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COMMUNICATIONS FROM THE MAMMAL SOCIETY-NO. 63 REFERENCES
Askew, R. R. & Yalden, D. W. (1985). The Woodchester Park Valley. In Case studies in population biology: 1-26. Cook. L. M. (Ed.). Manchester: Manchester University Press. Chitty, D. & Kempson, D. A. (1949). Prebaiting small mammals and a new design of live trap. Ecology 30: 536-542. Gurnell, J. & Flowerdew, J. R. (1982). Lire trapping small mammals. A practical guide. (Mammal Society, London.) Montgomery, W. I. (1979). An examination of interspecific sexual and individual biases affecting rodent captures in Longworth traps. Acta theriol. 24: 35-45. Stoddart, D. M. (1982). Does trap odour influence estimation of population size of the short-tailed vole, Microtus agrestis? 1. Anim. Ecol. 51: 375-386. Stoddart, D. M. & Sales, G. D. (1985). The olfactory and acoustic biology of wood mice, yellow-necked mice and bank voles. Symp. zool. Soc. Lond. No. 55: 117-139. Stoddart, D. M. & Smith, P. A. (1984). Wood mice (Apodemus syluaticusi can distinguish con specific from heterospecific odors in the field. J. Chem. Ecol. 10: 923-928. Stoddart. D. M. & Smith, P. A. (1986). Recognition of odor-induced bias in the live-trapping of Apodemus sylraticus. Oikos 46: 194-/99.
J. Zool., Land. (]991) 225, 662-665
Simultaneous suckling and suckling bout lengths in Oryx dammah
Department of Zoology, University oj Cambridge, Downing Street,
Analysis of data on lactation, during a study on the social behaviour of captive scimitar-horned oryx females (Oryx dammah), revealed an unexpected relationship between the degree of suckling synchrony observed and the duration of suckling bouts. Activity synchronization and social facilitation have been the subject of several studies (e.g. dogs, Scott, 1968; lactation in pigs, Kilgour, 1978; captive wildlife, Altmann, 1985; domestic stock, reviewed by Craig, 198]). In these studies, however, the duration of the activity concerned was not analysed in respect to the presence or absence of a simultaneously performing conspecific. In this report, I describe the duration of suckling bouts in oryx antelopes as a function of simultaneous suckling by nearby females. The suckling behaviour of three scimitar-horned oryx females was studied for 13 days in June ]987 at Whipsnade Zoological Park, England, where they were kept in a 1·13 ha enclosure together with seven non-lactating females, The oryx had free access to a sheltered yard. During the study period, the main food intake occurred in the open pasture of the enclosure. At the start of the observations the calves of the females F 1, F2 and F3 were 31,28 and nine days old, respectively. All calves were males. The herd was observed daily for two hours in the morning and six hours in the afternoon during which all suckling events were recorded. A suckling bout was regarded to have started if the calf had had contact with the udder for more than three seconds. The bout was considered as terminated when either the mother walked away (40 cases) or the calf stopped sucking for over 10 seconds (four cases). After an interruption of more than 10 seconds further solicitations by the calves were generally rejected. Mothers terminated the bouts by walking away more often than the calves by stopping sucking (Binomial-test, P < 0'00]). The duration of brief
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interruptions within one suckling bout were excluded from the bout duration. The suckling bout length only included time in contact with the udder. Simultaneous suckling events were defined as those in which the suckling bouts of two females overlapped. Isolated suckling bouts were defined as those which were neither preceded nor followed by another female's suckling within 30 min of the start of the bout. The mean suckling bout length was 60·2 sec (S.D.=31·7, range 3--126 sec, median=55·5 sec, II = 54, three females pooled). The duration of a suckling bout was positively correlated with the time elapsed since the last bout by the same female (Spearman's r=0'55, n= 19, P<O·OI). The mean interval length between two suckling bouts by the same female was 105·8 min (S.D. = 67,3, range 3-253 min, median = 98·0 min, n = 27, three females pooled). Neither the mean suckling bout length nor the mean interval between suckling bouts differed significantly between the females (two-sample r-test). Between two suckling bouts, calves spent most of the time hiding amidst bushes or in the tall grass (Tying-out' sensu Lent, 1974). The two older calves were still observed to lie-out at the age of eight weeks. The time interval between two consecutive suckling bouts by different females was in 61 % of the cases less than 5 minutes. The distribution of interval lengths (Fig. 1) differed significantly from a negative exponential distribution (X2 = 8· 356, df, = 2, P < 0'02). The fact that short intervals were over-represented among the observed values indicates that suckling bouts tended to be synchronized (Fig. I). The expected frequency of occurrence n of any interval between t and t+ w, given a random distribution of suckling bouts between females, was calculated from n = Nwl'F exp (- tiT), where N is the total sample size (31 observations), w is the category width (5 min), and Tis the mean observed interval length (10' 56 min). Out of 68 recorded suckling events 10 (15(/'0) occurred in the context of sim ultaneous suckling. Visual contact between the females was possible in all instances of simultaneous suckling. The simultaneously suckling females did not start or finish suckling at the same time. In general, as one
20 _ Observed
25 30 35 40 45 Length of interval (min)
FIG. 1. The synchronizaIion of suckling events. The histogram shows the frequency distribution of 31 time intervals between the suckling events of different females. A negative exponential distribution was used to calculate the expected values under the null hypothesis that suckling events by different females were randomly distributed in time.
COMMUNICATIONS FROM THE MAMMAL SOCIETY-NO. 63
The duration of suckling bout lengths as a function of the presence or absence" ofa simultaneouslv such ling female
Mean suckling bout length (sec)
(I) Three females during
simultaneous suckling events (2) Three females during isolated suckling events
Two sample r-test between (I) and (2). P < 0-002; d.f,
female was already suckling her calf another female started suckling hers close by. Suckling bouts during simultaneous suckling were almost twice as long as during isolated suckling events (Table I). Both the first and the second female prolonged their bout durations considerably above average, and finished suckling with a time delay which roughly corresponded to the delay between the start of the bouts. Simultaneous suckling bouts were also significantly longer than consecutive bouts occurring with no overlap but starting within five minutes of each other, as shown by comparing the bout length of the first female to start suckling in overlapping (mean = 99·2 see, S.D. = 23· 3, n = 5) and non-overlapping (mean = 58·7 sec, S.D. = 23,6, II = 10) sequences (two-sample z-test, P < 0,05, d.f, = 8). This difference in duration was not related to the activity performed prior to suckling. Neither is it explained by the positive correlation between time elapsed since the last bout and the duration of the suckling bout (see above), because there was no difference in the time elapsed since the prior suckling bout for simultaneous or isolated suckling bouts. The results suggest that scimitar-horned oryx increased their suckling bout length as a consequence of another female suckling simultaneously. On six occasions, as one calf started suckling, another immediately solicited milk from its own mother, but was effectively rejected by her. In these cases, the second calf had not had milk for at least 45 min. Female F2 accounted for four of these cases of rejection and female Fl for two. These observations indicate that, in spite of the solicitations of the calf and the current suckling by a nearby mother, female scimitar-horned oryx sometimes preferred not to start a simultaneous bout. The positive association between simultaneous suck ling and the duration of suckling bouts was unexpected. The adaptive significance of this behaviour is unknown. The following post hoc explanation is presented as a working hypothesis for future studies. The fact that suckling bouts during simultaneous suckling were almost twice as long as during isolated suckling events may be related to antipredatory behaviour. Some activities such as suckling young, giving birth and drinking at a water hole, appear to make ungulates particularly vulnerable to predator attacks. Close proximity to neighbouring conspecifics during such activities and, in particular, to individuals performing the same risky activity should decrease the average probability for each individual of becoming the prey, following the reasoning of Hamilton (1971) and Bertram (1978). The duration of a risky activity is generally constrained by an increasing risk the longer the activity is performed. A reduction of the vulnerability during such an action, either through the dilution effect or through increased vigilance in the group (Bertram, 1980), should allow an individual to prolong its duration. The assumptions above being true, one could predict that risky
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activities will have a longer duration when performed simultaneously by two neighbours than when performed by a single individual. Under the assumption that suckling is a risky activity, particularly for the calf, the longer duration of simultaneous suckling reported here is in accordance with that prediction. Two observations, however, appear to question this interpreta tion. First, if simultaneous suckling was advantageous, then one could ask why it did not occur more often, considering the overall synchronization of suckling. Secondly, in six cases a female appeared actively to avoid suckling simultaneously (see above). Replication of the approach presented and a field test of the proposed hypothesis are needed before any conclusions about the functional significance of the reported effect can be drawn.
I am grateful to the Zoological Society of London, to V. G. A. Manton, the Curator ofWhipsnade Zoological Park and to the keepers of the oryx section, Allan, Neil, Collin and Graham for permission to work in the zoo and providing invaluable logistical support, as well as information on the study herd. I thank Keith Lindsay and an anonymous reviewer for their comments on the manuscript, and Keith Eltringham for encouragement and the patient supervision of this study.
REFERENCES Altmann. D. (1985). Mach-mit-Verhalten bei Tieren. Milu, Berlin 61/2,3-10.
Bertram. B. (1978). Living in groups: predators and prey. In Behovioural ecologv: An etolutionarv approach: 64-96. 1st edn.
Krebs, J. R. & Davies. N. B. (Eds). Oxford: Blackwells. Bertram, B. (1980). Vigilance and group size in ostriches. Anim. Behar. 28: 278-286. Craig.T, V. (1981). Domestic anima! behaviour: causes and implicationsfor anima! care and management. Englewood Cliffs, New Jersey: Prentice Hall. Inc. Hamilton, W. D. (1971). Geometry for the selfish herd. 1. theor. Bio!. 31: 295-311. Kilgour, R. (1978). The application of animal behaviour and the humane care offarm animals. J. Anim. Sci. 46: 1478-1486. Lent, P. C (1974). Mother-infant relationships In ungulates. IUCN Publs (N. S.) No. 24: 14-55. Scott.T. P. (1968). Social facilitation and allelornimetic behavior. In Socia! facilitation and imit atite hehorior: Outcome of the 1967 Miami Unirersity symposium on social beharior: 55- 72. Simmel, E. C. Hoppe. R. A. & Molton, G. A. (Eds). Boston: Allyn & Bacon Inc.
J. Zool., Lond. (1991) 225, 665-670
The diet of Pipistrellus pipistrellus during the pre-hibernal period
L. R. HOARE*. Department of Science, Christ Church College, Canterbury, Kent
All British bats are insectivorous and insect abundance varies with, and depends on. climatic conditions. Summer densities of insects may provide sufficient food for bats and other insectivores. but in times of reduced densities in the prc-hibernal period, competition between bats and other
* Present address: Durrell Institute of Conservation and Ecology. The University. Canterbury. Kent CT2 7NX