Rangelands 6(6), December1984 261

Comparative Contribution of Grasses,

Forbs, and Shrubs to the Nutrition of Range
Ungulates
Jerry L. Hoiechek

Knowledge of the role of different forages in meeting the these substancesas gases and greatly reducestheir assimi-
nutritive needs of different range ungulates is critical for lationby theanimal's digestive system. Ifassimilatedat high
good range management. This knowledge is particularly levels, the volatile oils found in many sagebrushes,rabbit-
useful for decisions regarding brush control, range seeding, brushes, and Juniperscan be toxic to the animal.
grazing management, and forage allocation to different
ungulates. This paper will discuss the role ofgrasses,forbs, Theintermediate Feeders
and shrubs in meeting the nutritional requirementsof range Domestic sheep,burros, and caribou are consideredto be
ungulates. intermediatefeeders.Theseanimals have thegreatestcapa-
bility to adjust their feeding habits to whatever forage is
Forage Selection by Different Lingulates available.Domesticsheepare probably better adaptedtothe
forage resource in the Intermountain West than any other
Range ungulates can be divided into three groups based ungulate because they will readily use grasses, forbs, or
on their foraging habits. These groups include the grazers shrubs depending on availability. The primary problem with
which consume grass-dominateddiets, the browsers which domestic sheep is that their short legs and relatively large
consume primarilyforbs and shrubs, and the intermediate bodymake them very susceptible to predation.
feeders which use equal amounts of grasses, forbs, and
shrubs. Comparative Nutritive Value of Grasses, Forbs and
Shrubs
TheGrazers
Cattle,elk, bighornsheep,mountaingoats,musk oxen and On ranges of the western United States, the primary nut-
bison are North American ungulates consideredto begraz- rient constraints on ungulate productivity are inadequate
ers. However, on some ranges these ungulates, with the concentrations of energy, protein, phosphorus and vitamin
exception of bison and musk oxen, do consume large A in the diet. With a few localized exceptions, mineral defi-
amounts of forbs and shrubs. This occurs primarily when ciencies other than phosphorus are not a problem.
green grass is unavailable. These ungulates show a strong Various studies on forage nutritive quality show different
avoidance of shrubs high in volatile oils (junipers, rabbit- forages provide different levels of criticalnutrients at differ-
brush,various sagebrushes,etc.) becausetheylack mecha- ent times of the year. Therefore, ranges with the widest
nismsto reduce the toxic effects of these substances. diversity of plant species providethe best nutritional condi-
tionsfordomestic or wildungulates whenyear-long grazing
The Browsers is practiced.
Moose,pronghorn, mule deer,domestic goats, and white-
tailed deerfeed primarily on forbs and shrubsthroughout the TheGrasses
year regardless of location. Withthe exception of domestic Grassestypicallyhave lower crude protein, phosphorus,
goats,these ungulates experiencedigestive upsets if forced and lignin concentrationsand highertotalfiberand cellulose
to consume diets dominated by maturegrass. This groupof concentrations than do forbs and shrubs. Digestibility of
ungulates consumes a limited amount of grass in the spring grassesis generally lessthan forbsand shrubs.Atcompara-
when it is green and forbs and shrubs are unavailable.How- ble growth stages cool-season grassesare higher in crude
ever, dry mature grass is almost completely avoided. The protein, phosphorus,and digestibility and lowerin fiber than
smaller ruminants in this group can consume large amounts warm-season grasses. Plant fiber is digested more slowly
of forages high in volatile oils becausetheir small, pointed than the cell contents. Thehighcellulose (digestible portion
mouth parts enable them to select the portions of these offiber)concentration and high celluloseto lignin (indigest-
plants with the lowest levels of volatile oils. In addition, the ible portion of fiber) ratio makesgrassesbestsuited to large
small ruminants chew their food to a much greater extent ruminants such as cattle or cecum digestors (horse) that
than large ruminants or monogastric animals. Apparently have low nutrient requirementsper unitbodyweight. Leaves
finechewingof plantshighin volatile oils results in releaseof ofgrassesare nutritionally superiorto stems. Forthis reason
short grasses are nutritionally superior to mid and tall
The author is associateprofessor of range science,DepartmentofAnimal
and Range Sciences, New Mexico State University,Las Cruces 88003. grasses particularly during dormancy. Grasses are usually
the component of the forage resourceavailablein the great-
262 Rangelands 6(6), December1984

estquantityunlessovergrazing has beensevere.Theirhigh that explains why differentungulates select differentlevels

availability makes them important to large ruminants that of grasses,forbs, and shrubs in their diets.
have a total forage requirement. Ungulateshavetwobasictypesofdigestivesystemswhich
include the rumenand cecum systems. Both systemsevolved
TheForbs to enable ungulatesto digest plant fiber (plant cell walls) by
Forbs have higherlevels of crude protein, phosphorusand microbial (bacteriaand protozoa) fermentation.Thefermen-
digestibilityand lowerlevels of fiber when actively growing tation processes are quite similar in both the rumen and
than grassesor shrubs. Leaves fromdeciduous shrubs are cecum. Thesystems differ in that the rumen is an enlarged
similar toforbs in nutrientcontent. Whendormant, forbs and portion of the digestive tract that food must pass through
deciduous shrubs leaves rank intermediatebetweengrasses before entering the true stomach. Thececum occurs as an
and evergreen shrubs in nutritive quality. Becauseof their enlarged portion of the large intestine that food enters after
low fiber levels, forbs and deciduous shrub leaves break passing through the truestomach.
down quickly in the rumen and permit higher intakes than The rumen system has two advantagesover the cecum
grasses or evergreen shrub leaves during active growth. system. Theprocessofrumination (regurgitationand rechew-
Forbs and deciduous shrub leaves are critical dietary com- ing of forage) results in considerable reduction of particle
ponents to small ruminants such as white-tailed deer and sizethat providesmore surfacearea for microbial digestion.
pronghorn that require low fiber diets. Becausefood must be broken downto fine particle size to
It isimportant to recognize many forbs are poisonous. For leave the rumen, retention of fiber is longer than in the
this reason they have often been collectively considered cecum.Thisresults in morecomplete digestionoffiberinthe
undesirable by range managers. Studies conducted by rumen than cecum since fiber digestion isatimedependent
myself in Oregon with cattle and Leo B. Merrill in Texas process.A second advantage is that in the rumen system,
(JRM, 1978, 31 :351) with cattle, sheep and goats show that microbes are passed from the rumen into the abomasum
poisonous forb problems are minimal if the range is in good where they are digested and then absorbed, providing the
condition(a high diversity of palatable plants) and grazing animal with an important source of protein. Little microbial
intensity is moderate.The ranges I studied in Oregon con- protein is absorbed by cecum digestors because microbial
tained a wide variety of forbs. Many of these forbs were fermentation occurs afterthe food has passed through the
poisonous. During three years of study involving over 350 stomach. However,recent researchshows horseswill ingest
different cattle, no death losses due to poisonous plants their feces when their diet is low in protein, which partially
occurred although many poisonous plants occurred as compensates for the inefficient use of microbial protein
minor dietary species.These rangeswere in good condition associatedwith cecum digestion.
and stockedso approximately30%use oftheforage resource The primary advantage of cecal digestion of fiber is that
occurred. forage material can passeasily Out ofthe cecum withoutany
great reduction in particle size. Although fiber digestion is
The Shrubs lesscomplete by cecum digestors compared to ruminants,
Evergreenshrub leaves and buds from deciduous shrubs compensation occurs because they can consume a much
have highercrude protein, phosphorus, carotene (Vitamin
greater amount of forages as theydo not have to break fiber
A) and digestibility levels and lowerfiber levelsthan grasses down to a small particle size to pass it out of their system.
and forbs when forage is dormant. Woody material from On the basis of the previousdiscussion it isapparent that
shrubs is highly lignified and very low in nutritive value. cecum digestors can subsist on lower quality diets than
Therefore grazing animals are highly selective for leaves, ruminants. However,they must haveagreater forage supply
buds, fruits, and young twigswith low lignification. Rumi- since they use the forage lessefficiently. This explains why
nants with small mouth parts such as goatsor pronghorn can horses can survive on coarse, mature grasses better than
select against woody material much better than cattleor elk. cattle.
However,evergreen shrub leaves do provide an important Large ruminants can subsist on higher fiber diets than
crude protein, phosphorus,and carotenesupplementtocat- small ruminants becausethey have lower nutrient require-
tle and elk on many ranges when grasses are dormant. ments per unit body weight. Therefore a large portion of the
Deciduousshrubs with broad leaves such as snowberry and dietis typicallycomprisedof highly available forage such as
ninebarkare heavilyused bycattleduring periodsofdrought. grassesfor bison and cattleor woody material from shrubs
Manyevergreenshrubs such as the oaks, sagebrushes,and and trees as in the case of the moose.Thesmall ruminants
junipers havevolatile oils or tannins which bind up proteins,
such as white-tailed deer and pronghorn must consume
reducing the nutritive valueoftheforage. Animalswith small dietsdominated by leafy material and fruitsfromforbs and
mouth parts that canbe highly selectivesuch as goats, deer shrubs that have high levels of crude protein, phosphorus
and pronghorn use these plants most efficiently. and digestibilityand low levels of fiber. These animals can
General Discussion affordto beselectivefor thesematerials becausethey havea
low total forage demand.
Domestic sheep are small ruminants that subsist well on
Digestive Systems
In order to understand ungulate forage selection, some grassdominated diets. Thisis becausetheyhavea relatively
knowledge of their digestive physiology is necessary.I will large rumensize in relationtotheir body weight compared to
provide a discussion of the primary aspects of this subject other small ruminants like white-tailed deer or pronghorn.
Rangelands 6(6), December 1984
ManagementImplications
In the past, range managementpractices haveoften been
gearedtowards replacing forbs and shrubswith pure stands
of grasses.Thevast acreagesof crested wheatgrassin the
Great Basin and lovegrassesin the Southwest support the
above statement. Rangelands with a pure stand of grass
provide good forage forcattleorin somecaseselkand mule
deer during active growth but poorly meet the nutritional
requirements of large or small ungulates duringmost of the
year. Researchin the Great Basin and the Southwest show
that inclusion of palatable forbs and shrubs in seeding mix-
tures with grasses can greatly improve livestock perfor-
mance during forage dormancy and provide better habitat
for small wild ungulates and otherwildlifespeciesthan pure
stands of grasses.
Range condition is usually based on the density and pro-
duction of native, palatable, perennial grass.A better criter-
ion might bethe diversity ofpalatable forage species. Under
this criterionit mightbe desirable if up to 20%of the yearly
Viewpoint: Ecological Site/Range Site/Habitat Type
Rextord Daubenmire
My attention has recently been called to a note that
appearedin theAugust 1983 issueof Rangelands,p 187-188,
which contains some misconceptions as well as errors that
should be pointed out.
In espousing his preferencefor the term "range site,"the
author of that note, E. William Anderson, stated that "the
habitat type* does not have managementimplications" for
rangelands.Thisiscompletely atodds with much workpub-
lishedby range specialists.The latter haveworked out identi-
fying characters of habitat typesin westernrarigelands and
pointed out their managment implications in work done in
New Mexico (Francis and Aldon 1983), Colorado (Francis
1983), Idaho (Tisdale 1979, Hironaka,et al. 1983), Montana
(Jorgensen 1979, Mueggler & Stewart 1980, 1981) and Brit-
ish Columbia (McLean 1970). Other workers have adopted
the concept but have not used the term habitat type.
It iscuriousto note that Andersonhasseenthat essentially
the same type of virginvegetation (bluebunch wheatgrass/-
Idaho fescue) occurs on north-facing slopesin dry climates
at low elevation, but shifts to south-facing slopes under
higher rainfall at higherelevations.Thishe views as indicat-
ing different "ecological sites." To a plant ecologist this
vegetationis indicating the reappearanceof ahabitat typeat
places having essentially the same ecologic sum of climatic
and soil conditions, as a result of microclimates of the con-
trasted slopes compensating for differences in macro-
climates.
*Habitat type is a collective term for all parts of the earth's surface which
support,or are capableof supporting,thesamekind ofplant association,i.e.,
thesame climax. The plant associationmust have thesame potential domi-
nants in all layers.
263
forage production wascomprised of palatable annuals.It is
important to recognizemany annualgrassesand forbs grow
in periodswhen perennialsare dormant. Researchby Randy
Rosiere and others (JRM, 1975, 28:89) in south central New
Mexico and myself in northern New Mexico shows annual
forbs provide an important nutritional contribution to cattle,
sheepand pronghorn dietsand reducepressureon palatable
perennial grassesduring the growing season.In the North-
west, Martin Vavra's researchshows the introduced annual,
cheatgrass,provides green forage forcattle, mule deer, and
sheep in the fall, winter, and early spring when the native
perennial grassesare dormant.
A large number of studies involving both domestic and
wild ungulates in North America are consistent in showing
that forage selection changes tremendously within and
between years. The nutritive quality of various forage spe-
ciesalso shows greatfluctuations within and betweenyears.
Thegreater the degreeof forage selection a range provides
domestic or wild ungulates, the more likely they will be to
meet their nutrient needs. S
The concept of "range site" suggests a single-purpose
objective, i.e., managementof land for producing livestock
forage. Habitat type, in contrast, emphasizessimilarities and
differences in ecosystems which carry implications for a
variety of land management objectives such as livestock,
wildlifeandtimber production, for predicting diseasehazard,
for indicating hydrologic cycles, etc. It is not clear what
Anderson means by "ecological site," but if these words
carry their usual meaning,the concept should coincide with
habitat type, for the latter represents a narrow range of soil
plus climatic conditions, as indicated by the tendency for
this combination to favor a remarkably few species out of
hundreds in the flora, and to determine which among them
will dominate all others. The concept was first developed
solely as a basis for arranging a wide variety of disturbed
vegetationtypes in ecologically equivalent units. However,it
soon becameevident that these unitshad practical value in
forest managementwith respectto choiceof speciestofavor
maximum productivity in each habitat type, and which habi-
tat types are best suited for the growth of a particularly
desirable species. Habitat types provide a guide to distin-
guish between land where dwarf mistletoe can and cannot
infect ponderosa pine. Especially significant was the fact
thatwhen independentlychecked by field workersin applied
ecology, the system proved easy to use and capable of
adding materiallytothestock ofeconomicallyuseful informa-
tion.
Theclaim thatthe"range site concept iswidelyused in the
U.S. and is becoming internationally accepted as the basic
unit of resource inventory for the purpose of planning use