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The Use of Acoustical Cues for Prey Detection by the Indian False Vampire Bat, <Emphasis Type="Italic">Megaderma

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The Use of Acoustical Cues for Prey Detection by the Indian False Vampire Bat, <Emphasis Type= ,Journal of J Comp Physiol A (1987) 160:509 515 6ensory, Comparative and..~.,, Physiology A BehavioralPhysiology Springer-Verlag 1987 The use of acoustical cues for prey detection by the Indian False Vampire Bat, Megaderma lyra G. Marimuthu 1 and G. Neuweiler 2 1 Department of Animal Behaviour, School of Biological Sciences, Madurai Kamaraj University, Madurai 625 021, India 2 Zoologisches Institut der Universit/it Miinchen, LuisenstraBe 14, D-8000 Miinchen 2, Federal Republic of Germany Accepted December 18, 1986 Summary. 1. The response of the echolocating bat, capture prey from ground and water surfaces. Me- Megaderma lyra, was tested to different kinds of gaderma lyra eats the larger arthropods and verte- prey in an outdoor cage. The bats caught larger brates of suitable size e.g. frogs, geckoes, lizards, flying insects (moths, beetles, grasshoppers, and fish, mice, birds and other bat species (Advani cockroaches) on the wing and also picked up ar- 1981; Habersetzer 1983). M. lyra is a large bat thropods (solifugid spiders, beetles and cock- (bodyweight 30-38 g) with large, medially fused roaches) and small vertebrates (mice, fishes, frogs pinnae which form a unified auditory receiver di- and geckoes) from the ground. After touching the rected towards the ground. This species is also an prey with the muzzle, the bats were able to differ- excellent flyer capable of hovering and even of fly- entiate between species. Scorpions and toads were ing backwards. The False Vampire emits very brief not taken by M. lyra. (0.4-1.0 ms) broadband echolocation signals of 2. In lighted and in dark conditions, M. lyra low intensity (less than 80 dB SPL) consisting of detected and caught prey only when it moved. 3-6 harmonics, which cover a frequency range of Dead frogs briskly pulled over the floor were also 20-120 kHz. During obstacle avoidance and when detected and caught, whereas stationary dead frogs snatching mealworms in flight from forceps, M. were disregarded by the bats (Table 1). lyra consistently emits echolocation signals, even 3. When dead frogs were pulled over the wa- in bright daylight (M6hres and Neuweiler 1966). tered surface of a glass plate to eliminate noises However, in a more natural situation when M. lyra by motion, the motion no longer alarmed the bats. is required to detect and catch prey in complete From the results of these experiments it was con- darkness, it is capable of flying towards the target, cluded that M. lyra detects prey on the ground locating and successfully capturing it, often with- by listening to the noise of the moving target only, out emitting a single echolocation sound (Fiedler and not by echolocation (Table 1 C, Fig. 1). Fur- 1979). M. lyra are also attracted by tape-recorded thermore, M. lyra were not attracted by frog calls. rustling noises produced by moving mice on the 4. M. lyra differentiated between palatable floor, and will even attack the broadcasting loud- frogs and non-palatable toads only after touching speaker. the prey with the muzzle. In the natural habitat Macroderma gigas, the 5. Experiments with freshly killed frogs coated Australian relative of M. lyra, is attracted by a with toad secretions or covered with toad skins Audubon bird caller, and individuals may fly as indicate that M. lyra differentiates between frogs close as a few centimetres around the head of the and toads by chemical means. There was no evi- experimenter (Coles and Neuweiler, unpublished dence that these prey were differentiated by means observations). Antrozouspallidus, another gleaning of echolocation. bat species, is also attracted by the noises of prey moving on the ground and does not emit echoloca- tion signals during the approach and attack of prey (Bell 1982). Introduction All the above observations suggest that glean- The Indian False Vampire belongs to a heteroge- ing bats may detect and locate prey on the ground neous group of echolocating bats which are called facultatively or exclusively by listening to rustling gleaners (Fenton 1984). Gleaning bats prefer to noises produced by the targets, without the aid 510 G. Marimuthu and G. Neuweiler: Prey detection by Megaderma lyra of echolocation. The present study therefore re- potential prey started to move the bats directed ports observations and experiments on the prey- their heads towards the target. A few seconds later catching behaviour of M. lyra, in an attempt to the bats flew off, briefly hovered over the target answer the question of the sensory mechanisms by and then descended on the prey, grasping it in the which this gleaning bat detects prey. mouth. Then the bats immediately flew from the ground and returned to the roost to consume the prey. The freshwater crabs which ran over the floor Materials and methods were caught without hesitation in eight trials. In- All observations and experiments were performed in an outdoor terestingly, three other crabs were approached re- cage in the Botanical Garden of Madurai University, India, peatedly but not caught by M. lyra. After removal and the bats were maintained in the natural climate and diurnal of their claws, when these crabs started to run cycle. The flight cage measured 7.5 x 3.4 x 3.5 m and contained again they were quickly caught. It is not clear why a dark enclosure at one corner of the ceiling so that the bats could seek refuge during the day. On the floor of the flight the bats refused to catch clawed crabs in these three cage, an artificial pond was dug (4.15 x 2.30 x 0.64 m) in which cases whereas the other eight crabs were caught frogs were maintained. A total of 24 M. lyra (11 males and with intact claws. In this context, it is well known 13 females) were caught at the caves at Keela Kuilkudi and that the megadermatid bat Cardioderma cor can Pannianmalai, about 10 km from the University campus and prey successfully on poisonous scorpions housed in the outdoor cage. The bats were fed daily between 18 and 22 h, which corresponds to their natural activity period. (Vaughan 1976). However, M. lyra was observed The food for the bats consisted of frogs, mice and cockroaches, to land close to scorpions but never touched them, depending on availability. returning to the roost. All experiments and observations were conducted during Most of the vertebrates that the bats preyed the evening up to midnight using room light. The use of visual cues for prey detection was excluded by repeating all experi- upon were introduced into the cage by the experi- ments in darkness and monitoring the bats with an infrared menter. However, two geckoes, one running over night vision scope. the floor and another along a wall, were detected During each experiment, echolocation sounds of M. lyra quickly and captured by a single M. lyra. In further were detected by a QMC-ultrasound microphone and recorded on a high-speed tape recorder (Lennartz, 76 cm/s). tests, seven field mice (Mus booduga, bodylength up to 7 cm) and 15 laboratory mice placed on the floor of the cage were detected and caught by the bats within 1 min. Usually the bats grasped a Results mouse at the front and carried it off to the roost. There the bat manipulated the mouse with the help Prey-catching behaviour of the wrists in order to bite the ventral neck. The The initial observations concerned conditions bats then closed the jaws firmly around the throat under which M. lyra became alerted to and caught of the mouse until it suffocated. Also, three live prey that moved spontaneously in the cage or was fish (Ophiocephalus spp., bodylength 8 cm), which introduced by the experimenter. It was found that had been thrown on the floor and were flipping the bats were able to catch flying insects as well their tails, were caught swiftly and consumed by as arthropods and vertebrates moving on surfaces. M. lyra. Even when freshly killed bats (Hipposider- Flying insects were intercepted aerially, and after os speoris and H. bicolor) had been thrown on to capture the bats returned to their roost with the the floor of the cage, the bodies were immediately prey in the mouth and consumed it at the roost. picked up and consumed completely by M. lyra. A total of five moths, three beetles and one cock- Frogs were found to be a preferred food item roach with bodylength ranging between 2 and 4 cm for M. tyra, and as in all other observations, frogs were caught in this way. The bats emitted echolo- were detected and caught only when moving or cation sounds continuously during these insect after jumping, usually with no chance of escape. capture flights in the dimly lit outdoor cage. Experiments with the South American frog-eat- In addition, beetles, cockroaches and grasshop- ing bat Trachops cirrhosus have shown that frog pers were caught whilst walking or jumping on calls might attract bat predators (Tuttle and Ryan the floor of the cage. Furthermore, four solifugid 1981). On three occasions in the present study spiders and eleven freshwater crabs (bodylength frogs in the artificial pond of the cage called loudly approx. 2.5 cm) were caught quickly when they for several minutes. However, M. lyra showed no started to run over the floor. There was no indica- reaction to long duration and loud calling even tion that the bats paid any attention to stationary though the frogs were only a few meters away. arthropods, even if they were only a few centi- As soon as these frogs moved or jumped they were metres from their heads. However, as soon as these detected and caught immediately by the bats.
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