RESEARCH,
MANAGEMENT, AND ADMINISTRATION OF THE
RANGELAND
ECOSYSTEM
-by
C. Wayne Cook
A
NEW approach to rangeland management concerns the synthesis and use
of information dealing with the structure and function of the rangeland
ecosystem. Management and administrative decisions can be made more
logically if the entire ecosystem is understood. In addition, voids
in information are more readily discovered through this approach. Thus,
the need for and the role of research become more obvious.
Many flow charts have illustrated
the function of the natural and the agricultural ecosystems. The rangeland
ecosystem, since it is concerned largely with native vegetation, more
nearly resembles the natural ecosystem. However, since livestock are
introduced and manipulation procedures are frequently involved, it partly
resembles the agricultural ecosystem.
The rangeland ecosystem
differs from a natural ecosystem in that it will be purposely manipulated
for more effective production of salable produce. It differs from the
domestic crop ecosystem in that the total standing crop does not represent
the exported product.

|
Flow chart
showing structural characteristics of the ecosystem in six
compartmental areas and connecting interaction linkage among
compartments as a function of process.
|
|
STRUCTURE
OF THE ECOSYSTEM
All
physical features, along with expressions of flora and fauna, characterize
the structure of the ecosystem. Structure of the ecosystem is concerned
with the whole community complex of plant composition, plant aggregation,
life forms, layering, density, crown cover, leaf form, age classes, herbage
yield, constancy, fidelity, etc. Measurements describing the structure
of the ecosystem involve vegetation patterns, dispersion, heterogeneity,
population dynamics: in general, the synecology of each community and
the autoecology of each species in both above- and belowground environments.
Because both plant and soil development on any site is the product
of environment, both micro and macro climate, along with latitude, altitude,
and exposure, should be described.
The status of animal life is an important characteristic of the ecosystem.
This segment of the biotic community represents the consumers in the food
web. Structure of the ecosystem concerns the identification of dominant
species of fauna, their population dynamics, dietary habits, and behavioral
patterns.
New statistical methodology allows for the evaluation of the influence
of each independent factor upon any one of the several dependent factors
in the ecosystem. Such programming in ecosystem analysis has been rather
meager to date, but in the future such an approach shows great promise.
By such analysis, the effect of environmental factors on yield, frequency,
soil nutrients, plant chemical content, and many others can be assessed.
The effect of each environmental factor can be evaluated when operating
separately or when working together with all or any of the other designated
environmental factors.
FUNCTION
OF THE ECOSYSTEM
The function
of the ecosystem is rather complicated and involves many biotic and abiotic
interactions. These phenomena are not as well understood as the structure
of the ecosystem because of the lack of research in this area. The research
concerning functions of the ecosystem has been fragmented, and more work
remains to adequately explain the complete operations of the ecosystem.
Soil
Organisms
The importance of soil organisms
as decomposers and contributors to soil fertility is becoming more evident.
The addition of organic matter by fungi is believed to be important to
primary productivity on desert biomes. Soil crust algae and other cryptogams
may be major nitrogen fixers. Population levels of soil bacteria have
been identified with high soil fertility and optimum structure. Transfer
and conversion of soil nutrients for absorption by plants are of paramount
importance to nutrient cycling in the ecosystem. However, in arid range
soils their role has not been adequately studied or evaluated to the degree
that explains their full importance.
Energy Flow
A primary consideration
in the function of an ecosystem is the efficiency of the conversion of
solar energy to plant material by the primary producers. This is the first
step in evaluating efficiency of turnover of energy by the various trophic
levels and the efficiency of energy export from the ecosystem.
The conversion of solar
energy through photosynthesis by plants is governed largely by light,
water, and nutrients. On range ecosystems there is usually unlimited sunlight,
and the leaf area index is far from complete coverage of the surface area.
Therefore, there is little correlation between primary productivity and
incident solar radiation. Plant life may be dormant many months of the
year because low temperature, limited soil nutrients and inadequate moisture
reduce photosynthetic activity. Even at best, less than 2 percent of the
solar energy is converted to chemical energy. However, energy conversion
and transfer among different trophic levels in the rangeland ecosystem
is an important principle that can serve as a guide to management activities
for efficient use of all cycling nutrients in the system.
Duncan proposed a model for
predicting or evaluating efficiency of solar energy conversion by plant
foliage by considering the following canopy architecture: leaf area, leaf
angle, leaf position, reflectivity, transmissivity, brightness, and position
of sun. It does not appear that this model would optimize energy conversion
in rangeland ecosystems unless it included other limiting factors such
as soil nutrients and moisture, either of which should be as important
or more important than the leaf characteristics themselves.
Since the function
of the ecosystem actually starts with the process of photosynthesis, it
is important to understand this physiological phenomenon. Age of leaves
is important in the effectiveness of conversion of energy. As would be
expected, the more mature leaves are less efficient than the younger ones.
Life forms and species differ in the effectiveness of their chlorophyll-bearing
tissue in converting radiant energy to chemical energy.
Defoliation by animals, infestation
by parasitic insects that suck the life-giving saps from the plants, and
diseases that cause malfunction of the plant tissue all decrease the efficiency
of food manufactured by the plants.
In natural native ecosystems
the herbivore population is generally proportional to food produced or
food available for consumption. In any event, the entire consumer population
is directly or indirectly related to the ability of the plant population
to furnish food. Animal population dynamics, of course, depend on more
than food supply, because diseases, catastrophe, or social behavior may
also be regulatory.
Efficiency
of Energy Turnover
Efficiency
of solar energy conversion within the ecosystem has received much attention.
This is of paramount importance because of the transfer of solar energy
for sustenance of life in the ecosystem. The existence of the ecosystem
itself is dependent upon this phenomenon.
Biological
Efficiency
The herbivore
population represents the first major turnover in the use of fixed photochemical
energy in plants. This involves exchanges between plants and animals and
exchanges among animals with each other. The first interaction is obvious
since it concerns animal response from consuming plants and plant response
from being defoliated by animals. In this case, the stability of plant
cover and animal response to consumption of the primary producers is a
measure of efficiency of energy turnover.
In evaluating
consumer response, it is necessary to consider more than gross energy
content of the ingested forage. For instance, according to some studies,
maximum digestible energy (TDN X 2000) per unit area was produced by plants
that were harvested only after they completed their annual growth cycle;
but maximum digestible protein was produced when plants were harvested
some time during mid-growth and, again, at the end of the growing season.
Livestock responses on grassland ranges were satisfactory only when phosphorus
and digestible protein were high or added to the diet. This was true even
though digestible energy remained more than adequate.
It would seem that
the biological efficiency of the rangeland ecosystem could be based upon:
1) the amount of photochemical energy or nutrients produced per unit of
land surface area, and 2) the amount of this material that is transported
from the ecosystem in eatable produce, services, or wealth. The amount
of solar energy converted to chemical energy or nutrients and then to
the consumers serves as a measure of efficiency of the ecosystem, provided
limiting factors are corrected through manipulation consistent with all
uses to be made of the area and with reasonable cost and return relations.
It is generally
acknowledged that as energy passes down the food chain from producer to
consumer, the efficiency of use of solar energy diminishes. In each case,
however, management can increase the efficiency of energy utilization
in the transfer from one trophic level to another.
Species
Composition
The total
gross energy produced by plants has already been identified as a measure
of efficiency of the primary producers, but the transfer of this energy
to the consumers would, to a large degree, depend upon the chemical form
of the energy-furnishing constituents. For instance, many native browse
and forb species produce rather high quantities of resins, waxes, and
essential oils that represent chemical energy. However, energy in these
forms is not available to most consumers. In studies by Cook, it was found
that grass species produced approximately 2050 kcal of gross energy per
pound of dry matter during early growth, and browse species produced about
2322 kcal. The browse yielded only 1205 kcal of metabolizable energy for
animal use, compared to 1324 kcal of metabolizable energy per pound of
dry matter in grasses. Later in the season, when plants were mature, the
grasses contained 1859 kcal, and browse contained 1922 kcal of gross energy
per pound of dry matter. The grasses netted 42.9 percent usable energy,
but the browse netted only 33.4 percent usable energy. This was because
the essential oils in browse are not used by animals.
Animal
Nutrition
Nutritional deficiency among
animals is commonly a limiting factor in the efficiency of energy transfer
from the primary to the secondary producers. In most rangeland areas of
the world, the nutritional value of forage can be assessed on the basis
of digestible protein, phosphorus, vitamin A, and energy-furnishing constituents.
In local areas of higher precipitation and sandy soils, cobalt and copper
may be deficient in the forage. Effective conversion or transfer of energy
in the food chain is regulated by more than the mere supply of energy
in plant material.
It should also be noted
that the requirements of range livestock are different for various physiological
functions of the animal. For instance, the requirements for lactation
are about 25 to 30 percent higher than for gestation. For gestation
they are slightly higher than for maintenance; and for early lactation,
somewhat higher than for late lactation (Table 1).
| TABLE
1. RECOMMENDED
NUTRIENT REQUIREMENTS FOR CATTLE UNDER RANGE CONDITIONS DURING GESTATION
AND LACTATION ON A DRY MATTER BASIS. |
| Phase
of production |
Nutrient
|
| |
DP
(%)
|
TDN
(%)
|
DE
(kcal/lb)
|
ME
(kcal/LB)
|
P
(%)
|
C
(mg/LB)
|
| Gestation |
4.4
|
46
|
830
|
665
|
0.17
|
0.6
|
| Lactation |
|
First
8 weeks
|
5.4
|
57
|
1,120
|
900
|
0.22
|
1.6
|
|
Last
12 weeks
|
4.5
|
49
|
880
|
700
|
0.20
|
1.6
|
| *Nutrient
requirements are slightly higher for sheep because smaller animals
have a somewhat higher metabolic requirement per unit of body weight. |
| DP
represents digestible protein. |
| TDN
represents total digestible nutrients (calculated by deducting allowance
for higher ether extract in shrubs in the diet). |
| DE
represents digestible energy (calculated as for TDN). |
| ME
represents metabolizable energy. |
| P
phosphorus. |
| C
carotene. |
Providing
proper nutrition through the conjunctive use of seeded range with native
range for lactating animals in the Intermountain area of the western United
States has shown that more pounds of lamb and calf at weaning time per
breeding female can be obtained. In similar fashion, females in gestation
that were supplemented to correct protein and phosphorus deficiencies
produced more pounds of lamb and calf at weaning time compared with unsupplemented
females. Thus, it is demonstrated that a nutritionally balanced diet can
materially increase the efficiency of energy turnover in the ecosystem.
This would be true for all animal life in the ecosystem.
Supplements
not only furnish the deficient nutrient in the diet but, in many cases,
they enhance the value of the range forage by increasing daily intake
and digestibility of ingested range forage. In trials by Cook and Harris,
protein supplements consisting of cottonseed meal or soybean meal increased
digestibility of protein and cellulose in the range forage on desert ranges
used during the winter. Intake of digestible protein and metabolizable
energy from the range forage was increased by the protein supplements
fed to range sheep at either 0.5 or 0.33 pounds per day. Cottonseed meal
supplements, in most cases, increased daily intake of range forage. Energy
supplements consisting of barley or corn reduced the digestibility of
protein and cellulose in the range forage and in some cases decreased
daily intake of range forage (Table 2).
|
TABLE
2. AVERAGE DAILY INTAKE DURING WINTER GRAZING AS AFFECTED BY SUPPLEMENTS
OF PROTEIN AND ENERGY
|
|
|
|
Digestible
Protein
|
Metabolizable
Energy
|
|
Range
types and supplement
|
Daily
intake of forage (LB/day)
|
In
range forage (%)
|
Total
intake from forage (LB/day)
|
In
range forage (kcal/LB)
|
Total
intake from forage (kcal/day)
|
| Wheatgrass
pastures: |
|
|
Control |
2.31
|
2.8
|
.066
|
458
|
1058
|
|
|
Corn
supplement |
2.56
|
2.5
|
.064
|
423
|
1082
|
| |
CSM
supplement |
2.77
|
3.3
|
.091
|
544
|
1507
|
| Sagebrush-grass
pastures: |
| |
Control |
3.21
|
4.2
|
.135
|
318
|
1021
|
| |
Corn
supplement |
3.38
|
3.4
|
.115
|
271
|
916
|
| |
CSM
supplement |
3.69
|
4.6
|
.170
|
366
|
1351
|
| |
Barley
supplement |
2.70
|
3.8
|
.103
|
252
|
680
|
| |
SBM
supplement |
3.01
|
4.5
|
.135
|
358
|
1078
|
*
CSM - cotton seed meal
* SBM
- soybean meal
|
Proper supplements
fed to sheep on winter range during gestation have given as much as a
15 percent increase in lamb crop, 8 pounds more lamb per ewe, and a pound
more wool per animal. Replacement heifers, when supplemented on winter
range, gained 20 pounds more than the controls, and supplemented cows
weaned calves weighing 23 pounds more than unsupplemented cows.
Thus, it is shown that
a protein supplement, when supplied to grazing animals on dormant desert
ranges, actually increases the efficiency of energy turnover in the ecosystem,
even though protein may not be deficient in the forage according to common
recommendations.
Energy
and Productive Animal Physiology
It is generally
acknowledged that sheep are slightly more effective in converting energy
into grain and milk than cattle. There is, however, a rather marked difference
in the efficiency of energy conversion from range forage into gain of
young weaned lambs and steers compared with gain from lambs and calves
receiving milk during lactation. As shown in Table 3, about 40 percent
more gross energy is required per pound gain in sucking calves compared
with steers. By knowing the degree of utilization and the response of
the grazing animal, it is possible to calculate the energy production
of the ecosystem by the indirect method. Calculations for energy conversion
in the range ecosystem (Table 3) with respect to species of animal and
physiological function involved the following data:
|
|
450
to 500 kcal for heat increment per pound of dry matter consumed, |
|
|
33
kcal per 100 pound live weight for walking a mile, |
|
|
kcal required for steers per pound of gain is about 1900, and for
lambs on range forage about 1700, |
|
|
maintenance
was calculated by the formula 70Kcal/kg -0.75/d, and |
|
|
3280
kcal in whole milk was allowed per pound of calf gain, and 2650 kcal
in whole milk per pound of lamb gain is suggested. |
| TABLE
3. COMPARISON OF ENERGY CONVERSION BY LACTATING COWS AND GROWING STEERS
ON SPRING AND SUMMER RANGES |
| |
COW
|
STEER
|
| Animal
Unit Weight lb / individual |
900
|
650
|
| Acres
/ individual / month |
2.80
|
2.0
|
| Pounds
of oven-dry forage eaten / month |
795
|
564
|
| Gross
energy kcal / animal month |
1,570,488
|
1,114,793
|
| Digestible
energy kcal / animal month |
968,991
|
687,827
|
| Metab.
energy kcal / animal month |
772,680
|
564,780
|
| Energy
for maintenance kcal / animal month |
193,830
|
151,230
|
| Remainder
after maintenance / animal month |
578,850
|
413,550
|
| kcal
for 2 lb steer gain / day for month |
------------
|
114,000
|
| kcal
in milk produc. for 2 lb calf gain/day for 1 mo. |
196,800
|
------------
|
| kcal
remaining for heat increment and travel |
382,050
|
299,550
|
| Gross
energy kcal / lb gain |
26,174
|
18,580
|
Common Use
Even if we assume that sheep
and cattle convert plant nutrients with similar efficiency, we still have
marked differences in dietary habits between the two species on most rangeland
ecosystems. This is a result of the difference between the two in use
of plant species and varying topography. Some rangeland ecosystems are
better suited to cattle, and others are better suited to sheep.
In Table 4 it is shown that
a mountain rangeland was slightly better suited to sheep than cattle when
comparing the quantity of energy consumed. However, the two animal species
using it in common were more effective from the standpoint of both quantity
of herbage consumed and efficiency of transferring energy into meat produced
in the ecosystem.
| TABLE
4. AVERAGE CONSUMPTION OF ENERGY PER ACRE ON MOUNTAIN RANGE |
| |
UTILIZATION
|
| |
COWS
|
SHEEP
|
COMMON
|
|
Grass
500 lb / A
|
28%
|
14%
|
26%
|
|
Forbs
400 lb / A
|
6%
|
28%
|
28%
|
|
Browse
100 lb / A
|
5%
|
20%
|
18%
|
|
Pounds
consumed lb / A
|
169
|
202
|
260
|
|
Gross
energy kcal / lb
|
1916
|
1845
|
1886
|
|
Digestible
energy kcal / lb
|
924
|
702
|
856
|
|
Metabolizable
energy kcal / lb
|
748
|
555
|
685
|
|
Metabolizable
energy eaten kcal / A
|
126,412
|
112,110
|
178,100
|

|