Environmental enrichment, shortened here to enrichment, provides captive animals with stimuli needed to participate in species-typical behavior and increases the amount of time an animal spends in goal-directed behavior.1 It is often used in zoos to decrease the occurrences of stereotypical behaviors, or any repetitive behavior exhibiting no obvious goal or function. Enrichment is not one size fits all, and it is important to identify effective enrichment for individual species, and individual animals, in captivity.1 However, the fact that enrichments can be successfully used across species provides insight into the behavioral similarities and differences between species. This article explores responses to enrichment in two closely related species – the black-footed cat (Felis nigripes) and the domestic cat (Felis catus) – through video footage and analysis of each species’ interaction with three enrichment treatments (wand toy, foraging bags, and tube with prey).

The cats

Black-Footed Cats

The black-footed cat is a small wild cat native to South Africa, weighing an average 1.3 – 1.03 kg (2.8 – 4.3 lbs).2 They are listed as vulnerable on the IUCN Red List, which means their population is decreasing.3 Nicknamed the “world’s deadliest cat,” this species is most well known for their incredible hunting ability, with a 60 percent hunting success rate.4 By comparison, leopards and tigers have hunting success rates of 14 percent and 5 percent, respectively.5

Five black-footed cats housed in Lory Park Animal and Owl Sanctuary in Midrand, South Africa, were used in this comparison: Miley, 1-year-old female; Mowgli, 1-year-old male; and their three 2-month-old kittens, Nala, Nile, and Nizara. Miley and Mowgli were housed together prior to the kittens’ birth. At the time of this study Mowgli had an enclosure to himself and Miley shared an enclosure with her kittens. Each enclosure included opportunities for hiding, climbing, and other natural behaviors. All five cats were frequently handled and had positive associations with their caretakers prior to the study. Miley and Mowgli had a history of exposure to enrichments, but none of the five black-footed cats had previous experience with the enrichments used in this study.

Domestic Cats

Four cohabitating domestic cats located in New Hampshire, United States, were used in this comparison: Franky, 7-year-old neutered black domestic longhair; Mischa, 6-year-old spayed gray and white domestic shorthair (DSH); Stitch, 5-year-old spayed tabby and white DSH; and Connor, 3-year-old neutered buff tabby DSH. A fifth cat, Washburne, a 3-year-old neutered marbled tabby medium hair, was also in the home but chose to stay in a different part of the house when the enrichments were introduced.

This group lived together for three years prior to the study. The three males were preferred associates (i.e., they frequently exhibited affiliative behaviors including allogrooming, sleeping together, etc.). While the two females did not have preferred associates, it could be said the five cats cohabitated peacefully. The environment included multiple opportunities for hiding and climbing, and each domestic cat had a positive association with their caretakers prior to the study. There was a history of exposure to enrichments, including exposure to wand toys. None of the domestic cats had previous experience with the foraging bag or tube with prey enrichments used in this study.

Methods

The three enrichments used in this study (wand toy, foraging bags, and tube with prey) were designed as part of an animal behavior research study through Loop Abroad and Lory Park Animal and Owl Sanctuary. The wand toy and foraging bag enrichments were designed by Skylar Gomez and the tube with prey enrichment was designed by Melissa Craven. Each enrichment was introduced to the cats’ spaces, and the cats were allowed to investigate at will, except for the wand toy, which was manipulated by a human. The cats’ caretakers were present for the duration of each enrichment.

The enrichments were introduced to Lory Park’s resident black-footed cats in November 2020 and the interactions recorded by Kara Heynis and Tasch Sibilski. Similar enrichments were then introduced to the group of cohabitating domestic cats in December 2020, the interactions recorded, and both black-footed cat and domestic cat footage was analyzed by Skylar Gomez.

Wand Toy

Black-Footed Cats

Miley readily interacted with the wand toy, both visually tracking and chasing after it. She most often used a front paw to pull the colorful prey toward her mouth rather than pouncing. She would try to walk away with the prey, and when it inevitably bounced back, she immediately went after it again. Her ears were front facing while watching the toy and held flat to the side like airplane wings while actively interacting with it.

The three kittens exhibited neophobic behaviors when introduced to the wand toy. Instead of interacting with the toy, they huddled together in the corner of their box shelter and focused their attention on watching the toy and Miley. When the toy neared them, the kittens backed farther into the corner and hissed. After a while, one of the kittens left the box shelter to investigate. The kitten did not interact directly with the toy, still moving away if it came too close, but also did not hiss and was not solely focused on the toy. This indicated the kitten was in a more relaxed state than when they were in the box shelter.

Another of the kittens swatted at a leaf dangling from a branch like the colorful prey of the wand toy. This suggests the kittens may enjoy the wiggling movement caused by the wand toy but were neophobic of the toy itself.

Domestic Cats

Like Miley, Mischa readily interacted with the wand toy. She focused on the feathers, pupils dilated and body oriented toward the toy anytime it moved. She swatted at it with both paws and brought it to her mouth several times. After capturing the feather, Mischa alternated between releasing it, trying to walk away with it, and shaking it. Periodically she shifted focus from the toy to her surroundings. Stitch sat out of camera view and watched the interactions. This appeared to be her preferred method of play. Connor sniffed the feather, chasing it instead of attempting to capture it. Franky was not present for this enrichment.

Foraging Bags

Black-Footed Cats

Foraging is a natural behavior in which an animal obtains food. This intrinsically motivating and engaging behavior is frequently incorporated into enrichment treatments in zoos.6 Four paper bags filled with dry grass, three of which included an added scent (i.e., cinnamon, catnip, and mouse blood), were added to Miley’s enclosure. Miley jumped right into the first bag without hesitation. 

She demonstrated more cautious behavior around the other three bags. While investigating the second and third bags, she kept her muscles tensed, took deliberate steps, and was highly focused on the bags and easily startled. She flattened her ears and called to her kittens when she walked past where they were hiding in a large tunnel.

Miley stopped to clean her hind leg after investigating the third bag, a displacement behavior suggesting elevated stress from the new stimuli. In other words, this displacement behavior may have been Miley’s way of taking a moment to process the foraging bags.

The fourth bag collapsed under her paw when she stepped on it. Instead of retreating, she lifted her paw and paused for a moment, focusing on the bag as it lifted to its original position. She sniffed the small space between the bag and the large tunnel and extended her front leg, presumably to investigate if there was enough space to squeeze between the two objects. Miley stepped on the bag a second time and lowered her head to sniff it before backing up to continue her visual investigation.

She walked around the other side of the bag and ducked inside it, bringing her whole body in except for one hind leg and her tail. She stayed inside the bag for about eight seconds then backed out and looked around. 

Miley exhibited different responses to each of the bags but did not exhibit a strong preference for any specific bag.

Domestic Cats

A fabric ball was placed inside three paper bags, two of which were scented with cinnamon or catnip. The bags were placed on the floor and the cats allowed to explore on their own. Stitch remained on the couch just out of camera view and did not interact with the bags. Connor sniffed the middle bag (cinnamon) and Franky stood behind Connor, his head lifted and bobbing slightly as he scented the air. Mischa briefly sniffed the bag on the left (unscented) before going straight to the bag on the right (catnip), stopping in front of it and blocking access to it from the other cats.

Connor and Franky approached the bag on the right, Franky walking to the back of it and Connor to the side. While Mischa’s front end was lowered and her hind legs tucked under her body, Connor had a more relaxed stance with his body squared and tail arched in a typical posture for him. Mischa briefly touched noses with Connor, then returned her attention to the bag. When she walked to the side of the bag, Connor moved closer to investigate the opening, his body lowered and neck outstretched.

Mischa touched her paw to the top of Connor’s head, although he did not retreat, and they both continued sniffing the bag. The bag slid across the floor when Mischa pushed her head in, and Connor visually tracked its movement. Franky stood upright, looked around, then walked away; he had not been near the bag opening yet. Connor suddenly pulled away from the bag and walked toward the camera. Franky turned to take Connor’s place and Mischa backed out of the bag. She sat and shifted her weight away from Franky, licking her lip and lifting one paw toward him. The lip lick could be indicative of agitation, or it could have been because she got a piece of catnip.

Tube with Prey

Black-Footed Cats

A mouse was placed in a plastic tube and introduced to Mowgli to encourage foraging and cognition (problem solving). Mowgli investigated each end of the tube, alternating as the mouse moved. His ears were perked forward, and his gaze focused on the tube.

Periodically he walked away from the tube toward the keeper, his posture loose and steps slow, but quickly returned to the tube. Mowgli has a positive association with his keepers, and this may have been a “checking in” behavior similar to what is sometimes exhibited in domestic dogs and human children.

While interacting with the tube, Mowgli crouched low to the ground and took short, quick steps. He pawed at the outside of the tube but did not paw inside it, instead sticking his nose in the end of the tube. At one point he was so enthusiastic about sniffing and pawing at the tube that he flipped over the end of it.

Domestic Cats

Two fabric balls coated in catnip were placed inside a cardboard tube and introduced to the cats. Fabric balls were used instead of mice since the domestic cats were not accustomed to playing with or eating mice.

Connor stuck his nose in immediately after the tube was placed on the floor. His tail arched over his back, which is typical for him while investigating or otherwise aroused by the environment. He walked to the other side of the tube briefly before turning back and pawing at the opening. He bit the bottom and top edges of the opening, using a paw to hold it steady. While Connor interacted with the tube, Franky rubbed his body on some spilled catnip and Stitch watched from just outside of camera view. Mischa sat underneath a table and watched but did not directly interact with the tube at all.

Connor used his face to push the tube across the floor until it came up against the wall, where he lifted it off the ground for a moment. When it fell back to the floor, he used a paw to steady it and tried to bite the top of the tube. He moved to stand over the tube and used his paw to lift it toward his mouth. It fell a second time and he moved back to the front of the tube, stuck his head in for a moment, then pulled his head out to eat a few pieces of catnip. Meanwhile, Franky slowly came closer while sniffing the floor, his front end low to the ground and his tail held loosely away from his body.

Just after Connor bit the top of the tube again, he and Franky startled at a sound. Connor lowered his body and oriented toward the sound. Franky froze, front end still low to the ground, and head raised towards the sound. Once they watched Stitch get off the couch, Connor licked his nose and both walked away from the tube. Stitch walked up to the side of the tube and sniffed. While she investigated, Franky came back to sniff at the tube opening and Connor returned to stand on the other side of Franky.

This behavior is noteworthy as Connor is known to antagonize Stitch. Franky has an affectionate personality and seems to have a calming effect on the other cats. Connor’s decision to approach the tube with Franky between him and Stitch suggests that the tube was of higher value than displacing Stitch. The tube’s status as a high-value item was further supported by Stitch remaining in close proximity to Connor, with Franky as a buffer, for ten seconds instead of immediately leaving.

Conclusion

A meta-analysis found that many enrichments can be successfully used between species.1  This comparison of enrichments in five zoo-housed black-footed cats and four cohabitating domestic cats suggests these two groups exhibit similar behavior patterns and responses to enrichment, further supporting the findings of the meta-analysis. Current research most often judges the efficacy of an enrichment by the impact it has on the frequency of an animal’s stereotypic behavior,1  with enrichments frequently implemented based on perceptions of what one thinks would be enriching to an animal. Just as it is important to identify effective enrichments for a species, it is necessary to evaluate the efficacy of an enrichment for the individual. Enrichments should be designed for an individual animal by considering their natural history, current environment, personality, anatomical characteristics and current body condition, past history with enrichment items, and underlying health conditions, and whether the proposed enrichment poses a risk or provides a benefit to the individual.7

The goal or desired outcome of an enrichment must be identified to determine its success. The wand toy enrichment would have been deemed ineffective if the goal was for Miley’s kittens to exhibit active hunting behaviors. However, a goal of introducing the kittens to new items and decreasing neophobic behaviors may have been considered successful since one of the kittens approached the wand toy to watch their mother play. Alternatively, using natural materials may have been more effective in encouraging active hunting behaviors, suggested by the fact that one of the kittens was seen playing with a leaf in a similar manner as the wand toy.

The animal’s behavior should also be monitored, ideally before, during, and after the introduction and removal of the enrichment. Only behaviors observed during the introduction of the enrichments were reported in the current study. Close observations can provide insight into an individual animal’s preferred enrichments. Findings from this study suggest Stitch preferred the tube with prey over the wand toy and foraging bags. She physically interacted with the tube with prey instead of watching as she did with the other two enrichments. She also chose to continue interacting with the tube after Connor approached, even though she typically prefers a certain degree of space from him.

It is widely accepted that enrichment improves animal welfare,1 and research suggests that improved welfare leads to increased life expectancy.8 Black-footed cats have an 8-year life expectancy; however, in captivity the average life expectancy is only 5 to 6 years (K. Heynis, personal communication, November 29, 2020). Although only one study has been published on the efficacy of enrichment in black-footed cats,9 the findings from the current study suggest using domestic cats as inspiration for black-footed cat enrichment treatments may be a successful strategy. Further research is needed to evaluate the efficacy of specific enrichment treatments and if the improved welfare positively impacts the black-footed cats’ captive life expectancy. As a vulnerable species, this is critical to successful conservation efforts.

Special Thanks

Special thank you to Kara Heynis, park manager for Lory Park Animal and Owl Sanctuary and founder of The Black-Footed Cat Project, and Tasch Sibilski, Loop Abroad coordinator for Lory Park Animal and Owl Sanctuary. This project would not have been possible without your enthusiasm and dedication. The Black-Footed Cat Project is a nonprofit focused on conservation and education.

References

1. Shyne, A. (2006). Meta-analytic review of the effects of enrichment on stereotypic behavior in zoo mammals. Zoo Biology, 25(4), 317-337.
2. Wilson, B., Sliwa, A., & Drouilly, M. (2016). A conservation assessment of Felis nigripes. In The Red List of Mammals of South Africa, Swaziland, and Lesotho. South Africa: South African National Biodiversity Institute and Endangered Wildlife Trust.
3. Sliwa, A., et al, (2016). Felis nigripes. The IUCN Red List of Threatened Species.
4. PBS. (2020, April 21). Super Cats: A NATURE Miniseries | Meet the Deadliest Cat on the Planet. Nature. 
5. Davies, E. (2015, December 22). Which animal is the deadliest hunter on the planet? Retrieved from BBC website
6. Delgado, M., Bain, M. J., & Buffington, C. (2019). A survey of feeding practices and use of food puzzles in owners of domestic cats. Journal of Feline Medicine and Surgery, 22(2), 193-198.
7. Duncan, A. E. (1997). A veterinary assessment of the risks and benefits of environmental enrichment. The Shape of Enrichment.
8. Teng, K. T.-Y., et al (2018). Welfare-Adjusted Life Years (WALY): A novel metric of animal welfare that combines the impacts of impaired welfare and abbreviated lifespan. PLOS ONE, 13(9).
9. Wells, D. L., & Egli, J. M. (2004). The influence of olfactory enrichment on the behavior of captive black-footed cats, Felis nigripes. Applied Animal Behaviour Science, 85(1-2), 107-119.