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PRINCIPLES OF GRAZING ANIMAL NUTRITION "The range resource is by far the most variable commodity that is encountered in livestock

nutrition. It can vary in nutrient quality from a low comparable to sawdust to a high comparable to top quality alfalfa. It is capable of producing gains approaching 1.4 kg/day in growing classes of cattle at certain time of the year or of allowing starvation in some cattle at other times"... "Optimum range livestock production can only be achieved through compatible livestock, wildlife and forage management." (Raleigh and Lesperance 1973. In Church et. al Practical Nutrition Oregon State University Press). These authors should also have added compatible soil management to the list of practices for optimum range livestock production. Therefore, the Range Manager needs to be continually in touch with the whole range ecosystem because neglect of one part of the ecosystem can have serious consequences on the many other parts. Chemical Factors Influencing Nutrient Intake and Animal Productivity. The major nutrients required by ruminant animals, such as cattle, sheep and goats, through the herbage consumed are: 1) 2) 3) 4) 5) 6) Energy Protein (some of which can be in the form of non-protein N) Macro minerals - P, Ca, Mg, Na, S, K. Trace minerals - Cu, Se, Co, Zn, Mn, Mo, I Vitamins A and E Water

Of the first 4 categories listed above, energy (as digestible energy), protein, phosphorus and Vitamin A are the most likely nutrient deficiencies encountered by ruminant animals which will restrict intake and production. The macro minerals, other than phosphorus, such as K, Ca, and Mg are rarely in deficient amounts in grazed herbage. In some cases deficiencies of Na and S have been reported but of more significance is the possible high NaCl and Na 2S04 content of underground water supplies for livestock. High concentrations of these salts will render water unpalatable to livestock but much lower concentrations, particularly of Na2, S04, will interfere with the metabolism of some trace elements, notably Cu and Se, in animals thereby inducing deficiencies of these elements. The trace element content of herbage varies with soil type, the concentration of these elements in the soil and the ability of the plant to extract the element from the soil. Single or multiple deficiencies of all the listed trace elements have been reported in grazing animals. The trace elements are required for many of the enzymes used in metabolic processes within the animal and in many of the energy transfer pathways. Some plants are very efficient in extracting some trace elements from the soil and may accumulate them in toxic quantities. For example Astragalus bisulcata will accumulate selenium at levels up to ten times that which is toxic to sheep and cattle and is a frequent cause of death in these animals grazing range infested with this plant. Sweet clover (Melilotus spp) may accumulate molybdenum, if grown on soils of high Mo content, to levels that may become toxic to livestock. The mineral nutrition of livestock is very complex and goes beyond the scope of these lectures. However, it is important to understand that many of the minerals interact with each other. For example, the functions and requirements of an animal for P are very closely dependant on the amount of Ca in the diet. Ca requirement of animals is greatly increased when they are grazing plants which accumulate high levels of oxalates, such as Kochia (Kochia scoparia) or Greasewood
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(Sarcobatus vermiculatus). Adequate levels of Cu in plants may become inadequate for animal health and production if the diet also contains high levels of Mo, in which case insoluble copper molybdate is formed, or if the diet or water or both contain high levels of sulphates in which case insoluble copper sulphide is formed in the rumen. If both Mo and S04 are present, the Cu becomes bound as copper thiomolybdate and is almost totally unavailable to the animal. When this occurs, very high levels of Cu must be fed or Cu injected into the muscle of animals to prevent Cu deficiency occurring. These high levels of Cu would be toxic to animals eating herbage and/or water with low contents of Mo and/or S04. When in a deficient state, all minerals will reduce feed intake, animal growth, reproductive function and milk and wool production. In high concentrations all minerals can exert varying degrees of toxicity. Vitamin A deficiency does not occur when grazing animals consume green herbage but may be important on arid ranges or where animals are required to graze senesced herbage for long periods of time. Vitamin D rarely becomes deficient in grazing livestock because all animals, including man, can convert ultra-violet rays from the sun into Vitamin D in the skin. In ruminant animals, the B vitamins, which are a large and complex group of vitamins, are rarely a problem with respect to deficiency because the rumen micro-organisms can synthesize these vitamins in adequate amounts to meet the ruminant animal's requirements. The exception to this is vitamin B12 which can only be synthesized if adequate Co is present in the diet. The rest of the discussion in this section will be restricted to energy and protein because these are almost always the first two limiting factors in the nutrition of grazing animals and rarely is a mineral or vitamin deficiency uncomplicated by concurrent energy and/or protein deficiency. Energy Energy is measured as heat energy or as the heat of combustion of the feed. Energy in animal nutrition is partitioned into gross energy (GE), digestible energy (DE), metabolizable energy (ME) and net energy (NE). This is shown schematically in Figure 15. It should be noted that GE for all plant materials varies very little from 4.37 kcals/g, with the exception of plants that have high levels of oil in the seeds or leaves. The latter may have GE values of 5 to 7 kcals/g. However, the digestibility of plant materials varies considerably between and within plant species depending on soil type, climate and plant phenology. Therefore, the differences in DE between and within plant species can be very large. Because of the difficulty of measuring energy loss in urine, methane and heat increment in grazing animals the energy nutrition of range animals is usually described in terms of DE.

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Figure 15. Partition of feed energy within ruminant animals. The most important animal related factors limiting energy intake and utilization in rangeland animals are: a) digestibility of the herbage

Apparent Digestibility = Matter in Feed - Matter in Feces x 100 % Matter in Feed True Digestibility = Matter in Feed-Matter in Feces-Endogenous Matter x 100% Matter in Feed The greater the digestibility of the diet, the greater the amount of energy captured from the diet. b) rate of passage of the digesta through the intestine and particularly the rumen The faster the rate of passage, the faster the clearance of material from the rumen and hence the greater the ability of the animal to consume more feed. Some animals have adapted to low digestibility diets by increasing the rate of passage. For example, the digestibility coefficient of a herbage is greater when fed to cattle than deer but the rate of passage through deer is faster. This means that the rumen of deer is cleared more rapidly than cattle allowing more space for intake of additional feed. Because the coefficient of digestibility diminishes with increasing time of residency in the rumen, this means that the energy intake is usually greater for deer than cattle when consuming the same feed. Also, plants differ in their rate of passage through an animal For example, legumes have a faster rate of passage than grasses of the same digestibility.

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c) metabolites of digestion Nutrient energy in ruminants comes largely from the carbohydrates in the feed. Carbohydrates make-up the biggest proportion of substances present in plant matter. The carbohydrates are fermented by the rumen microflora and microfauna and the end products of this fermentation are the volatile fatty acids (VFA). The most important VFA's are acetic, butyric and propionic acids. The higher the fermentation to propionic acid the more efficient is the transfer of energy from plants (referred to as net primary product or NPP) to the animal (referred to as net secondary product or NSP) as shown below: Propionic acid TCA (2 mols of 3 C + 2H+)
109%

Forage

Cellulose
62%

Glucose (1 mol of 6 C) Grain Starch

Pyruvic acid (2 mols of 3 C)

Acetic acid (2 mols of 2 C) O2 78%

TCA

CO2 + CH4

Butyric acid (1 mol of 4 C)

Fat The efficiency of conversion of Pyruvic acid (CH3 - C0 - C00H) to: Acetic acid (CH3 - C00H) is 62% because CO2 and CH4 are lost Propionic acid (CH3 - CH2 - C00H) is 109% because 2 atoms of H are added Butyric acid (CH3 - CH2 - CH2 - C00H) is 48% because additional O2 is lost. The amounts and proportions of the volatile acids depend on the substrate (feed) and the varying populations of rumen micro-flora and fauna fermenting the feed. High cellulose diets, such as those consumed by range animals, lead to high proportions of acetic acid in the fermentation products. Thus the transfer of energy from plants to animals is inefficient at this point (Table 26). Table 26. Typical fermentation patterns of hay, grain and range herbage diets. Acetic (%) Good quality hay Grain Range Young green Mature dry 60 52 65-70 75-80 Butyric (%) 9 8 10 10 Propionic (%) 25 38 20-25 10-15

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Energy nutrition of ruminants is also greatly influenced by climate (see the section on winter feeding) and other environmental conditions such as the terrain of the range or pasture. For example, topography will influence the energy cost associated with walking. Table 27 shows the additional energy cost for an animal to walk in a vertical compared with a horizontal plane. Table 27. The energy cost of an animal to walk 1m in a horizontal and vertical plane. ___________________________________________________ Plane Energy cost (J/Wkg/m) ___________________________________________________ Horizontal 2 Vertical 26 ___________________________________________________ The energy costs or walking, grazing and ruminating are considerably higher for cattle on range or pasture compared with cattle in pens (Table 28). Table 28. % Increase in metabolic rate above resting for cattle in pens and on range while performing certain functions. ______________________________________________________ Function Pens Range ______________________________________________________ Walking 6% 45% Grazing/eating 12% 30% Ruminating 4% 9% ______________________________________________________ These increases are associated with the greater distance required for cattle to walk to obtain their feed and water on range compared with cattle in pens; the longer time required for prehending their feed; and the longer time spent ruminating because of the more fibrous nature of range forage compared with hay, silage and grains fed to penned cattle. These, and other factors, result in cattle on open pasture and rangeland having maintenance energy requirements which are almost twice those of cattle in pens (Table 29). Table 29. The maintenance energy requirements of cattle at pasture and in pens. ________________________________________________________ Maintenance Energy Requirements (KJ/W0.75/day) ________________________________________________________ Pens 480 Open pasture 740-830 Rangeland 740-990 ________________________________________________________ Protein Figure 16 provides a diagram of the pathways between dietary N intake and protein synthesis in the ruminant animal. Note this is somewhat more complicated than Figure 15 because, unlike energy which passes through the system only once, nitrogen re-cycles within the ruminant just as it recycles in soil-plant systems.

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Note that ruminants are unique in that they can utilize dietary non protein nitrogen (NPN) as a source of protein because of the ability of the rumen microbial population to convert NPN into microbial protein which is digested in the small intestine and becomes a very important source of amino acids for the ruminant.

Figure 16. The pathways between dietary protein intake and protein metabolism in the ruminant. Also note that urea can be recycled back to the rumen via the saliva and ruminants can partially offset a dietary protein deficiency by conserving urea excretion in the urine and diverting it back to the rumen via the saliva. Urea has a very high concentration of nitrogen. Feed grade (0 = C
NH2 NH2

) urea is 45% N or 281% CP equivalents.

However, urea contains no precursors for the formation of a carbon skeleton for protein so high energy feeds must be fed simultaneously to maximize microbial protein production if the animal is to receive maximum benefit from the urea. The energy must be readily available such as molasses or
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processed grain since, once in the rumen, the urea is rapidly hydrolyzed to NH 3 and absorbed through the rumen wall. The conversion of urea to protein is outlined below.
Microbial urease

1.

Urea

C02 + NH3
Microbial enzymes

2.

CH0

VFA + keto acids


Microbial enzymes

3.

Keto acids + NH3


Microbial enzymes

Amino Acids

4.

Amino acids

microbial protein
Enzymes in abomasum& S.I.

5.

Microbial protein

Free amino acids

6.

Free amino acids are absorbed.

Note also that protein can be used as an energy source by the conversion of amino acids to glucose. Distinguishing Between Protein and Energy Deficiencies It is very difficult to distinguish between deficiencies of energy and protein under range conditions. This is because the two are so closely correlated. For example: 1) Low protein herbage also has a low digestibility. 2) Protein deficiency in herbage leads to a protein deficit in the rumen which reduces microbial activity and herbage digestibility is reduced. This in itself reduces the transfer of energy from the plant to the animal but it also reduces the rate of passage of the digesta and herbage intake is reduced so energy intake is also reduced. 3) Apart from the influence of protein intake on rumen function, the amount of protein reaching the small intestine also influences the amount of herbage consumed. This has been demonstrated by the infusion of casein solution into the duodenum via a duodenal cannula. Infused animals consumed more herbage than animals infused with the same volume of liquid as water. This lead to an understanding of the role of 'by-pass' or non-degradable rumen protein in ruminant feeds. It is often claimed that energy is the first limiting factor to animal productivity on range. This has been based on studies which have indicated that the amount of nitrogen present as NH 3 in the rumens of animals on range was always in excess of the amount of N required by the rumen microorganisms to sustain the level of fermentation (i.e. VFA production) taking place. Thus it has been assumed that herbage intake, and subsequent productivity has been limited by the digestibility of the
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herbage consumed or the ability of the microorganisms to digest the herbage and liberate its energy to the ruminant animal in the form of VFA. This is sometimes supported by the general lack of response to urea supplements given to sheep and cattle on range compared with the positive response from urea supplements which occurs when sheep and cattle are fed high energy diets, for example in a North American feedlot. While this is true at the rumen level it is not the whole story. This was first demonstrated in a series of experiments with cattle on winter range in Australia. Cattle were given supplements of (1) sorghum grain (high energy/low protein) and (2) by-pass protein (protein which was not degraded in the rumen but which was digested and absorbed in the small intestine). The supplements were given in factorial combinations and the range herbage had an organic matter digestibility of 40% and crude protein content of 3.6%. The protein supplement was a mixture of 70% cottonseed meal, 20% meatmeal and 10% fishmeal. The results of those experiments are summarized in Figure 17.

Figure 17. The effect of feeding an energy (sorghum) or protein (cottonseed, fish and meat-meals) on herbage intake of grazing beef cattle. For every unit of ME provided in the sorghum supplement there was a reduction in ME intake from the herbage (i.e. the sorghum energy became a substitute for herbage energy). The corresponding increase in liveweight gain was small, indicating that the energy limitations of the range herbage were only small. On the other hand, there was a substantial increase in ME intake from the herbage when the protein supplement was eaten and a correspondingly large increase in liveweight gain indicating a major influence of protein deficiency on productivity of cattle on winter range. The same limitations no doubt apply to wildlife. The relationship between herbage organic matter intake (HOMI) and intakes of sorghum (S) and the protein supplement (P) is shown in the following equation.

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HOMI = 2.96 - 0.76 S + 2.33 P = 1.45 P2 (R = 0.79; P < 0.05). Production data are shown in Tables 29, 30 AND 31. Table 29. The effect of protein and energy supplements on weight gains of steers grazing winter range from 6 to 15 months of age. _________________________________________________________ ADG (kg/day) No supplement 560 g sorghum/day 600 g by-pass protein/day -0.04 +0.10 +0.35

Table 30. The effects of a protein supplement on weight gain and pregnancy rate of heifers grazing winter range from 0 to 15 months of age. ______________________________________________________ ADG Pregnancy (kg/day) % No supplement 800 g by-pass protein/day -0.08 +0.5 0 92

Table 31. The effects of protein supplements on herbage intake and liveweight change of cows 45-90 days post partum and on milk yield measured on day 71. (Calves were creep fed during the experiment so that production data are not applicable directly to cow supplementation and are not included). ________________________________________________________________ Amount of Protein Supplement (g/kg W0.75 0 5.3 10.6 15.9 21.2 Herbage Intake (kg DOM/day) Liveweight Change (kg/day) Milk Yield Day 71 (kg)

4.41 -2.40 2.3

5.41 -2.37 2.8

6.63 -0.99 3.6

7.97 -0.06 4.4

7.44 +0.08 4.5

The calves were creep fed so their weight changes relative to cow milk yield were not reported.

Similar data have more recently been reported from USA and Canada (Tables 32 and 33)

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Table 32. The effect of cottonseed cake supplement on intake and weight change of steers grazing blue grama grass in New Mexico from January to May. Control Intake (g/kg BW) Weight Change (kg/d) 10.8 -0.03 Cottonseed Cake 12.9 0.24

Table 33. The effect of canola meal supplements on cow weight changes when grazing rough fescue pasture in Alberta from December 1993 to January 1994. Canola Supplement (kg/d) 0 0.4 0.8 1.2 Weight Change (kg) -35.1 -46.1 -25.1 -19.5 Back Fat (mm) -1.05 -0.94 -0.56 -0.44

Relationship Between the Quantity of Intake and the Quality of the Herbage Consumed. Because of the difficulty of measuring how much a grazing animal eats and the quality of the herbage which it consumes while on range there are few relationships reported in the literature showing the effect that herbage quality has on the quantity of herbage consumed. One report for cattle is cited as an example: OMI/W0.75 = 0.107* DIG + 0.030 * N (R2 = 0.84; RSD = 0.022; P<0.001) where OMI is organic matter intake (kg/day) W is the liveweight of the animal (kg) DIG is herbage digestibility expressed as a decimal unit and N is the herbage nitrogen content (%).

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PASTURE MANAGEMENT TO ENHANCE THE NUTRITION OF CATTLE "The range resource is by far the most variable commodity that is encountered in livestock nutrition. It can vary in nutrient quality from a low comparable to sawdust to a high comparable to top quality alfalfa. It is capable of producing gains approaching 1.4 kg/d in growing classes of livestock at certain times of the year or of allowing starvation in the same cattle at other times." This quote is taken from Raleigh and Lesperance (1972) in Church et al. Practical Nutrition. It is also very important to remember that the beef cow has her greatest nutrient requirements when she is at pasture because this is when she is lactating and re-breeding (Figure 2). Also, the calf has increasing nutrient requirements that must be met from the pasture in order to maintain good growth rates. So, arguably, the grazing season is the most important time in a beef operation. However, unfortunately it is often the most neglected part of a beef operation. Factors affecting nutrient intake and productivity of grazing animals The factors influencing nutrient intake and productivity of cattle on range or pasture are summarized in Figure 17.

Chemical composition Nutritional Value Digestibility Type and proportions of endproducts of digestion Animal Productivity Nutrient Intake Forage yield and physical availability Physiological state of the animal Forage Intake Environmental factors such as temperature,humidity, topography Plant acceptability Digestibility and rate of passage Pasture structure Figure 17. Factors affecting the intake and productivity of cattle on range or pasture. As the grazing season advances plants mature with a subsequent decline in their nutritive value and the productivity (liveweight gain) of herbivores grazing them Figure 18).

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Figure 18. Effect of plant phenology on forage quality Figure 19 shows schematically how the digestible protein and energy contents of range and pasture declines with advancing maturity and the consequent decline in liveweight gains of yearling cattle and calves. Getting the most out of the grazing season The phenological cycle of growth is not the same for all species of plants. Some plants start growth early in the spring (cool season plants) and some do not commence growth until late in the spring or even early summer (warm season plants). The commencement of growth is temperature controlled, but it can be ameliorated by soil moisture and fertility. Similarly, not all plants enter the reproductive phase at the same time or senesce (death of the above ground parts leaves and stems) at the same time. Some plants have a short growing season while some plants have a long growing season. For example, crested wheatgrass starts growth early but has a short growing season and therefore matures early and loses quality rapidly so that its nutritional value is poor after mid July. In areas where late summer rainfall is reliable, crested wheatgrass will provide valuable forage from re-growth. However, in areas where summer is not reliable the reliability of re-growth is reduced and cannot be counted on providing useful grazing beyond mid-July. In contrast, Russian wild ryegrass, commences growth almost as early as crested wheatgrass but more slowly. However, it has a much longer growing season and holds its nutritional quality much longer than crested wheatgrass. Therefore, it is a good grass for late season grazing. This means that these two grasses complement each other and can be used in a complementary grazing system in which a field of crested wheatgrass can be used for early season grazing and a field of Russian wild

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Figure 19. The amount of digestible nitrogen and energy require for maintenance and growth of yearling steers (A,B,C) and cows & calves (D,E,F) on range.

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ryegrass can be used for late season grazing. Complementary grazing and/or various forms of rotation grazing such as those described in the Managing Saskatchewan Rangelands book (http://www.agr.gov.sk.ca/saf/default.htm) can be used to both lengthen the grazing season and to increase the quality of the forage available to the grazing animal, thus increasing animal production. Proper rotation grazing can achieve two important grazing management objectives. It can increase the period of vegetative growth by forcing the plant to produce more vegetative tillers and this in turn increases the quality of the herbage since new vegetative growth is always of greater nutritional quality than mature growth. The extent to which this can be used depends, however, on the availability of moisture. Without moisture plants do not grow. Therefore, the practice of rotation grazing usually is more beneficial in the more mesic (wetter) regions compared with more arid (dry) regions. Examples of the different growth curves of different species are given in Figure 20

Figure 20. Relative yield and period of growth of native grass and seeded pastures. Examples of how the information in Figure 20 can be used to design various complementary grazing systems for the different soil zones of Saskatchewan are given in Figure 21.

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Figure 21. Grazing systems for Saskatchewan soil zones.

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PRINCIPLES OF RANGE MANAGEMENT Basic terms in Range Science 1) 2) Climax communities. These are plant communities in equilibrium with their environment. Succession. There are four types of succession in range ecosystems:

a) Primary succession. This refers to the establishment of plants on land not previously vegetated. For example, on the edge of a gradually filling slough or a sand bar in a river. b) Secondary succession. This refers to the invasion of land that has been previously vegetated. For example after fire, logging, cultivation. c) Progressive succession. This leads to communities with greater and greater complexity and biomass and to habitats that are progressively less severe and more stable. d) Retrogressive succession. This leads backwards toward communities with lower diversity and less moderate habitats that are less stable. Caused by drought, fire or grazing. Avoidance of this type of succession is a primary objective of range management. "By far the most important factor contributing to retrogression on range is improper grazing" (Stoddart, Smith & Box, 1980) and this is the major problem facing the world's rangelands today. Stages in Grazing Retrogression There are two major stages in retrogressive succession: 1) 2) Physiological disturbances of the climax plants. For example, the most preferred climax plants are grazed heavily, lose vigor and produce less. Composition changes in the climax cover. For example, reduced photosynthesis in the climax species as a result of overgrazing encourages competition from other less palatable plants that may eventually eliminate the preferred palatable plants.

Those plants most susceptible to grazing pressure are called DECREASERS. Some examples of range plants that are decreasers are winter fat, rough fescue, northern wheatgrass, green needlegrass. The less preferred or more resistant plants which are at a competitive advantage are called INCREASERS. Some examples of range plants that are increasers are the grasses - Blue grama, June grass, needle and thread; forbs such as pasture sage, club moss, yarrow; and shrubby plants such as cinquefoil, silver sagebrush, western snowberry and rose. Continued grazing pressure may cause reductions in the increasers and lead to the invasion of weedy species called INVADERS. Some examples of common invaders are dandelion, Canada thistle, kochia, lambs quarters and leafy spurge. These changes are illustrated in Figures 22 and 23.

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SUCCESSIONAL STAGE
CLIMAX COMMUNITY

% CLIMAX SPECIES

CONDITION CLASS
EXCELLENT

75-100% GOOD

51-75%

FAIR 26-50%

POOR 0-25% WEED STAGE

Figure 22. Approximate relationship between range condition and degree of retrogression from a climax ecosystem.

Figure 23. The effect of grazing intensity on range condition and the relative proportions of decreasers, increasers and invaders.
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Several things can be done to reduce the extent of retrogressive succession on rangelands. 1. Practice Good Range and Pasture Management 3) Don't graze too early. Defoliation during the early growth period of plants reduces leaf area index that reduces the conversion of solar energy to plant energy by photosynthetic tissue in the leaves. This reduces plant vigor and the ability of the preferred (grazed) species to compete with the non-preferred (ungrazed) species. It also reduces energy storage in the roots. Therefore, the grazed plants do not compete well with non-preferred species for light, moisture and nutrients in subsequent years with a resulting long-term change in range botanical composition and condition. Cattle in Saskatchewan should not graze native range until mid-June. Although many rangelands in Saskatchewan are grazed in mid-May, research at Swift Current has shown that their yield (and hence carrying capacity) can be increased by 50% or more if grazing is delayed until early to mid June (Table 34). Table 34. The production of a mixed grass prairie at Swift Current based on clipped plot yields. ______________________________________________________________________________ __ Date of First Summer Yield % Increase in Yield Clipping Kg/DM/ha Due to Delayed Clipping ______________________________________________________________________________ __ May 16 664 June 5 1067 60.7 June 20 1137 71.2 July 5 1218 83.4 ______________________________________________________________________________ Delaying grazing also reduces the risk of ingestion of toxic plants since many toxic plants grow early in spring and at that time may make up a high proportion of the available forage. 4) Don't graze too heavily. Approximately 40-45% of the total herbage yield should be left to carry over through winter for a number of reasons: 1 allows storage of energy reserves in roots 2) allows trapping of snow and moisture penetration in spring is increased 3) ensures adequate seed production 4) reduces water evaporation from the soil 5) reduces wind erosion 6) reduces soil compaction and erosion from rain 7) protects the crown of the plant from winter kill 8) increases soil temperature in the winter thus increasing plant survival 9) decreases summer soil temperature which increases plant growth 3) Use Appropriate Grazing Systems (i) (ii) Rotational Grazing Short Duration Grazing - High Intensity/Low Frequency
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5) Deferred Grazing 6) Complementary Grazing These grazing systems 1) allow recovery of plant species which are susceptible to grazing 2) allow established plants to gain in vigor which increases survival 3) allow cattle to obtain more palatable herbage 4) allow more uniform utilization of the sward 5) range improvement techniques are easier to apply because of smaller areas 6) animal distribution is easier to control because of smaller areas 7) breeding efficiency increases due to smaller areas 8) possible internal parasite control. However, 9) must have sufficient water for each field 10) fencing costs are high - can reduce costs by using solar generated electric fences 11) more frequent handling of livestock 12) fire hazard of accumulated forage ?? 13) increased requirement for breeding bulls ?? 3. Range Improvement Seeded range and the use of several species of grass to complement native range can extend the grazing season and the nutritive quality of the herbage during the grazing season. Fire can be used to control undesirable and woody species. Although the use of fertilizer on native range is usually not economical, it will increase herbage yield and quality that result in increased carrying capacity and animal production. Fertilizer is more likely to be an economic proposition if applied to seeded pasture than to rangeland because seeded species have a greater potential to respond to increased fertility. Control of toxic plants and weedy invader species. 4. Livestock Distribution Proper distribution of stock-watering sites, salt, and shelter ensures better use of pastures. Grasslands more that 1km (3/4 mile) from watering sites will be under-grazed. Salt blocks, mineral feeders and oilers should be located away from water and occasionally relocated to attract livestock to under-grazed areas. Range riders can be used to ensure good animal distribution. Cross fencing, though expensive, will frequently provide economic returns affording better grazing control and carrying capacities. 5. Provision of Supplements Energy supplements such as hay, grain, range pellets, range cubes, etc. provide a substitute source of energy for grazing animals and as a result they will reduce their grazing activity and intake of range herbage. Thus if range or pasture is being overgrazed, cattle should be removed or if this is not possible, provision of an energy supplement will relieve the grazing pressure on the range. Protein supplements, particularly those with a high rumen by-pass value, on the other hand, stimulate herbage intake so should only be provided in situations where the range herbage has matured or senesced and is in abundant supply. Mineral supplements should be available at all times. These typically should contain calcium, phosphorus and trace minerals (Cu, Se, Zn, Mn) depending upon local soil types and deficiencies.
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Cobalt iodized (blue) salt should be available at all times and consumption calculated on the basis of 1.5 kg/cow/month in spring reducing to about 0.125 kg/cow/month in July-August. Creep Feeding Energy supplementation of calves on range and seeded pasture to alleviate their increasing inability to meet the calf's energy requirement for satisfactory growth (i.e. about 1kg/day). The creep consists usually of ground good quality hay and grain or straight grain, preferably oats, as oats provide fewer digestive upsets. Consumption will increase from about 0.25 kg/day at 1-2 months of age to 1.5 kg/day at 7 months of age. More creep will be required where the forage supply is severely limited. STOCKING RATE, GRAZING PRESSURE AND CARRYING CAPACITY The following definitions are taken from Terminology for Grazing Lands and Grazing Animals prepared by the Forage and Grazing Terminology Committee and officially adopted by the Range Management Society. A thorough understanding of the terminology is essential to the study of range management. Although officially adopted in 1991, there is still considerable confusion among range management personnel and who have adopted several variations of these definitions with detrimental consequences to range management. Stocking rate is the number of animals per unit area of land for a stated time. Stocking density is the number of animals on a specified unit of land at a particular point in time. Grazing Pressure or Grazing Intensity refers to the number of animal units or forage intake units per unit of available forage dry matter at any one point in time. An animal unit (AU) is one non-lactating bovine weighing 500 kg and fed at maintenance level. It is expressed as weight0.75 in other kinds of animals. A forage intake unit is an animal unit with a rate of forage consumption equal to 8 kg dry matter / day. This is based on the assumption that the maintenance requirement of a mature non-lactating bovine weighing 500 kg is 8 kg of forage dry matter / day whether the forage has a digestibility of 80% or 40% which is a false assumption. An animal unit month (AUM) is the amount of forage consumed by one animal unit for one month. Forage allowance is the amount of forage dry matter available per animal unit or per forage intake unit at one point in time. The Carrying capacity of a pasture is the maximum stocking rate that will achieve a target level of performance, in a specified grazing system, that can be applied over a defined time period without deterioration of the ecosystem. The carrying capacity of a pasture is not static from season to season or year to year. Therefore, the average carrying capacity refers to the long-term carrying capacity averaged over a number of years. The term Animal Unit Month is commonly used in Saskatchewan but its use is not without problems. Firstly it is not always clear what definition of animal unit is used and whether all parties are using the same definition. The old definition of an animal unit was a 1000lb (450kg) cow with calf at foot and that of a forage intake unit was 30 lb/day (13.5 kg). If this definition is still being used it is not appropriate since most cows in Saskatchewan now weigh at least 550kg with many
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weighing 600-700kg. The forage intake unit for a lactating cow weighing 550kg using the current definition would be about 13.5kg DM when allowance is made lactation for the additional weight of a mature cow. This is close to the old value of 13.7kg. However, it is not clear if allowance is made for pasture intake of the calf, which could average at least 3kg/day over the grazing season. Nor is it clear if allowance is made for the removal of only 55% of the available herbage that is recommended in order to sustain range condition and productivity. Current publications in Saskatchewan still define a yearling steer as 0.7 AU which suggests that the old definitions are still being used, at least by some agencies, since, in Saskatchewan, a backgrounded yearling steer going on to pasture in spring would probably weigh 300-350kg (700-750lb). That steer could be expected to gain 70-120kg (150-250lb) while on pasture. This means it would be equivalent to 0.7-0.95 AU. Effects of Stocking Rate on Pasture Pasture production is increased by defoliation. Defoliation stimulates the production of new growth (tillers). The rangelands of the Northern Great Plains and Prairies evolved under grazing. The macro-grazer during this evolution was the Bison. The macro-grazer of today is the beef cow. The deterioration of rangelands during the last 100 years is not the fault of the cow per se but the way in which we have allowed the cows to graze. Therefore, the lobby to remove cattle form western rangelands in the US is mis-guided since this will not result in the return of the original climax community but a different climax community. The encroachment of Aspen trees into the parkland regions of the prairies is a living example of progression towards a different climax community as a result of changing land use following settlement. Before settlement, the First Nations people burnt these areas to maintain them as grasslands in order to attract the Bison. Since settlement, these areas have not been burnt which has resulted in the encroachment of aspens on the unploughed areas of the parkland regions. However, pasture growth rate is reduced at high stocking rates. The reduction in leaf area following intense grazing pressure results in reduced interception of solar radiation by the leaf and hence lower photosynthetic activity in the plant. The carbohydrate reserves in the roots and crowns of heavily grazed plants are therefore reduced. Pasture growth at high stocking rates may also be reduced by exhaustion of soil moisture because of greater evaporative loss directly from the soil, if grazing increases the amount of bare ground, or through the plants, if the stimulation of new growth increases the amount of evapo-transpiration through the plant. However, pasture growth may also be reduced at low stocking rates because of increased senescence and a greater proportion of matured leaves that are photosynthetically less efficient. It, therefore, is very important to define these extremes if the most efficient use of the pasture is to be made. High stocking rates frequently increase the proportion of less palatable species (increasers). Also, reduction in the proportion of palatable legumes (decreasers) in a seeded pasture will not only reduce the overall palatability of the pasture but it will also reduce nitrogen fixation and hence overall pasture productivity. Excessive grazing creates an unstable pasture community and a depleted ecosystem. Stocking Rate and Soil Properties High stocking rates increase the bulk density of the soil and decrease pore space (Tables 35 and 36). This reduces the rate of water infiltration and increases the amount of run-off water thereby increasing the risk of erosion. This compaction occurs because of three things: i) the direct effect of treading or trampling by the animals which increases at high stocking rates. ii) a decline in root mass in the soil following the weakening of plants with high stocking rates and iii) reduced protection of the soil from the erosive action of wind and rain.
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Table 35. The effect of static loads exerted by vehicles and animals when stationary on range. LOAD kg/cm2 Crawler tractor Sheep Wheel tractor Horse and cow Truck 0.32-0.63 0.65 1.4-2.1 1.7 3.5-7.0

Table 36. The effect of degree of range use (stocking rate) on soil pore space and moisture infiltration rate. Degree of Range Use Moderately grazed Over grazed Depleted Soil Pore Space (%) 68.1 59.1 46.5 Moisture Infiltration Rate (cm/hour) 10.5 5.5 2.1

Stocking Rate and Animal Nutrition The direct effect of high stocking rates is to reduce the amount of herbage available to each animal on the pasture and so to restrict individual animal intake. This will result in a decrease in animal productivity. This is especially important at those times of the year when herbage is in short supply and/or when the physiological demands of the animals are high, such as in late pregnancy and early lactation. Increasing stocking rate to increase production per unit area of land may therefore require some adjustment of calving dates to match nutritive demands with feed supply. Stocking rate, if it affects botanical composition, which it invariably will, may also affect dietary digestibility and protein content and so have a further indirect effect on feed intake and utilization. Stocking Rate and Animal Production The effects of stocking rate on animal production are greatest at those times of the year when the feed supply is inadequate. In Canada this corresponds to late fall through early spring. High stocking rates result in reduced animal liveweight gains, particularly at these times of greatest feed shortage. This is particularly important at breeding since estrus and ovulation rates are closely associated with liveweight gain. As a result the number of offspring per 100 females is reduced at high stocking rates. Increases in stocking rate also cause reduced milk production with a resulting decrease in the growth rate of the offspring. Wool growth is also decreased with increasing stocking rate in sheep. In addition, at very high stocking rates the wool fibre may be weakened ("tender" wool) and more likely to break thus reducing the spinning quality of the wool. Finally, animal health may be adversely affected at high stocking rates. Increased mortality is likely at high stocking rates due to under-nutrition. Increased wear on the incisor teeth is common thus reducing the animals' foraging ability and shortening their life span. The increased wear on
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the teeth is caused by the shortness of the leaf length of overgrazed grass causing the animals to continually eat very close to the soil. There is also evidence that parasite burdens increase at high stocking rates and bacterial and viral diseases are more readily spread at high concentrations of animals. Quantifying Animal Production in Relation to Stocking Rate There is considerable debate among scientists over the relationship of stocking rate to animal production. It is unlikely that the shape of the curve will be the same for all pasture types and all animal production traits. There are three most likely shapes (Figure 24 a, b and c). ADG
(a)

ADG
(b)

ADG
(c)

Stocking Rate

Stocking Rate

Stocking Rate

Figure 24. The relationship between stocking rate and average daily gain (ADG) of beef cattle. In figure 24(a) the plant response to increasing stocking rate is very small at low levels of stocking rate. Thus the effects of stocking rate on herbage yield, availability, botanical composition and nutritional value are very small and barely detectable in terms of animal product. As the stocking rate increases, however, the effects on pasture botanical composition, yield, availability and nutritional value become increasingly important and more readily detectable in declining animal product. The rate of decline in animal production, therefore, increases at a more rapid rate for each incremental increase in stocking rate. In Figure 24(b) very low stocking rates have a detrimental effect on animal production. This may be caused by increased senescence and hence reduced nutritional value of the pasture at these low stocking rates. In theses cases a moderate increase in stocking rate will increase the rate of defoliation which will stimulate vegetative re-growth and tiller production. However, further more severe increases in stocking rate will cause more severe defoliation with resulting changes in botanical composition and productivity of the pasture which will ultimately reduce animal production. Another example is the development of wolf plants in bunch grass species such as crested wheatgrass, Russian wild ryegrass etc. at low stocking rates. At low stocking rates, cattle will not consume all plants in a pasture at the same rate. Thus some will be left ungrazed and will become coarse, senescent and unpatatable. Cattle will avoid these plants and prefer to return to previously grazed plants with new regrowth and tillers that are much more palatable and nutritious. These plants become overgrazed while the ungrazed plants become more coarse and less and less palatable. The leaves and stems of these unpalatable plants die but do not decay during winter. New growth the following spring commences from the crown and is protected by all the remaining dead, coarse unpalatable material. As a result, the new season growth remains untouched while the new season growth on those plants grazed the previous season becomes further overgrazed. Eventually, these ungrazed plants become large and rank and are termed wolf plants while the rest of the pasture becomes increasingly overgrazed.

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However, most experimental results indicate that the stocking rate curve is linear (24c). For example, Jones and Sandland (J. Agric. Sci. Camb. (1974) 83:335) examined data from a large number of stocking rate experiments and concluded that in all cases the relationship did not depart significantly from the linear model Y = a - bx where Y = animal production and x = stocking rate and a,b are the regression coefficients of intercept and slope. They therefore concluded that the relationship between animal production and stocking rate could be determined from just two stocking rates spaced anywhere along the line. It seems unlikely that this is true at the extremes of stocking rates and experimental linearity is most likely due to the choice of stocking rates by the researchers within the linear part of figures 22(a) and 22 (b). This choice has probably been a conscious one by researchers because: i) this part of the curve is of more interest and value in terms of predicting productive efficiency ii) very low stocking rates are expensive and impractical because they require a large area of land iii) very high stocking rates are also expensive because animal productivity is very low and may even result in animal mortality (death). Knowledge of the value of the regression coefficients a and b can provide the pasture manager or researcher with a considerable amount of information. For example, the intercept coefficient "a" provides a dimensionless estimate of the quality of the grassland herbage. This is because the line intercepts the Y axis (animal production) when x = 0 or at zero stocking rate. At zero, or more correctly in mathematical terms, infinitely low stocking rate, there is an infinite amount of herbage available per animal so animal production is not restricted by herbage availability. If herbage availability is not restricted then if animal production is less than the maximum genetic potential for the animal, production in a normal healthy animal must be restricted by the quality of herbage on offer. Similarly, the regression coefficient for slope, "b", provides a dimensionless estimate of the yield or growth response of the pasture to increasing stocking rate or the effect of stocking rate on the quantity of herbage available to the animal. For example, a high value for the "b" coefficient indicates a rapid and severe degeneration of the range or grassland with increasing stocking rate. A low value for "b" indicates only a small effect of stocking rate on the grassland ecology. Consider, for example, Figure 25 (a) and (b) showing the response in average daily gain (kg) of cattle grazing two grasslands A and B. In Figure 25(a), the slope of the regression of ADG (Y) on stocking rate (x) is the same for both pastures that is bA = b B). Thus both grasslands are showing the same response to increasing stocking rate. However, the intercept aA is much greater than the intercept aB indicating that at all levels of stocking indicating that the nutritional value of grassland A is greater than that of grassland B. In Figure 25(b) however, it can be seen that at an infinitely low stocking, or in the climax state, grassland C has a greater nutritional value than grassland D but it is more sensitive to increases in stocking rate than is grassland D.

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ADG (Y)

ADG (Y)

(a) aC aA bA A bB B Stocking Rate (x) bC aD

(b)

bD D

aB

C Stocking Rate (x)

Figure 25. Examples of possible responses of different pastures to increasing stocking rate. Furthermore, if the effect of stocking rate (x) an individual animal production (Y) is linear Y = a - bx then multiplying both sides of the equation by x gives Yx = ax = bx2 where Yx now = animal production per unit area of land. This relationship is now shown in Figure 26.

Figure 26. The relationship between stocking rate and average daily gain / animal and / ha.
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It can be shown mathematically that: 3) Maximum animal production per unit area of land (Ymax) occurs at a stocking rate equal to a/2b. This is defined as the optimum stocking rate since higher or lower stocking rates will result in decreased animal product per unit area of land. ii) Optimum stocking rate occurs when animal production per head is half that which is possible at zero, or infinitely low, stocking rate (a/2) iii) At twice the optimum stocking rate a/b, animal production per unit area of land becomes zero. This method of calculating optimum stocking rate is known as the half intercept method. However, it should be noted that it is only appropriate if the grasslands reach equilibrium at each of the stocking rates applied. That this may take several years. Further, the production per animal at Ymax (a/2) must be compatible with production goals. For example, stocking 100 steers on 100 ha may allow for the sustained expression of Ymax or maximum beef production per unit area of land but this may be achieved at individual animal gains which are less than the goals of the pasture manager. Therefore to meet these goals, the stocking rate x must be reduced. In addition, there are confounding factors when a multi-specific pasture is being considered because the optimum time to commence grazing will vary for each species of plant. Therefore, the pasture should be managed for the key species, which may or may not be the most susceptible decreaser species. Thus grazing multi-specific pastures will always involve an element of compromise between maintaining good range condition and maximum animal production. PALATIBILITY, PREFERENCE AND SELECTION Palatability and preference are plant/animal interrelated factors. For example, preferred plants are obviously palatable. In most cases preference refers to animal reactions to a plant and palatability refers to the plant characteristics that promote that reaction. Selection expresses the degree to which animals harvest plants or plant parts differently from random removal. The selectivity ratio expresses the dietary composition in proportion to range composition. The higher the selectivity ratio the greater the selection or preference (Figure 27). The answers to the following questions are important to the understanding of grazing animal behavior, nutrition, production and the effects of the grazing animal on range and pasture condition: Why does a single species of animal have a preference for particular plants or particular parts of plants? Why do different species of animals have different preferences? (Table 37) The answers are complex and by no means complete. While there is a great deal of information on what herbivores eat there is very little information on why they eat it and because of the complexity of the subject only a brief summary can be presented here. Some very broad generalizations are possible: 1) 2) 3) 4) Green plant material is preferred to dead plant material Grass and legume leaves are preferred to stems Different species of animals select different diets (Table 37) Within any one species of animal the diet selected varies with the season (Table 38).

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Selection SHEEP CATTLE

Legumes

2.2 2.0 1.8 Legumes

Forb Leaves Leaves

1.6 1.4 1.2 1.0 Grasses Leaves

Grasses

0.8 0.6 0.4

Forb Leaves

Figure 27. Selectivity ratios exhibited by sheep and cattle in California. Plants or plant parts at the top of the scale were highly preferred while those at the bottom of the scale were poorly preferred.

Table 37. The selectivity ratio for six different plant species grazed by sheep and cattle sharing the same rangeland in California, USA, during summer. _____________________________________________________ Plant Species Selectivity Ratio Sheep Cattle _____________________________________________________ 10.0 3.1 1.8 0.9 0.4 2.6 2.2 1.9 1.0 0.6

Phalaris aquatica Stipa pulchra Trifolium spp Bromus spp. (annual) Avena barbata

Aira caryophyllea 0.3 0.5 _____________________________________________________


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It will be noticed from Table 37 that sheep are more highly selective than cattle. The gradient from the most preferred species (Phalaris aquatica) to the least preferred species (Aira caryophyllea) is much wider for sheep than for cattle but the order of preference is the same. It is also apparent that sheep exert a much higher degree of grazing pressure on Phalaris aquatica than do cattle. This plant is well adapted to grazing but, if it were not, it is obvious that sheep would deplete the pasture of this species at a much faster rate than cattle. Table 38. The selectivity ratios for grasses, forbs and browse of goats grazing rangeland in Texas during the four seasons. ___________________________________________________________ Season Selectivity Ratio Grasses Forbs ___________________________________________________________ Winter Spring Summer 0.7 0.7 1.1 --8.3 8.0 Browse 1.6 1.0 0.7 0.8

Fall 1.0 4.0 ___________________________________________________________

It can be seen from Table 38 that goats selected grasses in close equilibrium (balance) with their presence on the range in all seasons. The selection of forbs was very high during spring, summer, and fall and non existent in winter, probably because after such high selection pressure very few forbs remained through winter. The preference of goats for forbs has lead to the practice of using goats to "clean up" weedy pastures. During winter the goats turned more selection pressure onto browse. There are three types of ungulate herbivores: 1. Bulk/roughage feeders such as cattle and bison. These animals graze comparatively indiscriminately, use the tongue to gather the food into the mouth followed by a short jerking motion of the head during the biting process. They take comparatively large quantities of herbage with each prehensile motion. They have a large rumen volume in proportion to their body weight and a long retention time of ingesta in the rumen and as a result intake is low in relation to their body weight. A relatively large proportion of the rumen biota are protozoa. 2. Concentrate feedrers such as deer, moose and horses. These animals characteristically have soft pliable lips and in the case of horses, upper and lower incisors. They are therefore able to be highly selective with a comparatively small bite size. They are able to eat very discretely selecting single leaves or even parts of leaves. In contrast to bulk/roughage feeders, concentrate feeders have a comparatively small rumen volume in relation to body weight, rumen retention time is low and as a result intake is high in relation to their body weight. Only a small proportion of the rumen biota is protozoa.

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3. Intermediate feeders such as goats, elk and carribou. The diets of these animals are characterized by variety and frequent compositional changes but they are also able to exert a high degree of selectivity. Sheep have many characteristics of both intermediate and bulk/roughage feeders. Other characteristic are intermediate to bulk/roughage and concentrate feeders. Palatability a. Physical Factors (i) Presence of awns, spines, hairs, stickiness and coarseness of texture all influence palatability. (ii) Growth stage. As herbaceous plants mature they generally decrease in palatability and nutritive value. The whole plant becomes higher in fibre content and the leaf : stem : fruit ratios change towards a higher proportion of stem. Succulence decreases and coarseness increases due to the position and extent of lignification. (Deer will eat twigs in spring and summer when they are succulent but avoid them in winter when they are mature). In rare instances, the palatability of some species, for example Medicago hispida (Burclover) increases as maturity progresses. However, this may be associated with a reduction in alkaloid content as the plant matures (see section b.(v). (iii) Environment - Climate, topography and soil all affect palatability of plants and animal preferences for plants. For example, a simgle plant species on different sites will vary in chemical composition, succulence, proportion of leaf (or leaf:stem ratio) and coarseness of the foliage. Different animals prefer different sites and site affects their selection of food (see later section on animal behavior). Moisture from rain or dew may increase the palatability of plants. (b) Chemical Factors

(i) Sugars - the literature is confusing on the influence of sugars on palatability. For every report of a positive correlation between total sugar content (or soluble carbohydrate content) and preference there is a report that they were not correlated. One of the problems in sorting this relationship out is that the methods of extraction between various studies has differed and there is usually some doubt as to what carbohydrates were actually being determined because of the complexity of plant carbohydrate chemistry. However there are several reports that when unacceptable herbage was sprayed with sucrose or molasses it was eaten and this was the basis of the early development of molasses/urea feeding or spraying dry standing pasture and crop stubble with urea/molasses to increase its value as a feed. (ii) Organic acids - Citric malic, malonic, succinic, quinic and shikimic acids have shown a close positive correlation to preference in sheep. (iii) Tannins - these substances have been negatively correlated with preference. (iv) Coumarins - To humans these have a sweet smell and bitter taste. The smell of coumarin is objectionable to sheep and some plants (eg. Melilotus spp such as sweet clover) contain high concentrations of coumarins and these may be the reason for poor acceptability of these plants by sheep. However, sweet clover is very palatable to cattle. v) Alkaloids - A large number of plants contain alkaloids in varying amounts (as much as 2.5% of the dry matter of some plants) and these substances reduce the palatability of plants. In most
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cases these alkaloids also have a physiological effect on ruminants either by impairing digestion or metabolism and this may be a case of being physiologically wise and, by experience, avoiding plants which cause discomfort. Alkaloids also have a bitter taste. For example, there are both sweet (alkaloid-free) and bitter (alkaloid-containing) strains of some lupins. Sheep avoid grazing the varieties containing alkaloids but readily graze the alkaloid free varieties. Fortunately the presence or absence of alkaloids in lupins appears to be controlled by a single gene which suggests that alkaloid may be the only factor influencing acceptability and it can easily be removed by breeding. Phalaris arundinacea (Reed canary grass) is another plant species that contains alkaloids and its palatability is negatively related to its alkaloid concentration. Alkaloid free varieties of this plant have also been bred. (vi) Fats - Work in South Africa has shown that sheep preferences for some shrub species was negatively correlated with the ether extract content, but they had no idea what fats or oils were involved. No doubt the above list is far from complete. The study of palatability, selection and preference crosses the boundaries of animal physiology and animal psychology and very little research has been done in this area. The physical aspects influencing diet selection are more easily studied and probably for that reason, better documented. The next section will discuss these physical aspects which include the influence of the availability of plants and their canopy structure on diet selection and animal productivity. Preference There are a number of factors influencing preference. These are: (a) The Herbivore's oral anatomy and prehensile technique (or method of gathering food into its mouth. For example, sheep may appear to prefer a particular species of plant to a greater extent than cattle but this may simply be that they are better able to graze it because of their smaller mouths and the difference between the two species of animals in prehensile technique. Sheep nibble, using their lips to a greater extent than cattle and use small jerking motions to prehend herbage. Cattle wrap their tongue around the herbage, taking much larger quantities of herbage into their mouth then use a slow conspicuous movement of the head to prehend the herbage. Because of these differences, sheep are more selective of their diet than cattle. (b) Plant availability. The availability of a plant species may be related to a number of factors. For example: (i) The accessibility of the plant. This may be dependent upon soil type, site, aspect, slope, drainage, and proximity to water or in countries where herding is used to control livestock such as China, proximity to the village. Thus some plants may appear to be of low preference simply because they are too far from the village or too far from water or too high up the mountain slope and so on. (ii) The amount of the preferred plant material present. A particular species of plant may appear not to be preferred simply because there is very little of this plant material present in the grassland. This will be dependent on the response of the plant to defoliation (ability to withstand grazing) and also on its ability to compete with other plant species for moisture, light, nutrients, and to withstand climatic stress of high and low temperature, drought and flood and so on.

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(iii) The structure of the plant canopy. For example, an experiment in Australia studied the reasons for a lower milk production when dairy cows grazed Setaria sphacelata compared with Digitaria decumbens. The two pastures were not different in the amount or nutritive value of the dry matter available to the cows. However, the cows grazing the Setaria sphacelata pasture took significantly more bites per kg of dry matter consumed and consumed significantly less dry matter in a day than did cows grazing the Digitaria decumbens. The cows grazing the Setaria sphacelata produced less milk because they consumed less forage and the researchers concluded that they ate less forage because the Setaria sphacelata was more difficult to prehend and the cows suffered from grazing fatigue. This phenomenon was attributed to the differences in canopy structure. Setaria sphacelata is a tall bunch type grass while Digitaria decumbens is aggressively stoloniferous, short and more dense, thus making it easier for the cows to prehend. Another example is that of a wolfy bunchgrass such as crested wheatgrass or Russian wild ryegrass pasture. For example it has been reported that as much as 60% of the total forage in a wolfy crested wheatgrass pasture on highway 19 south of Saskatoon was present as wolfy plants that were unacceptable to cattle. (c) The Role of Special Senses

The senses of sight, touch to the lips and/or tongue, taste and smell are all involved in diet preference and selection. Sight appears to be used primarily to orient animals to other animals and their environment. While animals do use sight to recognize conspicuous food plants this sense appears to be relatively unimportant in selecting a preferred diet. For example, one experiment compared the diets selected by blindfolded (sight impaired) and non-blindfolded sheep and it was found that there were no significant differences between the two groups of sheep with respect to the types or parts plants selected or the selectivity ratios (Table 39). Table 39. Effect of Sight on Herbage Selection by Sheep _________________________________________________________ % of Herbage Eaten +Sight _________________________________________________________ Phalaris tuberosa 65 Medicago sativa 95 Trifolium pratense 50 Festuca arundinacea 90 Eragrostis curvula 0 Dactylis glomerata 65 Stipa hyalina 55 Lolium perenne 45 _________________________________________________________ Pasture Species -Sight 55 85 60 85 10 75 55 45

The same researchers conducted a series of experiments in which sheep were treated surgically to produce single or multiple impairment of the senses of touch, taste and smell. They found that while all these three senses were used to determine preference, taste was the most important. For example, only in one of five cases did smell impaired sheep select a diet different from normal sheep while taste impaired sheep consistently selected a different diet from normal sheep and touch impaired sheep always selected a similar diet to normal sheep. In these experiments, 140 different
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plant species were available to the sheep so a large opportunity existed for the sheep to display changes of preference when sense impaired. Since taste appears to be the most important sense used in diet selection, it seems likely that a considerable amount of herbage that is prehended must be rejected and not consumed, since the material must first be taken into the mouth before being tasted. While this rejected material is returned to the pasture as plant litter and ultimately decomposed and re-cycled it may nevertheless represent a considerable amount of unnecessary defoliation of plants that are not consumed. (d) The Role of Physiological Status on Preference.

The animal's response to sense stimuli is moderated by its current nutritional status. For example, hungry animals usually have lower thresholds of rejection for taste. These thresholds (critical levels) are determined both by the number and kind of molecules transmitted from the plant to the animal and by the number and type of receptors in the animal. Thus, palatability is probably determined largely by the number and kind of molecules transmitted from the plant to the animal and preference by the number and type of receptors in the animal that can receive and disseminate information from these molecules from the plant. The number and kinds of sensory receptors differ between species of animals and this may partly account the differences between species of animals with respect to diet preference. There appear to be only small differences between lambs and older sheep or calves and cows with respect to preference. In addition, there is no evidence that pregnancy or lactation influence preference. (e) The Role of Previous Grazing Experience on Preference

Previous grazing experience, especially that learned when an animal is young, influences diet preference throughout the lifetime of the animal. However this can be modified by subsequent experience. However there are examples where preference is cannot be modified by subsequent experience in some herbivores such as with panda bears, koala bears and mountain gorillas. (f) The Role of Nutritional Wisdom on Preference

Does an animal prefer a particular plant because it is nutritionally good for it and avoid another plant because it is toxic? There is much debate over this question. This kind of nutritional wisdom is unlikely since death following ingestion of toxic plants is not uncommon among herbivores but some herbivores are undoubtedly wiser than others. However, most herbivores do not eat discrete meals of a single food but a meal can last for many hours, include a large number of foods that are regurgitated and chewed again. It therefore seems unlikely that animals can relate illness or benefit to a particular food component. It is more likely that "apparent" nutritional wisdom is related to palatability through the sense of taste. For example, alkaloids are bitter and unpalatable and for that reason are likely to be rejected unless the animal is hungry. Also, it has been suggested that phosphorus deficient sheep select a diet which is higher in phosphorus than that measured in cut herbage samples. However, it has also been shown that phosphorus adequate sheep select a diet which is higher in phosphorus than the cut herbage. It is likely that the sheep are not selecting for high phosphorus but for lower content of free phenols since phosphorus content and free phenol content is negatively related in plants and phenols such as tannins are negatively related to palatability. Nevertheless, it has been widely demonstrated that phosphorus deficient cattle will seek out and chew old bones (osteophagia); sodium deficient sheep and cattle will lick soil (geophagia); and coprophagia (eating of feces), which is commonly practiced in lagomorphs (rabbits), is known to be a means of compensating for diets deficient in protein &/or B vitamins. These examples of deviant feeding behavior are called pica. Factors affecting palatability are summarized in Table 40.
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Table 40. Factors that commonly improve or reduce palatability of plants to grazing herbivores. FACTORS WHICH IMPROVE PALATABILITY Plant physical factors of growth High succulence High leaf:stem ratio Seedstalks scarce New growth/regrowth abundant; long growing season Leaves fine, tender Twigs small, spaced Plant physical factors of morphology No thorns, awns spines etc. Leaves glabrous (smooth) Plant chemical factors Sugar, soluble carbohydrate, organic acid contents high Tannin, alkaloid contents low Environmental factors Weather promotes growth Previous growth normal Plant surface moist from rain, dew High sunlight except where maturation advanced Plant surface clean Plants not affected by disease, insects etc. Weather imposes dormancy Previous growth rapid & coarse Plant surface dry Low sunlight except where maturation retarded Plant surface covered with dust, mud, feces, urine Plants affected by disease, insects etc. Sugar, soluble carbohydrate, organic acid contents low Tannin, alkaloid contents high Thorns, awns, spines present Leaves pubescent (hairy) Low succulence, high dry matter Low leaf:stem ratio Seedstalks abundant Old growth abundant; plant dormant Leaves coarse, tough, rank Twigs large, compacted, sharp REDUCE PALATABILITY

Factors that influence diet selection in grazing herbivores are summarized in Figure 28.

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ANIMAL Motor outflow Reflexes of attention, approach, examination and consumption or rejection Senses of sight, smell, touch and taste PLANTS Plant species present and their chemical and physical characteristics and relative availability Modified by PLANT ENVIRONMENT Soil texture Soil fertility Plant community Modified by PHYSICAL ENVIRONMENT Topography (slope, aspect, site e.g. slough, rocky knoll etc.) Distance plant is from water Distance plant is from shade Modified by ANIMAL FACTORS Animal species Animal individuality Physiological status (food demand, disease etc.) Grazing behavior Social behavior Modified by PREVIOUS EXPERIENCE DIET COMPOSITION Figure 28. Schematic diagram of diet selection in the grazing herbivore.

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EFFECTS OF GRAZING ANIMALS ON RANGE AND SEEDED PASTURE LANDS Grazing animals have both physical and chemical effects on grassland. The interaction of the grazing animal on plant and soil ecology is summarized in Figure 29.

Figure 29. The soil-plant-animal ecosystem of range and pasture lands Physical effects Defoliation Effects of defoliation on plant morphology Defoliation of plants is not detrimental to plant survival but severe defoliation is detrimental. The severity of defoliation is defined by the frequency and timing of the defoliation and also by the amount of plant material removed. Correct defoliation is almost always beneficial and it is fallacious to assume that the removal of livestock from public lands will be the answer to the problems of land degradation. For example, grazing of most grasses stimulates new growth and increases the amount of tilling and grazing of shrubs (browsing) causes hedging (new growth of succulent stems and leaves from the old wood of these plants. In general, plant species that are resistant to defoliation have four main characteristics: (i) (ii) They maintain vegetative buds in or close to the soil surface They do not elevate the apical meristem more than 2 or 3cm until rapid elongation and flowering take place. (iii) They produce numerous fruiting stems. (iv) They have the capacity to initiate abundant new culm development (tillers from basal buds). The degree of development of the above characteristics in a plant species will determine its ability to withstand grazing and survive.

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Effects of Defoliation on Plant Growth The disappearance or persistence of grazed plants is largely regulated by the amount of energy reserves stored in the plant. Over-grazing generally reduces the non-structural carbohydrates in roots and perenniating stem bases and can have a detrimental effect on plant growth and survival. However, structural carbohydrates are not considered when discussing the effects of defoliation on growth because they rarely become energy sources for plant growth. The non-structural carbohydrates are the major sources of stored energy reserves and these are referred to as Total Available Carbohydrates (TAC). TAC compounds originate in the leaves and other sites of photosynthesis and are stored in the leaves, roots, rhizomes, stolons and lower stems (i.e. the points of origin of new growth). Variations in TAC between plants are related to: (i) (ii) (iii) (iv) The species of plant The plant growth cycle The part of the plant being examined The site where the plant is growing.

Because of these variations, TAC is not always a good measure of plant response to grazing but it can be used as an indicator. Management of grazing should not necessarily aim for maximum levels of TAC but should not exceed critical minimum levels. A lot of research is directed at determining these critical minimum TAC levels. Defoliation reduces the leaf area index (LAI) or the ratio of living leaf area to the ground surface, at least temporarily, and this reduces the conversion of solar energy to plant energy so the morphological response of a plant to defoliation has a marked effect on its growth. Effects of Intensity of Grazing on Plant Survival The intensity of grazing relates to the amount of material remaining after grazing not to the amount of material removed during grazing. The intensity of defoliation or grazing will influence plant survival depending upon the morphological response of plants to grazing. For example, alfalfa does not withstand high intensity grazing because it damages the crown whereas white clover (Trifolium repens) can withstand high intensity grazing because it producing rhizomes and under moderately heavy grazing it will actually be an increaser. Effects of Frequency of Defoliation on Plant Survival The frequency of defoliation refers to the interval of time between defoliations and the number of defoliations. Frequency is closely related to intensity of defoliation but is not the same, since repeated defoliation at the same height would hold grazing intensity constant but vary the grazing frequency. For example, consider two plant species that are grazed constantly to the same height. One species responds to defoliation by producing new growth twice as rapidly as the second species. The first species can then be grazed twice as frequently as the second species, but since the same amount of plant material is left in both species, the intensity of grazing is the same for both species. The ability of a plant to produce new growth is dependent on many things such as plant morphological response, soil fertility, soil moisture, temperature, day length, site aspect (north or south facing slopes of hills) and so on. However, plants within one ecosystem may respond differently to defoliation depending upon the season in which the defoliation occurs.

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Effect of Season of Defoliation on Plant Survival The definition of season of defoliation should be made in relation to the growth curve of the plant. For example, plants that commence growth very early in spring are more sensitive to defoliation in early spring than are plants which commence growth in late spring and are still dormant in early spring. In late spring, however, the situation is reversed. Some grasses and forbs, those that have elevated growing points, are highly susceptible to defoliation and lose vigor when growing points are removed at any time. Others, that do not have elevated growing points, show little effect as measured by dry weight and seed production. The sensitivity of some plants to defoliation increases rapidly when the flower stalks begin to develop and decreases rapidly as the plants approach maturity. Plant growth curves differ from species to species depending on such seasonal factors as day length and temperature and moisture, so the season during which defoliation occurs will influence plant response differently between species. However, all plants are most susceptible to defoliation during the early part of their growth. Nevertheless, very recent and unpublished research from Agriculture Canada Research Station at Swift Current has indicated that the productivity of upland and lowland mixed prairie native pasture and crested wheatgrass is unaffected by season of grazing provided grazing occurred only once/year and was restricted to 6-10 days. Prehensile Technique Animals frequently pull whole plants from the soil. This is a drastic form of defoliation but during milder defoliation, animals tear developing grass stems from this sheaths and remove pieces of plants which they do not eat but discard. Plants differ in their resistance to pulling from the soil depending upon root structure, soil type and soil moisture but an overgrazed plant with a depleted root system surrounded by bare and eroded soil is much more likely to be completely removed from the soil. Animals also differ in their eating action. Bark wounding (removal of bark from trees), which if severe will cause death of the tree, is a special form of plant damage. Most bark wounding is caused by wildlife such as elk and deer when removing velvet from their antlers. The most spectacular destroyer of woody plants is the African elephant that has eliminated whole forests by breaking limbs from trees, pushing down trees and tearing whole shrubs and small trees from the ground. Over a long period of time this has converted much woodland in Kenya into grassland. Treading The treading action of animals breaks plants. This can be both beneficial and detrimental. If treading breaks dead standing herbage it brings it into contact with the soil surface and in so doing it speeds the process of decay and re-cycling of organic matter and nutrients. Treading can also help to bring seeds into contact with the soil and thereby increase the rate of seedling germination and survival. However, if treading breaks green plant material or newly emerging tillers it not only renders this herbage unavailable to the grazing animal but also damages the plant. Treading compacts the soil which: Reduces seed emergence Reduces root growth which in turn reduces nutrient uptake by the plant and as a consequence plant growth is reduced Reduces moisture infiltration, increases moisture run-off which increases the risk of erosion and reduces plant growth and survival (see Table 35, page 68). The degree to which treading will compact soil depends on the stocking rate, texture, structure, porosity and moisture content of the soil.
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Fouling Fouling, or the covering of vegetation with feces, by livestock has both beneficial and detrimental effects. Feces and urine are important components in the re-cycling of nutrients on range and seeded pasture. This will be discussed in a later lecture. However, fouling can also damage the vegetation. For example one report from New Zealand indicated that dairy cows deposited feces at an average rate of 13.9 time/day and that each deposit covered 0.07m2. This amounts to 355m2/cow/year (in New Zealand grazing continues for 365 days/year) or 0.035ha/cow/year or 3.5ha for a herd of 100 cows. The experiment also found that 75% of the vegetation under the manure was killed and did not re-grow the following year. Beef cattle defecate less frequently than dairy cattle but in southern Saskatchewan where the rainfall is considerable less than in New Zealand, the cow pies take much longer to decay and the damage to the vegetation is probably much longer lasting, though there are no data available to confirm or deny this suggestion. The extent of this damage depends on the species of animal (sheep, goats, deer etc. cause less damage than cattle because their feces are deposited in small pellets and are much drier, allowing the vegetation to grow through. Re-distribution of plants by grazing animals Animals are important distributors of plants over native range and seeded pasture lands. They can distribute plants by the physical adherence of seed to the hair, wool, horns, hooves, etc. The presence of hooks, barbs, awns, and sticky exudates on seed will influence the physical distribution of plants in this way. Whether or not animals are cloven hoofed (divided feet such as cattle or sheep) or not (such as horses or mules) will affect plant distribution by this method. Sheep and cattle carry more seed between the claws of their feet than horses. Seeds are also redistributed in the manure. This has been recognized an important way of revegetating seeded pastures in Australia for 20 years. For example, white clover (Trifolium repens) has been re-introduced into pasture from small nursery areas. The nursery was located in a pasture where the clover seed was allowed to ripen and cows were then allowed access to both the nursery and the pasture. In 3 years the pasture had been re-vegetated. Manure provides a favorable environment for nutrients and moisture for the developing seedlings but not all seedlings survive. However, in that experiment, 30% of the white clover seed passed through the digestive tract of the cows and 85-90% of them germinated. Although this may be a slow way to introduce the legume into the pasture compared with re-seeding it was effective, economical and environmentally friendly. Birds are also important in redistributing seeds over rangelands. Of course not all seeds redistributed by animals and birds in this manner are desirable and in North America we have been perhaps pre-occupied with the negative effects of plant re-distribution. For example, a study 30 years ago in California suggested that in a single grazing season one cow could redistribute 36 species totaling more than 900,000 viable seeds of weed species. However a report from Utah in 1996 indicated that crested wheatgrass could be introduced into a pasture by feeding 1/3lb of seed/acre. Cows are more successful at re-distributing seeds on range and pasture than sheep of horses because their feces are more moist and they do not masticate their food to the same extent as sheep and horses. For these reasons, wildlife are also probably less likely to redistribute plants than cattle. Finally, the rejection of fouled areas by animals for sources of feed also provides an important source of parent material for seed setting and helps to ensure the survival of plant species.

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Chemical Effects Nutrient Cycling Figure 30 provides a generalized diagram of nutrient cycling in range and pasture systems

Figure 30. Generalized representation of nutrient cycling on grazed range and pasture Grazing animals return a large proportion of the consumed plant nutrients to the soil. A mature cow may produce as much as 25kg feces and 9kg urine daily. Average chemical composition on a percentage net weight basis of the excreted materials is shown in Table 40. Table 40. Average nutrient contents of feces and urine of beef cattle. _________________________________________________ Nitrogen % In feces 0.38 P 20 5 % 0.18 K 20 % 0.22

In urine 1.1 0.01 1.15 _________________________________________________ Most of the voided phosphorus occurs in the feces whereas the urine is rich in nitrogen and potassium.

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Table 41. Nitrogen, phosphorus and potassium excretion in the feces and urine of cattle grazing native range.

N In feces (%) In urine (%) Total (g/d) Fertilizer equivalent (kg/ha) 0.38 1.10 194 15.2

P 2O 5 0.18 0.01 46 1.6

K2O 0.22 1.15 159 7.7

Table 42 compares the amount of nutrients removed from the soil in a crop of spring wheat and a crop of calves. These data indicate clearly why it is not possible to maintain satisfactory yields of wheat without the application of fertilizer. In contrast, the grazing of cattle on range and seeded pasture is the most sustainable form of agricultural production because nutrient removal in minimal. Table 42. Amount of various elements removed from the soil by a crop of grain or calves (R kg) expressed as a proportion of those elements taken up by the plants (V t). N Wheat R/V (kg/t) Calves R/V (kg/t) 735 39 P 852 36 K 86 4 Ca 204 71 Mg 500 1

Without herbivores, nitrogen and minerals cycle from soil to plants, to litter and back to soil. Animals add many more pathways because the nitrogen and minerals pass through animals, may be changed in chemical structure, may be exogenous or endogenous and are returned to the soil in feces or urine. They are removed from the rangeland the form of meat, milk, wool, etc. and, while the yield of these animal products is very much lower than that of grain and oilseed products, the reduced efficiency of production translates into increased sustainability of production. For example, in the U.K. it has been reported that pasture yield increased from 10,000 kg/ha for cut and removed pasture to 12,000 kg/ha for grazed pasture with the same rates of fertilization. The increased production from the grazed pastures is caused by the re-cycling of nutrients as opposed to their removal from the cut and carry operation. The nutrients in urine are readily available to plants, though there is some loss, particularly of nitrogen, through volatilization. Rates of manure decomposition depend on the climate (rainfall and temperature) and coprophagous insects and other biota and micro-biota are also important. The activities of these organisms incorporate manure into the soil, reduce infective stages of parasitic worms, reduce breeding areas for flies, and increase rate of mineral cycling.

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GRAZING ANIMAL BEHAVIOR Animal Distribution Control of the distribution of animals on range is a means of increasing livestock productivity and therefore increasing the economic return. It is also important for the prevention of overgrazing in some areas and hence maintenance of range condition. However, uniform grazing may not always be conducive to wildlife management. For example, uniform removal of ground cover is detrimental to bird nesting grounds and small ground dwelling mammals such as mice, gophers etc. that are an important food source for game birds. It may also remove important food sources for large herbivores such as deer, elk, pronghorn etc. especially if the grazing by livestock occurs on the traditional winter grazing grounds of these wild herbivores. Factors Affecting Animal Distribution on Range: Forage type. The presence of preferred species influences animal distribution on range. Animals tend to congregate in those areas where the preferred species are most abundant. Vegetation Type. Vegetation type influences animal distribution. For example, domestic animals will not normally move into dense timber except for shade and then they usually remain close to the perimeter. For example, it has been reported that regrowth of trees in a mosaic with grassland after logging and fire reduce the area available for grazing because patches of trees restrict animal movement. Topography: In one study, 80% forage utilization by cattle was reported at the base of 20% slopes and 0% utilization at a point 1.6 km up the slope. Another example comes from a mixed bunchgrass area in Southern Idaho in which 38 different bunchgrass species were identified. 75% utilization occurred to a distance of 740m above the base of a 10% slope and continued a further 32m when the gradient changed to a 60% slope. However utilization decreased to zero a further 80m up the 60% slope. The positioning of water, which usually occurs at the base of the slope, may also be a complicating factor. Animal species and breed: Sheep prefer to graze into the wind and it has been reported in Australia that over grazing occurs in that part of a paddock that is nearest to the direction from which the prevailing winds come. This has not been reported for cattle even though they usually prefer to face into the wind. It has been reported that Santa Gertrudis cattle walk further than Herefords in Texas and thus utilize range more evenly. Deer will move through thicker undergrowth than elk, cattle and pronghorn. Water requirements of different species of animals vary and proximity to water will affect the distribution of species on range. Even different breeds within a species have different requirements. For example, an observation in Australia with Merino and Dorset Horn breeds of sheep suggested that differences in water intake may influence animal distribution and grazing behavior. Sheep have strong flocking instincts and it was frequently noticed that when the sheep traveled from the grazing grounds to water the Dorset Horn sheep drank while the Merinos did not. The Merino sheep waited for the Dorset Horn sheep to finish drinking and then traveled back to the grazing grounds with them. This was a waste of energy for the Merino sheep and they would have been better off separated from and so not influenced by the Dorset Horn sheep. (Sheep have strong flocking instincts and this improves control of movement, and control of predators but can have deleterious effects on range as a result of overgrazing and trampling unless the sheep are moved regularly). There are other examples. Camels range further from water than any other herbivore. Elephants on the Serengeti Plain of
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Tanzania range further from water than zebra or wildebeest. These behavioral differences reflect physiological differences between the various animals. Topography x Species Interaction: Different species of animals prefer different topographical sites. For example, in Utah, greatest summer use by deer occurred at slopes between 30 & 40%, by elk at slopes < 30%, by cattle at slopes < 10%, while mountain sheep and goats preferred slopes > 40%. Water: Water serves as a focal point for grazing animals. For example, in Northern Territory, Australia, the average size of a cattle ranch is 443,000 ha (1 million acres). There are no fences and cattle are watered at watering points serviced from underground water. These "water holes" serve as the fences as cattle don't graze any great distance from them. The distribution of water holes therefore controls the distribution of range utilization. Animals trail to and from water leaving stock trails caused by trampling the vegetation. These become sources of erosion. In one 260 ha paddock sheep left trails radiating from water a total of 32km. Forage utilization is influenced by distance from water. For example, in Montana, utilization of Agropyron smithii (Western Wheatgrass) by cattle was 100% at water, 54% 180m from water, and 28% 1460m from water and utilization of Bouteloua gracilis (Blue grama grass) was 100% at water, 38% 180m from water, 19% 1460m from water (Table 40). Table 40. Utilization of Agropyron smithii and Bouleloua gracilis by cattle as influenced by proximity to water. __________________________ A.smithii B.gracilis ___________________________ At H20 180 m from H2O 100 54 100 38 19

1460 m from H2O 28 ___________________________

Water is such a strong focal point for grazing animals that if water points are too far apart, severe overgrazing of range will occur at or near water while range distant from water may be totally ungrazed. Practices to reduce animal concentrations Development of water: Dug-outs, wells, springs, upland tanks into which water is pumped by electrical or solar energy should be developed in areas where grazing is lightest. If possible they should be located away from other minor focal points such as shade, entrance gates etc. Cattle should not have to walk more than 1 km to water from any point in a pasture. If spring water is used it is very important to prevent cattle entering the source as they will quickly compact the soil and slow or entirely stop the flow of water. Riparian areas, those areas surrounding water, are ecologically important for plant and animal survival but they are also ecologically fragile. Therefore care must be taken to ensure that these areas are protected from degradation by livestock. This can be achieved by fencing these areas and pumping the water to tanks, or fencing and allowing access to cattle only in designated areas.

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Fencing: Delineates ownership and allows confinement and direction of animals to areas to be grazed. Because of the high cost, fencing has to be effective. Solar generated electric fencing is now very popular and comparatively inexpensive. Wind generators can also be used. Salt, mineral supplements and oilers: Salt and mineral supplements and oilers can be used to increase range utilization and animal distribution. Because water is the focal point for animals on range, these are all minor focal points and should be placed away from water and in those areas of least utilization. Unlike water, the salt and mineral supplements and oilers can be moved to alter the distribution patterns of cattle. This is a very economical way of influencing range utilization. Salt consumption is about 1.5kgsalt/animal/month in spring reducing to 150250g/animal/month in August. Salt is not effective in controlling livestock distribution on saline range. Blue salt, (cobalt iodized) is recommended for the prairies because prairie soils are mostly marginal or deficient in these trace minerals. Livestock Trails: These are used in areas of dense timber and shrub stands to encourage animals to walk through these areas. Dikes over sloughs will encourage animals to range further. Both these techniques are commonly used in the higher rainfall areas of British Columbia. Herding and use of Range Riders: The range rider's (or herder's) job is to: i) Control the distribution of animals on the range by directing animals to under utilized areas ii) Repair fences, windmills and other hardware. iii) Maintain adequate water and salt and move salt and mineral supplements iv) Care for sick animals v) Keep bulls evenly distributed in breeding herds vi) Prevent death losses from toxic plants, predators, disease etc. vii) Look after animal husbandry such as dehorning, castrating, fly control, internal and external parasite control, etc. Fertilizer and range seeding: The use of fertilizer and seeded species may increase the palatability of the species on range, particularly if a large number of unpalatable increasers are present. Fertilizer and range seeding can also be used to encourage animals further away from water, or further up the slope or into other underutilized areas. Prescribed Burning: Burnt areas may also attract animals because burning stimulates new grass growth. Multiple use: Grazing range with two or more species of animals at the same time can increase range utilization without damaging the range. This topic will be the subject of a later lecture. Consequences of Animal Distribution Behavior Destruction of Vegetation Uneven distribution of cattle on range or seeded pasture will result in uneven defoliation with the result that some areas will be over-grazed and other areas will be under-grazed as discussed in the section on defoliation (pages 81-83). Re-distribution of nutrients The importance of encouraging livestock to consume forage away from focal points has been emphasized. However, by then congregating in large numbers at the focal where they also defecate and urinate, livestock cause a redistribution of minerals. One report found that sheep deposited about 1/3 of their excreta on only 5-7% of the total area of a 40ha pasture. About 30% of the
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pasture became enriched and 70 percent impoverished. In addition it was reported that the bedding sites when compared with areas 128m away had twice the total N and exchangeable Ca, 5 times the Mg, 14 times the available P and 130 times the amount of K. Diurnal Patterns of Grazing Behavior Most grazing periods begin near dawn and again in the late afternoon ending around sunset. As the days get shorter, the intervals between grazing also decrease so that at latitudes greater than 20, midwinter grazing, if practiced is almost continuous during the daylight hours. This may be due to 1) Photoperiod 2) Temperature, or 3) Reduced herbage availability but it does have some implications. For example, it leaves less time for drinking, resting and ruminating. This effect on rumination time may be most important since, during winter, herbage fibre is highest, requiring more not less frequent rumination periods for efficient digestion. In tropical areas, where day length varies little, some studies have shown that cattle graze predominantly at night. This may be due to high temperature and humidity during daylight hours. During summer, cattle time their grazing to reach water during the middle of the day when it is hottest and they rest by the water until it cools in the afternoon when they commence grazing again. This means that cattle are reluctant to graze more than 1 km or so from water. Thus unless watering sites are carefully planned part of the range will be undergrazed and the other part will be overgrazed. The diurnal pattern of grazing is altered continually to adjust for climatic conditions and herbage availability. For example, as herbage availability decreases grazing time increases and vice versa. However, there are other factors that influence grazing time. For example, it has been shown that grazing time of cattle decreased at temperatures below 5C and above 26C. However, provided there is adequate herbage available, bite size will increase so that intake may not be reduced but selectivity will be reduced. At very high or low temperatures intake may be reduced because of a reduction of time spent grazing Thus, unless feed availability is increased beyond the thermoneutral zone (for example, by moving cattle to pasture with a higher DM availability or providing supplementary feed such as hay or grain) then feed intake will be reduced with a subsequent reduction in productivity. Other Factors Affecting Feeding Behavior Pregnancy and lactation: Pregnancy and lactation increase the rate at which food is consumed. For example, rate of consumption in cattle was increased 27% in late pregnancy and a further 20% in early lactation in one experiment. Pregnancy however, did not alter the time spent grazing, suggesting that bite size increased and feeding was less discriminate, whereas lactation increased grazing time by 7-12%. This may represent slower grazing because of the presence of a calf. Body condition: Thin sheep and cattle eat more than fat sheep and cattle. This is due to a small increase in grazing time and a large increase in rate of consumption and bite size all of which reduce selectivity.
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Level of production: Grazing time does not vary with cow size (Breed) but increases with increasing milk production. Social facilitation: Groups of animals whose grazing behavior patterns differ may interact socially to reduce the difference in grazing times. (This has been previously discussed). Energy supplementation: Feeding energy supplements, such as hay, grain, range cubes etc., to grazing animals reduces the time that they spend grazing. Herbage availability: In general, when feed is short more time will be spent grazing, the number of bites per minute will increase but the bite size will decrease and intake per hour of grazing will decrease. This has important implications for animal energetics in that energy expenditure is increased for the same or less energy intake so energy utilization is less efficient. This also may lead to grazing fatigue.

THE CONCEPT OF ENERGY FLOW THROUGH A GRASSLAND SYSTEM Energy transformations are achieved in primary production (plant growth) by photosynthesis and in secondary production by the assimilation of these products of photosynthesis (plant matter) into animal growth (bone, muscle, fat, vicera, organs etc.), animal product (meat, milk, wool) and animal reproduction through the conception, gestation (pregnancy),parturition (birth) and nourishment (lactation) of offspring. Each level of energy transformation is referred to as a trophic level. Thus, energy transformations from radiant solar energy to plants is termed the Primary Trophic Level. Energy transformations from plants to herbivores are at the Secondary Trophic Level. Carnivores (meat eating animals including man) occupy the Tertiary Trophic Level and decomposers (soil biota such as insects, microrganisms, fungi) can occupy the secondary, tertiary or quaternary (4th) trophic levels depending on whether they are decomposing plant material (secondary level), dead herbivores (tertiary level) or dead carnivores (quaternary level). Activity at each trophic level depends upon: i) ii) iii) iv) The amount of energy flow through the system The cyclic flow of nitrogen and essential minerals The availability of moisture The availability of gases such as CO2, O2, etc.

At each trophic level, energy is partitioned and nutrients altered by assimilation, respiration, excretion and production. These transfers of energy and cycles of nutrients constantly change in rate and magnitude. The most important objective in range and pasture management is to maintain or increase the long term efficiency of energy flow through the ecosystem. Energy flow through an ecosystem obeys Newton's first law of thermodynamics which states that "Energy is neither created nor destroyed but may be transformed from one form into another". For example, energy passes through a system only once. It can never by recycled. Primary production transforms radiant energy from the sun into chemical energy by photosynthesis. This chemical
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energy is not created by the plant but transformed within the plant from radiant energy. Neither is this energy destroyed when the plant is eaten by an animal or the plant dies. It is merely transformed from plant chemical energy into animal chemical energy or from plant chemical energy into the chemical energy of the decomposers. Gross primary production includes the energy stored by the plant and the energy used in its own respiration. The more useful concept of net primary production (NPP) excludes the energy used in respiration and in fact is the accumulated plant biomass. Similarly, gross secondary energy includes the energy used and eliminated during digestion, and respiration while net secondary energy represents the energy used for maintenance of body weight, weight gain, wool growth, reproduction, lactation and so on. A simplified energy flow diagram is given in Figure 31. Each of these connecting processes can be described by mathematical relationships in order to predict the energy level of the next compartment. This is the basis of computer simulation models such as GrassGro that can be constructed to predict the production of meat, milk, wool etc. from the amount of solar radiation and precipitation received by a pasture. In reality, quantifying the energy flow through a grazed pasture system is much more complex because it is also influenced by many other factors, some of which are indicated in Figure 32. Once constructed, a computer simulation model can be verified from actual data and then used to make very quick and inexpensive predictions as to the outcome of various inputs or management practices. Only the most successful inputs or practices would then be put into practice saving years of costly experimentation.

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SOLAR ENERGY Unusable wavelengths Minerals, H2O, CO2 Respiration NET PRIMARY PRODUCTION Death Food ignored CONSUMPTION H 2O, O2, supplements Excreta ASSIMILATION Respiration NET SECONDARY PRODUCTION Weight loss, death BIOMASS CHANGE (YIELD) Trucking, packaging, marketing PREDATION FOOD Trim (bone, fat, offal, hide, hooves, horns, bruising etc.) Figure 31. A simple energy flow chart showing the sources, pools and sinks for a range or pasture ecosystem.
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Figure 32. Flow diagram indicating some of the major factors influencing the conversion of solar energy to liveweight of grazing sheep. When energy flow through a grassland ecosystem is studied, the most striking information learned is the realization of just how inefficient this flow of energy is. In one study it was estimated that rangeland in California received 1,600,000 Kcals/m2 of solar energy. 700,000 Kcals or 44% of this energy was in wavelengths that were usable by plants. Net primary production at the site amounted to 3,275 kg DM/ha or in energetic terms 14,150 kcals/m2. This level of production from natural grassland is quite high when compared to many other grasslands such as those in southwest Saskatchewan where NPP may be in the range of 1,000-2,500 kg DM/ha. Nevertheless NPP at the California site represented only 0.2% of the usable energy (UE) and 0.09% of the total energy (TE). Cattle production, or net secondary production, from the grassland amounted to 69 kcals/m2 or 4.9% of the energy in the NPP and 0.004% of TE. Assuming that 57% of the carcass remains after removing the hide, head, feet and intestines and that bone accounts of 30% of the remainder then only 40% of NSP is usable as food. This amounts to 27.6 kcals/m2 or 0.0017% of TE received at the grassland site. Thus, even though NPP at this grassland site was high in comparison to many areas in other parts of the world, the efficiency of converting total solar radiant energy and its conversion into food for human consumption was very low. Nevertheless, it must be remembered that rangeland is, by definition, non-arable for the growing of grain and vegetable crops so it is infinitely more efficient to graze it with animals that can be harvested for food than to leave it ungrazed in terms of its ability to produce food. Further, the low efficiency of production from rangeland is the basis of its high sustainability. However, both the efficiency of energy flow through and sustainability of rangeland are greatly reduced in poorly managed range.
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Similar studies have been carried out with wild animals to determine comparative net secondary production between wild and domestic animals. Cattle are comparatively efficient net secondary producers as can be seen from the following examples. Animal species Cattle Field mice African elephant White tailed deer Net secondary production (kcals/m2 ) 69.0 17.5 23.3 43.1

It is important to emphasize that the above ground animal species account for only a small proportion of net secondary production. Soil biota account for as much as 90% of the total secondary production because of their large numbers and rapid metabolic rates. Energy passes through a biological system only once with large unrecoverable losses at each transfer. Because energy is not recycled like nitrogen and minerals, increased efficiency of energy use depends greatly upon prevention of losses. There are many things that can be done to increase the efficiency of energy flow through a grassland ecosystem. For example, the flow of TE to NPP can be increased by proper grazing management to increase leaf area index (LAI) or perhaps genetic engineering may make it possible to develop plants which can use a broader spectrum of wave lengths of radiant energy. It may be possible to increase NPP by re-seeding the climax species in depleted areas if this is the most desirable species, or by seeding introduced (artificial species), by using fertilizer, irrigation, weed and brush (shrub) control, control of plant diseases, pest control (such as grasshoppers, rodents, rabbits, etc.). The efficiency of net secondary production can be improved by techniques such as providing range supplements of energy, protein and minerals at strategic times. Other practices include controlling animal distribution on the range, controlling animal diseases with vaccines, anthelmintic drugs, etc., predator control and using good animal husbandry such as superior genetic material and so on. Finally, the efficiency of energy flow from net secondary production to the tertiary (3rd) trophic level (food) can be increased by providing fast, efficient marketing methods, safe, humane and rapid slaughter and effective packaging, storage and retailing of meat products.

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MULTIPLE USE AND LIVESTOCK / WILDLIFE INTERACTIONS The practice of grazing two or more species of animals together on range is a form of multiple or common use. Attitudes towards common use have changed in recent years but even more influential in changing people's attitudes to common use is the growing interest in preservation of wild animals for game meat products and their recreational value. Multiple use can be an effective method of increasing the energy flow through a range ecosystem provided the species are complementary or synergistic and not competitive. Historically, domestic animals have been developed and used to provide human food. This is because the domestic animals are more efficient than are wild animals (Table 41). This should not be surprising considering the centuries of artificial genetic selection exerted by humans on domestic livestock. In general, animals separate themselves on range based on social tolerance between species and the proportions of grass, forbs and browse present on the range, the slope, elevation and density of trees etc. For example, pronghorn are socially compatible and will graze the same areas of a rangeland but bighorn sheep are not socially compatible with cattle and move off rangeland that is grazed by cattle. Cattle and horses prefer grass to a greater extent than do sheep and goats. Sheep and goats do well on grass but can also survive on browse. However, neither cattle nor horses can survive for long on a diet that is high in browse. Cattle make good use of tall coarse grasses while sheep make better use of the short fine grasses and broad-leaved weeds and so on. Table 41. Secondary productivity (yield/ha) of some large herbivores. Species Yield or Secondary Productivity (kg/ha/yr) ____________________________________________________________________________ Uganda Kenya Michigan Russia Michigan Tanzania Zimbabwe Tanzania, Native Range Kenya, Improved Range Australia Tropical Native Range Australia Tropical Seeded Range 2.4 1.1 4.6 1.4 1.4 19.0 22.4 76.0 473.0 36.6 676.0 Geographic Location

African Elephant Wildebeest White Tail Deer Saiga Antelope Moose Masai Cattle Native Cattle Beef Cattle Beef Cattle Beef Cattle Beef Cattle

Beef Cattle U.S.A. Temperate Native Range 46.4 Beef Cattle U.S.A. Temperate Improved Range 1000.0 ____________________________________________________________________________ Comparisons of the diets of cattle, elk and deer elk (Table 42 and Figure 33) indicate that cattle and deer and elk and deer are much less competitive than cattle and elk.

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Table 42. Spring/Summer Diets of Cattle, Deer and Elk. _____________________________________________ Grass Browse Forbs _____________________________________________ Cattle Deer 73 6 19 80 8 13

Elk 60 28 12 _____________________________________________

Figure 33. The effect of stocking by cattle on use of forest and grassland by deer and elk. It is not surprising to find that the introduction of cattle to range occupied by white tailed deer has little impact on continued use by the deer while the introduction of cattle to range occupied by elk significantly reduces elk use. Further, bighorn sheep appear to be relatively intolerant of livestock and will abandon areas occupied by livestock but pronghorn appear to have little aversion to grazing with cattle and sheep The similar dietary preference of cattle and elk has challenging implications to the management of range grazed by these two species. However, the common use of range by cattle and deer can considerably increase the productivity of the range because they graze different plants and often at different times of the year. The fact that several kinds of animals occupy the same region, pasture or even vegetation type does not mean that they necessarily occupy the same niche (microenvironment) and are in direct competition with each other. It is more likely that they occupy separate, but overlapping parts of the total area. There are six suggested ways that the separation of animal species is accomplished on rangeland: 1. Species concentrate in different parts of the vegetational mosaic. For example, deer feed and travel in areas where there is some brush present for shelter while cattle will feed and travel on the open range

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2. Species select different types of food. For example, pronghorn will eat pasture sage while cattle ignore it. 3. Species separate topographically. For example, cattle, deer, elk, bighorn sheep and mountain goats occupy areas of increasing slope and elevation or, species select the same area but at different seasons. For example, in BC, elk graze high range in spring/summer which cattle do not graze but graze lower range in fall/winter which is the spring/summer range of cattle. 4. Species separate on the basis of availability of water. For example, Elephants range 40 km away from permanent water so are not in competition with other species during the dry season but may be during the wet season when other species will range, more widely due to the presence of temporary water). Elk will range further from water than moose. 5. Species select different feeding levels in the vegetation For example, cattle will graze on the taller grasses while sheep prefer the shorter grasses. Cattle will not browse unless feed is in very short supply while goats are much stronger browsers. 6. Species separate on a vegetational basis according to season. For example, on the slopes of B.C. and Alberta during summer, cattle prefer to graze the lower slopes while elk will move to the higher slopes. However, in winter, the elk move down to the lower slopes. Thus cattle summer grazing grounds are elk winter grazing grounds though they are not in direct contact. As the number of herbivore species increases, the spectrum of consumed forage broadens and increasingly overlaps. A change in the population of one herbivore species influences the population of another species because it changes the vegetation. There are three fundamental resources available to grazing herbivores; space, forage and time. Common use grazing can expand all three of these resources. Examples of common use grazing increasing range productivity are difficult to find in the scientific literature. However, there are some examples. In an experiment in which sheep and cattle grazed a seeded phalaris grass/white clover pasture in Australia, the productivity of sheep was greater when they grazed with cattle compared with productivity when they grazed without cattle (Table 43). This was attributed to a higher yield of total green dry matter, clover and seeded grass and to fewer invader grasses under common grazing (Table 44). Table 43. Effect on sheep production when sheep grazed with cattle (SC) or separately without cattle (S). Data are mean of 3 years. Cattle production was not altered if they grazed alone or with sheep. _____________________________________________________________________________ Fleece Wt. Ewes Preg. Lambs Weaned Weaning Wt. (kg) (%) (%) (kg) S CS S CS S CS S CS _____________________________________________________________________________ 14.8 4.2 5.0 79.0 87.7 66.3 93.3 19.33 23.0 Difference/ha 11.9 1.3 4.0 128.1 _____________________________________________________________________________ Stocking Rate Sheep Equiv/ha

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Table 44. Botanical composition at end of the experiment summarized in Table 43. ________________________________________________ S CS _________________________ Green DM (kg/ha) % Clover % Phalaris grass 4544b 36b 20b 5480a 46a 30a C 5406a 57a 29a 14b

% other grasses (invaders) 44a 24b ________________________________________________

a and b - different letters following a value indicate the differences are significant (p<0.05) In another experiment, also in Australia, nematode parasite populations in sheep were shown to be reduced if they grazed pasture in rotation with cattle compared to continuous grazing of the pasture by sheep alone (Table 45). Table 45. The effects on mean worm counts in sheep grazing the same pasture continuously (S) or in rotation with cattle on either a 6-month rotation (SC 6) or a 12-month rotation (SC 12). __________________________________________________________________ Hc Oc T ax Nem Tc ________________________________________________________ S SC 6 1560 92 1382 679 2977 1120 1603 689 15627 247 Co 21 26 26

SC 12 1576 790 408 337 395 __________________________________________________________________ H c = Haemonchus contortus O c = Ostertagia circumcinata T ax = Trichostrongylus axei Nem = Nematodirus spp. T c = Trichostrongylus colubriformis C o = Cooperia oncophora

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Other interactions Disease Wild animals are susceptible to many of the diseases of domestic livestock and act as reservoirs. For example, Brucellosis (Brucella abortus), which causes abortion in cattle also occurs in deer, elk and bison. The incidence of Brucellosis in white tailed deer is very low but leptospirosis (Leptospira spp. ), which also causes abortion in cattle, is widespread in deer. Tuberculosis is another disease that crosses species barriers and can be carried by wild and domestic species. Foot and mouth disease can be carried by all ungulates. Predation Predators (carnivorous or meat eating animals) occur on most rangelands where wild &/or domestic herbivores are abundant. For example, coyotes can be found in almost all areas of the North American continent. Wolves are now mostly confined to the northern latitudes of Canada and recent attempts to re-introduce them to the northern United States has met with strong opposition from many ranchers who see their livelihood threatened and is still the subject of legal and legislative discussion in both U.S.A and Canada. Bears are now mostly confined to the mountain and wooded regions throughout the continent. All these animals can be a problem if they occupy rangeland that domestic livestock graze. In some states in the U.S.A., compensation payments are made to farmers and ranchers who lose livestock to predators. No province in Canada currently offers compensation for coyote predation, though Alberta compensated for losses in excess of $100 prior to 1993 when the program was discontinued. Alberta currently offers compensation for wolf and bear predation. Saskatchewan and Manitoba are currently considering compensation programs for coyote predation and Manitoba is also considering compensation for wolf predation. In all cases, compensation is based on the commercial value of the animal and in some instances this is limited to $400/animal. No additional compensation is paid for purebred animals. The compensation, when it is paid, is therefore insufficient to alleviate the losses. The issue is therefore a very emotional one involving heated debate between environmentalists and ranchers. Legislation concerning coyote control varies between states and provinces. In Saskatchewan, a rancher is allowed to exterminate any coyote that preys on livestock by any method that is humane and legal. Similar legislation exists in Alberta. Coyotes are mostly a problem for sheep, rarely attacking cattle. Control methods include: a) Husbandry practices such as removal of carcasses, shed lambing, night corralling, lighted corrals, removing natural cover adjacent to sheep pastures and corrals. b) Frightening tactics such as sirens, strobe lights, propane exploders, bells etc., however, these are often only effective for a short time until coyotes become accustomed to them. Coyotes are extremely adaptable to human habitation. c) Fencing such as high netting fences which may sometimes be electrified. This is expensive however. d) Aversion tactics which involve the conditioning of coyotes to avoid sheep meat. The most common aversion substance is lithium chloride that is placed on a fresh carcass. Lithium chloride when ingested makes the animal violently ill without killing it and the coyote learns to avoid the source of the problem, that is sheep. The technique is not very successful, however, probably because the coyote population is constantly changing and so new coyotes continually need conditioning. e) Use of guarding animals such as dogs. The most popular guard dog is the Great Pyrenees. These dogs are trained to live with the sheep day and night, leaving the flock only once a day at a pre-conditioned time to return to the homestead for feeding. If two dogs are used they can be trained to feed at different times so that the flock is always protected. Other
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effective guarding animals include donkeys and llamas, both of which are aggressive towards canids. Cattle grazing with sheep are also effective in small pastures. Ewes tend to turn and run from a predator leaving the lamb vulnerable. Cows, on the other hand, turn and chase a predator being much more protective of their young. This technique is commonly used in Australia where most farms have both sheep and cattle. f) Hunting and trapping can be effective but in Saskatchewan this requires a permit_ g) Denning is effective and involves the fumigation of pups before they leave the den. Carbon monoxide from vehicle exhaust fumes in commonly used as the fumigant. Frequently, the adults are also killed but sometimes they may get away and breed again. h) Toxicants such as cyanide, strychnine and 1080 (sodium monofluoroacetate) are all extremely toxic and can be very effective for coyote control. These substances can be used in baits or on collars with an inflatable rubber bladder that is strapped to the neck of sheep and bursts when the coyote attacks the sheep. Although effective, these collars are expensive and cannot be strapped to all sheep in the flock so their effectiveness depends on the coyote choosing the right sheep to attack. Rules governing what can be used, by whom, when and how vary from province to province and anyone contemplating this method of control should first contact the relevant Department of Agriculture. If the elimination or control of predators is contemplated however, it must be clear that the predator is causing a problem. For example, is the predator attacking and killing livestock or simply scavenging on dead livestock? Is the attack an isolated incident or is the predator attacking at random? Competitive advantage between livestock and wild herbivores Where there is competition for forage between livestock and big game, livestock have several advantages: 1. In times of feed shortage (winter, drought etc.) livestock can be removed from the range or fed on the range. Range feeding of wildlife is possible but less successful. 2. Livestock are less selective in their foraging habits than wildlife and accept alternate feeds more readily. 3. Many big game species become habituated to certain range areas and do not readily move to other areas during a feed shortage. Consequently the competition exerted on wild animals by livestock is generally more severe than the reverse. While most ranchers are intolerant of wild herbivores if they cause excessive destruction of stored winter forage such as haystacks, they also welcome the presence of these animals on their pastures. This is not only because of their intrinsic beauty but also because they know that, their presence is an indicator of good range management. This is because these animals will disappear from depleted range. Impact of small animals and insects on rangelands Grasshoppers are very large consumers of vegetation on many of the world's rangelands. In fact it has been claimed that they are the most important herbivore in terms of consumption. They are capable of totally denuding all vegetation in their path when they reach plague proportions. However, they are more abundant on heavily grazed pasture than moderately or lightly grazed pasture. They are difficult and expensive to control on range and pasture and therefore attention to proper grazing management is the most effective method of control. The elimination of grasshoppers is not an option as an alternative to control through good management because they
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are an important, and at times the main or only, food source for many species of birds and therefore play an important role in the maintenance of rangeland ecology. Small animals also occupy almost all of the world's rangelands and their impact can be substantial. For example, 380 gophers are equivalent to 1 steer and 5.8 Jack rabbits are equivalent to 1 sheep. One estimate in U.S.A. indicated that rodents consumed 28.7% of all the vegetation on range and 38.8% of the most valuable grasses for livestock. Examination of the stomachs of prairie dogs in Montana, Wyoming, Colorado and Arizona indicated that 78% of the food was of valuable forage species and only 19% was low value plants for livestock. However, it must also be remembered that these animals are also a very important source of food for many species of game birds and they are therefore an important component of range ecology. Their presence on overgrazed range is likely to increase providing yet another example that moderately grazed range is not only beneficial to the range ecology but also, in the long term, to the economics of ranching.

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