Sward height and bite size affect the functional response of barnacle geese Branta leucopsis

Intake rate, the rate in which herbivores can process their food, is presumed to be an important factor in habitat selection down to the scale of the foraging patch. Much attention has been given to the selection of swards of high nutritional quality, but much less has been given to the influences of sward structure on patch selection in small herbivores. In this study we tested the effects of sward density and height on the functional foraging response of barnacle geese, Branta leucopsis. The functional response curve for herbivores describes how intake rate is affected by food availability. We conducted feeding trials to determine intake rate and bite size of barnacle geese on experimentally manipulated swards. Results indicate that intake rate is mainly dependent on sward height and that there is a strong correlation between bite size and intake rate. Sward density does not influence the rate of food consumption; it is, however, a crucial parameter affecting potential total yield. We conclude that bite size is the crucial parameter influencing intake rate. Bite size is explained both by sward height and individual differences in bill morphology. Furthermore, intake rate seems to be dependent on the physical structure of the grass species consumed.     

The allometry of patch selection in ruminants

An axiomatic feature of food consumption by animals is that intake rate and prey abundance are positively related. While this has been demonstrated rigorously for large herbivores, it is apparent from patch selection trials that grazers paradoxically tend to prefer short, sparse swards to tall, dense swards. Indeed, migratory herbivores often shift from areas of high to low sward biomass during the growing season. As nutritional quality is an inverse function of grass abundance, herbivores appear to sacrifice short-term intake for nutritional gains obtainable by eating sparse forage of higher quality. Explicit models of this trade-off suggest that individual ruminants maximize daily rates of energy gain by choosing immature swards of intermediate biomass. As body mass is related positively to both ruminant cropping rates and digestibility, there should be an allometric link between grass abundance and energy maximization, providing a tool for predicting patterns of herbivore habitat selection. We used previously published studies to develop a synthetic model of trade-offs between forage abundance and quality predicting that optimal sward biomass should scale allometrically with body size. The model predicts size-related variation in habitat selection observed in a guild of grazing ungulates in the Serengeti ecosystem.     

Body size dimorphism and sexual segregation in polygynous ungulates: an experimental test with Soay sheep

Sexual segregation in Soay sheep (Ovis aries) was investigated using an experimental approach in order to test the sexual dimorphism-body size hypothesis. Two corollaries of the sexual dimorphism-body size hypothesis were tested: (1) in dimorphic species males, the larger sex, have relatively smaller bite sizes on short swards because of the scaling of incisor arcade with body weight, and (2) they move off earlier to feed on taller but poorer-quality swards when such swards are patchily distributed on a scale which enables the spatial segregation of individuals. Patch choice between sexes was estimated using a matrix of grass patches which differed in both quality and biomass of grass on offer (HQ: high-quality-low-biomass; LQ: low-quality-high-biomass). Sex differences in patch choice and grazing behaviour were tested in short-term preference trials. Incisor breadth showed no significant difference between sexes. On the other hand, muzzle width was dimorphic, with females having a narrower muzzle than males. Bite size was significantly different between the sexes, being smaller in females than in males, although it was not significantly different between sward types. Females had a higher bite rate than males and the bite rate was higher in the HQ sward type than the LQ sward type. When the effect of body mass was removed, no sex differences in muzzle size, bite size or bite rate were found. The intake rate did not differ between the sexes or between sward types. Whilst both sexes preferred the HQ sward type, females spent a significantly longer time feeding on the LQ sward type than did males. The difference detected between the sexes in patch choice was not consistent directly with the sexual dimorphism-body size hypothesis. Alternative explanations based on sex differences in foraging behaviour in relation to body mass sexual dimorphism are discussed to explain the result. Received: 1 February 1999 / Accepted: 12 May 1999     

Foraging behaviour of wild impala (Aepyceros melampus) and Burchell's zebra (Equus burchelli) in relation to sward height

Foraging behaviour plays a key role in the interaction between herbivores and vegetation, their predominant food source. Understanding this interaction is crucial to providing information that is useful for conservation of herbivores. The objective of this study was to determine how sward height influences functional response and movement patterns of free ranging wild impala and zebra at the Kenya Wildlife Service Training Institute. The study was conducted for 3 months. Sward height is an important parameter that indicates how intensive a sward has been grazed and it influences intake rate through its effect on bite size. Bite size, instantaneous intake rate, specific mass intake rate and feeding station interval for impala and zebra increased with sward height. Sward height in combination with an animal's sex was found to have a profound effect on specific mass intake rate in impala. Zebra had a longer feeding station interval and lower stepping rate in tall swards compared to impala. Despite differences in their specific body mass and digestive strategies, impala and zebra maximized their intake rates in tall swards as a trade‐off among the swards. Tall swards are therefore critical in the study area and should be protected from bush encroachment which is a persistent problem.     

Foraging behaviour of brent geese, Branta b. bernicla, on grasslands: effects of sward length and nitrogen content

Feeding behaviour and preferences of brent geese, Branta b. bernicla were observed on pastures of different sward lengths and nitrogen contents. On swards of 2.0-6.0 cm captive geese took larger bites and had a higher intake rate when feeding on 6-cm swards than when feeding on shorter ones. In the field wild geese chose 6-cm swards in preference to both shorter and longer ones. For unfertilized pastures there was a strong negative relationship between nitrogen content and sward height: 11-cm swards contained 2.8% nitrogen, 3.5-cm swards 4.2% nitrogen. Application of 75 kg N ha^-1 organically based fertilizer at the end of October eliminated this relationship between nitrogen content and sward height, swards of all heights then having a mean content of 4.1% N. On fertilized plots the geese preferred swards longer than 6 cm with no indication of a decrease in preference up to the maximum height investigated, 16 cm. Breaking strain of grass laminae was measured to give an indication of the proportion of strengthening tissues in the leaves and hence their digestibility. Apical laminae from longer swards had a higher breaking strain than those from shorter swards. Unfertilized swards had a higher breaking strain than fertilized swards but the difference in breaking strain between long and short swards was the same on fertilized and control treatments. These results are discussed in relation to the forage maturation hypothesis and are interpreted as indicating that the primary constraint on maximizing energy intake rates is not the reduced energy digestibility of older foliage but the reduced nitrogen content in the higher-biomass swards. We conclude that it is the balance between maximizing energy intake and nitrogen absorption rates which results in the observed preference for intermediate-height swards.     

Feeding patch selection by herbivorous Anatidae: the influence of body size, and of plant quantity and quality

Recent findings suggest that herbivores select feeding sites of intermediate biomass in order to maximise their digestible nutrient intake as the result of the trade-off between forage quality and quantity ('forage maturation hypothesis'). We propose a reformulation of this hypothesis which recognises this trade-off, but also underlines that constraints due to body mass (i.e. metabolism and digestive constraints, size of the feeding apparatus) can lead to variations in grazing patterns. We tested this latter hypothesis experimentally in three species of herbivorous Anatidae of different body mass: the wigeon Anas penelope (in our study c. 620 g), the barnacle goose Branta leucopsis (c. 2000 g), and the greylag goose Anser anser (c. 3500 g). Each species was tested separately from 0600 to 0930 hours, in an enclosure with a mosaic of patches of grass of three different heights: short, medium and tall. The behaviour, and the location (i.e. patch) of each individual were recorded every 5 minutes. Our results show important interspecific differences in intake rates resulting in different feeding site selection: wigeon and barnacle goose fed fastest on the shortest swards, and selected short grass which was also of higher quality. Tall grass provided the highest dry matter intake rate and digestible protein intake for greylag geese, and they preferred these swards. These choices allowed the birds to maximise their digestible nitrogen intake rate rather than dry matter intake rate and our results thus underline the importance of nitrogen as a major currency for foraging decisions in herbivorous Anatidae. Since the birds selected the two extreme sward heights (instead of the medium one), the results give support to our hypothesis and underline the role of body size as an important cause of variations in patch selection in herbivorous Anatidae.     

Ecological Engineering by a Mega-Grazer: White Rhino Impacts on a South African Savanna

Herbivory can change the structure and spatial heterogeneity of vegetation. We ask whether all species of grazers in a savanna ecosystem can have this effect or whether megaherbivores (>1000 kg) have a ‘special’ role that cannot be replicated by other species of grazers. We performed a replicated landscape scale experiment that examined the effects of White Rhino on the grass sward, on other species of grazing mammals and on the movement of fire through the landscape. White Rhino maintained short grass (‘lawn’) patches in mesic areas (∼750 mm pa) with increases in grass sward height when they were removed. Other species of grazers were unable to maintain short grass communities when White Rhino were removed. In semi-arid areas (∼600 mm pa) other, smaller grazers were able to maintain short grass communities in the absence of White Rhino and sward height did not increase. White Rhino removals affected fire by increasing fuel loads and fuel continuity. This resulted in larger, less patchy fires. We propose that the White Rhino acts as an influential ecosystem engineer, creating and maintaining short grass swards, which alter habitat for other grazers and change the fire regime. These results indicate the existence of context-dependent facilitation between White Rhino and other grazers in mesic, but not in semi-arid, savannas. Such top down effects on the ecosystem may have been much more widespread before the extinction of large grazers in the Pleistocene.     

Foraging decisions underlying restricted space use: effects of fire and forage maturation on large herbivore nutrient uptake

Recent models suggest that herbivores optimize nutrient intake by selecting patches of low to intermediate vegetation biomass. We assessed the application of this hypothesis to plains bison (Bison bison) in an experimental grassland managed with fire by estimating daily rates of nutrient intake in relation to grass biomass and by measuring patch selection in experimental watersheds in which grass biomass was manipulated by prescribed burning. Digestible crude protein content of grass declined linearly with increasing biomass, and the mean digestible protein content relative to grass biomass was greater in burned watersheds than watersheds not burned that spring (intercept; F_1,251 = 50.57, P < 0.0001). Linking these values to published functional response parameters, ad libitum protein intake, and protein expenditure parameters, Fryxell's (Am. Nat., 1991, 138 , 478) model predicted that the daily rate of protein intake should be highest when bison feed in grasslands with 400–600 kg/ha. In burned grassland sites, where bison spend most of their time, availability of grass biomass ranged between 40 and 3650 kg/ha, bison selected foraging areas of roughly 690 kg/ha, close to the value for protein intake maximization predicted by the model. The seasonal net protein intake predicted for large grazers in this study suggest feeding in burned grassland can be more beneficial for nutrient uptake relative to unburned grassland as long as grass regrowth is possible. Foraging site selection for grass patches of low to intermediate biomass help explain patterns of uniform space use reported previously for large grazers in fire‐prone systems.     

Repeated grazing of a salt marsh grass by moulting greylag geese Anser anser– does sequential harvesting optimise biomass or protein gain?

The effects of simulated goose grazing on common saltmarsh-grass Puccinellia maritima plants were tested on a Danish salt marsh during the flightless moulting period of greylag geese Anser anser (3–21 June 1998). Plants in an area exclosed from the influence of grazing and the nutrient effects of goose faeces were subject to removal of youngest lamina at 3-, 6-, 9- and 18-day intervals during this period. Average biomass and protein accumulation between harvests was highest at defoliation intervals of 9 days or more. Field observations from two separate study areas demonstrated geese returned to regraze the Puccinellia sward after 6–8 days and oesophageal contents from feeding geese showed selection for lamina lengths consistent with the results of clipping every 6 days. Geese therefore regrazed Puccinellia patches at shorter intervals than expected were they to maximise their intake of biomass or protein at each visit. However, total cumulative lamina elongation, equivalent to the long term gain during the entire moult period, showed no significant difference between the three most intensive defoliation treatments, which were significantly greater than those of plants defoliated at 18 day intervals. Highest overall lamina protein levels were maintained at 6- and 9-day defoliation intervals. This suggests geese regrazed Puccinellia patches at a rate that maximised their number of harvests during the flightless period, but maintained highest protein levels and overall biomass in the sward. This suggests, in line with earlier studies, that moulting greylag geese combine dietary selection, reduced nitrogen excretion and regrazing patterns to meet protein demands during regrowth of flight feathers.     

Patch depression in grazers: the roles of biomass distribution and residual stems

1. In this paper the results are presented of two experiments designed to examine the relationship between sward structure variables and the nature of patch depression experienced by a large grazing herbivore, Bos taurus, grazing from small patches of forage. In experiment 1 the spatial distribution of leaves within patches was varied. In experiment 2 the role of reproductive stems of different lengths as potential intake rate inhibitors was examined.
2. Both experiments were conducted by offering cattle hand-constructed swards of Orchardgrass ( Dactylis glomerata ) leaves attached to plywood boards. Each of three experimental animals was allowed to take 10, 20, 30, 45 or 60 bites from a patch and intake was estimated on the basis of pregrazing and postgrazing dry-matter. All trials were videotaped and the trial length in seconds was determined from playback of the tapes.
3. In both experiments patch depression (reduction in instantaneous intake rate) was due to decreases in bite mass concomitant with patch depletion. Handling time per bite was unaffected by depletion level. In experiment 1, different spatial configurations resulted in markedly different gain functions ranging from linear to sigmoid. In experiment 2, the presence in the sward of long (15 cm) reproductive stems slowed cumulative gains by restricting bite mass and by increasing the per-bite handling time relative to patches with shorter (5 cm) or no stems.
4. The study of patch depression mechanisms now needs to be generalized over a range of herbivore body sizes and feeding styles.     

Ecology of grazing lawns in Africa

Grazing lawns are a distinct grassland community type, characterised by short‐stature and with their persistence and spread promoted by grazing. In Africa, they reveal a long co‐evolutionary history of grasses and large mammal grazers. The attractiveness to grazers of a low‐biomass sward lies in the relatively high quality of forage, largely due to the low proportion of stem material in the sward; this encourages repeat grazing that concomitantly suppresses tall‐grass growth forms that would otherwise outcompete lawn species for light. Regular grazing that prevents shading and maintains sward quality is thus the cornerstone of grazing lawn dynamics. The strong interplay between abiotic conditions and disturbance factors, which are central to grazing lawn existence, can also cause these systems to be highly dynamic. Here we identify differences in growth form among grazing lawn grass species, and assess how compositional differences among lawn types, as well as environmental variables, influence their maintenance requirements (i.e. grazing frequency) and vulnerability to degradation. We also make a clear distinction between the processes of lawn establishment and lawn maintenance. Rainfall, soil nutrient status, grazer community composition and fire regime have strong and interactive influences on both processes. However, factors that concentrate grazing pressure (e.g. nutrient hotspots and sodic sites) have more bearing on where lawns establish. Similarly, we discuss the relevance of enhanced rates of nitrogen cycling and of sodium levels to lawn maintenance. Grazer community composition and density has considerable significance to grazing lawn dynamics; not all grazers are adapted to foraging on short‐grass swards, and differences in body size and relative mouth dimensions determine which species are able to convert tall‐grass swards into grazing lawns under different conditions. Hence, we evaluate the roles of different grazers in lawn dynamics, as well as the benefits that grazer populations derive from having access to grazing lawns. The effects of grazing lawns can extend well beyond their borders, due to their influence on grazer densities, behaviour and movements as well as fire spread, intensity and frequency. Variation in the area and proportion of a landscape that is grazing lawn can thus have a profound impact on system dynamics. We provide a conceptual model that summarises grazing lawn dynamics, and identify a rainfall range where we predict grazing lawns to be most prevalent. We also examine the biodiversity associated with grazing lawn systems, and consider their functional contribution to the conservation of this biodiversity. Finally, we assess the utility of grazing lawns as a resource in a rangeland context.     

The effect of differences in herbage height on the grazing behaviour of lactating Bennett's wallabies (Macropus rufogriseus rufogriseus)

The grazing behaviour of lactating Bennett's wallabies was studied on a number of different pasture types. Increases in mean sward height resulted in a greatly reduced bite rate and an increase in the search time per bite. All wallabies showed a diurnal rhythm in grazing activity irrespective of habitat type, with least grazing activity at midday and most at night. Animals maintained on very short grass swards compensated for the reduced herbage mass by extending their grazing activity into the midday period. Lactating animals grazed for longer, had higher biting rates and a greater number of estimated bites per day than non-lactating wallabies (17,700 vs. 7600 bites per day, respectively). In general, the response of wallabies to changes in herbage availability is similar to domestic ruminants, although wallabies may be able to graze for longer each day than ruminants.     

Competition between grass and clover in spring as affected by nitrogen fertiliser

Detailed measurements of irrigated ryegrass/white clover swards growing without interruption, with or without nitrogen fertiliser in spring, showed that the relative growth rate of clover was as great as that of grass, in the + N sward, and considerably greater than grass, in the – N sward. Clover leaves were not overtopped by grass leaves. Indeed, in both swards, clover had a greater proportion of its leaf lamina area in the upper, well lit, layers of the canopy than grass did. Consequently, clover had a greater mean rate of leaf photosynthesis in situ in the sward than grass. Clover's advantage in photosynthetic rate per leaf area was offset to some extent by its smaller ratio of leaf area to total above-ground dry weight than grass. The consequences of these results for our understanding of competition between grass and clover in mixed swards are discussed.     

Dietary and microtopographical selectivity of Greenland white-fronted geese feeding on Icelandic hayfields

The feeding ecology of Greenland white-fronted geese Anser albifrons flavirostris was studied during .spring staging in Iceland 1997. Geese feeding on Poa pratense dominated hayfields (> 80% cover) were highly selective, selecting for Deschampsia caespitosa which comprised only 10% of the sward. Geese fed most on the south-facing fringes of Deschampsia tussocks. Subsequent analysis showed that the southern fringes of Deschampsia tussocks supported significantly greater biomass (27% greater mass of green material) and that leaves growing on the southern faces had significantly higher protein content than those on the northern faces (33.9% vs 30.5%)_- It appears that the geese maximise their nutritional intake in spring by selecting the grass species of highest quality and taking the most nutritious parts of the plants.     

The herbivores’ dilemma: trade-offs between nutrition and parasitism in foraging decisions

An experiment was carried out using a trade-off framework to determine the rules of sward selection, in relation to gastrointestinal parasite dispersion, used by mammalian herbivores, and the effect of level of feeding motivation and parasitic status on these rules. Twenty-four sheep divided into four animal treatment groups resulting from two levels of feeding motivation (high and moderate) and two parasitic states (parasitised with Ostertagia circumcincta and non-parasitised) were presented with pairs of experimental swards which varied in N content (high and low), sward height (tall and short) and level of contamination with faeces and thus parasites (contaminated and non-contaminated). The selection for tall swards outweighed both the selection for N-rich swards and the avoidance of faecal contaminated swards. The selection for N-rich swards could not completely overcome faecal avoidance. Parasitism in animals with a moderate feeding motivation reduced their bite rates and grazing depths, thereby probably reducing the rate of ingestion of parasitic larvae. In contrast, highly feeding-motivated animals (including those parasitised) increased their bite rates and grazing depths, thereby increasing the rate of ingestion of parasites. The inclusion of parasite distributions, both in the environment and within herbivore host populations, is likely to advance optimal foraging theory by enhancing its predictive power. Received: 30 November 1999 / Accepted: 14 February 2000     

Dark-bellied Brent geese aggregate to cope with increased levels of primary production

We report on an aggregative response of Dark-bellied Brent geese to increased productivity of the vegetation during the growing season on agricultural fields on the island of Schiermonnikoog, the Netherlands. Plant standing crop was found to be maintained at low levels in the fields where geese activity focussed, whereas the remainder of the fields escaped herbivore control and developed a high standing crop. This pattern can be explained by a decreased efficiency of grazing in vegetation with a high standing crop. In other words, the functional response of the geese is not monotonically increasing but dome-shaped. As a consequence, continuously grazed swards are more suitable for feeding than temporarily ungrazed swards. We present a model showing that, for a dome-shaped functional response, optimal foraging under increasing primary productivity leads to spatial heterogeneity in standing crop. Beyond a certain threshold value, a further increase in productivity leads to a progressive release of vegetation from herbivore control and to the development of a high standing crop. Interestingly, our model suggests that only in a stable and predictable environment the aggregative behaviour of herbivores is able to maintain the intake rate close to its potential maximum. Misjudgement of patch quality by the herbivore or any other process disrupting the match between local primary production and consumption leads to a less than optimal intake, as suitable vegetation becomes depleted. This has important implications for ecological inferences, such as the prediction of carrying capacities in herbivore-dominated ecosystems.     

Grazing decisions of Soay sheep,Ovis aries,on St Kilda: a consequence of parasite distribution?

Relationships between the height of grass swards and herbage intake have been established for a number of large herbivores and sward height plays a major role in determining the selection of herbivore diets. However, tall grass swards also represent a more stable damp microclimate for gastrointestinal parasite larvae and tall swards contain generally greater numbers of parasites than short swards. Herbivores may then trade-off the need to maximise nutrient intake through the selection of tall swards with the need to avoid parasite ingestion. Stratified sward sampling techniques were used to determine the distribution of nutrients and parasites in a heterogeneous sward structure on the island of Hirta, St Kilda, which is grazed by a population of feral Soay sheep. The disparity between the short gap vegetation and the tall tussock vegetation was greatest in spring, when gap vegetation was some three-fold shorter than tussock vegetation; this led to tussocks offering greater nutrient and energy intake rate when compared to gap vegetation in spring. Parasites were concentrated in tussocks in spring, thus creating trade-offs. Such trade-offs were not present in summer when parasites were more evenly distributed across the gap/tussock sward structure and the nutritional advantages associated with grazing tussocks were diminished. The diet selection of the resident population of sheep was determined by recording the number of bites taken from gap and tussock vegetation by randomly selected focal animals over repeated 5-min grazing periods. Overall, all animals avoided grazing tussocks, which were most strongly avoided in the spring, and older animals avoided grazing tussocks to a greater extent. Overall, females with lambs and males avoided tussocks to a similar degree and both avoided them less than barren females. Faecal egg counts (FEC) of female sheep were negatively correlated with tussock selectivity and vice versa for male sheep. The interaction between the grazing behaviour of each sex and FEC is discussed in relation to the immunocompetence and nutrient requirements of the different sexes. The maximum disparity between the costs and benefits of the trade-off studied occurred in late winter/early spring which also represents the time of greatest nutrient demand in the Soay sheep, since many are close to starvation and/or are in the advanced stages of pregnancy. Grazing decisions of individuals at this time, determining relative nutrient and parasite intake, may then be related to subsequent fitness and survival.     

Forage quality and patch choice by wapiti (Cervus elaphus)

Recent models suggest that herbivores might optimize energygain by selecting patches of intermediate vegetation biomass.We tested this hypothesis in wapiti (Cervus elaphus) by estimatingdaily rates of energy gain in relation to grass biomass andby measuring patch choice in experimental pastures in whichgrass biomass was manipulated by mowing. The digestible energycontent of grasses declined with increasing biomass due to maturationalchanges in fiber and lignin content. Daily rates of dry matterintake by wapiti increased with grass biomass at a deceleratingrate, implying a Type II functional response. Linking thesevalues to published ad libitum energy intake and energy expenditureparameters, Fryxell's (1991) model predicted that the dailyrate of energy gain should be highest when wapiti feed in grasslandswith 1000–1100 kg/ha. In trials in which grass biomasswithin a mosaic of patches was manipulated experimentally between800–2900 kg/ha, wapiti preferred patches of 1200 kg/ha,close to the value predicted by the energy gain model. Our resultssuggest that the rate of energy gain by wapiti is constrainedby both grass biomass and grass fiber content, the latter ofwhich varies inversely with grass biomass. Behavioral preferencefor grass patches of intermediate biomass and fiber contentcould help explain patterns of aggregation and seasonal migrationreported previously for wapiti.     

Seasonal and interannual variations in feeding station behavior of cattle: effects of sward and meteorological conditions

A feeding station is the area of forage a grazing animal can reach without moving its forefeet. Grazing behavior can be divided into residence within feeding stations (with bites as benefits) and movement between feeding stations (with steps as costs). However, relatively little information has been reported on how grazing animals modify their feeding station behavior seasonally and interannually in response to varying environmental conditions. The feeding station behavior of beef cows (Japanese Black) stocked on a tropical grass pasture (bahiagrass dominant) was monitored for 4 years (2010 to 2013) in order to investigate the association of feeding station behavior with meteorological and sward conditions across the seasons and years. Mean air temperature during stocking often exceeded 30°C during summer months. A severe summer drought in 2013 decreased herbage mass and sward height of the pasture and increased nitrogen concentration of herbage from summer to autumn. A markedly high feeding station number per unit foraging time, low bite numbers per feeding station and a low bite rate were observed in summer 2013 compared with the other seasons and years. Bite number per feeding station was explained by a multiple regression equation, where sward height and dry matter digestibility of herbage had a positive effect, whereas air temperature during stocking had a negative effect (R²=0.658, P<0.01). Feeding station number per minute was negatively correlated with bite number per feeding station (r=–0.838, P<0.001). It was interpreted that cows modified bite number per feeding station in response to the sward and meteorological conditions, and this largely determined the number of feeding stations the animals visited per minute. The results indicate potential value of bite number per feeding station as an indicator of daily intake in grazing animals, and an opportunity for livestock and pasture managers to control feeding station behavior of animals through managements (e.g. fertilizer application, manipulation of stocking intensity and stocking time within the day).