Social relationships in N'dama cattle during supplementary feeding

Applied Animal Behaviour Science, 34 (1992) 285-290

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Elsevier Science Publishers B.V., Amsterdam

Social relationships in N'dama cattle during supplementary feeding J.D. Kabuga Department of Animal Science, University of Science and Technology, Kumasi, Ghana (Accepted 13 February 1992)

ABSTRACT Kabuga, J.D., 1992. Social relationships in N'dama cattle during supplementary feeding. Appl. Anita. Behav. Sci., 34: 285-290. Social relationships of 30 N'dama cows during supplementary feeding, post-grazing, were studied over a period of 1.5 years. Dominance ranks determined during idling and feeding periods were strongly correlated (Spearman's rank correlation (rs = 0.964, P< 0.01 ). The number of animals dominated by a cow during feeding was strongly (P< 0.01 ) related to liveweight (r = 0.822) and age (r= 0.755 ). Low status cows ate less frequently than medium and high status animals, while middle ranking cows were ejected more frequently from the feed trough than other dominance groups. Animals had preferences in the use of feed troughs, with social rank being the dominant factor determining the location of feed trough space used. Cows of similar status were generally preferred feeding and movement neighbours and antagonists. However, the dominance rank of an animal and its preferred neighbour during idling were not significantlycorrelated (rs= 0.220, P> 0.05 ). Voluntary leadership ranks into and out of the pen were, respectively, related closely (P< 0.01 ) to feeding dominance ranks (r~=0.661,0.640) and idling dominance ranks (rs=0.621,0.669).

INTRODUCTION

It is generally acknowledged that when feed, water and space resources are limited, the influence of social dominance becomes important, with high ranking animals having priority to these resources (Friend and Polan, 1974; Metz, 1983 ). This therefore implies that when enough resources are available the influence of social status will be at a minimum. It appears, however, that this may not be the case in all situations. For example, Wagnon (1965) observed in range cattle that even when enough feed and feeding space were available at one location for the whole day, social rank effects were still important during feeding. This was because when not feeding, dominant cows still prevented subordinates from feeding. Wagnon (1965) thus concluded Correspondence to: J.D. Kabuga, Department of Animal Science, University of Science and Technology, Kumasi, Ghana.

© 1992 Elsevier Science Publishers B.V. All rights reserved 0168-1591/92/$05.00

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that the age composition of a herd had a strong effect on the use of supplementary feed on the range. These results have been confirmed from studies with dairy cattle (Metz, 1983), and have led Metz to argue that the social determinants for feed competition is not rank order per se, but rather social relationships among members of the herd such as non-related individuals, strangers, kin relationships and c o m m o n rearing experience. Family relationships and c o m m o n rearing experience result in less aggression and competition at the feed troughs. There is, therefore, the need for other studies to help clarify the effects of social status and other social relationships on feed competition. Most studies on social relationships at feeding have been obtained with Bos taurus cattle breeds. It is not clear if Bos indicus type cattle similarly behave in the same way as European type cattle during feed competition. This is partly because Jewell and Nicholson (1989) found that severely dehydrated Zebu cattle which had been without water for 3 days had the remarkable ability to wait patiently for their turn to drink. Also Kabuga et al. ( 1991 ) could not find any influence of social status in use of shade and drinking frequency in grazing N ' d a m a and West African Shorthorn cattle. The objective of this study was to investigate the social relationships of N ' d a m a cattle during supplementary feeding post-grazing and the influence of social status in the use of feed trough space. ANIMALS, MATERIALS AND METHODS

Thirty (20 multiparous and 10 primiparous) N ' d a m a cows, average age 75.9 (range 36-185 ) months and weight 239 (range 200-350) kg, were used for the experiment. All experimental animals, except two unrelated cows which were brought in from the Livestock Section of the Department of Animal Science, had been handled as a single herd (of about 200 animals) for 9 years at the University of Science and Technology Dairy/Beef Cattle Research Station at Boadi. The experimental area has previously been described (Kabuga et al., 1991 ). The experimental animals included five m o t h e r - d a u g h t e r pairs, three sister pairs, six pairs of animals which were born within 30 days of each other, and 11 animals unrelated as kin or by time of birth. All cows were dehorned at birth and were individually ear tagged during the experiment with large yellow ear tags for easy identification. The experimental animals were managed with the main herd except when separated daily at 15 : 00 h for the study. The cows were observed once a week for 2 h from December 1989 to June 1991, a total of 156 h. In each weekly observation, a 1 h observational session was allocated both to a non-feeding (idling) period ( N F ) when no supplementary feed was provided and to a feeding period (F) when 5 kg of wet brewer's spent grain per cow d a y - 1 was provided. The feed was provided in

SOCIAL RELATIONSHIPS 1N N'DAMA CATTLE

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six feeding troughs (Troughs 1-6) in a 20 m × 10 m pen. Troughs 1 and 5 were 1.9 m long and Troughs 2, 3, 4 and 6 were 3.6 m long. All troughs were 0.6 m wide. Both the NF and F took place in the feeding pen with the NF always preceding the F. During the NF, agonistic interactions (butting, threatening and avoidance) were recorded as well as the identities of the performing and receiving animals. At 10 min intervals the herd was scanned and animals standing almost touching were recorded as neighbours. During the F, agonistic interactions at the feed troughs were recorded as described for the NF. In addition the herd was scanned every 3 min and the feeding position at each trough of each cow was noted. An animal was said to be feeding when it had its head either in or above the feed trough and was manipulating feed in the mouth. The order of entry and exit of the animals from the feeding pen was recorded once a month. Dominance ranks were determined for the NF (idling dominance) and the F (feeding d o m i n a n c e ) from the n u m b e r of animals dominated based on the n u m b e r of wins of agonistic interactions. A total of 3210 and 4470 agonistic interactions were used in the NF and the F, respectively. From the dominance ranks the herd was divided into three equal groups of high, m e d i u m and low ranking cows. The orders in which cows entered or exited from the feeding pen were used to derive separate leaderships for passage into and out of the feeding pen. From the leadership ranks the preferred leader of an animal was determined as the most frequently followed conspecific by the animal. Animals with the highest observed associations during feeding together at the same trough, less than 0.6 m apart (adjacent) or more than 0.6 m apart (nonadjacent), were classified as preferred feeding partners. Cows with the highest n u m b e r of emitted agonistic interactions between them were classified as preferred antagonists. Data on various social and physical characteristics were analysed by analysis of variance (Sokal and Rohlf, 1981 ) to determine the relative importance of dominance groups. All frequency data were log transformed before analysis although the means quoted are actual values. Z 2 analysis was used to determine if the proportions of the dominance rank groups using the same feeding troughs were similar. Correlations of ordinal ranks were determined by Spearman's rank correlations (rs) (Siegel, 1956) while those of interval ranks were compared with Pearson's product m o m e n t correlation (r). RESULTS

The number of animals dominated by each cow in the NF and F were closely correlated (r=0.977, P<0.01 ) as also were the dominance ranks (rs=0.964). The low status cows were the youngest and smallest while the high status animals were the oldest and heaviest (Table 1 ). Thus there was a significant

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TABLE 1 Differences b e t w e e n low, m e d i u m a n d high s t a t u s a n i m a l s in physical a n d b e h a v i o u r a l traits Trait

Liveweight (kg) Age ( m o n t h s ) F r e q u e n c y o f eating at 3 rain intervals ( per cow h - , ) Agonistic interactions initiated ~ ( per cow h - ~) Agonistic interactions received ~ ( per cow h - t ) F r e q u e n c y o f ejections ~ ( per cow h - , )

Status

SE

Low

Medium

High

179.5 a 38.8 a 4.91 a

239.5 b 87.4 b 7.19 b

295.0 c 106.8 c 8.81 c

3.51 2.64 0.478

0.49 a

1.96 b

3.48 ~

0.177

2.05 a

2.63 b

1.25 c

0.155

1.94 a

2.60 b

1.28 c

0.188

M e a n s in the s a m e row with different superscripts are significantly ( P < 0.05 ) different. ',-Xt the feed trough.

( P < 0.01 ) positive and strong correlation between the number of animals dominated during feeding and idling, respectively, with age (r = 0.755, 0.607) and liveweight (r=0.822, 0.828). Feeding dominance rank significantly (P<0.05) influenced the number of agonistic interactions initiated and received at the feed trough, the frequency of interruptions at the feed trough (Table 1 ) and was linearly related to the frequency of feeding (rs=0.806). Except for three cows, all animals showed highly significant (Z2= 12.7-18.9 5 d.f., P < 0.05 ) preferences for the same feeding troughs throughout the study. The high status group generally used Troughs 4, 5 and 6 while the medium status group preferred Trough 1 (Table 2). Feeding and idling dominance ranks were respectively correlated ( P < 0.01 ) with average leadership ranks into (rs=0.661, 0.669) and out of (r~=0.640, 0.621 ) the feeding pen. The rank of the animals based on the frequency of eating and agonistic interactions initiated at the feed troughs were respectively correlated (rs= 0.558, 0.534, P < 0.01 ) with the average leadership ranks into the pen. The ranks of preferred conspecifics to an animal during idling and during feeding adjacent (rs = 0.688 ) and non-adjacent (rs = 0.401 ) were significantly (P<0.05) related. All mother-daughter pairs, except one pair, and one sister pair were preferred idling partners while the two cows introduced into the herd were preferred feeding and idling partners. The feeding dominance ranks of animals were significantly (P<0.05) correlated with the dominance ranks of their

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SOCIAL RELATIONSHIPS IN N'DAMA CATTLE

TABLE 2 Percentage mixture of animals of different social rank groups using each feed trough Trough No.

1 2 3 4 5 6

;(2

Status Low

Medium

High

15.1 25.7 35.4 20.6 11.5 18.2

58.8 38.9 29.9 32.6 14.0 31.0

26.1 35.4 34.7 46.8 74.5 50.8

;(2=5.99, P< 0.05; zz=9.21, P< 0.01;

30.95 2.80 0.54 10.31 76.43 16.21

;(2= 10.60, P< 0.005.

preferred associates as antagonists (rs = 0.412 ), as leaders ( rs = 0.611-0.622 ) and as neighbours during eating adjacent (rs=0.528) and non-adjacent (r~=0.655). DISCUSSION

In line with other studies (reviewed by Hafez and Bouissou, 1975; Syme and Syme, 1979) there was a close relationship between social dominance and liveweight and age. However, contrary to these reviews liveweight rather than age showed a stronger relationship with dominance in this trial. Perhaps the reported (Reinhardt and Reinhardt, 1975) decline in rank for very old cows may have influenced the present results; 20% of the cows were over 9 years old. In this investigation low ranking animals were observed to eat less frequently than dominant cows. Also, the more aggressive cows were generally the first to enter the pen to eat, confirming the results of Schmisseur et al. ( 1966 ). It is therefore likely that the low ranking animals fed less frequently to avoid aggression from dominant herdmates. This probably explains why cows closer in rank, although involved more in aggressive encounters than cows further apart in social status, were preferred neighbours during feeding and movement. This effect of social status on feeding frequency has also been reported previously in European type cattle (McPhee et al., 1964; Friend and Polan, 1974). The present results therefore suggest that the N'dama, an indigenous taurine cattle breed, may behave differently from Zebu cattle (Jewell and Nicholson, 1989 ). The preference of animals for certain feed troughs suggests some sort of resource protection, with dominance rank being the major factor determining the preferred feeding location. These results thus imply that under high stock-

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ing rates dominance hierarchy could result in inefficient use of feeding space probably with cows overcrowding in some feed troughs. It would seem that the animals in this study were similar to those studied by Reinhardt and Reinhardt ( 1981 ) and Kiley-Worthington and de la Plain ( 1983 ) in that they were prone to form relationships based on preferential attachments. However, except during idling, these attachments were not strongly based on kin relationships or peer associationships but probably to interindividual preferences related to similarity in social status and farm of origin. ACKNOWLEDGEMENTS

The author wishes to thank the management of Boadi Dairy/Beef Cattle Research Station for the facilities for the research, and K. Awunyor, P. Yawkpe, Owusu Ansah, D. Akumsa, M. Dassah and C. Kabuga for help with observations. REFERENCES Friend, T.H. and Polan, C.E., 1974. Social rank, feeding behaviour and free stall utilization by dairy cattle. J. Dairy Sci., 57:1214-1220. Hafez, E.S.E. and Bouissou, M.F., 1975. The behaviour of cattle. In: E.S.E. Hafez (Editor), The Behaviour of Domestic Animals. Bailli6re Tindall and Cox, London, pp. 203-245. Jewell, P.A. and Nicholson. M.J., 1989. Strategies for water economy amongst cattle pastoralist and wild ruminants. Symp. Zool. Soc. London, 61: 73-87. Kabuga, J.D., Gari Kwaku, J. and Armor, S.Y., 1991, Social status and its relationships to maintenance behaviour in a herd of N'dama and West African Shorthorn cattle. Appl. Anita. Behav. Sci., 31: 169-181. Kiley-Worthington, M. and de la Plain, S., 1983. The Behaviour of Beef Suckler Cattle (Bos taurus). Birkhauser, Basel, 195 pp. McPhee, C.P., McBride, G. and James, J.W., 1964. Social behaviour of domestic animals. III. Steers in small yards. Anim. Prod,, 6: 9-15. Metz, J.H.M., 1983. Food competition in cattle. In: S,H. Baxter, M.R. Baxter and J.A.D. MacCormack (Editors), Farm Animal Housing and Welfare. Marlinus Nijhoff, The Hague, pp. 164-170. Reinhardt, V. and Reinhardt, A., 1975. Dynamics of social hierarchy in a dairy herd. Z. Tierpsychol., 38: 315-323. Reinhardt, V. and Reinhardt, A., 1981. Cohesive relationships in a cattle herd (Bos indicus). Behaviour, 77: 121-151. Schmisseur, W.E., Albright, J.L., Dillon, W.M., Kehrberg, E.W. and Morris. W.H.M., 1966. Animal behaviour responses to loose and free stall housing. J. Dairy Sci., 49: 102-104. Siegel, S., 1956. Non-Parametric Statistics for the Behavioural Sciences. McGraw-Hill, New York, 312 pp. Sokal, R.R. and Rohlf, F., 1981. Biometry. W.H. Freeman, New York, 859 pp. Syme. G.J. and Syme, L.A.. 1979. Social Structure in Farm Animals. Elsevier, Amsterdam, 200 PP. Wagnon, K.A., 1965. Social dominance in range cows and its effect on supplemental feeding. Calif. Agric. Exp. Stn., Bulletin 819: 1-32.