Above: A male puma growling (photo by Max Allen)
Communication is an important component of animal behavior, but is difficult to study in the wild. This is especially true for cryptic wildlife species, such as carnivores, that are difficult to observe. Recent advances in the technology of motion-triggered video cameras now enable researchers to remotely record intimate behaviors, such as vocalizations, in cryptic species. My research focuses on solitary carnivores, and one subject I study is how they use scent marking to communicate indirectly (e.g., Allen et al. 2014, 2016a). I often use motion-triggered video cameras to record carnivore behavior, and have compiled thousands of videos documenting different behaviors. One of the most interesting behaviors to record are vocalizations because of their complexity and varied functions.
Pumas are large, solitary felids that range widely across North and South America. Pumas primarily use scent marking for communication, but also use other forms including visual, olfactory, auditory, and tactile cues. Vocal communication is generally limited to family groups (i.e., between mothers and kittens, because only females exhibit parental care) and to mating pairs. Females give birth to 1-4 kittens in each litter and raise them for 12-24 months before the kittens disperse, and vocalizations may be an important aspect of communication between mothers and their young before dispersal. Because these cryptic carnivores are rarely observed directly, little is known about their vocalizations in the wild, and most information on their vocalizations has been based on captive animals. However, captivity often alters an animal’s behaviors, making it important to collect observations of puma vocalizations in the wild.
A trio of puma kittens mewing while nursing at their den.
Over the course of 5 years I recorded a variety of puma vocalizations in the Santa Cruz Mountains of California among females and family groups by opportunistically deploying motion-triggered video cameras with microphones at sites of interest. By better understanding acoustic communications, we can shed light on the evolutionary basis of these calls, as well as provide a more complete picture of animal behavior. The acoustic structure of puma vocalizations reflect specific adaptations to their function and behavior, and I hypothesized that the acoustic characteristics of each of these vocalizations have likely been optimized through natural selection to communicate most effectively at different distances (short-, medium-, and long-range) while minimizing exposure to predation risks. I therefore attempted to categorize and understand the function of each vocalization, and in the end, I placed five types of vocalizations into two categories: 1) attention-attracting, and 2) calls.
An uncollared female puma lounging at a community scrape and caterwauling.
Vocalizations to attract attention include caterwauling and mewing by kittens, as shown in the videos above. By placing a camera at community scrape I was able to record female pumas caterwauling on multiple occasions. Caterwauling is a long series of calls used to attract mates from a distance, and is primarily used by females. To document vocalizations by kittens we set a camera at a nursery of puma 23F captured her interacting with three neonatal kittens. The kittens do not vocalize before their mother’s return, but begin mewing as they start to nurse and shift their positions.
Vocalizations designed to attract attention (caterwauling and mewing) were characterized by broad frequencies, and caterwauling was characterized by long durations. The broad frequency ranges allow the vocalizations to carry long distances and attract possible mates that are in the vicinity. Similarly, mewing is a vocalization aimed at gaining attention, as the immediate reward of food greatly outweighs any risks they observe. Young siblings compete with each other, and the more insistent individuals often obtain more nutrition, allowing them to grow larger and out-compete their siblings increase their chances of survival. The direct benefits of attracting a mate or caregiver apparently outweigh any potential risks posed by predators.
A mother puma giving a contact call to gather her dallying kittens.
I split calls into 3 vocalizations (contact, agitated and alarm) that are short, narrow-frequency vocalizations that are similar in structure, but that varied in intensity along a scale. In each case calls were used to communicate with other nearby individuals. In the video above, a mother puma gives a contact call, presumably to gather her three kittens who had paused to investigate a community scrape. In the video below, I recorded a young kitten (5-7 months old) investigating a scent marking site alone. When the kitten seems to realize it is alone it gives an agitated call, presumable to locate its mother. Last, I recorded the alarm call of 33M, a 13-month-old uncollared male kitten of 19F, when he was trapped at one of 19F’s kill sites. 33M triggered the trap instead of his mother, and he was visibly startled immediately upon capture, and he vocalized a series of short whistles 15 seconds later.
A puma kitten giving an agitated call, most likely to try locating its mother.
Contact calls are vocalizations that are theoretically used for communication while limiting danger, and are characterized by pure tones and high pitches. This allows for immediate communication, but also limits the ability of predators to easily locate the sender or receiver of vocalizations. It is the nature of puma family groups to travel large distances, and my observations suggest they use vocalizations to stay together, especially after kittens begin traveling their mothers. Agitated and alarm calls were short and structurally similar to contact calls, but they incorporated slightly broader frequencies, possibly signifying higher urgency and potential danger. The broader frequencies likely made the kitten more immediately locatable than contact calls, but the increased risk of these calls might be acceptable because these calls were produced in response to a perceived immediate threat or danger.
These are among the first published recordings of vocalizations from wild pumas, and provide new insight into the structure and possible adaptive significance of puma vocalizations. Vocal communication entails risk, as it can attract the attention of predators or competitors and increase the risk of injury or mortality for both the sender and receiver of communications. My findings show that these vocalizations fall into two types: 1) vocalizations that are used to attract attention of other pumas with little regard to cost, and 2) contact and alarm calls which are short calls that vary in intensity and are used to communicate with nearby pumas. Pumas vocally communicate in numerous circumstances, including at nurseries, to maintain contact between family groups while travelling or in distress, and when trying to locate mates. The use of vocalizations by pumas demonstrates that acoustic communications may provide benefits that outweigh their risks, and highlights the importance of the structure of vocalizations used during different behaviors.
This study explores just a small part of puma communication with a small sample of vocalizations (see Allen 2016b for details), and much more work is needed to begin fully understanding these behaviors. Keep up to date with my research as I explore these and other questions at https://www.facebook.com/thewildlives. I would like to express many thanks to my collaborators on this project: Chris Wilmers and Yiwei Wang, as well as Dan Gardoqui and Abe Borker for discussions and feedback, and Paul Houghtaling, Yasaman Shakeri, and many field technicians and volunteers for their contributions to the project.
Allen, M. L., H. U. Wittmer, and C. C. Wilmers. 2014. Puma communication behaviours: understanding functional use and variation among sex and age classes. Behaviour 151: 819-840.
Allen, M. L., H. U. Wittmer, E. Setiawan, S. Jaffe, and A. J. Marshall. 2016a. Scent marking in Sunda Clouded Leopards (Neofelis diardi): novel observations close a key gap in understanding felid communication behaviours. Scientific Reports 6: 35433.
Allen, M.L., Y. Wang, and C.C. Wilmers. 2016b. Exploring the adaptive significance of five types of puma (Puma concolor) vocalizations. Canadian Field-Naturalist 130: 289-294.