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Home , Brown palm civet, Paradoxurinae, Paradoxurus

Diet of the Common Palm hermaphroditus in a rural habitat in Kerala, India, and its possible role in seed dispersal

Abstract P. S. JOTHISH

Paradoxurus hermaphroditus

Common Palm Civet diet in a rural habitat in India was studied by faecal analysis. Observations were made from January 2008 to May 2010, although no faeces wereCarica found papaya after January was the 2010. most In common total 94 fruitfaeces of werewhich collected. remains Of these, vegetable matter alone was found in 82% and matter alone in 4%; 14% held both plant and animal matter. Fruits were the predominant vegetable matter in the faeces. Papaya were found and was eaten year round. No seasonal change in proportion or dietary shift between vegetable and animal matter- was observed. Seeds from faeces were viable and a high percentage of defecated seeds germinated. Considering the movement pattern of Common Palm studied elsewhere and the high seed germination rate, Common Palm Civet is plausibly an ef fective seed disperser of fleshy-fruited plants in the study area. Common Palm Civet often attacks poultry and is perceived as a Keywordsthreat by the poultry farmers.

: Balaramapuram, faecal analysis, frugivory, poultry raiding, seed germination Introduction Methods

- - Order includes a wide range of character Food composition was studied by analysing faeces, a technique ised by a diversity of diets. Studying diet in carnivores helps- widely used to study small carnivore diet (Corbett' 1989,' E)Rab of understand their influence on distribution and abundance of inowitz 1991, Grassman 1998). Faeces were collected from a other including plants, and their population dynam rural house terrace in Balaramapuram (8°42 N, 77°04 largeics, habitat carnivores, use and information social organisation on the diet (Rabinowitz of tropical 1991,Asian owner,Thiruvananthapuram, up to three animals Kerala, were India. present, The terracealthough was it was near not a smallChuang carnivores & Lee 1997). is limited Despite (e.g. extensive Joshi et literatureal on diet of house where civets were observed to roost. According to the et al et al. 2010). Diets of small . 1995, Grassman confirmed that more than one provided the faeces analysed 1998, Zhou . 2008, Mudappa et al. here. Observations were made every week from January 2008 2010).carnivores, especially civets, in India have been studied few stillto May slept 2010, at the although same place. no faeces were found after 24 January times (Krishnakumar & Balakrishnan 2003, Mudappa - 2010, and it was not possible to confirm whether the civets ridae) occur across southern and some of eastern Asia in a The palm civets (subfamily , family Viver as theFaeces roosting were animal(s), identified shape as (including from Common elongated Palm nature) Civet Paradoxurus hermaphrodi- based on their occurrence in more-or-less the same location- tuswide lives variety in tropical of habitats forests, (Pocock plantations, 1933, Lekagulfruit orchards & McNeely and 1977). The Common Palm Civet and composition of undigested plant or animal matter. Col lected faeces were examined by naked eye and Carica with papayaa hand, human-settled areas (Prater 1971), often residing in eaves of lens—seekingArtocarpus seeds, pericarps, heterophyllus hairs, ,bones, Jungle feathers,Jack A. hirsutus insect, houses or outbuildings, across the Indian subcontinentet and al. exoskeleton, insect Annona wings reticulata etc. Seeds of Papaya Caryota southern China to Southeast Asia including the Philippines urensCommon Jack and east to New Guinea, in part by introduction (Patou Custard-apple and Fishtail Palm - 2010). In Kerala, a coastal state of peninsular India, Common ecated collected seeds; no from anti-seed-predator the civet faeces wereenclosures sown directlywere used. and Palm Civets are usually found in well-wooded areas and in without any treatment in backyard soil to test viability of def open places including areas adjacent to human habitations - (Balakrishnan 1997).et al Seeds were checked daily for germination and predation. Common Palm Civet is an omnivore that feeds on fruits- Results are presented as percentage frequency of occur and flesh (Joshi . 1995, Grassman 1998, Krishnakumar & rence (number of faeces with the particular item/total number Balakrishanan 2003). In forested areas, it is primarily frugiv animalof faeces matter). x 100) of a particular matter in the faeces. A chi-square orous, feeding on berries and pulpyet al fruits including those of- test was done to investigate the dietary preference (fruits vs. figs and palms and is an effective seed disperser (Rabinowitz Results 1991, Corlett 1998, Nakashima . 2010). It is often consid- ered a pest because of its raiding of coffee plantations, other faeces contained fruit crops and poultry, even though the coffee beans recov ered from its faeces are used to make a high-value speciality In total, 94 faeces were collected. Most (81) faeces coffee (Prater 1971). The present paper reports the diet of either plant or animal matter, but 13 contained both (Table 1). Common Palm Civet in a rural area of Kerala, India, and the Vegetable matter predominated: more than 95% of the animal’s possible role in seed dispersal. contained vegetable matter, alone or with animal matter. Only Small Carnivore Conservation 14

, Vol. 45:14–17, December 2011 Common Palm Civet diet in Kerala, India

Table 1. Vegetable and animal matter composition of the faeces Table 2. Fruits and seeds present in Common Palm Civet Paradoxurus of Common Palm Civet Paradoxurus hermaphroditus, collected hermaphroditus faeces collected from a rural site in Kerala, India, from a rural site in Kerala, India, during 2008–2010. during 2008–2010. Item description Number of Percentage of Plant type Frequency Percent Total faeces occurrence of occur- occur- number of Vegetable matter alone 77 81.91 rence rence seeds recovered Animal matter alone 4 4.25 Carica papaya 24 25.53 346 Vegetable and animal matter 13 13.83 Papaya (Caricaceae) Vegetable matter with or with- 90 95.74 Artocarpus hetero- 13 13.82 19 out animal matter phyllus Animal matter and with or 17 18.10 Common Jack ­without vegetable matter (Moraceae) A. hirsutus 10 10.63 107 Jungle Jack (Moraceae) 4.3% of faeces contained only animal matter. Fruit materials Caryota urens 6 6.38 42 Fishtail Palm were the dominant remains (90.1% of the faeces)2 year round. χ df (Arecaceae) Intake of animal prey was low. The faeces held fruits much P < 0.001). Musa spp. 3 3.19 0 more frequently than they did animal matter ( = 23.62, = 1, with animal matter) and cooked rice were also found in the Plantain (Musaceae) Grass plugs, i.e. partly chewed grass leaves (mostly faeces. - Mangifera indica 2 2.12 0 Mango (Anacardi- A single food item was recovered from 76 (80.8%) fae aceae) ces, 17 (18.1%) contained two food items and one contained- Capsicum annuum 2 2.12 14 three food items (1.1%). There was no evidence of a seasonal Chilli (Solanaceae) shift in relative importance of animal material in the diet. Fae Annona squamosa 5 5.31 68 ces composed entirely of animal matter were found in July- Custard-apple 2008 (13 and 18 July; rodent skull and hairs and millipede (Annonaceae) remains), February 2009 (frog skin and bones) and Novem Annona reticulata 2 2.12 11 ber 2009 (millipede remains). Faeces containing both animal Bullock’s heart and plant matter were obtained in February (twice), May, June (Annonaceae) and November 2008; January, March (twice), July, September Annona muricata 1 1.06 2 with(twice) Carica and papayaOctober 2009; and January 2010. . (Annonaceae) Eighteen species of fruits and/or seeds were collected, Passiflora edulis 2 2.12 56 and defecated undamaged the most seeds.frequent Although fruit recovered remains (Table of Arto 2)- a passion fruit (Passifloraceae) carpusIn most heterophyllus cases, the civet(s) ingested fruit pulp along with seeds Flacourtia jangomas 3 3.19 47 (Flacourtiaceae) were observed 13 times in faeces, seeds C. papaya, Ananas comosus were found on only four occasions, probably reflecting their 2 2.12 0 A. heterophyllus, A. hirsutus, Caryota urens and Annona squa- Pineapple mosalarge size. The five predominant fruits found were (Bromeliaceae) Syzigium cuminii 1 1.06 18 . Vertebrate matter, including rodents, birds, reptiles and Jamun (Myrtaceae) amphibians was found in seven faeces. Invertebrate remains- Psidium guajava 2 2.12 92 tained(insects, chilli millipedes, Capsicum beetles annuum and snail shell) were found in 11 Guava (Myrtaceae) Annonafaeces. Cooked squamosa rice was found in four faeces of which two con Ficus bengalensis 2 2.12 148 pericarp and seeds. Excepting Fig (Moraceae) (62.2%), all plant species showed higha direct rates Cucurbitaceae fruit 2 2.12 35 sightingof germination) of (Table 3). - Unidentified 1 1.06 17 During the study period, seven incidents (onerecorded . animals assumed to be Common Palm Civets tak- ing live chickens from unattended poultry sheds were Viverricula indica in the All incidents occurred between 18h15 and 19h30. In the di - rect sighting (18h25), the fowl escaped when the civet was that they never see threatened. In all other incidents, the animal killed the fowl area. Thus, attribution of these kills to Common Palm Civet re and only the carcass of the fowl was found, without head and- Discussionquires confirmation. neck portion. These were identified as Common Palm Civet Accordingkills because to thisthe farmers,mode of domesticattacking dogspoultry on itsfamiliaris head is andbe among the more frugivorous viverrids lieved by localFelis catuspoultry farmers to be peculiar to the species. Common Palm Civet is - stray usually tear the bird at the site itself and (Corlett 1998). No systematic studies on its diet have been carry the carcass away, while apparently carried out in India (Singh 1982, Krishnakumar & Balakris attack only chicks and not adult birds. The local farmers felt hanan 2003) but fruits have been recorded as its major food 15 Small Carnivore Conservation

, Vol. 45, December 2011 Jothish

Table 3. Germination percentage of seeds collected from Common Palm Civet Paradoxurus hermaphroditus faeces collected from a rural site in Kerala, India, during 2008–2010. Faeces with vertebrate and invertebrate remains showed- that the animal supplements its carbohydrate-rich fruit diet Species Number of Number Percentage of - with protein-rich animal matter. Su Su & Sale (2007), in My seeds sowed of seeds germination mains in the diet of P. hermaphroditus - anmar, found only very few vertebrate and invertebrate re germinated Bekele et al . Cooked rice in the fae Carica papaya 78 69 88.46 ces showed the scavenging natureCivettictis of the civetta Common in Ethiopia Palm Civet. and Artocarpus 10 10 100.0 . (2008) observed high rate of scavenging in human ­heterophyllus habitats by the Artocarpus 40 36 90.00 Balakrishnan & Sreedevi (2007) observed that faeces of Small ­hirsutus Indian Civet collected near human habitats often containing Annona 45 28 62.22 cooked rice and fish bone. - ­squamosa In the present study, no seasonal dietaryet alshift between Caryota urens 30 22 73.33 fruits and animals was observed, in contrast to studies in Ne pal and Thailand (Rabinowitz 1991, Joshi . 1995). The English names and botanical families for each species are given in Table 2. year-round availability. of one or more types of fruits in the studyAll area seven might explain the absence of seasonal dietary shift et al - in this part of Kerala poultry-raiding incidents during the two-year in countries such as Nepal (Joshi . 1995), Thailand (Rab period occurred between 18h15 and 19h30, the time that inowitz 1991, Grassman 1998) and Myanmar (Su Su & Sale civets leave their day roost. The civet may have drunk the- 2007). A recent investigation of Common Palm Civet feeding blood by removing the head, as there were no signs of blood- ecology in semi-urban. Joshihabitats et al in Trivandrum revealed fruit on the incident site or neighbouring areas. A recent observa as the predominant (about 82%) component (Krishnakumar tion of BalakrishnanTurdoides & Sreedevi (2007) on the stomach con & Balakrishnan 2003) . (1995) observed that fruits tent of Small Indian Civets showed that only the head region constituted about 84.5% of the total faeces, with only 15.5% of a babbler (a bird) was chewed and feathers and animal matter in Nepal. The present observations corroborateA. hirsutus, body were left intact. Similarly, in all these attacks apparently Ficusthe earlier and Mangifera studies. Balaramapuram indica being a rural area, every by Common Palm Civet, the headless carcass of the bird was home yard has one or more fruit plants. Excepting found. The animal perhaps leaves the body of poultry because , these fruit plants have no specific of its weight.et al Common Palm Civets become active around time of fruiting, so fruits were available all year. - 18h00 and activity decreases later in the night (Rabinowitz- Most faeces contained a single fruit species/food item, 1991, Joshi . 1995, Su Su & Sale 2007): thus, in the present showing that the civet(s) ate a single source in bulk at a par- study, the civet raided poultry in its peak activity time. Al ticular feeding time. The frequent occurrence of grass leaves in- though the species responsible for this raiding behaviour are the faeces (10.6% of total faeces) is consistent with other stud menace.not confirmed, the local belief that Common Palm Civet is the ies, mostly concluding a possible role of grass leaves in scour culprit means that the poultry farmers consider the species a ing the intestineet and al. in2010). the digestion process (Grassman 1998, et al. 2010). When an animal ingests fruits, Krishnakumar & Balakrishnan 2003, Balakrishnan & Sreedevi Frugivorous carnivores may disperse seeds (Herrera- 2007, Mudappa et al. 1989, Nakashima 2006)Common. Palm Civet(s) ate at least 18 fruit species in andthe successfulgut passage dispersal and the movementof the seeds of depends animals. onThe feeding seeds colbe- the present study, mostly from non-native plants (Nayar lectedhaviour from of the the frugivores, faeces were the undamaged seed viability and after did notconsumption lose their Krishnakumar & Balakrishnan (2003) identified only- - 10 fruit species from Common Palm Civet faeces in two semi- Pagumaurban habitats larvata in Trivandrum, Grassman (1998) 13 fruit spe viability after the gut passage.et al Daily movement patterns of civ cies in faeces pooled from Common and etets al in the present study area were not studied, but elsewhere in Kaeng Krachan National Park (Thailand) (Rabinowitz 1991, Joshi . 1995, Su Su & Sale 2007, Koike and Su Su & Sale (2007) about 31 Caricatypes of papaya fruits wasin Common the pre- . 2008) Common Palm Civets move long distances so may Palm Civet faeces analysed from Hlawga, Myanmar, with little transport seeds equivalently. Carnivores may produce seed seasonal fluctuation in totalCarica intake. papaya is a common fruit tree shadows differinget al. 2010).qualitatively Even though and quantitatively faeces collected from in those this dominant fruit found in the present study, as by Krishnakumar produced by other dispersers such as birds and primates- & Balakrishnan (2003). (Nakashima found in almost every home yard in Kerala, which fruits year study were from a terrace and thus have low chance of germi seedsround; of and Artocarpus civets have heterophyllus easy access, toA. these hirsutus fruits., Annona Most faecessqua- nation in situ, other sites that the animal(s) defecated at may mosacontaining and Caryota papaya urens remains showed also hadhigh viableseed germination seeds. Similarly, rate well have been more suitable for germination. It is plausible Artocarpus to assume that Common Palm Civets in the study area may be rural and forested parts of acting as a disperser of fruit plants in the locality. - (see Table 3). Both species are abundant across In this study the faeces were collectedMoreover, from only although a single Kerala. Besides providing fruits, locality and may have all come from only one animal, forestall they are important timber species. By defecating viable seeds, ing firm generalisation to the regional diet. Common Palm Civets may help the effective dispersal of these faecal analysis proved the consumption of some species, some economically important trees. dietary components may have been overlooked if they are well Small Carnivore Conservation 16

, Vol. 45, December 2011 Common Palm Civet diet in Kerala, India

Paradoxurus hermaphroditus P.digested hermaphroditus or otherwise leave no readily identifiable remains. Krishnakumar, H. & Balakrishnan, M. 2003. FeedingSmall Carnivore ecology of Con the- Nonetheless, the present study provides a clue about the diet of Commonservation Palm Civet (Pallas) in in rural India and its role in seed dispersal of semi-urban habitats of Trivandrum, India. of Thailand . Association Acknowledgementseconomically important plants. 28: 10–11. The author thanks the reviewers and the editor for their critical and Lekagul, B. & McNeely, J. A. 1977. useful comments and suggestions that helped a lot to revise this pa- for the Conservation of ParadoxurusWildlife, Bangkok, jerdoni Thailand., a viverrid endemic per. Mudappa, D., Kumar, A. & Chellam, R. 2010.Tropical Diet and Conservation fruit choice Sci of- encethe References to the rainforest, India. . 3: 282–300. Nakashima, Y., Inoue, E., Inoue-Murayam, M. & Abd. Sukor, J. R. 2010. Balakrishnan, M. 1997 Mammalian diversityProceedings in Kerala: of the a ninth sustainable Kerala Functional uniqueness of a smallOecologia carnivore as seed dispersal Science­resource Congress for human welfare. Pp. 454–459 in Iyenkar, P. K., Nair,- agents: a case study of the Common Palm Civets in the Tabin K. S. S. & Pushpangadan, P. (eds) FloweringWildlife Reserve, plants Sabah,of Kerala Malaysia. – a handbook 164: 721–730. - . Kerala State Science, Technology and Environ Nayar, T. S., Rasiya Beegam, A., Mohanan, N. & Rajkumar, G. 2006. ment Committee, Thiruvananthapuram,Viverricula indica India. . Tropical Botanic Gar- Balakrishnan, M. & Sreedevi,Small M. B. Carnivore2007. Husbandry Conservation and management of den and Research Institute, Thiruvananthapuram, India. the Small Indian Civet (E. Geoffroy Saint-Hillaire, Patou,Paradoxurus M.-L., Wilting, A., Gaubert, P., Esselstyn, J. A., Cruaud, C., Jen 1803) in Kerala, India.Civettictis civetta 36: 9–13. Journalnings, A. of P., Biogeography Fickel, J. & Veron, G. 2010. Evolutionary history of the Bekele,forest, T., Afework, Ethiopia. B. Small & Balakrishnan, Carnivore Conservation M. 2008. Feeding ecology of palm civets – a new model for Asian biogeography.Para- the African Civet in the Menagesha-Suba State doxurus and Paguma 37: 2077–2097. Journal of Bombay (Viverricula indica, Herpestes urva and Melogale 39: 19–24.moschata) in Pocock,Natural R. I. 1933.History The Society palm civets or toddy cats of the genera Chuang, S.-A. & Lee, L.-L. 1997. Food habitsJournal of ofthree Zoology, carnivore London species The book of inhabiting Indian animals British India. 36: 855–877. Fushan Forest, northern Taiwan. 243:- Prater, S. 1971. , 3rd edn. Bombay Natural ison71–79. of 3 methods. Journal of Wildlife Management History Society and Oxford University Press, Bombay, India.Journal Corbett, L. K. 1989. Assessing the diet of Dingoes from feces: a compar Rabinowitz,of Zoology, A. R. London 1991. Behaviour and movements of sympatric civet Biological 53: Reviews 343–346. species in Huai Kha Khaeng wildlife sanctuary, Thailand. Para- Corlett, R. T. 1998. Frugivory and seed dispersal by the vertebrates doxurus hermaphroditus 223: 281–298.Journal of Bombay Natural His- in the Oriental (Indomalayan) region. 73:- Singh,tory L. A.Society K. 1982. Stomach contents of a Common Palm Civet 414–448. (Pallas). Grassman,Small CarnivoreL. I. 1998. Conservation Movements and fruit selection of two Paradox Paradoxurus 79: 403–404. hermaphroditus Viver- urinae species in a dry evergreen forest in southern Thailand. Su Suricula & Sale, indica J. 2007. Niche differentiation between Common SmallPalm 19: 25–29. CarnivoreCivet Conservation and Small Indian Civet Herrera, C. M. 1989. FrugivoryOikos and seed dispersal by carnivorous in regenerating degraded forest, Myanmar. mammals and associated fruit characteristics in undisturbed- 36: 30–34. Mediterranean habitats. 55: 250–262. Zhou, Y. B., Zhang, J., Slade, E., Zhang,Paguma L., Palomares, larvata F., Chen, J., Wang, Joshi,civets. A. R., JournalSmith, J. of L. Mammalogy D. & Cuthbert, F. J. 1995. Influence of food dis JournalX. & Zhang, of Mammalogy S. 2008. Dietary shifts in relation to fruit availability tribution and predation pressure on spacing behaviour in palm among Masked Palm Civets ( ) in Central China. 76: 1205–1212. Tropical Botanic Garden 89: and 43–437. Research Institute, Palode, Koike,cool-temperate S., Morimoto, deciduous H., Goto, Y.,forests. Kozakai, Journal C. &of Yamazaki,Forest Research K. 2008. Thiruvananthapuram – 695 562, Kerala, India. Frugivory of carnivores and seed dispersal of fleshy fruits in Email: [email protected] 13: 215–212.

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