatlands, Central Kalimantan, Indonesia. Sci Rep. 2015;5:14152. https://doi.org/10.1038/srep14152 PMID: 26369444 49. Lestari DP, Darusman T, Harsanto F, Arriyadi D, Ginanjar G. Understanding natural regeneration in burned tropical peatland: a strategy to acceler- ate the forest recovery process. Biotropia. 2022;28(3):204–13. https://doi.org/10.11598/btb.2021.28.3.1330

50. Alcocer I, Lima H, Sugai LSM, Llusia D. Acoustic indices as proxies for biodiversity: a meta-analysis. Biol Rev Camb Philos Soc. 2022;97(6):2209– 36. https://doi.org/10.1111/brv.12890 PMID: 35978471 51. Nabila RA, Imron MA. Deteksi vokalisasi gajah sumatra (Elephas maximus sumatranus) menggunakan passive acoustic monitoring di resort Tangkahan, Taman Nasional Gunung Leuser. Faculty of Forestry, Universitas Gadjah Mada; 2022. https://etd.repository.ugm.ac.id/penelitian/ detail/214159 52. Sueur J, Pavoine S, Hamerlynck O, Duvail S. Rapid acoustic survey for biodiversity appraisal. PLoS One. 2008;3(12):e4065. https://doi. org/10.1371/journal.pone.0004065 PMID: 19115006 53. John F, Sanford W, Brad P, Daniel A, Douglas B, Gabriel BB. Comparison to applied regression. 2022. https://r-forge.r-project.org/projects/car/ 54. Russell VL, Ben B, Paul B, Iago GV, Maxime H, Maarten J. Estimated marginal means, aka least-squares mean. 2023. https://rvlenth.github.io/ emmeans/ 55. Jari O, Gavin LS, Guillaume F, Roeland K, Pierre L, Peter RM. Community ecology package. https://github.com/vegandevs/vegan 56. Benedek AM, Sîrbu I, Lazăr A. Responses of small mammals to habitat characteristics in Southern Carpathian forests. Sci Rep. 2021;11(1):12031. https://doi.org/10.1038/s41598-021-91488-6 PMID: 34103594 57. Berriozabal-Islas C, Badillo-Saldaña LM, Ramírez-Bautista A, Moreno CE. Effects of habitat disturbance on lizard functional diversity in a tropical dry forest of the pacific coast of Mexico. Trop Conserv Sci. 2017;10. https://doi.org/10.1177/1940082917704972

58. Shilereyo MT, Magige FJ, Ogutu JO, Røskaft E. Small-mammal abundance and species diversity: land use and seasonal influences in the Seren- geti Ecosystem, Tanzania. Front Conserv Sci. 2023;4. https://doi.org/10.3389/fcosc.2023.981424 59. Pekin BK, Jung J, Villanueva-Rivera LJ, Pijanowski BC, Ahumada JA. Modeling acoustic diversity using soundscape recordings and LIDAR-derived metrics of vertical forest structure in a neotropical rainforest. Landscape Ecol. 2012;27(10):1513–22. https://doi.org/10.1007/s10980-012-9806-4 60. Ariano-Sánchez D, Mortensen RM, Reinhardt S, Rosell F. Escaping drought: Seasonality effects on home range, movement patterns and habitat selection of the Guatemalan Beaded Lizard. Global Ecol Conserv. 2020;23:e01178. https://doi.org/10.1016/j.gecco.2020.e01178 61. Papageorgiou D, Rozen-Rechels D, Nyaguthii B, Farine DR. Seasonality impacts collective movements in a wild group-living bird. Mov Ecol. 2021;9(1):38. https://doi.org/10.1186/s40462-021-00271-9 PMID: 34238382 62. Short JC. Density and seasonal movements of forest elephant (Loxodonta africana cyclotis, Matschie) in Bia National Park, Ghana. Afr J Ecol. 1983;21(3):175–84. https://doi.org/10.1111/j.1365-2028.1983.tb01179.x 63. Hou Y, Zhang M, Meng Z, Liu S, Sun P, Yang T. Assessing the impact of forest change and climate variability on dry season runoff by an improved single watershed approach: a comparative study in two large Watersheds, China. Forests. 2018;9(1):46. https://doi.org/10.3390/f9010046

64. Page SE, Rieley JO, Wüst R. Chapter 7 Lowland tropical peatlands of Southeast Asia. In: Developments in earth surface processes. Elsevier; 2006. 145–72. https://doi.org/10.1016/s0928-2025(06)09007-9

PLOS One | https://doi.org/10.1371/journal.pone.0334014 October 8, 2025 21 / 23 65. Hodgkins SB, Richardson CJ, Dommain R, Wang H, Glaser PH, Verbeke B, et al. Tropical peatland carbon storage linked to global latitudinal trends in peat recalcitrance. Nat Commun. 2018;9(1):3640. https://doi.org/10.1038/s41467-018-06050-2 PMID: 30194308 66. Wong GX, Hirata R, Hirano T, Kiew F, Aeries EB, Musin KK, et al. Micrometeorological measurement of methane flux above a tropical peat swamp forest. Agricul Forest Meteorol. 2018;256–257:353–61. https://doi.org/10.1016/j.agrformet.2018.03.025 67. Spooner FEB, Pearson RG, Freeman R. Rapid warming is associated with population decline among terrestrial birds and mammals globally. Glob Chang Biol. 2018;24(10):4521–31. https://doi.org/10.1111/gcb.14361 PMID: 30033551 68. Ficetola GF, Maiorano L. Contrasting effects of temperature and precipitation change on amphibian phenology, abundance and performance. Oecologia. 2016;181(3):683–93. https://doi.org/10.1007/s00442-016-3610-9 PMID: 27008454 69. Blaustein AR, Walls SC, Bancroft BA, Lawler JJ, Searle CL, Gervasi SS. Direct and indirect effects of climate change on amphibian populations. Diversity. 2010;2(2):281–313. https://doi.org/10.3390/d2020281 70. Inkley DB, Anderson MG, Blaustein AR, Burkett VR, Felzer B, Griffith B, et al. Global climate change and wildlife in North America. Wildlife Soc Tech Rev. 2004;1–26. 71. Porter WP, Budaraju S, Stewart WE, Ramankutty N. Calculating climate effects on birds and mammals: impacts on biodiversity, conservation,

population parameters, and global community structure. Am Zool. 2000;40(4):597–630. https://doi.org/10.1093/icb/40.4.597 72. Hanley TA. Potential management of young-growth stands for understory vegetation and wildlife habitat in southeastern Alaska. Landscape Urban Plan. 2005;72(1–3):95–112. https://doi.org/10.1016/j.landurbplan.2004.09.015 73. Paoletti A, Darras K, Jayanto H, Grass I, Kusrini M, Tscharntke T. Amphibian and reptile communities of upland and riparian sites across Indone- sian oil palm, rubber and forest. Global Ecol Conserv. 2018;16:e00492. https://doi.org/10.1016/j.gecco.2018.e00492 74. Ari Janiawati IA, Kusrini MD, Mardiastuti A. Structure and composition of reptile communities in human modified landscape in Gianyar Regency, Bali. Hayati J Biosci. 2016;23(2):85–91. https://doi.org/10.1016/j.hjb.2016.06.006 75. Pilliod DS, Bury RB, Hyde EJ, Pearl CA, Corn PS. Fire and amphibians in North America. Forest Ecol Manag. 2003;178(1–2):163–81. https://doi. org/10.1016/s0378-1127(03)00060-4 76. Earl JE, Harper EB, Hocking DJ, Osbourn MS, Rittenhouse TAG, Glennie M, et al. Relative importance of timber harvest and habitat for reptiles in experimental forestry plots. Forest Ecol Manag. 2017;402:21–8. https://doi.org/10.1016/j.foreco.2017.07.014 77. Simonetti JA, Grez AA, Estades CF. Providing habitat for native mammals through understory enhancement in forestry plantations. Conserv Biol. 2013;27(5):1117–21. https://doi.org/10.1111/cobi.12129 PMID: 24033701

78. McCarthy BC, Magurran AE. Measuring biological diversity. J Torrey Bot Soc. 2004;131(3):277. https://doi.org/10.2307/4126959 79. Wee YC. Observations on the behaviour of the yellow-vented bulbul, Pycnonotus goiavier (Scopoli) in two instances of failed nesting. Nat Sing. 2009;2(August):347–52. 80. Amit B, Tuen AA, Haron K, Harun MH, Kamarudin N. The diet of yellow-vented bulbul (pycnonotus goiavier) in oil palm agroecosystems. J Oil Palm Res. 2015;27(4):417–24. 81. Scarpelli MDA, Liquet B, Tucker D, Fuller S, Roe P. Multi-index ecoacoustics analysis for terrestrial soundscapes: a new semi-automated approach using time-series motif discovery and random forest classification. Front Ecol Evol. 2021;9. https://doi.org/10.3389/fevo.2021.738537 82. Sousa-Lima RS, Ferreira LM, Oliveira EG, Lopes LC, Brito MR, Baumgarten J, et al. What do insects, anurans, birds, and mammals have to say about soundscape indices in a tropical savanna. J Eco Acous. 2018;2(1):1–1. https://doi.org/10.22261/jea.pvh6yz 83. Metcalf OC, Barlow J, Devenish C, Marsden S, Berenguer E, Lees AC. Acoustic indices perform better when applied at ecologically meaningful time and frequency scales. Methods Ecol Evol. 2020;12(3):421–31. https://doi.org/10.1111/2041-210x.13521 84. Chen Y-F, Luo Y, Mammides C, Cao K-F, Zhu S, Goodale E. The relationship between acoustic indices, elevation, and vegetation, in a forest plot network of southern China. Ecol Indic. 2021;129:107942. https://doi.org/10.1016/j.ecolind.2021.107942

85. Zwerts JA, Stephenson PJ, Maisels F, Rowcliffe M, Astaras C, Jansen PA, et al. Methods for wildlife monitoring in tropical forests: comparing human observations, camera traps, and passive acoustic sensors. Conservat Sci Prac. 2021;3(12). https://doi.org/10.1111/csp2.568 86. Chandra Kasih R, Simon O, Ansori M, Porkab Pratama M, Wirada F. Rewetting of degraded tropical peatland by canal blocking technique in Sebangau National Park, Central Kalimantan, Indonesia. In: 15th International Peat Congress. 2016. 467–71. 87. Karimi S, Maher Hasselquist E, Salimi S, Järveoja J, Laudon H. Rewetting impact on the hydrological function of a drained peatland in the boreal landscape. J Hydrol. 2024;641:131729. https://doi.org/10.1016/j.jhydrol.2024.131729 88. Castro M, Novita N, Ritonga R, Erbaugh J, Yeo S, Masuda Y. Has peat rewetting reduced and prevented fires in West and Central Kalimantan?. 2023. https://meetingorganizer.copernicus.org/EGU23/EGU23-10738.html 89. Sutikno S, Rinaldi R, Saputra E, Kusairi M, Saharjo BH, Putra EI. Water management for hydrological restoration and fire prevention in tropical peatland. IOP Conf Ser: Mater Sci Eng. 2020;933(1):012053. https://doi.org/10.1088/1757-899x/933/1/012053 90. Hooijer A, Vernimmen R, Mulyadi D, Triantomo V, Hamdani, Lampela M, et al. Benefits of tropical peatland rewetting for subsidence reduction and forest regrowth: results from a large-scale restoration trial. Sci Rep. 2024;14(1):10721. https://doi.org/10.1038/s41598-024-60462-3 PMID: 38729962

91. Jati AS, Samejima H, Fujiki S, Kurniawan Y, Aoyagi R, Kitayama K. Effects of logging on wildlife communities in certified tropical rainforests in East Kalimantan, Indonesia. Forest Ecol Manag. 2018;427:124–34. https://doi.org/10.1016/j.foreco.2018.05.054

PLOS One | https://doi.org/10.1371/journal.pone.0334014 October 8, 2025 22 / 23 92. Rahman DA, Gonzalez G, Haryono M, Muhtarom A, Firdaus AY, Aulagnier S. Factors affecting seasonal habitat use, and predicted range of two tropical deer in Indonesian rainforest. Acta Oecologica. 2017;82:41–51. https://doi.org/10.1016/j.actao.2017.05.008 93. Rahman DA, Santosa Y, Purnamasari I, Condro AA. Drivers of three most charismatic mammalian species distribution across a multiple-use tropical forest landscape of Sumatra, Indonesia. Animals (Basel). 2022;12(19):2722. https://doi.org/10.3390/ani12192722 PMID: 36230461 94. Ritzema H, Limin S, Kusin K, Jauhiainen J, Wösten H. Canal blocking strategies for hydrological restoration of degraded tropical peatlands in Central Kalimantan, Indonesia. Catena. 2014;114:11–20. https://doi.org/10.1016/j.catena.2013.10.009 95. Page S, Hosciło A, Wösten H, Jauhiainen J, Silvius M, Rieley J, et al. Restoration ecology of lowland tropical peatlands in Southeast Asia: current knowledge and future research directions. Ecosystems. 2008;12(6):888–905. https://doi.org/10.1007/s10021-008-9216-2 96. Ellis PW, Page AM, Wood S, Fargione J, Masuda YJ, Carrasco Denney V, et al. The principles of natural climate solutions. Nat Commun. 2024;15(1):547. https://doi.org/10.1038/s41467-023-44425-2 PMID: 38263156 97. Cook-Patton SC, Drever CR, Griscom BW, Hamrick K, Hardman H, Kroeger T, et al. Protect, manage and then restore lands for climate mitiga- tion. Nat Clim Chang. 2021;11(12):1027–34. https://doi.org/10.1038/s41558-021-01198-0

98. Siregar M, Sambas EN. Floristic composition of peat swamp forest in mensemat-sambas, west kalimantan. In: Proceedings of the International Symposium on TROPICAL PEATLANDS. 2000. 153–64. 99. Shiodera S, Atikah TD, Apandi I, Seino T, Haraguchi A, Rahajoe JS, et al. Peat-fire impact on forest structure in peatland of central kalimantan. In: Osaki M, Tsuji N, eds. Tropical peatland ecosystems. Springer Japan; 2016. 197–212. https://doi.org/10.1007/978-4-431-55681-7_12 100. Blackham GV, Webb EL, Corlett RT. Natural regeneration in a degraded tropical peatland, Central Kalimantan, Indonesia: implications for forest restoration. Forest Ecol Manag. 2014;324:8–15. https://doi.org/10.1016/j.foreco.2014.03.041 101. Page S, Hosciło A, Wösten H, Jauhiainen J, Silvius M, Rieley J, et al. Restoration ecology of lowland tropical peatlands in Southeast Asia: current knowledge and future research directions. Ecosystems. 2008;12(6):888–905. https://doi.org/10.1007/s10021-008-9216-2 102. Kobayashi S. Initial phase of secondary succession in the exploited peat swamp forest (Shorea albida) at Sungai Damit, Belait in Brunei Darus- salam. In: Proceedings of the International symposium on TROPICAL PEATLANDS. 2000. 205–14. 103. Healy W. How degradation type impacts natural regeneration in a tropical peat-swamp forest. University of Exeter; 2020. 104. Wijedasa LS, Vernimmen R, Page SE, Mulyadi D, Bahri S, Randi A, et al. Distance to forest, mammal and bird dispersal drive natural regenera-

sity and decomposition rates. Front For Glob Change. 2018;1. https://doi.org/10.3389/ffgc.2018.00010 112. Budiadi, Susanti A, Marhaento H, Ali Imron M, Permadi DB, Hermudananto H. Oil palm agroforestry: an alternative to enhance farmers’ livelihood resilience. IOP Conf Ser: Earth Environ Sci. 2019;336(1):012001. https://doi.org/10.1088/1755-1315/336/1/012001 113. Luskin MS, Brashares JS, Ickes K, Sun I-F, Fletcher C, Wright SJ, et al. Cross-boundary subsidy cascades from oil palm degrade distant tropical forests. Nat Commun. 2017;8(1):2231. https://doi.org/10.1038/s41467-017-01920-7 PMID: 29263381 114. Susanti A. Jangka benah: alternatif solusi persoalan keterlanjuran kebun kelapa sawit monokultur di kawasan hutan. Jur Ilm Kehut. 2021;15(1):1– 3. https://doi.org/10.22146/jik.v15i1.1566 115. Phalan B, Onial M, Balmford A, Green RE. Reconciling food production and biodiversity conservation: land sharing and land sparing compared. Science. 2011;333(6047):1289–91. https://doi.org/10.1126/science.1208742 PMID: 21885781 116. Hidayat NK, Offermans A, Glasbergen P. Sustainable palm oil as a public responsibility? On the governance capacity of Indonesian standard for sustainable palm oil (ISPO). Agric Hum Values. 2017;35(1):223–42. https://doi.org/10.1007/s10460-017-9816-6 117. Wösten JHM, Clymans E, Page SE, Rieley JO, Limin SH. Peat–water interrelationships in a tropical peatland ecosystem in Southeast Asia. Cat- ena. 2008;73(2):212–24. https://doi.org/10.1016/j.catena.2007.07.010

118. Agus C, Azmi FF, Widiyatno, Ilfana ZR, Wulandari D, Rachmanadi D, et al. The impact of forest fire on the biodiversity and the soil characteristics of tropical peatland. In: Climate change management. Springer International Publishing; 2018. 287–303. https://doi. org/10.1007/978-3-319-98681-4_18

PLOS One | https://doi.org/10.1371/journal.pone.0334014 October 8, 2025 23 / 23 119. Nurulita Y, Adetutu EM, Gunawan H, Zul D, Ball AS. Restoration of tropical peat soils: the application of soil microbiology for monitoring the suc- cess of the restoration process. Agricul Ecosyst Environ. 2016;216:293–303. https://doi.org/10.1016/j.agee.2015.09.031 120. Hansson A, Dargusch P. An estimate of the financial cost of peatland restoration in Indonesia. Case Stud Environ. 2018;2(1):1–8. https://doi. org/10.1525/cse.2017.000695 121. Hayasaka H. Peatland fire weather conditions in Sumatra, Indonesia. Climate. 2023;11(5):92. https://doi.org/10.3390/cli11050092 122. Sulaiman A, Osaki M, Takahashi H, Yamanaka MD, Susanto RD, Shimada S, et al. Peatland groundwater level in the Indonesian maritime continent as an alert for El Niño and moderate positive Indian Ocean dipole events. Sci Rep. 2023;13(1):939. https://doi.org/10.1038/s41598-023- 27393-x PMID: 36653400 123. Austin KG, Mosnier A, Pirker J, McCallum I, Fritz S, Kasibhatla PS. Shifting patterns of oil palm driven deforestation in Indonesia and implications for zero-deforestation commitments. Land Use Policy. 2017;69:41–8. https://doi.org/10.1016/j.landusepol.2017.08.036 124. Tata HL. Genetic diversity in peatland restoration: a case of Jelutung. In: Mizuno K, Kozan O, Gunawan H, eds. Vulnerability and transformation of Indonesian peatlands. 1st ed. Springer Nature Singapore; 2023. 185–96.

125. Saito H, Koizumi A, Gaman S, Yuda P, Penyang, Shibuya M. Tropical peatland forestry: toward forest restoration and sustainable use of wood resources in degraded peatland. In: Osaki M, Tsuji N, eds. Tropical peatland ecosystems. Springer Japan; 2016. 513–49. https://doi. org/10.1007/978-4-431-55681-7_35 126. Azhar B, Saadun N, Prideaux M, Lindenmayer DB. The global palm oil sector must change to save biodiversity and improve food security in the tropics. J Environ Manage. 2017;203(Pt 1):457–66. https://doi.org/10.1016/j.jenvman.2017.08.021 PMID: 28837912