EARTH SYSTEM: HISTORY AND NATURAL VARIABILITY - Vol. III - Grasslands and Savannas - Ford P.L.

GRASSLANDS AND SAVANNAS

Ford P.L.
USDA Forest Service Rocky Mountain Research Station, Albuquerque, New Mexico,
USA

Keywords: grassland, savanna, ecology, nutrient cycling, productivity, biodiversity,

disturbance, fire, grazing, drought, climate change

Contents

1. Introduction
2. Evolution
3. Distribution

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4. Ecology

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4.1. Climate
4.1.1. Temperature

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4.1.2. Precipitation
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4.2. Soil
4.3. Productivity
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4.4. Disturbance
4.5. Wildlife
5. Temperate Grasslands
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5.1. Tropical Grasslands and Savannas

6. Selected Grasslands and Savannas
6.1. Africa
6.1.1. The Sahel
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6.1.2. Southern Africa

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6.2. Australia
6.3. Eurasia
6.3.1. Southern Asia
6.4. North America
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6.5. South America

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7. Conclusion
Acknowledgments
Glossary
Bibliography
Biographical Sketch

Summary

Grasslands are typically defined as lands on which the existing plant cover is dominated
by grasses. Savannas are broadly defined as grasslands with scattered trees. Grasslands
and savannas together cover approximately 5% of Earth’s surface. Grasslands evolved
during the Cenozoic era, in the course of a period of cooling and drying of the global
climate. Evolution of savanna ecosystems and their constituent organisms may have
already been in progress in the Eocene and Oligocene epochs. By definition, grasslands
and savannas can be either anthropogenic or natural. Anthropogenic grasslands and

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savannas require some form of disturbance, such as cultivation, heavy grazing, burning,
or mowing to persist. Natural grasslands occur across a wide range of climatic and
geological conditions, with soil and climate being the major limiting factors to their
distribution. Natural savannas, conventionally considered tropical savannas, occur in the
seasonal wet–dry zones sandwiched between the humid equatorial zones and the arid
zones in the mid latitudes, between lat 10° and lat 30° north and south of the equator.
Since for thousands of years both savannas and grasslands around the world have been
manipulated by humans for their benefit, it can be difficult in some areas at times to
separate the influence of humans from that of nature in the formation or maintenance of
these ecosystems. Today we have the potential to use knowledge and technology to
sustain these ecosystems indefinitely and without environmental deterioration.
However, management tools and decisions must be tailored to the specific type of
ecosystem and the management objective.

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1. Introduction

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Grasslands and savannas together cover approximately 5% of Earth’s surface, and make

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up an estimated 23% of Earth’s terrestrial biomes (see Figure 1). Grasslands are
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typically defined as lands on which the existing plant cover is dominated by grasses.
Savannas are broadly defined as grasslands with scattered trees. The two biomes are
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distinct, though they commonly grade into each other. In addition, they have both
unique and shared ecological processes, structures, and biotic assemblages.
Distributions of both grasslands and savannas are regulated by climate and soils, and
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modified by disturbance (natural and/or anthropogenic). Grasslands are categorized on a

broad scale as either temperate or tropical. Savannas are conventionally considered an
exclusively tropical biome, and the terms tropical grasslands and savannas are often
used interchangeably.
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Figure 1. Natural grasslands and savannas of the world

2. Evolution

Grasslands evolved during the Cenozoic Era (66.4 million years ago to the present), in
the course of a period of cooling and drying of the global climate. During this era, the

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continents assumed their modern configuration and geographic positions, and Earth’s
flora and fauna evolved toward those of the present. The grass family (Poaceae or
Graminae) evolved early in the Cenozoic era, an era commonly referred to as the Age of
Mammals due to the rapid spread and diversification of mammals during that time. The
date of the earliest appearance of grasslands varies from region to region. In several
regions, a succession of vegetation types can be recognized in the Cenozoic fossil
record as the climate progressively dried out. For example, in central Australia during
the past 50 million years tropical rainforests gave way successively to savanna,
grassland, and finally, desert. In some places expansions of grasslands to something
approaching their modern extent occurred only during the past two million years in the
extremely cold, dry intervals, called Ice Ages. See Tertiary, Quaternary.

Evolution of savanna ecosystems and their constituent organisms may have already

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been in progress in the Eocene (38–54 million years BP) and Oligocene (26–38 million
years BP) epochs. If savannas are considered exclusively tropical, and are defined as

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possessing a more-or-less continuous grass cover, it is thought they cannot have come

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into being before the Eocene, because the first records of pollen grains of grasses from

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the tropics date from the Middle Eocene. However, low open vegetation types
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constituted by herbs other than grasses may have existed before that time. Savanna
evolution continued during Miocene (5.1–23 million years BP) and Pliocene (1.8–5.0
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million years BP) times, with extreme changes during the Quaternary period (2 million
years BP), when climates in the tropics seemed to have alternated between rainy periods
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and drier spells. During these time periods, there was possibly large-scale and repeated
expansion of savanna vegetation during dry climatic intervals. It is also possible that
present areas of savanna were considerably smaller during periods of general increases
of precipitation in ecotones between savanna and rain forest, due to the invasion of
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savannas by rain forest.

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3. Distribution

By definition, grasslands and savannas can be either manmade or natural.

Anthropogenic grasslands of cereal crops, pastures, playing fields, or other types require
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some form of disturbance, such as cultivation, heavy grazing, burning, or mowing to

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persist. Human-made grasslands exist the world over, and are central to human welfare,
as they provide a major source of food when managed as cropland or rangeland.
Anthropogenic savannas, historically maintained by fire, also provide humans with
food, pasture for grazing animals, and recreational areas, in addition to being a source of
wood products. Since for thousands of years both savannas and grasslands around the
world have been manipulated by humans for their benefit, it can be difficult in some
areas at times to separate the influence if humans from that of nature in the formation or
maintenance of these ecosystems.

Natural grasslands occur across a wide range of climatic and geological conditions, with
soil and climate being the major limiting factors to their distribution. Natural savannas,
conventionally considered tropical savannas, occur in the seasonal wet–dry zones
sandwiched between the humid equatorial zones and the arid zones in the mid latitudes:
between lat 10o and 30o north and south of the equator. In general, natural grasslands
and savannas can be considered intermediates in an environmental gradient, with forests

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at one end and deserts at the other. Forests primarily occupy environments where the
nature of soil and amount of moisture are conducive to growth and survival of tall,
dense vegetation dominated by trees. Savannas are found where moisture, soil texture,
nutrients, herbivory, and fire allow codominance of grasses and trees. Deserts are found
where moisture is so lacking that a continuous permanent vegetation cover cannot be
maintained. A dynamic balance commonly exists between grasslands and related
vegetation types (i.e., desert, forest, savanna). Changes in the severity or frequency of
disturbance events (i.e., grazing, fire, drought, flooding) can cause a change from one
vegetation type to another.

Distribution of grasslands and savannas could also conceivably be influenced by global

climate change. Many models of the atmosphere and global climate suggest that
substantial warming of the atmosphere (1.5–5.5 °C) will accompany increasing

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concentrations of greenhouse gases: carbon dioxide (CO2), nitrous oxide (N2O),
methane (CH4), and chlorofluorocarbons. Carbon dioxide, the major greenhouse gas, is

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a principal by-product of the combustion of anything containing carbon, particularly

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coal and petroleum. Methane is produced by grazing livestock and rice paddies, and is a

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by-product of the combustion of wood, natural gas, coal, and oil. Nitrous oxide comes
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from chemical fertilizers and automobile emissions, and chlorofluorocarbons are
synthetic chemicals that until recently were used for a variety of uses including
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refrigeration and air conditioning. All of these gases have been released at an
accelerating rate in recent years. The warming results from the absorption of infrared
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(heat) radiation emitted from the surface of Earth (greenhouse effect). Most models
predict that higher concentrations of CO2 and other trace gases in the atmosphere will
make Earth a warmer and more humid planet in the future, and that temperature and
precipitation patterns will change. Some scenarios predict that heat and drought would
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become more prevalent in much of the midlatitudes, and milder temperatures would
prevail in higher latitudes. Some arid lands might receive more rainfall, ice caps could
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melt, and global sea levels would rise. If this occurs, it will most likely influence the
distributions of many terrestrial plants and animals. See Greenhouse Gasses.
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4. Ecology
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4.1. Climate

The ecological components of grasslands and savannas vary according to climate, soil,
and biotic factors. Climate provides a source of energy and water for ecosystems. The
sun is the main source of energy for Earth’s biota-energy flows into the biological world
from the sun, which supplies heat and light by solar radiation. Solar energy warms the
earth’s surface making it habitable, in addition to being captured by green plants and
converted into chemical forms of energy that power the growth, maintenance, and
reproduction of most living things (see Figure 2).

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Figure 2. Ecology of grasslands and savannas
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The way Earth revolves about the sun and rotates on its axis influences the distribution
of solar radiation at its surface, and the quantity of heat absorbed. As a result of Earth’s
curvature, the low latitudes or tropics receive more solar radiation than do middle and
higher latitudes. The polar regions receive only about 40% of the solar energy received
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at the equator. To balance this unequal distribution of energy, there is a large-scale

transfer of heat poleward, through atmospheric and oceanic circulation. These
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circulations moderate the buildup of heat in equatorial regions and the lack of heat in
polar regions, thereby making both those latitudinal zones more habitable than they
would be otherwise. The frictional effects of the rotating Earth’s surface on airflow
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cause these circulations to be relatively complex. Nevertheless, the solar energy and
atmospheric and oceanic circulations are distributed over the planet in an organized
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fashion. These factors, in turn, produce recognizable world patterns of temperature and
precipitation, the two most important climatic elements. See Atmosphere and Climate.

4.1.1. Temperature

Temperature is a measure of the amount of heat energy within a substance

(internationally measured in °C). The temperature of air and soil is important to biotic
distribution patterns. Fewer species of both plants and animals can survive in cold
regions than in areas of more moderate temperatures. Plants, in particular, have a
limited tolerance for low temperatures because they experience tissue damage and other
physical disruption when their cellular water freezes. Many animals have evolved
protection against low temperatures, and are able to avoid cold by moving around to
seek warmth or shelter. Variations in temperature on and within the surface of Earth
have a variety of causes, including the effects of latitude and altitude; continental,
seasonal, and diurnal effects; microclimatic effects; and, in soil and water, the effects of

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depth.

The angle at which Earth is tilted relative to the sun changes over an annual cycle, and
this drives some of the main temperature differentials on Earth’s surface (recall the
angle of incoming radiant energy relative to the planet's surface affects the quantity of
heat absorbed). This differential heating produces the seasons. The seasons are also
characterized by different lengths of day and night. In general, heating is most intense
when the sun is directly overhead, so that incoming solar radiation strikes perpendicular
to Earth’s surface. Therefore, the higher latitudes are cooler than the tropics because the
same quantity of solar radiation is dispersed over a greater surface area and passes
through a thicker layer of filtering atmosphere. Seasonality of climate increases with
increasing latitude due to variations in sun angles and length of day. For example,
tropical latitudes are always warm/hot because they always have high sun angles and

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consistent day lengths that are close to 12 h long. Conversely, the polar regions are
consistently cold because they always have low sun angles in spite of 24-h days in

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summer. Only the areas poleward of 66.5° have 24-h days and nights. See Insolation.

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4.1.2. Precipitation
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Precipitation is defined as all forms of condensation of atmospheric water, including
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rainfall, snowfall, and icefall. The most important geographic aspect of atmospheric
moisture is the spatial distribution of precipitation. The broad-scale pattern is based on
latitude, but many other factors are involved and the overall pattern is complex. The
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amount of precipitation on any part of Earth’s surface is determined by the nature of the
air masses involved and the degree to which the air is uplifted. The humidity,
temperature, and stability of the air masses are mostly dependent on where the air
originates (over land or water, in high or low latitudes) and on the trajectory it has
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followed. The amount of uplifting is determined largely by zonal pressure patterns,

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topographic barriers, and storms and other atmospheric disturbances.

The most conspicuous feature of the worldwide annual precipitation pattern is that
tropical latitudes contain most of the wettest areas. Dry lands are most prominent on the
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western sides of continents in subtropical latitudes (centered at lat 25° or 30°). Dry
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zones are most extensive in North Africa and Australia, primarily because the blocking
effects of landmasses or highlands to the east prevent moisture from coming in from
that direction. Dry regions in the midlatitudes are most extensive in central and
southwestern Asia, but they also occur in western North America and southeastern
South America. In each case, the dryness is due to lack of access to moist air masses. In
the high latitudes, there is not much precipitation anywhere. Areas of open water are
scarce and cold, so little opportunity exists for moisture to evaporate into the air. As a
result, polar air masses have low absolute humidity and precipitation is slight. These
regions are referred to as cold deserts. In addition, because continental coastal regions
are closer to moisture sources than interior regions, they usually receive more
precipitation.

Precipitation levels vary seasonally. Over most of the globe, the amount of precipitation
received in summer is considerably different from the amount received in winter. This
variation is most pronounced over continental interiors, where most of the year’s

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precipitation occurs during summer months. Coastal areas often have a more balanced
seasonal precipitation regime, due to their nearness to moisture sources. In general,
summer is the time of maximum precipitation over most of the world. Northern
hemisphere regions experience heaviest rainfall in July, and southern hemisphere
locations receive most precipitation in January. The only important exceptions to this
generalization occur in narrow zones along western coasts between lat 35° and 60° in
the United States, South America, New Zealand, and southernmost Australia. The most
conspicuous variation in seasonal precipitation is found in monsoon regions (principally
southern and eastern Asia, northern Australia, and West Africa), where summer tends to
be very wet and winter is generally dry. See Precipitation.

4.2. Soil

Soil, the outer, highly weathered layer of the earth’s crust, is the substrate in which

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nearly all plants grow. Soil’s main constituents are (a) mineral matter, which makes up

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the bulk of most soils; (b) organic matter, in the form of living or dead organisms; and
(c) pore spaces, filled with a mixture of water and air, which make up about half the

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total volume of most soils. Millions of years of weathering of rocks by physical and
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biological processes have produced the particles from which soils are formed. Physical
processes include freezing and thawing, and water and wind erosion. Biological
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processes include microorganism effects on the weathering of rock (e.g., lichens);
addition of organic material through decomposition of plants, animals, and microbes;
and plant root and microbe alteration of the chemical composition of soil. Plant roots
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also aerate, mix, and drain the soil (as do burrowing animals), in addition to providing
nutrients from the soil to the growing plant (see Figure 2).

Temperature and moisture are the climatic variables of greatest significance to soil
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formation. In general, both the chemical and biological processes in soil are usually
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accelerated by high temperatures and abundant moisture, and are slowed by low
temperatures and lack of moisture. The effectiveness of soil as a growth medium is
based largely on the availability of nutrients. In general, tropical soils are prone to
nutrition depletion because, where annual precipitation exceeds annual
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evapotranspiration, water movement is predominantly downward in the soil, and

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leaching is a pronounced process. In addition, temperatures in the tropics are relatively

high throughout the year. High temperatures coupled with dense rainfall cause organic
matter in the tropics to decompose rapidly, allowing only low levels of humus (organic
matter) to accumulate. Soils rich in humus are generally more productive than those
poor in humus. However, plant nutrients are not completely removed by leaching, as
natural vegetation can also quickly absorb many nutrients in solution. In addition, if
vegetation is relatively undisturbed, nutrients will cycle rapidly, and the soil will not be
totally impoverished by the speed of mineral decomposition and leaching. In temperate
regions, where annual precipitation is generally less than potential evapotranspiration,
principal soil moisture movement is upward, and leaching is limited. Materials that
would be carried downward in other moisture regimes, instead become concentrated in
the soil. Therefore, temperate soils generally have high productivity due to high levels
of humus. See Soil.

Humus contributes to the fertility or productivity of soil through its positive effects on

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the chemical, physical, and biological properties of the soil. It has a nutritional function
in that it serves as a reservoir of nitrogen, phosphorous, and sulfur for plant growth; a
physical function in that it promotes a good soil structure as it increases the ability of
soil to resist erosion and enables the soil to hold more water (when humus is lost, soils
tend to become hard and compact); and a biological function in that it serves as a source
of carbon and energy for soil organisms. Black or dark brown soils usually indicate a
considerable humus content; the blacker the soil, the more humus it contains. Tropical
and subtropical soils are commonly reddish and yellowish in color owing to low organic
content from leaching. See Macronutients and Micronutrients.

4.3. Productivity

Plants, algae, and some bacteria have the ability to harvest sunlight’s energy, along with
water and carbon dioxide, to make organic compounds and release oxygen, a process

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called photosynthesis (see Figure 2). It has been estimated that ~1–5% of the solar

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energy that falls on a plant is converted to organic material. Primary productivity is a
measure of the amount of sunlight converted to plant growth, and is considered a basic

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indicator of ecosystem functioning. Primary productivity includes photosynthesis and
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chemosynthesis, the chemical oxidation of simple inorganic compounds like
ammonium, nitrite, and sulfide. Secondary productivity refers to the rate that consumers
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store energy.

Productivity is not evenly spread across the earth, because solar energy can be
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efficiently captured only when water and nutrients are available, and when temperatures
are in the range suitable for plant growth. There is a general trend (with much variation)
of increasing productivity with decreasing latitude. However, this trend does not hold in
areas of land that receive abundant solar radiation, but lack adequate water (e.g., the
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continental interior of Australia). The amount of rainfall in a region is closely correlated

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with its productivity. For example, in North America the amount of annual rainfall
influences the height of grassland vegetation, with taller grasses generally occurring in
wetter regions (i.e., tallgrass prairie) and shortgrass occurring in drier regions (i.e.,
shortgrass steppe; see Figure 2). In addition, small differences in topography, and
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microclimatic conditions can result in large differences in productivity.

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Primary productivity is a term used to describe the amount of organic matter produced
from solar energy in a given area during a given period of time. While sunlight, carbon
dioxide, water, and soil nutrients are the resources required for primary production,
temperature has a strong influence on the rate of photosynthesis. Therefore, the length
of the growing season is a function of both precipitation and temperature variation.
Growing seasons in hot climates vary between 120 and 190 days each year. In more
temperate climates, the growing season commences when the average temperature
reaches about 5 °C to 10 °C and extends from 100 d to more than 165 d. During the
course of a year, the productivity of a community may be limited by many factors,
including drought, low rates of nutrient cycling (i.e., carbon and nitrogen cycles), or by
grazing animals reducing leaf area available for capturing sunlight for photosynthesis.
See Productivity.

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Bibliography

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Biographical Sketch

Paulette Ford is a Research Ecologist with the USDA Forest Service Rocky Mountain Research Station
in Albuquerque, NM. She holds a B.S. (1989) in Biology and Psychology and an M.S. (1992) in Biology
from the University of New Mexico, in addition to a Ph.D. (2000) in Renewable Natural Resources from
the University of Arizona. She has worked extensively in Latin America and the American Southwest on
research ranging from the systematics of parasites and amphibians, to small mammal and amphibian

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community structure in deserts, grasslands, and tropical deciduous forests. Her current research interests

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include the role of disturbance in structuring grassland and desert communities, and scale and ecosystem
resilience. Results of her research have been published in numerous scientific journals and both national

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and international symposium proceedings. Paulette’s ongoing research in northeastern New Mexico uses

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a long-term (18-y) experimental framework to analyze the effects of season and frequency of fire on
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shortgrass steppe.
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