Influence of sward height on diet selection by horses

https://doi.org/10.1016/j.applanim.2004.08.001Get rights and content

Abstract

Foraging herbivores are often faced with spatial and temporal heterogeneity within the vegetation they have available to graze and therefore have to make decisions where and when to graze. The study reported in this paper investigated the influence of sward height on diet selection by horses grazing perennial rye-grass swards. The study comprised two experiments. In Experiment 1, perennial rye-grass paddocks were mown to four sward heights (heights: 3.5, 4.5, 7.5 and 15 cm) to create a patchy environment. Within each paddock one horse grazed for a period of 1 h during which residence time, number of bites and frequency of visits per patch were recorded. This was replicated with all seven horses used in the experiment. The same experiment was repeated in Experiment 2, but without mowing the field and allowing 1 week of re-growth for each paddock. During both experiments horses entered equally often but resided significantly longer on patches with long grass (15 cm) than on those with short grass (below 4.5 cm; P < 0.05). Grazing time and number of bites on a patch were highly correlated. The number of bites on patches with the highest sward height was greater than that on short patches (P < 0.05). Horses behaved as selective grazers, feeding mainly on grass taller than 7 cm. In Experiment 2, re-growth of the sward reduced the difference in sward height between the patches. Time spent per patch and total numbers of bites taken were less affected by sward height than in Experiment 1. It is suggested that horses behaved as energy maximisers (residing longer periods on patches and increasing number of bites taken). These data complement previous findings that bite dimension and bite mass increase with increasing sward height. When grazing, a horse rarely resided on a preferred patch for a long duration of time (maximum 305 s, median 79 s), but moved on after a few minutes. They sampled their environment continuously, but almost exclusively returned to long patches for feeding.

Introduction

Diet selection is defined as an animal's choice of food from a range of different food items (Norbury and Sanson, 1992) influenced by many factors such as nutritional demands, toxic plant compounds, forage availability, social interactions and predator risks (Krebs and Davis, 1993). Selecting the ‘right’ food is a trade-off between costs and benefits and has a short- and long-term effect on the animal's fitness (Krebs and Davis, 1997). Foraging herbivores are often faced with spatial and temporal heterogeneity within the vegetation they have available to graze (Senft et al., 1987) and therefore have to make decisions where and when to graze. Optimal foraging models (Belovsky, 1986, Stephens and Krebs, 1986) predict that animals forage in ways that optimise the rate of food intake. Patch models like the marginal value theorem (Charnov, 1976) even suggest that ‘when the intake rate, in any patch, drops to the average rate of the habitat, the animal should move on to another patch’. If animals do behave in such a way, they must be able to gather and interpret information about their environment in order to modify their grazing behaviour to increase the rate of reward (Bailey et al., 1996). Studies on grazing behaviour by sheep, cattle and horses (Black and Kenney, 1984, Laca et al., 1992, Ungar and Ravid, 1999, Ungar et al., 2001, Illius et al., 1992, Naujeck and Hill, 2003) showed that they discern differences in grass height and that they increased bite dimensions with increasing grass height. To maximize voluntary intake, foraging models suggest that herbivores should select from tall, rather than short, grass. However the feeding site may be characterized by intermediate biomass in order to maximize intake of digestible nutrients as a result of the ‘trade-off’ between herbage quality and quantity (forage maturation hypothesis; Durant et al., 2004). Such behaviour was confirmed in studies on sheep and cattle (Bazely, 1990, Diestel et al., 1995), but little is known about diet selection by horses (Duncan, 1992, Fleurance et al., 2001, Putman et al., 1987, Gordon, 1989).

The present study aimed to investigate the influence of sward height on diet selection by horses grazing perennial rye-grass swards. For this purpose, patchy paddocks were created by mowing. Behaviour such as time spent on patches, number of bites per patch and frequencies of visits per patch were recorded.

Section snippets

Materials and method

The study was conducted at Writtle College (latitude 51°44′N, longitude 0°26′E, 32.5 m OD), UK in August 2002. It comprised two experiments. Experiment 1 investigated the grazing behaviour by horses on a heterogeneous paddock in which patches were cut to four different sward heights the day before measurements were made. In Experiment 2 grazing behaviour by horses was recorded on two of the same paddocks used in Experiment 1, but after one week of sward re-growth.

Vegetation

The data for sward height and dry matter mass are shown in Table 2, Table 3 (Experiments 1 and 2, respectively). The variability associated with sward height and dry matter yield are typical of that of perennial rye-grass swards after cutting to a series of heights (Experiment 1; Table 2). After 1 week of re-growth, there was a considerable increase in the variability of sward height and dry matter yield, reflecting an increase in heterogeneity in herbage production (Experiment 2; Table 3).

Number of visits per patch

The

Discussion

The results of this study suggest that horses behave as selective grazers when offered a choice in grass height. Previous studies on the influence of sward height on bite dimensions of horses showed bite depth, weight, volumes and bite area were not a fixed size, but increased with sward height (Naujeck and Hill, 2003). The selection for a particular sward height in a patchy environment would therefore have an influence on bite dimensions and total intake of herbage by the horse.

In the first

Acknowledgement

The authors wish to thank Writtle College for funding this study, which is a part of a wider investigation on grazing behaviour of horses.

References (32)

  • D.W. Bailey et al.

    Mechanisms that result in large herbivore grazing distribution patterns

    J. Range Manage.

    (1996)
  • D.R. Bazely

    Rules and cues used by sheep foraging in monocultures

  • G.E. Belovsky

    Optimal foraging and community structure: implications for a guild of generalist grassland herbivores

    Oecology

    (1986)
  • J.L. Black et al.

    Factors affecting diet selection by sheep II. Height and density of pasture

    Aus. J. Agric. Res.

    (1984)
  • E.L. Charnov

    Optimal foraging, the marginal value theorem

    Theor. Popul. Biol.

    (1976)
  • R.A. Diestel et al.

    Patch selection by cattle: maximization of intake rate in horizontally heterogeneous pastures

    Appl. Anim. Behav. Sci.

    (1995)
  • P. Duncan

    Horses and grasses

    The Nutritional Ecology of Equids and their Impact on the Camargue

    (1992)
  • D. Durant et al.

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

    J. Avian Biol.

    (2004)
  • G. Fleurance et al.

    Daily intake and the selection of feeding sites by horses in heterogeneous wet grasslands

    Anim. Res.

    (2001)
  • D. Frape

    Equine Nutrition and Feeding

    (1998)
  • A.F. Fraser et al.

    Farm Animal Behaviour and Welfare

    (1990)
  • I.J. Gordon

    Vegetation community selection by ungulates on the Isle of Rhum III. Determinants of vegetation community selection

    J. Appl. Ecol.

    (1989)
  • W.B. Healy

    Ingestion of soil by sheep

    Proc. NZ Soc. Anim. Prod.

    (1967)
  • D. Heath

    An Introduction to Experimental Design and Statistics for Biology

    (1995)
  • J. Hodgson

    Grazing Management

    (1990)
  • A.W. Illius et al.

    Constraints on diet selection and foraging behaviour in mammalian herbivores

  • Cited by (33)

    • Development of an apparatus for evaluating the visual and olfactory discrimination ability of cattle under controlled target visibility

      2021, Behavioural Processes
      Citation Excerpt :

      Discrimination and selection of preferred patches against less-preferred and non-preferred patches are crucial for animals foraging in grasslands to maximize energy and/or nutrient intake and better fit the environment (i.e. better growth, survival and reproduction). Patch discrimination and selection have been shown in a range of domestic and wild ungulates such as cattle (Bos taurus: Hirata et al., 2012, 2017; Wallis de Vries and Daleboudt, 1994), sheep (Ovis aries: Garcia et al., 2003; Prache and Damasceno, 2006), goats (Capra hircus: Illius et al., 1999), horses (Equus caballus: Edouard et al., 2010; Naujeck et al., 2005), wapiti (Cervus elaphus: Wilmshurst et al., 1995), Svalbard reindeer (Rangifer tarandus platyrhynchus: Van der Wal et al., 2000), wood bison (Bison bison athabascae: Bergman et al., 2001) and African grassland species (Bukombe et al., 2019; Venter et al., 2014). The process of patch discrimination by ungulates involves the senses of vision (sight), olfaction (smell), tactility (touch) and gustation (taste) (Arnold, 1966a, 1966b; Ginane et al., 2011; Hirata et al., 2015; Krueger et al., 1974; Vallentine, 1990).

    • Forage yield and its determination

      2018, Horse Pasture Management
    • Introduction to pasture ecology

      2018, Horse Pasture Management
    • A Review of Equine Grazing Research Methodologies

      2017, Journal of Equine Veterinary Science
    View all citing articles on Scopus
    1

    Current address: Institute of Land and Food Resources, University of Melbourne, Victoria 3010, Australia.

    View full text