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The surprising diversity of Sunda clouded leopard communication behaviors

Above: A portrait of a Sunda clouded leopard (Photo by Max Allen)   Sunda clouded leopards (Neofelis diardi) are part of the Panthera lineage of felids that includes African lions (Panthera leo), tigers (Panthera tigris), and jaguars (Panthera onca). These are among the most charismatic wildlife species, but Sunda clouded leopards are the least understood...

Above: A portrait of a Sunda clouded leopard (Photo by Max Allen)


Sunda clouded leopards (Neofelis diardi) are part of the Panthera lineage of felids that includes African lions (Panthera leo), tigers (Panthera tigris), and jaguars (Panthera onca). These are among the most charismatic wildlife species, but Sunda clouded leopards are the least understood and studied of this group. Sunda clouded leopards are found in Borneo and Sumatra, while the similar species clouded leopards (Neofelis nebulosa) are found in mainland Asia. Sunda clouded leopards are very cryptic in nature and are considered a threatened species, but little is known about their ecology or behavior. Because of this, new research is shaping the way we understand this fascinating species.

A Sunda clouded leopard family group (a mother and two kittens) in Borneo


One area of my scientific research is how animals communicate, including scent marking, which helps us understand behavioral ecology and inform conservation efforts. Previous to our study no scent marking by Sunda clouded leopards had ever been recorded, which led some researchers to conclude that they do not scent mark and instead have other means of communicating (e.g., Rabinowitz et al. 1987). Consider the table below, however, which shows that studies of communication exist for 29 of the 39 felid species, with each of the studies mentioning scent marking as a primary form of indirect communication. This makes it seem implausible that Sunda clouded leopards don’t scent mark, and with field effort we found that their scent marking has just previously gone undetected by scientists.

The communication behaviors reported for each felid species and lineage, with no known behaviors for Sunda clouded leopards.

To explore the scent marking behavior of Sunda clouded leopards I teamed up with Andy Marshall and his long-term wildlife population ecology project in Gunung Palung National Park, West Kalimantan, Indonesia. Andy and his collaborators Heiko Wittmer, Endro Setiawan have been running this project since 2000, and have been doing continuous monitoring of wildlife since 2007. The aims of the project are to explore how changes in climate and the productivity of different habitats affect the spatial and temporal variation of wildlife populations. Sunda clouded leopards are a species of interest for the project because they are one of the largest carnivores in the area.

A map of our study area highlighting the location of our camera traps and habitat types.

In order to determine the scent marking behaviors used by Sunda clouded leopards we used motion-triggered video cameras. I prefer to use non-invasive methods, such as motion-triggered cameras, for research when possible because the animals are not harmed. In addition, their behaviors are not changed, which allowed us to get an accurate assessment of potential scent marking behaviors of Sunda clouded leopards in a natural setting. We set the 28 cameras across the study area over the course of 9 months, and I was fascinated by what we found here. We found that Sunda clouded leopards use a large diversity of communication and scent marking behaviors. In total we documented 10 different behaviors, including: scraping, urine spraying, claw marking, cheek rubbing, olfaction, and vocalizations.

A male Sunda clouded leopard scent marking through scraping.


A male Sunda clouded leopard scent marking through urine spraying.


We documented scent marking occurring through three different types of marking: urine spraying, deposition of feces, and scraping. Urine appears to be primarily sprayed backwards onto a tree trunk or other vertical surface, while scraping is where Sunda clouded leopards rake their hind feet into the ground and then sometimes urinate and/or defecate on the mound of material that has been scraped together. The scraped mound of material acts as a visual cue that makes the scent mark easier for other individuals to find (sensu Allen et al. 2014). These behaviors are the same as those used by other solitary felids, including tigers and leopards (Panthera pardus).

We also documented four types of body rubbing behaviors, including one that has never been previously documented for any felid species. Claw marking is where Sunda clouded leopards rake, gouge, and/or grip a tree trunk with their claws, leaving visual and potentially scent cues. Cheek rubbing is where Sunda clouded leopards rub their cheek on a tree trunk or other object, leaving scent cues from their sebaceous glands. Rolling (also called ‘vegetation flattening’ in some felid studies) also occurs, where Sunda clouded leopards roll back and forth on the ground, but the reasons behind this behavior are not yet known.

A male Sunda clouded leopard exhibiting cheek rubbing.


The fourth body rubbing behavior we documented was tail wrapping, where the Sunda clouded leopard wrapped its tail around a tree trunk. Sunda clouded leopards are known for their arboreal behaviors, including hanging from tree limbs from their claws and using their notably long tails for balance. The wrapping of their tails around objects on trails may be related to this, or it may be used intentionally to leave scent, as with other forms of body rubbing. We only documented this behavior once during the sampling period used for our first paper – although our ongoing camera sampling in the study area has since led to additional observations of this behavior. Tail wrapping does not appear to have been previously described in literature for any other felid species, and may be unique to Sunda clouded leopards.

A Sunda clouded leopard exhibiting olfaction and flehmen response.


We also documented two investigative behaviors, which are also used by many mammals and most felids. Olfaction (also called ‘sniffing’ in some studies) is where Sunda clouded leopards smell to investigate cues and signals. The flehmen response is more dramatic, where the Sunda clouded leopard lifts their head and curls back their upper lip, sometimes arching its neck backwards. This is used to expose their vomeronasal organ, which allows for greater investigation of scent.

The frequency of communication behaviors observed in clouded leopards, divided into four types of behaviors. Within each group, frequencies did not differ significantly between behaviors sharing the same symbol to the left of the bars; behaviors with different symbols differed significantly.

Despite the relatively short duration of our study, we documented Sunda clouded leopards using many different communication behaviors. The use of diverse communication behaviors in Sunda clouded leopards is consistent with observations from other species of the Panthera lineage (Figure 1), and overall appear most similar to leopards or snow leopards (Panthera uncia). The important finding is that while many other felids use these behaviors, very few use so many. These findings solidify the findings of how felids communicate by showing that there is a notable similarity in the form and function of scent marking behaviors used by felids. Because they are similar in form and frequency to other felids, including the Panthera lineage, the functional ecology of Sunda clouded leopard scent marking and communication behaviors are also likely similar. Specifically, Sunda clouded leopard scent marking is likely used for intraspecific communication, with functions including both territorial marking and finding and selecting mates.

The communication behaviors reported for each felid species, showing the diversity of behaviors used by Sunda clouded leopards.

Our discovery of Sunda clouded leopard scent marking, their fidelity to identified scent marking areas and the use of the same scent marking areas by different males has the potential to help overcome existing limitations studying the ecology of Sunda clouded leopards. Specifically, repeated visits of identifiable individuals to scent marking areas should help improve accuracy of population estimates using non-invasive remote cameras that are needed to develop effective conservation strategies for this endangered carnivore in Borneo.

Our findings in this study are just the beginning. We documented behaviors, such as vocalizations, that we have not yet begun to interpret. Keep up to date with my research as I explore these and other questions. I would like to express many thanks to my collaborators on this project, and as a group we would like to thank the State Ministry of Research, Technology and Higher Education (KEMENRISTEKDIKTI), the Directorate of Natural Resources and Ecosystems (KSDAE), and the Gunung Palung National Park Bureau (BTNGP).



  1. Allen, M.L., H.U. Wittmer, and C.C. Wilmers. 2014. Puma scrape and communication behaviors: understanding functional use and variation by sex and age. Behaviour 151: 819–840.
  2. Allen, M.L., H.U. Wittmer, E. Setiawan, S. Jaffe, and A.J. Marshall. 2016. Scent marking in Sunda clouded leopards (Neofelis diardi): novel observations close a key gap in understanding felid communication behaviours. Scientific Reports 6: 35433.
  3. Rabinowitz, A., P. Andau, and P.P. Chai. 1987. The clouded leopard in Malaysian Borneo. Oryx 22: 107–111.

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Meet the Author

Author Photo Max Allen
Max Allen is a carnivore ecology professor at the University of Illinois and Illinois Natural History Survey. He completed his Ph.D. in Conservation Biology from Victoria University, Wellington in 2014, with his dissertation entitled: The Ecology and Behaviour of Pumas (Puma concolor) in Northern California. Max has since published over 45 peer-reviewed articles in scientific journals, with a focus on using camera trapping to understand solitary carnivores and ecological interactions. He is currently working on felid conservation projects on four continents, including pumas and bobcats in North America, leopards and cheetahs in Africa, tigers and clouded leopards in Asia, and lynx in Europe. In addition to research, Max enjoys running, exploring wild places, and using photography to connect with wildlife.