Decomposition Dynamics of an Orangutan (Pongo pygmaeus morio) Carcass in a Tropical Forest: Implications for Conservation Practices

Decomposition Dynamics of an Orangutan (Pongo pygmaeus morio) Carcass in a Tropical Forest: Implications for Conservation Practices | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL Ecology and Evolution This is a preprint and has not been peer reviewed. Data may be preliminary. 30 July 2025 V1 Latest version Share on Decomposition Dynamics of an Orangutan (Pongo pygmaeus morio) Carcass in a Tropical Forest: Implications for Conservation Practices Authors : Sui Peng Heon 0000-0002-4075-737X [email protected] , Henry Bernard , and Robert Ewers Authors Info & Affiliations https://doi.org/10.22541/au.175387568.83231838/v1 Published Ecology and Evolution Version of record Peer review timeline 394 views 197 downloads Contents Abstract Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract Over the past decade, more than 600 rehabilitated Bornean Orangutans (Pongo pygmaeus morio) have been released into protected forests in Borneo. Releasing rehabilitant Bornean Orangutans into the wild is a standard conservation practice, yet monitoring post-release survival remains a challenge. Limited data exist on post-release survival, with many individuals classified as “missing but presumed dead” due to the absence of a carcass for confirmation. Detecting carcasses in tropical forests is particularly difficult due to dense vegetation and the narrow time frame for observing remains before complete decomposition or scavenger removal. Here, we report the first documented observation of an adult female Bornean Orangutan carcass decomposing process in the Danum Valley Conservation Area, Malaysian Borneo, on 21 May 2023. The approximately 30 kg carcass was monitored using camera traps and field observations. Decomposition was assessed using Payne’s (1965) decomposition framework, the Total Body Score (TBS) system, and Accumulated Degree Days (ADD) to evaluate the influence of ambient temperature on decay. Decomposition progressed to the dry-remains stage within six days, primarily driven by vertebrate scavengers such as the Asian water monitor lizard (Varanus salvator) and Blow flies (Calliphoridae spp). This rapid decomposition rate challenges existing knowledge on the rate of decomposition of medium-sized carcasses (>10 kg) and suggests that the common practice of weekly monitoring for post-release Orangutans may be insufficient. Understanding decomposition processes and scavenger activity in tropical forests can improve carcass detection, refine mortality estimates for released Orangutans and other endangered species, and enhance conservation strategies for this critically endangered primate. Introduction Translocation and reintroduction are key conservation strategies for the critically endangered Bornean Orangutan ( Pongo pygmaeus ), widely implemented in Malaysia and Indonesia (Meijaard et al., 2012; Sherman et al., 2025). These great apes face severe threats, including habitat loss, illegal pet trade, hunting, and forest fragmentation (Meijaard et al., 2012). A common conservation practice involves rehabilitating displaced and confiscated Orangutans in rescue facilities before releasing them into protected forests to support population recovery. Best-practice guidelines for wildlife rescue programs emphasize post-release monitoring to assess survival and adaptation (IUCN, 2013). However, tracking released Orangutans remains challenging. Many individuals are fitted with implanted radio transmitters, yet detection rates are low, with 40–95% never re-encountered (Sherman et al., 2021). Reported survival estimates vary from 6% to 80%, but reliable long-term records are lacking (Meijaard et al., 2012; Sherman et al., 2021). Some centres conduct intensive tracking for a few weeks before shifting to infrequent surveys, while others lack monitoring programs altogether (Sherman et al., 2020). The difficulty in confirming mortality further complicates survival estimates, as carcasses are rarely recovered. For example, in one of the rescue centres, 66% (25 of 38) of radio-collared Orangutans were unaccounted for three years post-release and presumed dead, despite no carcass sightings (Sherman et al., 2020). One explanation for the lack of carcass recoveries—beyond predation—is the rapid action of scavengers and decomposition in tropical forests. Globally, vertebrates remove up to 75% of carcasses within a short time (DeVault et al., 2003). In Borneo, Malay civets ( Viverra tangalunga ) and Asian water monitors ( Varanus salvator ) are key vertebrate scavengers (Lim et al., 2015; Twinning et al., 2017), while Calliphoridae blowflies are dominant invertebrate scavengers. Carcasses that escape vertebrate removal undergo six decomposition stages, from fresh to skeletal remains, driven by microbial and invertebrate activity (Payne, 1965). This framework is widely used in carcass ecology and forensic studies, yet scavenging dynamics in tropical forests remain poorly documented. Fewer than 20 Southeast Asian studies have examined carcass decomposition in tropical forests, with most focused on forensic applications, excluding vertebrate scavengers or examined only large-bodied scavengers while neglecting invertebrate activity. To our knowledge, no tropical study has simultaneously examined the roles of both vertebrate and invertebrate scavengers. To date, only one report has documented the decomposition of a wild Orangutan carcass by a forest scavenger, the Bearded pig ( Sus barbatus ; Galdikas, 1978). In our study, we document the complete decomposition process of a wild Orangutan carcass in a tropical forest. Although our observations are limited to a single individual, the opportunity to record such a process is rare – just two carcasses were observed in more than 8,000 hours of field observation over more than five years (Galdikas, 1978) – and holds significant ecological value. Experimental studies on the decomposition of wild animal carcasses, particularly of critically endangered species, are neither ethical nor feasible. Taking advantage of this unique event, our study addresses two key questions: (1) Which scavenger species interacted with the carcass? and (2) What was the rate of decomposition of the Orangutan carcass? We then (3) compare that rate to other published and unpublished reports and data to assess the extent to which our single observation might be indicative of decomposition processes more generally. Understanding these processes can improve post-release mortality assessments and enhance Orangutan conservation strategies, while also contributing new insights into carcass removal and scavenger dynamics in tropical ecosystems. Methods This study was conducted in primary lowland Dipterocarp forest at Danum Valley Conservation Area (4.96474 N, 117.80385 E), Sabah, Malaysian Borneo, in March 2023. On May 21, a known adult female Bornean Orangutan ( Pongo pygmaeus ) was found dead ~120 m from the field centre after distress calls from her infant were heard. The cause of death was unknown but presumed to be illness or senescence, as she had been observed spending unusual time on the forest floor prior to death (J. Anson, pers. comm.). She was last seen alive on May 20 with no visible injuries. The site was closed to visitors within 24 hours due to strong putrefaction odour. A Reconyx HC500 motion-triggered camera (Reconyx, USA) was deployed at the carcass, set to five-image bursts every 15 minutes and upon motion. Independent scavenger visits were defined as detections >30 minutes apart. On Day 4, the camera was repositioned ~2 m after an Asian water monitor ( Varanus salvator ) moved the carcass. A phone camera (Samsung, South Korea) supplemented monitoring in later stages when the limbs detached. The observer conducted daily visits to assess carcass condition and observe invertebrate scavenger activity, using randomized varying times to reduce disturbance. We assessed decomposition using two standard methods. First, we qualitatively recorded decomposition stages following Payne’s (1965) framework, as weighing the carcass was not feasible. These stages were compared to published decomposition studies in tropical forests. Second, we applied the Total Body Score (TBS) system and Accumulated Degree Days (ADD) method (Megyesi et al., 2005) to quantify decomposition relative to temperature. The TBS was calculated daily based on the head/neck, trunk, and limbs. After scavenger removal of the limbs, limb scores (10 points) were excluded, reducing the maximum TBS from 35 to 25. Camera trap images were used to validate decomposition stage classification (Wilson et al., 2019). We also obtained temperature data from the Danum Weather Station (O’Brien et al., 2025) to examine whether the rate of decomposition was comparable to that observed of 15 g small mammal in the same forest (Heon et al., 2025). We calculated Accumulated Degree Days (ADD) for each day of decomposition as follows: ADD = ∑ (T avg – T base ), where T avg = (T max + T min )/2. We set T bas e at 9.5°C, the minimum developmental threshold for Chrysomya rufifacies (Diptera: Calliphoridae), the dominant dipteran scavenger in Malaysian rainforests (Ahmad & Omar, 2018; Yanmanee et al., 2016). T max is the maximum temperature of the day while T min was the minimum temperature of the same day. Results The Orangutan carcass progressed through all stages of decomposition—from fresh to dry —in just six days (Figure 1). This decomposition rate was notably faster than that observed in other taxa in the region, despite the Orangutan’s relatively larger body size (Figure 2A). The carcass rapidly transitioned through the bloat (Day 1–2) and active decay stages (Day 2–3), whereas in other studies, these stages lasted longer. This accelerated decomposition appeared to be facilitated by the combined activity of both vertebrate and invertebrate scavengers, in contrast to other taxa where invertebrates dominated the process. The first scavengers to arrive were blowflies ( Chrysomya rufifacies and Chrysomya megacephala ), which were dominant from Day 1 to Day 3, corresponding to the fresh and active decay stages. By Day 3, dipteran larvae had consumed much of the soft tissue, with the Orangutan’s skull already exposed. Time-lapse footage revealed that bloat occurred twice during the early decomposition phase—specifically in the late evening and early morning—likely due to gas accumulation and subsequent rupture (Figure 1). The first vertebrate scavenger was detected only after the bloat stage, likely responding to the release of putrescent gases during carcass deflation. Camera traps recorded three vertebrate scavenger species. The Asian water monitor ( Varanus salvator ) appeared seven times, repeatedly visiting the carcass during the active and advanced decay stages. While we were unable to quantify the biomass removed by each scavenger—since weighing the carcass in situ would have disrupted invertebrate activity—camera observations showed two water monitors dismembering and carrying off the Orangutan’s limbs. The remaining trunk and head region decomposed in place, primarily via dipteran larval activity. Predatory staphylinid beetles ( Staphylinidae ) were observed feeding on the fly larvae during this stage. A Malay civet ( Viverra tangalunga ) was also observed on Day 2, engaging in scent-rubbing behaviour—likely a form of territorial or social communication—but it did not return in subsequent days. By Day 4, fly pupae were visible on the Orangutan’s hair and carcass. On Day 5, some pupae had dropped to the surrounding soil, and by Day 6, the carcass had been reduced to bone fragments, with no further visible scavenger or invertebrate activity. Three years after decomposition, the bones were distributed across an area of roughly 10 m 2 and remained largely undisturbed. The skull was still visible, while other bone fragments were partially buried under dense leaf litter. There were no signs of bone gnawing, but algal growth and tiny cavities, presumably created by termites, suggested post-decomposition decay processes were underway. Our analysis of the relationship between time and decomposition phases (Figure 2B) showed that both the Orangutan and small mammal carcass (Heon et al., 2025) reached their maximum Total Body Score (TBS)—25 for the Orangutan and 35 for the small mammal—within a range of 100–140 accumulated degree days (ADD). The most rapid decomposition occurred within the first 100 °C days, after which the rate notably decreased. Figure 1. Decomposition stages of a wild Orangutan carcass: (A) Fresh stage: Day 0-1; (B) Bloat:Day 1-2; (C) Active decay with skull emerging: Day 2-4; (D) Advanced decay:Day 4; (E) Dry - limbs removed and skeletal elements visible ( inset ):Day 6; (F) Skeletal remains: Day 10 onwards. Figure 2. (A) Decomposition rate (in days) of a wild Bornean Orangutan carcass by stage, following the classification of Payne (1965), compared with the decomposition rate of other species in Southeast Asian tropical forest landscapes: rabbit (Oryctolagus cuniculus) (2 kg; Ahmad et al., 2011), long-tailed macaque (Macaca fascicularis) (5.7 kg; Silahuddin et al., 2015), and domestic pig (Sus scrofa) (35 kg; Rimbin et al., 2020). For the domestic pig, the active and advanced decay phase were combined as “Decay” in the source study. (B) Decomposition curve of the Orangutan carcass and a small mammal carcass ( Mastomys natalensis ) using the Total Body Score (TBS) method and Accumulated Degree Days (°C days) over time. The maximum TBS for the Orangutan was 25 and for the small mammal was 35. Discussion The rapid decomposition of the Orangutan carcass in our study was likely due to the combined scavenging activity of both invertebrates and vertebrates. Dismemberment by Asian water monitors increased carcass accessibility, thereby accelerating decomposition by invertebrates. In comparison, carcasses in previous studies decomposed more slowly, likely because dipteran larval development requires 2–3 days before reaching optimal feeding levels (Ivorra et al., 2023). Although there are no obligate scavengers in Bornean tropical forests, our findings support the role of the Asian water monitor ( Varanus salvator ) as a key scavenger of Orangutan carcasses. This observation aligns with previous reports by Lim (2015) and Twining et al. (2017), who found this mesopredator frequently scavenging on small mammal and bird carcasses in Borneo. As generalist feeders with highly developed sense of smell, Asian water monitors are adept at detecting carcass (Cairncross et al., 2024). The scent of putrescence, strongest after the bloat stage when invertebrate activity has ruptured the carcass, likely facilitates this detection. Interestingly, the monitors did not consume the entire carcass, possibly due to the carcass size which is nearly double the body weight of an adult monitor. Removed limbs were not consumed on site, likely due to intraspecific competition. Contrary to expectations, no other vertebrate scavengers were detected, despite the nutrient-rich resource. Notably, this study took place shortly after an African Swine Fever outbreak in Sabah (Daniel et al., 2024; Ewers et al., 2021). The bearded pig ( Sus barbatus ), a crepuscular species and presumed scavenger, may have competed for the carcass under normal conditions, as noted by Galdikas (1978). However, its absence from the landscape post-outbreak leaves its current role as a scavenger of large carcasses uncertain. The consistently warm temperatures (25–29 °C) in Danum’s forests likely supported high invertebrate activity, rapid dipteran larval development, and microbial decomposition. Chrysomya rufifacies is known to thrive between 16–34 °C, with oviposition occurring within 0.5 days and a full life cycle completed in approximately 8 days (Zhang et al., 2018). Consistent with our findings, Ivorra et al. (2023) reported that Chrysomya larvae require 61.9–132 °C degree days to complete the larval feeding stage. Beyond 132 °C degree days, larvae transition to the pupal stage and burrow into the surrounding soil. Comparing our results to other published records show that decomposition rates and scavenger outcomes may vary considerably even under similar climatic and forest conditions. This variability likely reflects the limited number of studies conducted in natural environments, where carcass detection is inherently challenging (Turner et al., 2017); even long-term field studies have struggled to monitor mortality in the wild due to the difficulty of detecting freshly deceased animals (Galdikas, 1978; Milton, 1990). Experimentally placing carcass proxies provides the most practical approach for studying mortality and scavenging dynamics, but the large majority of published studies have excluded either vertebrates or invertebrates from the decomposition process, leaving a gap in our understanding of natural carcass removal when both classes of scavenger openly compete. Overall, our study suggests that the rapid decomposition of carcasses may partly explain the lack of confirmed mortality cases among rehabilitated Orangutans. This highlights the importance of reassessing the frequency and methods of monitoring released individuals. Using weekly monitoring to ascertain the fate of released Orangutans may be too infrequent to adequately detect mortality events. We found almost all evidence of a mortality event had disappeared within just six days, with even the smell of the carcass having dissipated and the only remains being inconspicuous bones on the forest floor. We do, of course, acknowledge that our study is an opportunistic one and based on a single carcass, which restricts generalizing our findings regarding Orangutan decomposition, scavenger behavior, and removal rates in tropical forests. However, our results are reasonably consistent with observations from other reports and data on both large and small mammal carcass decomposition rates and provides some guarantee that what we observed was not an aberrant event. Importantly, this study is among the first to document the decomposition of an Orangutan in the wild, and our results highlight the need to re-evaluate current monitoring practices for rehabilitated Orangutans. Acknowledgements We would like to thank to Jeffry Anson, nature guide at Danum Valley, for providing the information and location of the Orangutan carcass, Dr. Hollie Folkard-Tapp and Mr Yehezkiel Jahuri for their field support. We are grateful to Dr Felicity Oram and Dr Michael J. W. Boyle for their invaluable inputs to the study. We like to acknowledge the Sabah Biodiversity Centre, Danum Valley Management Committee and Yayasan Sabah for granting the necessary permits (JKM/MBS.1000-2/2/1 JLD.1 (213)) and permissions to conduct this work. This research was supported by the Association for Tropical Biology and Conservation (ATBC) through the ATBC Seed Grant program (2023). Data Availability: The data used in this study is available on http://doi.org/10.5281/zenodo.16250654 Competing interest statement The authors declare no competing interests. Author Contributions: Conceptualization: SPH RME ;Data curation: SPH; Formal analysis: SPH; Funding acquisition: SPH; Investigation: SPH; Methodology: SPH RME; Resources: SPH RME; Supervision: RME HB; Validation: SPH RME HB; Visualization: SPH ;Writing –original draft: SPH; Writing – review & editing: SPH RME HB; Project administration: SPH, All authors have read and agreed to the published version of the manuscript. References Ahmad, N. W., Lim, L. H., Dhang, C.C., Chin, H.C., AG, A, Wan Mustaffa, N.W., Kian, C.W., Jeffery, J., Hashim, R. & Azirun, S.M (2011) Comparative insect fauna succession on indoor and outdoor monkey carrion in a semi-forested area in Malaysia. 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Information & Authors Information Version history V1 Version 1 30 July 2025 Peer review timeline Published Ecology and Evolution Version of Record 11 Dec 2025 Published Copyright This work is licensed under a Non Exclusive No Reuse License. Collection Ecology and Evolution Keywords community ecology natural history terrestrial vertebrate Authors Affiliations Sui Peng Heon 0000-0002-4075-737X [email protected] Universiti Malaysia Sabah View all articles by this author Henry Bernard Universiti of Malaysia Sabah View all articles by this author Robert Ewers Imperial College London - Silwood Park Campus View all articles by this author Metrics & Citations Metrics Article Usage 394 views 197 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Sui Peng Heon, Henry Bernard, Robert Ewers. Decomposition Dynamics of an Orangutan (Pongo pygmaeus morio) Carcass in a Tropical Forest: Implications for Conservation Practices. Authorea . 30 July 2025. 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