Animal Science 2006, 82: 271276 Q 2006 British Society of Animal Science

ISSN 1357-7298 DOI: 10.1079/ASC200517

The effect of sward surface height on the response to mixed grazing by cattle and sheep
I. A. Wright1,, J. R. Jones2, D. A. Davies2, G. R. Davidson1 and J. E. Vale2
Macaulay Institute, Macaulay Drive, Craigiebuckler, Aberdeen AB15 8QH, UK Institute of Grassland and Environmental Research, Bronydd Mawr Research Centre, Trecastle, Brecon, Powys LD3 8RD, UK
2

E-mail: I.Wright@macaulay.ac.uk

Abstract
An experiment was conducted to examine the effects of mixed grazing of sheep plus cattle under continuous stocking of permanent pasture at different sward heights. The experiment had a 2 3 factorial design, with two sward surface heights (4 5 and 8 10 cm) and three combinations of animal species viz., sheep only, cattle only and sheep plus cattle. There were two replicate plots of each treatment combination and the experiment was conducted over 2 years consecutively. The sheep were Beulah Speckled Face ewes and their single Suffolk-cross lambs while the cattle were yearling Charolais-cross steers. Six core steers and six core ewes and their lambs grazed plots, as appropriately, while additional, non-experimental steers and ewes and their lambs were used to maintain sward heights. Each year the steers and the ewes grazed the pastures from May to October, while lambs were weaned and removed each year from the experiment in July. There was no signicant effect of mixed grazing on live-weight gain of steers, but ewes had signicantly higher live-weight gains on the sheep plus cattle treatment than on the sheep-only treatment (82 v. 61 g/day; P , 0001). The live-weight gain of the lambs was higher on the mixed grazing treatment than on the sheep only treatment on the 8 10 cm sward height treatment (243 v. 212 g/day; P , 005) but there was no signicant difference on the 4 5 cm sward height treatment (260 v. 250 g/day; P . 005). The total output of live-weight gain per ha from steers, ewes and lambs was not signicantly affected by animal species combination. It is concluded that while output per ha is not enhanced by mixed grazing by sheep and cattle when sward height is controlled, the live-weight gain of ewes is increased and the live-weight gain of lambs can be increased on taller swards. Keywords: cattle, grass sward, mixed grazing, sheep.

Introduction
Mixed grazing involves the concurrent grazing of an area of pasture by more than one animal species, and previous studies have shown that in some cases the performance of one, or both of the species involved is greater than when only a single species grazes, or that the total output of animal live weight per unit area is higher (see reviews by Nolan and Connolly (1977) and Lambert and Guerin (1989)). Other advantages claimed for using mixed as opposed to mono-species grazing include better matching of the animals seasonal energy requirements to herbage production, reduced gastro-intestinal parasite burdens, diversication of animal production, manipulation of botanical composition of swards (Lambert and Guerin, 1989) and a more balanced use of vegetation resources, promoting ecological stability and reducing the risk of landscape degradation (Nolan and Connolly, 1992). Many experiments have been conducted in which the effects of mixed grazing have been studied. However it is

difcult to draw general conclusions from these studies. The problem of ensuring that differences between treatments are due to the effects of mixed grazing per se and not due to some artifact of the experimental design were highlighted by Nolan and Connolly (1977). In their review they categorized mixed grazing experiments with cattle and sheep depending on the basis for comparison. These included trying to equalize grazing pressure in mono- and mixed-grazing treatments, choosing livestock equivalents across species, choosing a range of ratios of different species or choosing a range of different stocking rates. All have their disadvantages and can make interpretation of results difcult. Nolan and Connolly (1989) used large-scale eld experimentation on permanent pasture with a range of stocking densities and ratios of sheep and cattle and then used response equations (Connolly, 1987) to relate individual animal performance or output of animal product per unit area to stocking density and the ratio of the two animal species. Such an empirical approach, while providing quantitative information, is limited in that the values of the

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Wright, Jones, Davies, Davidson and Vale parameters in the response equations are likely to be specic to the situation in which they were derived. In most natural grassland ecosystems, different species of grazer co-exist. Theoretical studies initially indicated that species can only co-exist if they do not have completely similar niches (Volterra, 1928; Levin, 1970) although more recently this hypothesis has been called into question (Vance, 1984; Farnsworth et al., 2000). Within grassland ecosystems it has been argued that animal species of different body size can co-exist by grazing grass patches of different heights. Larger animals have a preference for taller patches, related to the rate at which they can harvest material of different heights in relation to their body weight. For example, mature ewes can obtain near maximal rates of intake from ryegrass swards of 4 to 6 cm (Penning et al., 1991) whereas cattle do not achieve their maximum intake rate until sward heights reach 10 to 12 cm (Wright, 1988). Furthermore the grazing of tall patches by a larger species may create shorter patches which, although being suboptimal for the larger species may still allow the smaller species to achieve near maximum rates of intake (McNaughton, 1984). To date no studies of mixed grazing have been conducted in which sward height has been used as a controlling factor in the experimental design, despite the fact that it has been recognized for over 20 years that sward height is a major determinant of herbage growth and utilization (Bircham and Hodgson, 1983; Parsons et al., 1983) and that sheep and cattle have different optimum sward heights in relation to rate of intake of herbage. The present experiment was designed to examine the effects of mixed sheep plus cattle grazing under continuous grazing of permanent pasture at different sward surface heights. The experiment was conducted at Bronydd Mawr Research Station, Brecon, Wales at an altitude of 300 to 320 m above sea level. The soils were brown earth soils (Milford series) derived from Old Red Sandstone. The pastures were improved permanent pastures which had been re-seeded approximately 30 years previously. The predominant species was perennial ryegrass (Lolium perenne) with small proportions of white clover and other grass species. Six core ewes (Beulah Speckled Face) and their single Suffolk-cross lambs and six Charolais-cross yearling steers were allocated to the sheep-only and cattle-only treatments respectively each year. On the mixed cattle plus sheep treatments there were six core steers and six core ewes and their single lambs, although the ratio of ewes and steers were kept at 3:1 by the addition of non-experimental ewes and lambs, since Nolan and Connolly (1989) suggested that this was the ratio that was likely to give the greatest response to mixed grazing. The average age of the lambs at the start of the experiment was approximately 6 and 5 weeks in 1996 and 1997 respectively. Lambs were weaned on 23 July and 22 July in 1996 and 1997 respectively. Routine veterinary care was carried out during the experiment, including periodic dosing with an anthelmintic. Adequate clean water was available to the animals at all times. No supplementation was given during the experiment. The experiment started each year when the sward height on the 810 cm sward height treatment plots reached 8 cm (21 May and 15 May in 1996 and 1997). Prior to the start of the experiment each year the 45 cm treatment plots were grazed with the appropriate animal species. Sward surface height was maintained at the required height by the addition and removal of non-experimental animals of the same breed and class as the core animals. The ratio of ewes to steers on the mixed sheep plus cattle treatment was maintained at 3:1 throughout. The experiment ended on 10 October and 25 September in 1996 and 1997 respectively. All swards were then grazed down to 4 cm prior to the winter, by grazing with non-experimental sheep. Nitrogen (N) was applied at 50 kg/ha (as 345% N) each year to all plots in spring (24 and 10 April in 1996 and 1997 respectively). A further application of 25 kg/ha was applied in late summer (7 and 12 August in 1996 and 1997 respectively). This was applied as 345% N fertilizer in 1996 and as a complete fertilizer (20.10.10) in 1997 to maintain P and K status.

Material and methods

Experimental design, site and animals The experiment was a 2 3 factorial design, with two sward surface height treatments (4 5 and 810 cm) and three combinations of animal species viz., sheep only, cattle only and sheep plus cattle. There were two replicate plots of each treatment combination and the experiment was conducted over 2 years consecutively (1996 and 1997). To minimize plot effects and increase the efciency with which treatment effects could be estimated, a partial crossover design was used. In each of the two replicates, there were four blocks of pasture, each of approximately 3 ha. In 1996 half of the blocks had the cattle plus sheep treatment, while the other half were divided into two areas, one of 05 ha and one of 25 ha. The different areas were used so that approximately equal numbers of sheep and equal numbers of cattle would be accommodated on the relevant treatments. The smallest of each pair was allocated to the sheep-only treatment while the larger was allocated to the cattle-only treatment. In 1997 the blocks which had the monospecies grazing were allocated to the mixed grazing treatment (the dividing fence in each block was removed) and vice versa.

Measurements Sward surface height was estimated on each plot twice per week using a HFRO sward stick (Barthram et al., 1986), by taking 40 measurements per plot. Live weight of core sheep and cattle was recorded at the start and end of the experiment each year and every 4 weeks, including at the time of weaning of lambs.

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Sward surface height and mixed grazing The live-weight gain per ha per day was calculated from the live-weight gain of the core animals and the number of animal grazing days, including non-experimental animals which grazed the plots to maintain sward height. The total liveweight gain per ha comprises the total live weight of steers, ewes and lambs. On four occasions in year 1 (30 May, 16 July, 27 July and 14 October) and on ve occasions in year 2 (13 May, 8 July, 12 August, 2 September and 23 September), the botanical composition of the pastures was estimated. On each occasion, six quadrats (025 050 m) were selected at random in each plot. The herbage in each quadrat then cut to ground level using a scalpel. The herbage was subsampled and separated into live grass lamina, live grass pseudostem, live grass reproductive stem, live clover ower, other live species and dead material. Following washing and drying (oven drying at 808C) the dry weight of all components was measured. Live-weight gain Live weights and live-weight gain are shown in Table 3. The live-weight gain of the steers was higher on the taller sward height treatment (960 v. 862 g/day; P , 005), but there was no difference in live-weight gain between the steers on the cattle-only treatment and the sheep plus cattle treatment. Sward height treatment did not affect the live-weight gain of the ewes (71 g/day for both sward height treatments). The ewes on the mixed, sheep plus cattle treatment gained more live weight than those on the sheep-only treatment (82 v. 61 g/day; P , 0001) and as a consequence were, on average, 36 kg heavier at the end of the experiment each year. The live-weight gain of the lambs up until weaning, when they were removed from the experiment, was signicantly higher on the shorter of the two sward height treatments (255 v. 228 g/day; P , 005). There was no effect of animal species treatment on the 4 5 cm sward height treatment, but on the 810 cm sward height treatment the lambs on the mixed grazing treatment had signicantly (P , 005) higher live-weight gains. They were therefore 20 kg heavier at weaning.

Statistical analysis All analyses were performed using Genstat (Lawes Agricultural Trust, 1990). Analysis of variance was performed using year, animal species combination and sward height treatment as main effects. Live weight and live-weight gain were analysed using individual animals as the experimental unit and output per ha was analysed using the value per plot as the unit of observation.

Output per hectare The number of animal grazing days for each species and the live-weight gain per ha per day is given in Table 4. Within a species and class of stock the number of animal grazing days per ha reects the mean stocking density for that class of stock. The total live-weight gain per ha was higher (although not signicantly so) on the 45 cm sward height treatment because of the higher stocking densities. The total output per ha was similar for all species combinations, there being no signicant differences (P . 005) between the cattleonly, sheep-only and sheep plus cattle treatments.

Results
Sward height, herbage mass and sward composition Overall the mean sward heights were within the required range (Table 1) except for a short period in June when they tended to be higher than required. The 810 cm sward height treatment had a signicantly higher (P , 0001) mean herbage mass (1831 kg dry matter (DM) per ha) than the 45 cm sward height treatment (1114 kg DM per ha), a higher proportion of dead material (0229 v. 0182; P , 005) and a greater proportion of grass pseudostem (0013 v. 0007; P , 005) and grass reproductive stems (0104 v. 0060; P , 001) (Table 2). The only effects of animal species were that grazing by cattle resulted in signicantly higher proportions of clover leaf plus petiole (0048, 0034 and 0028 for the cattle-only, sheep plus cattle and sheep-only treatments respectively; P , 001) and clover stolon (0009, 0006 and 0005; P , 005). However it should be noted that the proportions of clover in all the swards were low.
Table 1 Mean sward surface height

Discussion
This is the rst reported experiment in which mixed grazing by sheep and cattle has been compared with mono-species grazing when sward height has been controlled to levels that are known to give high levels of animal performance. It has been shown previously that sward surface heights of 45 cm and 810 cm provide near maximum intakes of herbage for sheep and cattle respectively (Wright, 1988; Penning et al., 1991).

Sward height treatment 4 5 cm Animal species Mean (cm) s.e. Cattle only 518 0093 Cattle plus sheep 480 0060 Sheep only 468 0066 Cattle only 951 0166 8 10 cm Cattle plus sheep 879 0136 Sheep only 838 0131

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Wright, Jones, Davies, Davidson and Vale

Table 2 Sward composition (proportion of dry matter (DM)) and mass (kg DM per ha) Sward height treatment 45 cm Animal species Live grass Leaf Pseudostem Reproductive stem Live clover Leaf and petiole Stolon Flower Other species Dead material Herbage mass Cattle only 0601 0007 0063 0050 0010 0001 0028 0241 1030 Sheep plus cattle 0616 0011 0062 0043 0009 00 0027 0232 1033 Sheep only 0602 0005 0088 0029 0007 00 0050 0224 1279 Cattle only 0502 0017 0087 0047 0009 00 0030 0309 1784 8 10 cm Sheep plus cattle 0547 0010 0136 0024 0003 00 0036 0245 1933 Sheep only 0545 0012 0135 0028 0004 00 0026 0251 1778 s.e.d. 00325 00037 00240 00076 00025 000042 00092 00226 1360

From known relationships between sward surface height and herbage intake and live-weight gain of cattle (Wright et al., 1986; Wright, 1988; Wright and Whyte, 1989), it would be expected that the 45 cm sward height treatment would lead to a reduction in herbage intake and live-weight gain. This was indeed the case for live-weight gain but no estimates of herbage intake are available in the present study. For ewes there was no effect of sward height treatment on live-weight gain but the lambs on the 8 10 cm sward gained live weight at a slightly lower rate than those on the short sward height treatment. It is possible that the digestibility of the herbage selected by the lambs on the taller treatment was lower, and this would account for the apparent discrepancy in the two variables. Previous detailed studies on the relationship between sward height and structure have shown that swards that are grazed by sheep at a sward surface height of 810 cm have a much higher proportion of reproductive tillers and a higher mass of reproductive stems and pseudostem than 4 5 cm swards (Penning et al., 1991; Wright et al., 2000). These sward components have a lower digestibility than vegetative leaf and if they had been selected by the lambs then this might explain the lower liveweight gain on the taller swards.

There was a considerable advantage in the mixed sheep plus cattle treatment in terms of ewe live-weight gain, on both sward height treatments, similar to that reported in other experiments of mixed grazing by sheep and cattle (see review by Nolan and Connolly (1977)). The 36 kg advantage in live weight at the end of the experiment in October is equivalent to about one-third of a unit of body condition score (Russel et al., 1969). If this advantage was maintained until the start of the mating period, it might be expected to result in a higher ovulation rate and an increase in lambing rate of about 10 lambs per 100 ewes mated (Gunn, 1983). The nding that mixed grazing had a greater effect on the live-weight gain of the sheep than the steers is in accordance with previous studies on the mixed grazing by sheep and cattle. Although in the study of Nolan and Connolly (1989) the live-weight gain of both steers and lambs was greater under a mixed grazing regime, the proportional benet to the lambs was greater. Dickson et al. (1981) recorded greater live-weight gain of steers when they grazed with ewes and lambs, although it is not clear if this was due to a general change in grazing pressure rather than the mix of species per se. Other studies have shown enhanced performance of sheep but not cattle, under mixed grazing (Bennett et al.,

Table 3 Mean live weight and live-weight changes Sward height treatment 4 5 cm Animal species Steers Initial live weight (kg) Final live weight (kg) Live-weight gain (g/day) Ewes Initial live weight (kg) Final live weight (kg) Live-weight gain (g/day) Lambs Initial live weight (kg) Final live weight (kg) Live-weight gain (g/day) Cattle only 346 463 850 Sheep plus cattle 345 466 875 411 531 85 167 344 260 402 484 57 172 342 250 Sheep only Cattle only 347 475 928 8 10 cm Sheep plus cattle 346 482 991 406 517 78 171 337 243 401 493 64 172 317 212 Sheep only s.e.d. 64 95 62 112 162 87 077 122 14

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Table 4 Output per ha Sward height treatment 45 cm Animal species Steers Animal grazing days per ha Live-weight gain (kg/ha per day) Ewes Animal grazing days per ha Live-weight gain (kg/ha per day) Lambs Animal grazing days per ha Live-weight gain (kg/ha per day) Total live weight gain (kg/ha per day) Cattle only 602 490 Sheep plus cattle 326 284 974 84 517 133 502 2941 145 1259 305 450 Sheep only Cattle only 466 437 810 cm Sheep plus cattle 241 237 722 60 405 100 396 2312 145 1224 258 403 Sheep only s.e.d. 748 555 3618 506 1467 334 786

490

437

1970; Hamilton and Bath, 1970; Van Keuren, 1970; Boswell and Cranshaw, 1978; Abaye et al., 1994). Studies from ecology can help explain why small grazers may benet more than large grazers from mixed species grazing. Large species tend to be less selective in their grazing behaviour (Gordon and Illius, 1988) and select plant material of a lower digestibility or protein content (OwenSmith and Novellie, 1982). Larger ruminants can also digest poorly digested forages better than smaller ruminants because of a longer retention time in the rumen (Blaxter and Wainman, 1966; Poppi et al., 1980). Thus cattle tend to select taller, less digestible patches of grass than sheep, especially if these patches are 810 cm tall or above, as this will tend to maximize their intake (Wright 1988), although they tend not to graze tall grass that surrounds cattle dung pats. However, grazing by one species can affect the structure and composition of the sward to the benet of another, termed facilitation by ecologists. The grazing of tall grass by large herbivores has been shown to result in the creation of pasture which is more conducive to grazing by smaller herbivores in, for example, the Serengeti Plains of Africa (McNaughton, 1984) and on grass/white clover swards in the UK (Wright et al., 2000). In the latter case, there was a reduction in the proportion of reproductive stems, as was also the case in the present experiment (Table 2). The greater response of the lambs to mixed grazing on the taller, 810 cm sward height treatment suggests that the facilitation effect was greater on the taller swards. Since their live-weight gain was lower on the tall sward height treatment than on the short treatment, this suggests that due to their smaller body size they were more disadvantaged than the ewes (whose live-weight gain was unaffected by sward height) by the taller, more reproductive and therefore lower digestibility sward. Thus the greater response of the lambs to mixed grazing at the taller sward height suggests that for these small animals the disadvantage of the taller sward is reduced by the facilitatory effect of grazing with cattle. The exact reasons for this are not clear. It is well recognized that grazing by cattle, especially at taller sward height, leads to patches of infrequently and frequently grazed areas (Gibb and Ridout, 1988 and 1986; Wright and Whyte, 1989). The frequently grazed areas are

shorter and the re-growth will have a higher proportion of leaf and probably be more attractive to grazing sheep, since they will be able to achieve a higher rate of intake of digestible organic matter than when grazing the tall patches. Although the short sward height treatment will also have had short vegetative swards there may have been more competition for them from the cattle. An alternative explanation is that tall patches tend to be initiated due to cattle not grazing close to cattle dung pats, and the mixed grazing treatment may have allowed sheep to graze these areas which, due to their proximity to the dung, may have had a higher nutrient content (Ronnel et al., 1980). This may have been more marked in the 810 cm treatment as there would have been less pressure for cattle to graze close to the dung pats. Although the clover content of the sward was low, grazing by cattle resulted in slightly higher clover contents than on the sheep-only swards. Although higher clover content in the diet can result in greater live-weight gain of lambs (Thomson, 1984) it appears that this is not the reason in the present experiment for the greater response to mixed grazing at the taller sward height: the clover content of the cattle plus sheep and the sheep-only treatment on the taller sward height was very similar. Although some studies of mixed grazing have shown increased output of live-weight gain per unit area (see review by Nolan and Connolly (1977)), this was not the case in the present experiment. However, in many cases it is not clear if the changes in output per ha are due to the way in which the studies were designed. In the present study, since stocking rates were adjusted to maintain constant sward surface heights, any difference in herbage growth rates or levels of utilization would have been reected in the total output per unit area. Although not signicantly different (P 012) there was a tendency for output per ha to be slightly higher on the 4 5 cm sward height treatment. This would be predicted by the results of Bircham and Hodgson (1983) who showed that as sward surface height increased, a maximum level of utilization was reached at a height of 3 to 4 cm, with a slight decline at heights between 4 and 8 cm. However, there was no indication of any difference in output per hectare and therefore efciency of utilization due to

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Wright, Jones, Davies, Davidson and Vale animal species combination. This suggests that when the level of utilization is controlled there is no inherent increase in efciency of use of pastures due to mixed grazing per se, and that in experiments where such an increase has been claimed, this may be due to an artifact of the experimental design.
Lawes Agricultural Trust. 1990. Genstat 5 reference manual. Oxford University Press. Levin, S. A. 1970. Community equilibrium and stability and an extension of the competitive exclusion principal. The American Naturalist 104: 413-423. McNaughton, S. J. 1984. Grazing lawns: animals in herds, plant form, and consolidation. The American Naturalist 124: 863-886. Nolan, T. and Connolly, J. 1977. Mixed stocking of sheep and steers a review. Herbage Abstracts 47: 367-374. Nolan, T. and Connolly, J. 1989. Mixed v. mono grazing by steers and sheep. Animal Production 48: 519-533. Nolan, T. and Connolly, J. 1992. System approach to soilplant-animal interactions. In Natural resource development and utilization (ed. R. de Jong, T. Nolan and J. van Bruchem), pp. 12-30. Wageningen. Owen-Smith, N. and Novellie, P. 1982. What should a clever ungulate eat? American Naturalist 119: 151-178. Parsons, A. J., Leaffe, E. L., Collet, B. and Stiles, W. 1983. The philosophy of grass production under grazing. Journal of Applied Ecology 20: 127-139. Penning, P. D., Rook, A. J. and Orr, R. J. 1991. Patterns of digestive behaviour of sheep continuously stocked on monocultures of rye grass or white clover. Applied Animal Behaviour Science 31: 237-250. Poppi, D. P., Minson, D. J. and Ternouth, J. H. 1980. Studies of cattle with sheep eating leaf and stem fractions of grasses. The voluntary intake, digestibility and retention time in the reticels-rumen. Australian Journal of Agricultural Research 32: 99-108. Ronnel, F., Nolan, T. and Connolly, J. 1980. Some effects of mixed grazing on pasture growth and chemical composition. In Proceedings of workshop on mixed grazing, Galway 1980 (ed. T. Nolan and J. Connolly), pp. 127-139. An Foras Taluntais. Russel, A. J. F., Doney, J. M. and Gunn, R. G. 1969. Subjective assessment of body fat in live sheep. Journal of Agricultural Science, Cambridge 72: 451-454. Thomson, D. J. 1984. The nutritive value of white clover. In Forage legumeso, ccasional symposium no. 16 (ed. D. J. Thomson), pp. 78-92. British Grassland Society. Van Keuren, R. W. 1970. Symposium on pasture methods for maximum production in beef cattle: pasture methods for maximizing beef cattle production in Ohio. Journal of Animal Science 30: 138142. Vance, R. R. 1984. Interference competition and the co-existence of two competitors on a single limiting resource. Ecology 65: 13491357. Volterra, V. 1928. Variations and uctuations of the number of individuals in animal species living together. Journal du Conseil. Conseil Permanent International pour lExploration de la Mer 3: 3-51. Wright, I. A. 1988. Suckler beef production. In Efcient beef production from grass, occasional symposium no. 22 (ed. J. Frame), pp. 51-64. British Grassland Society. Wright, I. A., Jones, J. R. and Parsons, A. J. 2000. Effects of grazing by sheep or cattle on sward structure and subsequent performance of weaned lambs. Grass and Forage Science 56: 138150. Wright, I. A., Russel, A. J. F. and Hunter, E. A. 1986. The effect of winter food levels on compensating growth of weaned, suckled calves grazed at two small heights. Animal Production 43: 211-227. Wright, I. A. and Whyte, T. K. 1989. Effects of sward heights on the performance of continuously stocked spring-calving beef cows and their calves. Grass and Forage Science 44: 259-266.

Acknowledgements
This work was funded by the Ministry of Agriculture Fisheries and Food and the Scottish Executive Environment and Rural Affairs Department. We are grateful to the staff of the Bronydd Mawr Research Centre for the care of the experimental animals.

References
Abaye, A. D., Allen, V. G. and Fonttenot, J. P. 1994. Inuence of grazing cattle and sheep together and separately on animal performance and forage quality. Journal of Animal Science 72: 10131022. Barthram, G. T., Elston, D. A. and Bolton, C. R. 1986. A comparison of two methods and quadrant mixing for determining the vertical distribution of herbage waste in ryegrass contaminated pastures. Grass and Forage Science 55: 193-200. Bennett, D., Morley, F. H. W., Clark, K. W. and Dudzinski, M. L. 1970. The effect of grazing cattle and sheep together. Australian Journal of Experimental Agriculture and Animal Husbandry 10: 694709. Bircham, J. S. and Hodgson, J. 1983. The inuence of sward conditions on rates of herbage growth and senescence in mixed swards under continuous stocking management. Grass and Forage Science 38: 323-331. Blaxter, K. L. and Wainman, F. W. 1966. The fasting metabolism of cattle. British Journal of Nutrition 20: 103-111. Boswell, C. D. and Cranshaw, L. J. 1978. Mixed grazing of cattle and sheep. Proceedings of the New Zealand Society of Animal Production 38: 116-120. Connolly, J. 1987. On difculties with replacement series methodology in mixture experiments. Journal of Applied Ecology 23: 125137. Dickson, I. A., Frame, J. and Arnot, D. P. 1981. Mixed grazing of cattle and sheep versus cattle only in an intensive grassland system. Animal Production 33: 265-272. Farnsworth, K. D., Focardi, S. and Beecham, J. P. 2000. Grassland herbivore interactions: how do grazers co-exist? American Naturalist 159: 24-39. Gibb, M. J. and Ridout, M. S. 1988. Applications of double normal frequency distributions tted to measurements of sward height. Grass and Forage Science 43: 131-136. Gibb, M. J. and Ridout, M. S. 1986. The tting of frequency distributions to height measurements on grazed swards. Grass and Forage Science 41: 247-249. Gordon, I. J. and Illius, A. W. 1988. Incisor arcade structure and diet selection in ruminants. Functional Ecology 2: 15-22. Gunn, R. G. 1983. The inuence of nutrition on the reproductive performance of ewes. In Sheep production, 35th Easter school in agricultural science (ed. W. Haresign), pp. 99-110. University of Nottingham Press. Hamilton, D. and Bath, J. G. 1970. Performance of sheep and cattle grazed separately and together. Australian Journal of Experimental Agriculture and Animal Husbandry 10: 19-26. Lambert, M. G. and Guerin, H. 1989. Competitive and complementary effects with different species of herbivore in their utilization of pastures. Proceedings of the XVI international grassland congress, Nice. 1989, pp. 1785 1789.

(Received 27 June 2005Accepted 3 November 2005)

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