Journal of Wildlife and Parks (2014) 29 : 37-44 37

FOOD CHOICE AND FEEDING HABITS OF THE FLAT-HEADED CAT (Prionailurus planiceps) IN CAPTIVITY

John Rasmussen*

Institute of Biology, University of Southern Denmark, Denmark

*Corresponding author: [email protected]

ABSTRACT

The flat-headed cat, Prionailurus planiceps, is a small Asian felid, whose ecology is not very well documented. This study aimed to uncover the food choice and feeding habits of P. planiceps in captivity. Two P. planiceps living in a small enclosure in Peninsular Malaysia were fed different types of food including fish, frogs and birds, to see which ones they prefer. They were also tested to see if they prefer living prey or dead food items. It was noted where the prey was consumed, how much was consumed and whether or not the cats would “play” with the living prey before eating it. Results indicate that P. planiceps is an opportunist and will take food in, or close to water, especially fish. It seems that P. planiceps prefers living prey over dead food items. Most food was consumed less than 1 meter from the water, however this may be only a matter of the behavior of P. planiceps in captivity. P. planiceps would always play with the living prey before consuming it. Nothing conclusive could be said about the degree of fish consumption, as the head of the fish was left uneaten in roughly half of the feeding events.

Keywords: Behavior, Dead food item, Ecology, Enclosure, Fish consumption, Living prey

INTRODUCTION

The family Felidae comprises all living cat species of the world. Today there are approximately 37 species of Felidae, of which almost half are considered threatened by the International Union for Conservation of Nature (IUCN, 2011). The flat-headed cat, Prionailurus planiceps, is an endangered cat species native to Peninsular Malaysia, Borneo and Sumatra (Sunquist & Sunquist, 2002; Esabii, 2011).

Adults range in size between 446-521 mm, with a tail length between 128-169 mm, and its weight is approximately 1.6 kg (Sunquist & Sunquist, 2002). The head is longer compared to other Malaysian cats, and the sheaths of the claws are reduced. The eye placement of flat-headed cats is more anterior in the head in comparison to other cats. The second upper premolar is relatively larger, however, the upper molars are smaller in comparison to the leopard cat, P. bengalensis. The flat-headed cat has short legs, tail and ears, and its color is brownish, with a white underbelly (Muul & Lim, 1970). The cats' mouth has evolved to become better at seizing wet and slippery prey compared to the fishing cat,P. viverrinus, and it probably has greater biting power (Sunquist & Sunquist, 2002).

In the wild, few studies have been done on the ecology of the flat-headed cat, and much less on the natural diet of the species. Flat-headed cats in the wild will probably hunt for fish and they seem to feed on a range of food types. The stomach contents of a dead P. planiceps contained flesh and some fish vertebrae (Muul & Lim, 1970). P. planiceps will also hunt for frogs and crustaceans (Salakij et al., 2008), and has also been observed killing and eating small rodents (Sunquist & Sunquist, 2002). 38 John Rasmussen

When a kitten was presented with live sparrows, it would completely ignore this type of food (Muul & Lim, 1970). However, a captive adult P. planiceps caught and consumed live quail (B. Simpson pers. comm.). P. planiceps has even been said to prefer fruit (Anderson & Jones, 1984).

A kitten studied in captivity would enter water when looking for food and grab pieces of fish, by completely submerging its head and grabbing the desired piece. It would also “wash” its food in the water, almost like a raccoon (Procyon lotor). After the kitten had collected a piece of food, it would drag the food at least 2 meters away from the water before consumption. The kitten captured live frogs by snarling and jumping upon them while grabbing them (Muul & Lim, 1970). Currently only 12 flat- headed cats are kept in captivity in Thailand (Thongphakdee et al., 2009), and fewer still in Malaysia. In 2002, less than 20 specimens had been collected from the wild (Sunquist & Sunquist, 2002).

The purpose of this study was to investigate the food choice and feeding behavior of flat-headed cats in captivity.

The following aspects of the cats’ food choice and feeding behavior were examined: 1. What food items did the cats prefer? 2. Did the cats prefer living or dead food? 3. Where would the cats eat the food?

METHODOLOGY

Study site and experimental animals One pair of flat-headed cats (an adult male and female) were kept in a 77 m2 enclosure (Figure 1), located at Sungai Dusun Wildlife Reserve, Selangor, Malaysia, which is located in an almost undisturbed peat forest area. The area is under the supervision of the Department of Wildlife and National Parks (DWNP), Malaysia. The enclosure contains 2 small pools (roughly 10 cm deep) and 2 small streams (water to the streams was turned off during the whole investigation), a hollow log and a few artificial platforms on which the cats can rest.

The cats were caught in the wild a few years ago and handed over to the DWNP. They have been living in the Sungai Dusun enclosure since January 2010, and are by the date of this study, used to their current environment.

Data collection and recording equipment Six Keep Guard camera traps (model number: KG-680V) were used to collect photos of the cats throughout all 24 hours of the day, and 1 camera trap (same model) was used to collect video of pool 1 where the food items for the cats were placed. Pictures and video were recorded on replaceable SD memory cards and were copied to a laptop computer for analysis each day at 7:00 pm. The cameras had a 1 second response time, so if an event was shorter than this (eg. a cat runs in and out of the screen), it would not be recorded. The cameras traps used heat signatures to identify when something living moved in front of the camera. Video recordings were set to record 15 second videos for each event. Camera trapping was chosen as the only method of data recording for this experiment, since the cats were (at least initially) scared of the observers, and would not come out of hiding until no one was around. Food Choice and Feeding Habits of The Flat-Headed Cat 39 (Prionailurus planiceps) in Captivity

Figure 1. Sketch of the cat enclosure with measurements of the walls' dimensions. Interior items are not to scale, but serve to illustrate the relative positions of various items, including water pools and the hollow log in which the cats would sleep during a large part of the day. The yellow dots with black arrows indicate the placement and viewing direction of the camera traps inside the enclosure. The transparent yellow zone around the pool indicates the approximately 1 meter zone around pool 1 within which events is categorized as being close to water, as described below. Chicken and frogs would be placed on the indicated feeding spots. North is pointing downwards.

Animal feeding trials The study took place from 8th February to 4th March 2011. The cats were fed once a day during the experiment, and were given several hours in which to finish their meal. Most of the time the cats were fed at 9 am, however initially feeding was at 3 pm. Leftover food was always collected at 7 pm. 40 John Rasmussen

The cats were fed combinations of 3 basic food items; dead frogs (3-5cm, locally caught, species unknown), fish (17-20cm, catfish Clarias batrachus and Indian mackerel, Rastrelliger kanagurta), and chicken pieces without feathers (two chicken piece was roughly equal to volume of 1-2 average catfish). The cats were fed different types of live or dead food items, with live and deadfish placed in the pool, and dead frogs and chicken on plates within 10 cm of the pond edge. More trials with frogs would have been helpful, but according to the local staff who bought the food for the experiment, frogs were more difficult to acquire, and thus sometimes trials were affected by the availability of food types. The cats have always been fed on feeding plates since moving to Sungai Dusun.

On a few of the trials with chicken, a bit of carnivore supplement powder was added to the chicken, as the staff wanted them to have this with their meals. It has a scent of meat in higher concentrations, but in the amounts put on the chicken bits (1 teaspoon for the trials) it is more or less odourless (to humans). Since the cats behaved no differently with or without the supplement, the presence of the supplement on the chicken is considered to be of no consequence for this study.

When analyzing where the cats would eat the caught fish, it was decided that feeding events within a zone of approximately 1 meter from the water (and including the pool itself) was categorized as “pool 1”, which is either close to water or in water. The 1 meter zone was decided upon because the kitten described in Muul and Lim (1970) would drag the caught fish 2 meters from water, but given that this enclosure is rather small, 1 meter was decided to be enough (going more than 2 meters from one side of the pool would almost make the cat hit the walls of the enclosure). Whether or not the cats were within the zone was estimated by the observer when viewing the video recordings, and so was done in the same way by the same person each time. For data analysis Chi2-tests was performed, using a table for critical values found in Samuels and Witmer (2003).

RESULTS

Preferred types of food During the trials, food items were tested against each other to see which ones the cats preferred. The different combinations and number of times were successfully tested is displayed in Table 2. There was one unsuccessful trial with dead fish and living fish, where the camera did not show which type of food was eaten first. Whenever the cats attacked the live fish to pull it out of the water with their mouths, they would submerge part, if not all, of their face when trying to grab the fish.

Table 2. The different combinations of food tested against each other during the trials and the number of trials each combination was used in.

Combination Number of succesful trials Dead fish & chicken 10 Dead fish & living fish 6 Dead fish, dead frog (2 units) & chicken 2 Dead frog (3 units) & chicken 1

Dead fish vs chicken The cats ate the dead fish first in all 10 trials. The chicken was always eaten only after the fish was consumed. There is a clear selection and preference for fish over chicken. Food Choice and Feeding Habits of The Flat-Headed Cat 41 (Prionailurus planiceps) in Captivity

Dead fish vs living fish The cats ate the living fish first in 5 of the 6 trials. The dead fish was eaten first in only 1 trial. The Chi2-test (at α = 5% level) did not show a significant difference of choosing between live or dead fish, however there was a difference shown at the 10% level (χ2 = 2.67, df = 1, p = 0.10). It would seem that the cats prefer live fish over dead fish, however further trials would be needed to confirm this using a larger sample size. It was noted that the cats often left the heads of the fish uneaten – in about half of the trials. The heads were left from both live and dead fish.

Other combinations There was not enough time to run enough trials to satisfy the Chi2-test sample size for fish vs frog vs chicken, or frog vs chicken. However, in the both trials of fish vs frog vs chicken, the fish was eaten first then the frog, and the chicken was eaten last. In one trial the chicken was not eaten at all. In the trial of frog vs chicken, the frog was eaten first and the chicken was eaten last. It seems that flat-headed cats prefer fish, then frogs, and lastly chicken.

Figure 2. Flat-headed cat catching a catfish photographed with a handheld camera during one of the last feeding trials.

Preferred place to eat Feeding events were recorded a total of 32 times near the feeding pool (pool 1). Feeding events were recorded a total of 5 times at a distance of more than 1 meters from water. There was a significant statistical difference of where the cats ate their food (χ2 = 19.7, df = 1, p < 0.0001). The cats preferred to eat close to the pool of water. When they did not eat close to the water, they were eating a fish. 42 John Rasmussen

DISCUSSION

P. planiceps would eat any of the food types presented to it during the trials, as could be expected from an opportunistic hunter. Both frogs and fish were readily eaten, and the cats rarely refused the chicken presented to them, though they would usually leave a few chicken leftovers, presumably because the cats were full and simply did not want to eat any more food that day. Even the Indian Mackerel was eaten with no left overs, despite this being a saltwater fish to which P. planiceps would not ordinarily have access to under natural conditions.

The results of the testing of fish vs chicken are pretty conclusive and show that the cats definitely prefer fish over chicken, but also show that flat-headed cats will also readily eat bird-type food (chicken). Flat- headed cats were said to catch and eat live quail when presented to them (B. Simpson pers.comm.), which is different in other studies where living bird prey was completely ignored (Muul & Lim 1970). One could imagine that the cat seems to have no problem eating many kinds of food, but just happen to prefer food occurring naturally in or close to water, as this is also where they presumably may hunt in the wild.

The results suggest that P. planiceps may have a preference for living prey, as shown by the results of a Chi2-test at the 10% significance level, although the Chi2-test showed no significant divergence from the null hypothesis at the 5% level. This ambiguity may result from a low number of trials (n = 6). The cats may not distinguish between living or dead fish as shown by the fact that when the living fish was taken by one cat, the other one would immediately grab the dead one and eat that one instead, not trying to fight the other cat for the living fish. One could argue that because the cats always play with a living fish before catching it, they do not prefer living fish to dead. However, in captivity, playing with food is one of the only activities the cats have available other than marking territory, bathing in the pools, or inert activities like sleeping. If the cats do not feel like playing with the fish, they have no reason to take the living one, as the dead one offers an easy meal. In spite of speculations, one can safely conclude that the cats readily eat either, but may have a slight inclination to choose the living fish. The problem might be, that there was simply not enough data points collected in the study due to time constraints and for other reasons explained above. In the future it might be worth doing the dead fish vs living fish experiment a few more times and get more data points. I am inclined to believe that the cats prefer living food, but no definitive conlusions can be made because more data is probably required. In addition, the cats preferred frogs over chicken. This might be because frogs are usually found around water, and so it is probably true that the cats would eat more of this in the wild as well.

Sometimes the cats would leave the head of a fish in the enclosure when they were done eating. Perhaps we were feeding them too much food over too short an interval, so they only ate the most tasty bits. This could not have been changed though, do to a tight study schedule.

It was found that the cats prefer to consume the fish less than 1 meter from the water. In fact many observations were of the cat eating directly next to the water, perhaps no more than 10 cm from the edge of the pool. This seems to be contrary to the few other studies, and may be a case of behaviour adapted to life in captivity. As noted in other studies, other cats would drag food at least 2 meters away from water before consuming it. However, the cats in this experiment were accustomed to living in captivity and may have learned that the risk of food escaping them in a 10 cm deep pond with no water flow or escape routes is not existent. Considering these factors, while it is certain the cats prefer to eat next to the pool, this preference may not represent a natural behavior. This statement might be backed by the occurence of "playful" activity while hunting the live fish, as they would "play" with the living prey every time it was presented to them. In the wild this probably would not happen, as the chance Food Choice and Feeding Habits of The Flat-Headed Cat 43 (Prionailurus planiceps) in Captivity of prey escaping the cat will be high if the cats simply released it every time they caught a fish. In conclusion, it is probably uncertain if the species as a whole prefers to consume their food right next to the water in which it is caught. To get more realistic studies of this, it would probably have been better to study newly caught cats, that have not had a chance to adapt to life in captivity.

CONCLUSION

The flat-headed cat, P. planiceps, is an opportunist as far as food choice is concerned, and will eat many types of food as long as it is available to it, even food types not naturally available to it, such as mackerel. It seems that flat-headed cats prefer fish over frogs over chicken, but will eat all readily. It seems that P. planiceps prefers living fish over dead fish, but is also happy to scavenge dead prey items. The flat-headed cat prefers to eat its food less than 1 meter from the water, but this might be a consequence of changed behavior patterns from living in captivity. Further study is needed for this endangered species.

ACKNOWLEDGEMENTS

I would like to thank the Department of Wildlife and National Parks, Malaysia, for allowing me to use their facilities and for being so generous in providing accomodation and helpful staff, and for allowing me to study their animals. Without this, the study would have been impossible to do. I would also like to thank Copenhagen Zoo / Carl Traeholt for being my external supervisor in Malaysia, who provided the camera traps and was very helpful, and also Boyd Simpson for being very helpful and for making our first days in Malaysia as comfortable as possible. Boyd Simpson also provided comments on earlier drafts of this paper. Last but not least a thank you to Ole Næsbye Larsen who has been my supervisor at the University of Southern Denmark and helped me throughout this entire project.

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Muul, I. & Lim, B.L. (1970). Ecological and morphological observations of Felis planiceps. Journal of Mammalogy, 51: 806-808. 44 John Rasmussen

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