Aggressive behaviour of captive Bandicota bengalensis

Proc. Indian Acad. SCI. (Anim. Sci), Vol. 92, Number 3, May 1983, pp. 185-191 c> Printed in India Aggressive behaviour of captive Bandicota bengalensis 1. Introduction SHAKUNTHALA SRIDHARA and RV KRISHNAMOORTHY Departments of Vertebrate Biology and Zoology, University of Agricultural Sciences, G KV K, Bangalore 560 065, India MS.received 18 October 1982; revised 3 February 1983 Abstract. The aggressive behaviour rhythm of Bandicota bengalensis under different social conditions were studied in the laboratory. The number, intensity and duration of aggressive behaviour peaks varied depending on the number, sex and social status of the interacting conspecific Is. Significantly the peaks of aggression were recorded during the dark period. In heterosexual conflicts both sexes were equally belligerent. However in male interactions with more females, the former emerged dominant always. Similarly one male dominated another in male-male encounters. Confrontation amongst several males in the presence offemale indicated the existence of only alpha and omega males. Keywords. Bandicota bengalensis; aggressive behaviour; male dominance; alpha males; omega males. Circadian rhythms of rodents have been wen documented for behaviour under natural conditions (Marten 1973), for activity movement in cages and mazes (Barnett et ai1975), for feeding and drinking (Possidente and Birnbaum 1979) and for aggressive behaviour(landau 1975; Lerwill1977). Desynchronizationof both metabolic and behavioural rhythms of Peromyscus maniculatus occurred due to social stress (Parr and Andrews 1978). Similar dissociation of activity rhythms due to crowding has been observed by several others (Calhoun 1975; Nygren 1978). In-this article we report alterations in the rhythms of aggressive behaviour of lesser bandicoot rat, Bandicota bengalensis under different social situations. 2. Material and methods Bandicota bengalensis, weighing 0-500 g were housed singly in metal cages (35 x35 x 50 cm) and were fed on rat and mouse feed (Hindustan Lever, India) supple- mented with vitamins and vegetables once a week with ad lib supply of water. Photoperiod was regulated at 12 hr light and 12 hr darkness with the light period beginning at 0600 hours. For recording behaviourthe animals were transferred to an observation chamber, details of which have been describej earlier (Sridhara and Krishnamoorthy 1982). Aggressive behaviour of individual members during encounters between male-female, male-male, female-female, one male-two females and one male-three females were studied. Attempts tc study agonistic behaviour in groups involving' more than one male were unsuccessful as submissive males died within 24 hr after being grouped with dominant males. One member of the pairimale in pair Igroup combinationwas restricted to one portion of the chamber with the other portion housing the 185

1 2M. &crd-r1' I"ll 2 '2 3 LIGHT 9, 01. ; ((I(III!I (I 111116 0600 2400 h<'jr. 52 2 2:f = " i " I cnrcbn, r:erm, il: :I:...dT1IlJ o 4 60 401.!: E S2 3 01 I! I b =, ==rl I I I 1 I 01 0=,,--eO cd] 1 1 Ch-dTI I CD (9 (9 <l 60 2 1 o 5 40-6\ 1(1! d I I I II I I I! I I I I I I I 0600 10 1800 2400 I hrsl 0 1 I I I I I I 1 1I I I 1[b:b--.J I I 1 I I! I I 0600 10 1800 2400 Ihrs) Figures 1-5. Each histogram represents the oscillation of an interacting individual in the cage over a 24 hr period...- 00 0\ <:I) ;J:r s::... ::l i:r <:I)... ::l $::l.. ;;;-... ::l cc-e... '<::

Aggressive behaviour of captive B. bengalensis 187 second member I females. By removing the partition the animals were allowed to interact for 10 min during each hour of the LD cycle. The aggressive behavioural acts and postures namely threat, crawling under lover lunge, chase,attack, fight, boxing and bite (Grant and Mackintosh 1963) were scored for each member of the pair Igroup during these observations. The data were collected under natural daylight during day time and under dim red light during nights. Four replications were. carried out for each type of encounter over a seven day schedule and histograms were computed from the combined means of different individuals. The aggressiveness of either sex was established by student t tests. To recognise ranks of aggressive behaviour, if any, during group encounters, Kruskal-Wallis one-way analysis was carried out (Seigel 1956). 3. Results The results presented in figures I-S illustrate that aggressiveness of bandicoots, motivated through the presence of other conspecifics, is rhythmic, The rhythmicity is either bimodal or unimodal depending on the sex, status and number of interacting conspecifics. However, a 7-day long study indicated that the animals have shown a fairly consistent rhythmicity (9S% rhythms) in their behaviour. Males exhibited a single peak at 0400 hr in male-female encounters (figure 1), at 0100 hr in one male-three females interaction (figure S), but two peaks of equal intensity at 10 and 2100 hr in male-male confrontation (figure 2) and three peaks at 0100, 1300 and 2100 hr in one male-three females group encounter, the last mentioned peak being the most intensive (figure 4). The second male in male-male pair exhibited a single peak of aggressive behaviour at 1700 hr (figure 2). The female in male-female pair and one of the females in the female- female encounter displayed two peaks of aggression, at 0400 and 2100 hr in the former case (figure 1) and at 1300 and20 hr in thelatter(figure 2). Maximum aggressive behaviour was scored foj.: the second female of the female- female (figure 2) and third female of one male-three females encounter at 0100 hr(figure 5). One female of one male-two females conflict exhibited peak aggressive behaviour at 0400 hr while the second female shifted her intensity to 1900 hr (figure 4). Similarly the first and second females of one male-three females interaction spaced out theirpeak aggressive behaviour rhythm at 20 and 1800 hr (figure S). Majority of peaks for aggressive behaviour occurred during the dark phase. Only one of the males of the male-male pairand one of the females of the femalefemale pair displayed afternoon peakswhile the othermale of male-male pairand second female of one male-three females interaction displayed intensive aggressive behaviour at twilight hours. Comparison of aggressive behaviour between the sexes indicated male and female as equally belligerent (P< O.OS, table 1) in male-female clashes while in male-male and female-female clashes, one of them established dominance over the other (P<O.Ol and <0.02, table 1). The male was significantly more aggressive than females in encounters involving more than onefemale (p < 0.01, table 1). No rank order of aggressiveness could be established in the two group clashes (H=3.021 and 4.32,P<0.104). Confrontation ensued between two equally matchedmales immediatelyafter their placement in the observation chamber(figure 6), continued violently(figure 7) till one of them was defeated and assumed submissive posture (figure 8). The latter, though not wounded fatally, succumbed to death within 24 hr. Thus we

188 Shakunthala Sridhara and R V Krishnamoorthy Table 1. Comparison of aggressivebehaviour counts scored by two sexes during different social interactions. Confrontation between Scored by Mean score ± S.E. Cumulative score for the combination Mand F M 1±28 F 233±7.353 MandM M1 741±51* M2 514±35 1255 Fand F F 1 446±48* F2 173±26 619 1Mand2F M 358±29* F 1 66±8 F2 58±6 482 1Mand3F M 487±37* F 1 58±16 F2 73±14 F3 21±23 639 M - male; F = female; * Significantly higher aggressive behaviour counts. Figure 6. Both male rats are equally aggressive. The one crouching in the corner is omega.

Aggressive behaviour of captive B. bengalensis 189 Figure 7. 'Boxing' posture of me conflicting rats. Submissive posture by an omega male in the background, Figure 8. The male on the right is alpha; the one on the left is displaying submissive posture and is omega. Another omega is crouching in the background.

190 Shakunthala Sridhara and R V Krishnamoorthy can assume the winner and loser of this encounter as equivalent to alpha and omega of Rattus noruegicus (Barnett 1975).The third male crouching in a corner (figures 6-8) displayed submissive posture even in the absence of threat! attack and avoided the clashing males, but still died without even being attacked, thus also qualifying for omega status. No betas were identified, which could be either due to the unnatural cage conditions of experimentation or due to the extreme violence of alpha B. bengalensis which does not allow other males to survive in his presence. 4. Discussion Parrack (1966) noticed two peaks of locomotor activity in B. bengalensis, one soon after the dark phase and the other early in the morning. Spillet (1968) failed to establish any rhythm for the agonistic behaviour in grouped B. bengalensis although such activity increased between 00 and 0530 hr. However, Parrack and Thomas (1970) noticed aggressivepeaks for dominant B. bengalensis at 0600 and 0900 hr. The occurrence of nightly peaks of aggression may be a natural corollary of the enhanced general activity of this nocturnal species during dark phase. The present results illustrate both rhythmic and non-rhythmic circadian oscillations of aggressive behaviour of B. bengalensis, which are not stable but are influenced by the number, status and sex of the interacting conspecifics. Similar phase dissociation ofmetabolic and behavioural rhythms of P. maniculatus occurred when intraspecific interactions were enhanced by crowding (Farr and Andrews 1975).Such failure of maintenance of phase relationships was drastic in subordinate animals which could be due to avoidance of more aggressive and dominant cohorts. The great variation seen in the aggressive behaviour rhythms in the present study is a reflection of social interactions! adjustments in the different stress situations of the experiments. The social organization of lesser bandicoots differs from that of Rattus noruegicus in that the females are also aggressive in heterosexual paired encounters. Males can be categorised into alpha and omega as in R. noruegicus. Acknowledgements The authors are indebted to the late Dr KRamakrishnan and Dr R Narayana for their constant encouragement and facilities. Technical assistance of MIs..Govindaraju and Rajanna and financial aid of Ford foundation (Grant No. 660 019), New Delhi, are acknowledged. References Barnett S A 1975 The rat: A study in behaviour, (Chicago: The University of Chicago press) pp. 124-125 Barnett S A, Cowan P E and Prakash I 1975 Circadian rhythms of movements of the house rat, Rattus rattus L.; Indiant. &&p. Biol.13 153-155 Calhoun JB 1975 Social modification of activity rhythms in rodents; Chronobiol. suppl. 1 11 Farr L and Andrews' RV 1978 Rank-associated desynchronizatlon of metabolic and activity rhythms of Peromyscus maniculatus in response to social pressure; Ccmp. Biochem. Physiol.A61 539-542 Grant E C and Mackintosh JH 1963 A comparison of the.social postures of some common laboratory rodents; Behaviour 21 246-259 Landau 1T 1975 Light-dark rhythms in aggressive behaviourofthe male golden hamster; Physiol. Behau. 14767-774. Lerwill C 1977 Diurnal variations in the agonistic behaviour of the golden hamster (Mesocricetus auratus); t. Zool182 159-161

Aggressive behaviour of captive B. bengalensis 191 Marten P 1973 The patterns of Peromyscus activity and their correlation with weather; J. Mammal 54 159-188 Nygren J1978 Inter individual influence on diurnal rhythms of activity in cycling and noncycling populations of the field vole, Microtus agrestis L., Oecologia 35 231-239. Parrack D W 1966 The activity cycle ofthe lesser bandicoot rat Bandicota bengalensis; CU". Sci. 35 544-545 Parrack D Wand Thomas J1970 The behaviour of the lesser bandicoot rat, Bandicota bengalensis (Gray and Hardwickeh J. Bombay Nat. Hist. Soc. 67 67-80 Possidente B and Birnbaum S 1979 Circadian rhythms for food and water consumption in the mouse, Mus musculus; Physiol. Behav. 22 657-660 Siegel S 1956 Non-Parametric Statistics for the behavioural Sciences (New York: McGraw-Hili) pp. Spillet JJ1968 The ecologyof the lesser bandicoot in Calcutta (Bombay: Bombay natural History Society) pp.136-144 Sridhara Sand Krishnarnoorthy RV 1982 Rhythmic oscillations in non-aggressive social behaviour in Bandicota bengalensis; Proc. Indian Acad. Sci. (Anim. Sci.) 91317-322