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FOOD HOARDING BY 'LESSER' BENGALENSIS (GRAY)

DISSERTATION SUBMITFED FOR THE DEGREE OF MASTCR OF PHIIiOSOPHY TO THE ALIGARH MUSLIM UNIVERSITY,

.•}•'•' ••'.«•.••••• J

BY Mrs. Anjum Siddiqui M.Sc.(Alig)

DEPARTMENT OF ZOOLOGY, ALIGARH MUSLIM UNIVERSITY, r ALIGARH. November 1988 ;.Vit.J«U""^ • ICJ i^ to Go«it»ttt«

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Mictions ; Ref. No.: •« ENTOMOLOGY 2 PARASITOLOGY 29.11.1988 3 ICHTHYOLOGY b FISHERIES Date. 4 AGRICULTURALNEMATOLOGY 5 GENET/CS

This is to certify thtit tne preoent work

entitled "Pood Hoaraing by 'Lesser' xjanaicoot

Rat, Baridicota bengal en sis (Gray)" Wasdone unaer

li.j oupervision.

The v/ork is '^ri,_inal ana duVuiices in luy

view Knov.ieafe about Inciari ro.;ent sy.-.ecies.

^iK,

(JAlVilL AHIviAlr KHAN)

i-roiessor oi ^ oology CONTENTS

Page Nps

ACK N 0 WL EDG EM EN T S

KISTORICAL R ESUM E 2-4

INTRODUCTION

MATERIAL AND METHODS 6-8

R ESULTS 9-lD

DISCUSSION 11-13

SLT-IMARY 14

REFERENCES 15-19

TABLES I I ' » ! f e « » 20-23

FIGURES 24-25

•k * J^ -A :* * ***;?:- t •k-k-k-k-k-k-k-k-k ******* ***** *** * (1)

ACKNOWLEDGEMENTS

I arn grateful to Professor Jamil Ahmad Khan for supervising this vvork.

I am thankful to Professor M.S. Jairajpuri, Chairman, Department of Zoology, for facilities.

Acknowledgement is also made to Lab. colleagues and staff.

(Anjum Siddiqui)

...2 (2)

HISTORICAL RESUME

The of the genus Bandlcota are widely distributed in India and much of the south-eastern

(Spill et, 1968). Of the two species, the 'lesser' bandicoot rat is represented by five geographical forms or sub-species (Ellerman, 1961). The sub-species

Bandicota bengal en sis bengal en sis (Gray) is common form found in North India.

The lesser 'bandicoot' is typically a field or rural rat (Blonford, 1888-1891; Ellerman 1961, Prater

1965). However, glfice th« beginning of this century, it has successfully epleniaed urban areas in India (Spillet,

1968). This is not only true of larger metropolitan urban centres as Calcutta, Bombay, Delhi and Madras (Rao,

1947; Seal, i960 b; Deoras, 1960-1963); but also of smaller as Aligarh. 'fhig shows tiie magnitude of change that has oGGUfred in urban rat population of

India.

The is a highly fossorial rat (Kamath, 1961; Rao, 1965; Roy, 1974). Sut besides other behaviour pattern, it also shows great propensity

...3 (3)

to 'hoard' food in its (Kamath, 1961; Deoras, 1964; Hoy, 1974) . Thus its burrowing habits give it both added protection as well as provide it with re­ serve food supply (Spillat, 1968) .

The lesser bandicoot rat is thus the most important agent of food destruction; and that is a matter of serious concern (Spillet, 1968). It is thus estim.ated that 25% of food grown' in. India is destroyed by in the field and an additional 25-30% in storage (Warshof sky, 1967) . This amount is larger than the buffer stock maintained by the Indian Government. It is quite obvious that if the'bandicoot' is controlled, losses of food to wOUld be drastically reduced.

The f;o©d hoarding by lessor bandicoot is, however, not being extensively studied. To-date, the most important vork is of Parrack (ige?'). It demons­ trates that the lesser bandicoot rat eats less, damages ' more; er.d hoaJ^'Cls^ faybstsntially., The total food lost to them is perhaps much more than that caused by 'non- hoarders' dS 'reef rat, Rattiie r.|_g..tus, or 'hoarders' as gerbils Tat era Indica. (Kiomari and Khan, 1978). (4)

The factors influencing food 'hoarding' behaviour of 'lesser' bandicoot rat, have also not been ainalysed. It is generally believed that it is a compulsive 'hoarder' and can even hoard many objects other than food. However, the facts are otherwise, as the present vx^rk shows.

This work has been done in line with similar work done on many other rodent species that hoard food as , . Albino rat, mice and gerbils, (Smith and Koss, 1950; and Terman, 197 2; KcCarty and Southwick, 1974; Rieh&rd, 1976, 1978; Tadlock and Klein, 1979j Kumafi ^d Khail, 1979; Begineux et. al., 1980; Etienne, et. al., 1983; Roderick, 1984).

Cbviously, jtigre investigations on food hoarding behaviour of economically important rodent species in India are required (Barnett and Prahssh, 1975). But this vTork, it is hoped; SuFi serve as the basis for fur­ ther investigation. (5)

INTRODUCTION

Food crops are infested by a number of rodent pests; and exploited at all stages of growth (Barnett

and Prakash, 1975). But harvest time is the season of

plenty; when large amounts of grains are carried and

'stored' by them in their burrows (Prasad, 1954; Prater,

1965) . Of these pests, the lesser bandicoot rat,

Bandicota bengal en sis (Gray), is the most important in

many parts of India (Barn©tt and Prakash, 1975).

Estimates of food 'hoarded' by B. bengal en sis,

have been few, but wider in the that observations

have elso been made in the field (Parrack, 1969; Roy,

1974), Hov/ever, more studies of it are obviously requir­

ed for use in |?©st-horvest technology, and protection of

stored grains (Spillet, 1969; Barnett and Prakash, 1975).

Accordingiy, 'hbatdihg' of food by adult B.

bengal en sis of both sexes, was observed in laboratory en­

closures. The r^Sults °^^- '^l'B^^'\^tssed here.

... 6 (6)

MATERIAL AND METHODS

The , B. bengal en sis, were trapped from Aligarh Railway Station. Traps baited with bread, were placed on tracks of main platform; v^ich was dug over most of its length. Traps were sprung within an hour at most of being set near what appeared to be the main 'entrances' to burrows; practically at all hours of the day.

Trapped bandicoots were brought to the labora­

tory; sexed, weighed and housed in wire-mesh enclosures,

0.82 X 0.32 X 0,31m. Each enclosure had a layer of sand mixed with insecticide at the bottom. Paper was given

for nesting. The bandicoots were maintained on a diet of bread and rat 'chow' and vegetables; water was given

ad lib. Arrangements made inside the enclosures are

shown in Fig, i.

Experiments were started after acclimiatization of bandicoc3t.? to laboratory conditions, for at least 15 days. Subjects were divided into four groups; each group was given the following treatinent:

...7 (7)

Group 1: All subjects were weighed, and given a surplus of wheat (Triticum aestivum) . Cages were checked daily for the next 15 days. Grains found in nest-boxes, were taken out and weighed.

Group 2: After "weighing, bandicoots were starved for 3 days; and then weighed again. They were then given a surplus of wheat; amounts hoarded each day were recovered and weighed over the next 15 days.

Group 3: Subjects were weighed and offered only brinjal (Sol angrn meloqena) for next 7 days. They were then weighed again, end given wheat in surplus arnQunts. Grains 'hoarded' were recovered and weighed daily for nfxt: 15 days.

Group 4; Bandicoots were weighed, and given 20g of wheat fpr 3 days; and then lOg* 8g, 4g and finally 2g wheat also for 3 days each. Subjects were then v;eighed again and given surplus amounts of wheat. ^©unts QS. gfains hoarded by each sub­ ject was recorded daily as before. Observations wat-e e^fentinued f§? a§ lofig as the behaviour lasted.

. • . S (8)

Results were statistically analysed according tx5 methods described by Bailey (1959) and Lehner (1979) . (9)

RESULTS

The results are summarised in Tables 1 to 4; and illustrated in Fig. 2.

The bandicoots of group 1 hoarded food, but in small amounts (Table 1). One female proved, however, an exception. But differences on account of sex were not discernible (t = 1.405, df = 4. P^O.2). Hoarding also lasted for variable durations.

Bandicoots of group 2 lost 10 to 12% of their body-weight due to starvation. But original body- weights were rapidly regained within 7 days of wheat becoming available. During the period of weight- recovery, ajid even afterwa:E'd5; all subjects hoarded food, but in smill amounts liXe the bandicoots of group 1 (Table 2) .

Group 3 bandicoots als© lost weight (15-20%) o n restricted dist sf bfifljal. But it was also rapidly regained. All subjects then hoarded food. But it was hoarded in small amounts, as Jay bandicoots of group 1

& 2 (Table 3) .

... 10 (10)

lost Group 4 bandicoots also/v/eight ^10% ) with gradual reduction of daily rations. The original weights were regained v;ithin 6 days of wheat becoming available again in surplus amounts. During the short period of weight recovery, and then over the next 59 to 9 2 days; all subjects hoarded wheat in very large amounts (Table 4). But one male carried it in excep­ tionally high amounts - 233g wheat was recovered from nest-box each day (Fig. 2). In case of all subjects, hoarding levels gradually increased to a 'peak' before declining. Afterwards, It stopped altogether.

...11 (11)

DISCUSSION

Several rodent species are found to 'hoard' food, both in natural as well as under laboratory con­ ditions (McCarty and Southwick, 1974; Barnett and

Prakash, 1975). >^parently, females 'hoard'more than males (Niorgan et. al. 194 3; Dewsbury, 1970; Nyby et. al., 1973). Although a complex activity, food hoaird- ing is quickly released by varying degrees of food dep­ rivation (Morgan et. al., 194 3; Smith and Ross, 1950a; iMcKelvey and Marx, 1951; Nyby et. al. 1973). Our observations on bandicoots also confirm this.

It is thus clear that bandicoots regularly

'hoard' food, under laboratory conditions too; but ordinarily in small amounts (Table 1). Methods of food deprivation as starvation or restriction to an exclusive diet of vegetables* do not influence hoarding levels

(Tables 2, 3). But bsfidicoots subjected to a schedule of reduced rations, start hoarding food in large amounts

(Tat)le 4, Fig. 2). The grains are carried by mouth and stored in next-boxes. The behaviour also lasts for long durations; and attains 'peaK' levels in male subjects

(Table 4, Fig. 2) .

...12 (12)

Since hoarding behaviour in bandicoots is also released by food deprivation, it is obvious that nutritional factors supply the drive for it (Morgan et. al., 1943; Kuinari and Khan, 1979). But unlike gerbils, Tatera indica in die a Hardwicke, both sexes in bandicoots hoard food; and hoarding also occurs over extended periods (Table 4, Fig. 2). Bandicoots are 'solitary', while gerbils are 'colonial' forms (Barnett and Prakash, 1975); and this may account for the differences observed between the hoarding behavi­ our of the two species.

Sex differences in such behaviour are observed in 'Mongolian' gerbils unquiculatus too; and have been attributed to its hormonal control, and in­ fluence of temperate climate. 'Hoarding' by males follows the fall in androgen levels, and coincides with term.ination of breeding season (Nyby et. al., 1973). Pood acquisition by them is essential for over­ winter survival of the colony. Consistently high levels of 'hoarding' by females ensures nourishment specifically for care and development of young (McCarty and Southwick, 1974).

. .. 13 (13)

Thus hoarding by males serves the important function of obtaining reserves of food for the entire, colony; and hence its adaptive value increases with social organisation (Kurnari and Khan, 1979). Hoarding by females has different function; also of great adap­ tive value. Since bandicoots are 'solitary', such dif­ ferences as may arise with social organisation, are obviously not observed. Both sexes gather their ' stores' of food separately, to tide over periods of scarcity. These may continually arise as for example with rotation of crops in the fields (Kumari and Khan, 1979). Accordingly, food hoarding by bandicoots in the natural environment also coincides with harvesting of crops (Prater, 1965).

..14 (14)

SUMMARY

Subjected to food deprivation by gradual reduction of daily rations, the lesser bandicoot rat, Bandicota bengal en sis (Gray), started 'hoarding' cereal grains then in very large amounts from a sur­ plus offered. Alternative methods of food deprivation as starvation or restriction to an exclusive diet of vegetables, failed to similarly provoke the behaviour in laboratory enclosures.

Although both sexes 'hoarded' food, hoarding attained 'peak' levels in male subjects. The behaviour in than also lasted for longer durations.

Food hoarding in natursl environment coincides with harvesting of crops. Grains 'hoarded' then, help individuals to survive ifl the coming 'lean' periods; when food obviously becomes scarce.

. ,. 1: (15)

REFERENCES

Bailey, N.T.J. (1959) Statistical Methods in Biology.

English University Press, London.

Barnett, S.A. and Prakash, I. (1975) Rodents of Economic

Importance in India. Arnold Heinanann Press, New

Delhi.

Begineux, F, J.M. Lassalle and G. Le Pape (1980) Hoarding

Behaviour of AKR, C 5 78L/6 mice and their F, in

their home living space, An automatic recording

technique. Physiol Behav 24(6); 1191 - 1194.

Elanford, Vv'.T. (1S88) The Fauna of British India, Mammalia.

Taylor and Francis, London.

Deoras, P.J. (i960) Studies on Bombay rat; a note on the

probable resistance of B. bengal en sis to .

Curr. Sci.,29:47 5.

Deoras, P,J, (1963) Studies on Bombay : frequency of

rat population. Curr.Sal,, 32; 153 - 165.

Deoras, P.J. (1964) Rats and tli©lr control. Indian J.

Eht., Silver Jubilee, 1964, 18 pp.

Dewsbury, D.A, (1970) Food hoarding in rice rats and

Cotton fats. Psychol. Rep., 26 (1): 174.

16 (16)

Ellerman, J.R. (l96l) The fanua of India, including Pakistan, Burma and Ceylon. Vol. 3 Rodentia Edited by M.L. Roonwal.

Etienne, A. S. Ernmanueli and M. Zinder (198 2) Ontogeny of hoarding in the golden Mesocricetus auratus. The development of motor pattern and their sequential coordination. Dev. Psycho biol 15(1); 33 - 46.

Kannath, M.K. (I96l) Studies on Bombay rats, rat burrows and ectoparasites in Mahim area (unpublished M.S. dissertation, Dept. of Ehtomology, Haffkine Inst., Bombay). 133 pp.

Kumari, P.V, and Khan J.A. (1978) Food preferences of gertoil Tajtera, Indies* Indie a Hadwicke. Zooi. J.

Kuinari, P.V. and Khan, J.A. (1979) Food hoarding by Indian ^atbil, Tatera Indi^Jc IsJiSJ* Proc. Indian Acad, Sci., 88(6), |t 131 - 135.

Lehner, P.M. (1979) Handbook of Ethological Methods. Garland TPM Press, Ktfv,* 'ibrk & London.

McCarty? R. and Southwick, CH (1974) Food hoarding by Southern grass hopper onychomys tprridus in labo­ ratory eiclOGures. J. .., 56; 708 - 712.

...17 (17)

Mckelvey, P-.K. and Marx, M.H. (1951) Effects of infantile food and water deprivation on adult hoarding in rat. J. comp. Physiol., 44; 168 - 177.

Morgan, C.T.; Stellar, E. and Johnson, 0. (1943) Food depri­ vation and hoarding in rats. J. comp. Psychol.,

3^1 215-291.

Nyby, J.; V.'allace, P.K. and Thiessen, D.D. (1973) An influ­ ence of hormones on hoarding behaviour in the Mongolian gerbil Meriones unquiculatus. Horm. Behav., 4: 28 3 - 288.

Parrack, D.W. (1969) A note on the loss of food to the lesser bandicoot rat, Bandicota benglaensis. Curr. Sci., J8: 93-94.

Prasad, M.R.N. (1954) Food of the Indian gerbil Tatera In die a. (waterhouse) jL_..Bombay Nat. Hist. Soc, 52i 321 - 325.

Prater, S.H. (1965) The book of Indian . Bombay Nat. Hl-Stj.. Soc, Bombay, 2nd ed,

Rao, S.K. (1947) Role of field rats in the epidemiology of plague in H.E.H. Nizam.'s dominion. Indian Med. Gaz,

8_2J 96 '= 99.

Rao, S.R. (1965) op.cited Spillet (1968).

^'l^^ "^.* .18 > •;:„ ^ !^i "li^d (18)

Rice L. and Terman C.R. (197 2) Food hoarding in freely growing laboratory population of prairie deer mice peromyscus imanicul atus bairdii . Paper presented at the Virginia Academy of Science meeting.

Roy, S.K. (1974) Pre-harvest loss of rice due to field rodents. Economic and Political Weekly, 9_: 66 - 67.

Seal, S.C. (1960) The changing pattern of rodents and in Calcutta and other cities. Hull. Wld. Hlth. Org. _23s 283 - 292.

Spill at, J.J. (1968) The of tbie lesser Bandicoot Rat in Calcutta. Bombay Nat. Hist. Soc.

Smith, V>J.I. & Ross, S. (1950) Wie hoarding behaviour in the hamater. J. Genet. Psychol., 77; 211-215.

Tadlcck, C.C, Klein, H.G. (1979) Nesting and food storage behaviour of Peromyscus manicul atus gracilis and P. leucopus no vebo r ac en si s. Can. Field Nat, 9JjJi: 239 - 24 2 En, m,

Wallace, Richard J. (1976) hoarding in the Albino-rat. Pjt\lm seV^ay,. Mlii^ 1"& - 180.

19 (19)

Wallace, Richard J. (1978) Hoarding of inedible objects by Albino rats. Behav. Biol., 23( 3) ; 409 - 414.

Warshofsky, F. (1967) Most destructive creature on earth. Reader' s Digest, Sept., 1967: 17 5 - 180.

VJong Roderick (1984) Hoarding versus the immediate con­ sumption of food among hamsters and gerbils. Behav. Proc. , 9 (1) : • 3 - 11.

. . .20 TABLE 1

Miounts of wheat hoarded by bandicoots of group 1 in their nest-boxes. (20)

Table 1

Description Length of Hoarding Anount of Food Bandicoot (Body-Vveight Observation Period Hoarded No q/Sex) (Days) (Days) q/day

1 365 i 15 5 4.00

2 452 15 7 41.40 $ 3 219 J 15 8 1.8 4 297 S 15 0 0 5 400 ? 15 4 9.0

6 207 J 15 7 5.00

. .. 21 TABLE 2

Amounts of food hoarded by bandicoots of croup 2 after starvation for 3 days. (21)

Table 2

Description Length of Anoiint of Food Bandicoot (Eody-•'.^; eight Observation Hoarded No (g/Sex) (Days) g/day

1 365 15 4.0 % 2 325 I 15 7.0 3 37 5 ^ 15 11.0

4 310 t 15 3.0 TABLE 3

Wheat hoarded daily by bandicoots of group 3 after receiving an exclusive diet of vegetable for 7 days. (22)

Table 3

Description Length of Amount of Food Bandicoot (Eody-V.'eight Observation Hoarded No (g/Sex) (Days) g/day

1 365 15 4.3 ^ 2 219 15 3.0 % 3 415 15 7.0 « 4 400 t 15 6.0

5 207 t 15 2.0 TABLE 4

Food hoarded daily by bandicoots of group 4; subjected to a schedule of reduced rations. (23)

Table 4

Amount o f Food DescriDtion Hoar( ied an di coot (Body-Weight Ho arclin g Period g/day No (g/Sex (Days from Start) (Mean + S.E.)

1 300 87 33.100 + 9.959 * 2 350 ? 79 39.4 37 + 9.401 3 350 t 77 12.375 + 3.184

4 400 ? 61 114.285 + 4.466

5 410 ? 92 193.800 + 33.387

...24 FIGURE 1

Arrangements inside the enclosures housing bandicoots. F = Food tray/ VJ = V^Jater tray, N = Nest-box. (24) FIGURE 2

Amounts of food hoarded dally by Band! coo "t

No. 6 of group 4. (2 5)

AMOUNT OF FOOD HOARDED, 9/day — l\) u» ^ Wl o o O o O o 0 «d 0 O - 1 1 i 1 t

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