Annals of Plant Sciences I SSN: 2287-688X OPEN ACCESS www.annals ofplantsciences.com Original Research Article Foraging Behaviour of Indian Flying Fox P. giganteus in Prasad ER*, Dileep P, Aryasree MP, Anjana K and Sunojkumar P Department of Botany, Univers ity of C alic ut, Malappuram, Kerala-6 7 3 63 5 , .

Received f or publication: September 0 5 , 20 1 4 ; Accepted: October 17, 2014.

Abstract: Over one year period, field study was conducted for the identification of food plants and s eed dispersal of giganteus by direct observations or indirectly by the analys is of ejec t found on ground. Observation were made on seven colonies of day roos t in undis turbed areas , amongst which six of them were in sacred grooves one in the protected area. O ver 2 2 different plant seeds were collected from the ejected materials as chewed , fres h s eeds or faec al materials. The result indicates that Pteropus is playing a vital role as a seed disperser in plants like Anacardium occidentale, Areca catechu, Manilkara achras, and Psidium gujava whic h are highly demanding plants in the market. The study concluded the mutual relations hip between plants and , flying foxes depends plants for food and roost while plants need bat as dis pers er and pollinator.

Key Words: P. giganteus , Kerala, field s tudy

Introduction Feeding behaviour of was The treatment on smaller fruits a gain influenced by fruit traits and it varies a cross depends o n bat size, smaller bats usually bat species. Feeding behaviour of fruit bat pluck a fruit, fly away to nearby fe eding was greatly influenced by fruit quality, roosts (20-200m away from the fruiting tree) especially, the fruit hardness. It may also be where they process the fruits (Marshall 1983; expected to vary significantly among bats and Phua and Corlett 1989) such fe eding roosts to be an important aspect of their resource are known as night roost. However, larger bat partitio ning strategies (Dumont 1999; Aguirre species seem more likely to process fruits in et al., 2002). Fruit bats have an unusual the fruiting tree itself. At fe eding areas, mode of feeding in which they consume the pteropodids usually defend small territories of fruits in small bite s, pressing out and fruits or in the canopy (Neuweiler swallowing the juice and spitting o ut rind a s 1969; Gould 1977; Richards 1990; Banack compact pellets (Morrison 1980). 1998; Brooke 2001). There is little or no may or may not remove fruits from a tree and report on Indian flying fox P. giganteus about they typically cling on to flowers a nd fruits, the resource defe nse. while lapping up or ingesting soft pulp and fruit juices (Fleming 1982). T hey may Materials and Methods consume large fruits in situ (Van der Pijl The fruits and leaves consumed by P. 1957). Ho wever, smaller fruits a re usually giganteus were collected once in a fortnight in carrie d away from the trees to special feeding roosting as well as in the foraging areas. The roosts fo r consumption (Ela ngovan et al., collected fruits and le aves were identifie d 1999). The strategy be tween bat species in taxonomically and a lso the discarded parts their treatm ent on the same fruits is were recorded. The weight of the fruits influenced by bat size. A larger Pteropus sp. carrie d to the roosting area is also me asured can fly off with a whole while a smaller using balance. Am ong the discarded fruits the sp. simply bites off pieces of the economically important fruits to the humans flesh in situ (C orlett 1998). There are were recorded. relative ly fe w data concerning the feeding behaviour of the Indian flying fox Pteropus The fo raging a ctivitie s o f these fruit giganteus and the already known information bats were also m onitored in sele cted trees on this species is mostly centered on foraging like Bassia latifolia, Ceiba pentandra and Ficus trees (McCann 1934, 1941) and behaviours religiosa. The fo raging activitie s of P. they exhibit at day roosts (Neuweiler 1969). giganteus were monitored from 18h to 6h until the returning of the bats from the *Corresponding Author: Prasad Erancheri, Research Scholar, Department of Botany, University of Calicut, Malappuram, Kerala-673 635, India. Page| 883

Pras ad ER et al., Annals of Plant Sciences , 2014, 3 (11), 883-887 fo raging trees. These bats were o bserved and P. guajava these bats came a round 20h using red filtered torchlight since it gives little and spent about 10 to 20 minutes in the tree or no disturbance to the foraging bats. The and removed the fruits and move to the number of bats visiting a particular tree was nearby large trees for feeding. Once carrie d counted using mechanical counter. Since the to the fruit it started consuming the fruits bit bat spent most time in the foraging tree the by bit and chews a nd spits o ut the fibrous number of bats present in the foraging tree remains. It uses its fo rearm to m ove the fo r every 30 minute s were recorde d. In the fruits to desired direction. It usually fo raging tree whether the bat fe ed the fruits consumes entire fruit of ripe m ango e xcept in the same tree o r carrie d the fruits to the the seed but it consumes the outer fleshy part neighbouring feeding roosts were monitored. of guava and voids the inner seeded part. The interspecific and intraspecific interactions After consuming the entire fruit it grooms of these bats in the foraging trees were a lso itself and taken rest for 10 to 15 minutes. The recorded. fe eding roosts used by these ba ts a re Albizia lebbek, Tamarindus indica, Borassus Results flabelliber, Azadirachta indica, Syzigium Fig trees, plantain field, and guava and cumini and Eucalyptus spp. Feeding roosts sapota orchards surround most of the P. used by these bats located usually next to the giganteus roosting places in Kerala. So they fo raging tree some time it may located 10 to get year round supply of fo od. These bats 200m from the feeding tree. Feeding roosts consumed fruits, nectar and leaves o f about are individual specific and e ach bats has its 22 species of plants in the study area (Table own specific fe eding roosts. 2). The die t of P. giganteus consisted predominantly the fruits of fig trees. During Table 1: Foraging plants the le an period it also consumed the leaves of S. No Name of the plants Parts consumed 1 Manicra achras Fruits Ficus religioa and Erythrina indica. Mo st of the 2 Areca catechu Fruits fruiting trees visite d by these bats a re tall. 3 Artocarpus heterophilus Fruits Ho wever it also feed on fruits from shrubs like 4 Bassia latifolia Fruits Achras sapota, Psidium guajava and Musa sp. 5 Borassus flabellifer Fruits, (toddy plant) 6 Carica papaya Fruits which are m ostly grown in the o rchards. In 7 Ceiba pentandra Nectar the tall trees like F. religiosa, F. bengalensis, 8 Erythrina indica Leaves Syzigium cumini these bats forage in groups 9 Eugenia jambolana Fruits of about 10 to 12 individuals. But in shrubby 10 Ficus bengalensis Fruits 11 Ficus religiosa Fruits, leaves trees it fo rages solitarily. 12 Ficus virens Fruits 13 Ficus glomeratus Fruits The small fruits and flowers such as F. 14 Mangifera indica Fruits religiosa, F. bengalensis and B. latifolifa it 15 Musa paradisiaca Fruits 16 Parkia biglandulosa Nectar fe ed the flowers in situ i.e. in the same 17 Pithecolobium dulsi Fruits fo raging tree itself. But the larger fruits such 18 Polyalthia longifolia Fruits as Mangifera indica and Psidium guajava, P. 19 Psidium guajava Fruits giganteus removed these fruits from the 20 Tamarindus indicus Leaves 21 Tectona grandis Fruits parent tree and ex situ fo raging was 22 Terminalia catappa Fruits observed. Likewise it also removed the fruits of areca nut tree and feed its o uter rind a nd Discussion dropped beneath the day roost. These a reca Constantine (1970) reported that the nut were collected from day roost and sold in fruit-eating bats could survive only in the the local market and the bat bitten such fruits areas where fruits a re constantly available . are readily brought by local people. Some o f Sim ilarly my observation shows that the the big fruits like M. indica and P. guajava were partially consumed and dropped fo raging trees, which give year round, supply of fo od to these foraging bats surround m ost beneath the fe eding roost. In the fruiting of the roosting places of P. giganteus. season of ‘big bang’ trees like F. religiosa, F. According to Carpenter (1986) and T homas bengalensis and B. latifolia, P. giganteus (1975) the energy cost of flight of pteropodid arrive d to the trees at 19h a nd remained in and phyllostomid bats a re 15 tim es higher the same tree up to 3h consuming the fruits and flowers a nd rest at the same tree a nd than that of resting. So the bat must gather fruits, which give the energy necessary to some bats rest at neighbouring trees a fter cover the cost of flight, in addition to fe eding. But in the small trees lik e A. sapota www.annalsofplantsciences.com P age | 884

Pras ad ER et al., Annals of Plant Sciences , 2014, 3 (11), 883-887 maintenance and breeding e ffort (Charles- evolve d a strategy fo r both maxim izing Dominique 1993). My observation showed nutrient intake and minimizing gut retention that it consumed a bout 22 plant species in tim e and thus can be expected to reduce wing the surrounding area to meat out its e nergy loading and energy expenditure while in flight requirements. (Kunz and Ingalls 1994). R elative to fruits, which have patchy distributions in space a nd Fig fruits dominated the die t tim e (Fleming 1992; Terborgh 1986) le aves consumption of P. giganteus. Various spe cies of some tropical plant species are ubiquitous of pteropodid bats have been reported and available year round (Ganzhorn 1992). fo raging on the fruits o f more than 30 fig Plant visiting bats that sele ctively fe ed o n species in tropical and subtropical Asia, Africa le aves should gain a higher yield o f protein and Australia (Bhat 1994; Fujita a nd T uttle per unit of fo raging effort, compared to those 1991; Marshall and McWilliam 1982; Thomas that feed o nly on protein poor fruits (Kunz 1984). P. giganteus chews the fruits and spits and Diaz 1995). The le aves also contain out the fibrous remains along with the seeds higher amount of calcium compared to fruits in the same tree and sometim es consumes (O ’ Brien et al., 1998). Early report on low the whole fruit and defecates the seeds. Since survival rates fo r Old World fruit bats fo od transit tim e in frugivorous bats are maintained in captivity were attributed to relative ly rapid; generally le ss than 30 die ts that consisted largely of cultiva ted minutes (Laska 1990; Tedman and Ha ll fruits. However the diets supple mented with 1985). Several studie s have demonstrated green leaves and multivitamins increases the that se ed germination was either enhanced or survival rate of these bats (Rasweiler 1977). unaffected after passage through the So the le aves not only helpful as die tary digestive tract of bats (Figueiredo a nd P erin supple ment but also provide o ther useful 1995; Fleming and Heithaus 1981; Liberman nutrients to the foraging pteropodid bats. and Liberman 1986). The small sized fruits lik e F. religiosa P. giganteus was also observed and F. benghalensis were eaten in the same fo raging in other trees such as Bassia latifolia, tree (in situ fo raging) by P. giganteus. Ceiba pentandra and Mangifera indica during Whereas the larger fruits like Mangifera indica their flowering and fruiting season. This were taken away from the foraging tree (ex shows that it is an opportunistic fe eder. situ fo raging) to the feeding roosts by these Gardiner (1977) reported that most fruit bats bats. But in the smaller sympatric fruit bat C. are fo od generalists rather than fo od sphinx the fo raging pattern is e ntirely vice specialists. Sim ilar fe eding pattern was versa and my observation corroborate that of observed in the neotropical Carollia the fo raging pattern of these bats in e arlier perspicillata. It consumes the fruits o f about report (Ela ngovan et al., 2001). The main 37 plant species and it concentrates o n the reason for the ex situ foraging is 1. to reduce fruits of various piper species based o n their fe eding interference by other bats a nd 2. to availability (Fle ming 1982). reduce predation risk (Fle ming 1982). Transport of fo od to a fe eding roost is Since many fruits consumed by bats common in small sized bats; P. giganteus are rich in soluble carbohydrates and poor in mostly feed the small sized fruits in the same crude protein bats may need to o ver ingest tree. It is possible that the size of the fruits and dump carbo hydrate to m eet out Pteropus makes them vulnerable to predators their protein requireme nts (Thomas 1984). waiting at rich patches of food (Fenton et al., Ho wever plant visiting bats may supple ment 1985). their fruit diet with protein rich leaves (Kunz and Diaz 1995; Zortea and Mendes 1993) and In the ‘big bang’ trees like Bassia insects (Courts 1998). O ther than fruits P. latifolia, F. benghalensis and F. religiosa giganteus also consumed leaves of F. religiosa groups of P. giganteus visite d these trees and Erithrina indica to supple ment their during the flowering season and fighting a nd protein requirements. Folivo ry and frugivory vocalization between these bats for acquiring by wedging (ex traction o f liquid fraction by the resource rich patches. This is in che wing fruits and leaves a nd e xpelling the consistence with other Pteropus spp. (Brooke oral spat) combined with rapid food transit 2001). We have also observed that there is tim e allows bats to process greater amount of interspecific fight between the sympatric fruit fo od per unit time. Thus, some bats have bat C. sphinx. To avoid this fighting and www.annalsofplantsciences.com P age | 885

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