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Biological Soil Crusts

What They Are and A Few Interesting Facts

 Ever walked across the rangeland and heard a little crunch under your feet? If so,
you’ve just discovered the biological soil crust. It’s a complex mosaic of living
organisms—algae, cyanobacteria (blue-green algae), bacteria, lichens, mosses,
liverworts, and fungi—that grow on or just below the soil surface. Biological soil
crusts are common worldwide in arid and semi-arid shrublands, grasslands and
woodlands.

 Biological soil crusts are highly variable in appearance. Which organisms

dominate the crust is determined by several factors, including soil chemical and
physical characteristics, and weather patterns. Biological soil crusts are
distinguishable from bare soil by a bumpy appearance, forming sort of a mini-
landscape on the soil surface complete with hills and valleys. They tend to be
dark in color, especially when dry. However add a little moisture and you may be
surrounded by vivid greens, yellows, and oranges as the mosses and lichens
come out of dormancy and spring to life.

 Biological soil crusts are known by lots of names:

 
o Microbiotic
o Cryptogamic
o Cryptobiotic
o Microphytic
o Microfloral

Don’t get confused. The names all refer to the same thing

What They Are Not

Biological soil crusts are not physical or chemical crusts. Physical and chemical crusts
are formed by different processes and tend to form a hard, impermeable layer on the
soil surface. They also lack the biological characteristics that make biological soil crusts
unique.

Why They Are Important

 Biological soil crusts are, literally, a carpet of photosynthetic life. Remember when
we talked about a whole living community being right under your foot? That is one
way to describe a healthy biological soil crust. In addition, they are also habitat for
fauna that, in turn, contribute to the development of the crust.
  They stabilize the soil. Some of the organisms secrete sticky substances
(polysaccharides), which hold soil particles together.

 Biological soil crusts make the soil more fertile. Most of the organisms associated
with the biological soil crust are photosynthetic, particularly during cold, wet
seasons when most plants are dormant. This means that the biological soil crust
increases the length of the time during which organic carbon is added to topsoil.
In addition, some cyanobacteria and lichens fix atmospheric nitrogen, even during
the winter. Biological soil crusts can make other nutrients more available for use
by grasses, forbs, and shrubs, as nutrients adhere to the aforementioned sticky
substances, and are prevented from leaching

 Biological soil crusts may help the soil to retain more moisture. The extent to
which this function occurs is highly dependent on both the composition of the
crust and soil characteristics.

 The nature of the crust itself can keep unwanted plants, such as exotic weeds,
out. Some of these exotic weeds—which include cheatgrass and medusahead
wildrye—can invade and dominate rangeland communities within relatively short
time periods. Native plants, which evolved with biological soil crusts, may have
developed mechanisms to allow seeds to penetrate the crust (e.g. small size, or
structures that "drill" them into the crust). Seeds of some exotic species are quite
large and may not have a way of getting through the crust.

 Because of their functions in rangeland systems, biological soil crusts have been
adopted by scientists and land management professionals in the U.S., Australia,
and South Africa as a visible indicator of rangeland health.

Learn More!

Belnap, Jayne, Kaltenecker, Julie Hilty, Rosentreter, Roger, Williams, John, Leonard,
Steve, and Eldridge, David. 2001. Biological Soil Crusts: Ecology and Management.
USDI, Bureau of Land Mangement. Technical Reference 1730-2. Denver, CO. 110pp.

Harper, K. T., and J. R. Marble. 1988. A role for nonvascular plants in management
of arid and semiarid rangelands in Vegetational Science Applications for Rangeland
Analysis and Management. P. T. Tueller (ed.). Kluwer Academic Publishers. Dordrecht,
Netherlands.

Johansen, J. 1993. Cryptogamic crusts of semi-arid and arid lands of North

America. Journal of Phycology 29: 140-147.

Johnston, Roxanna. 1997. Introduction to microbiotic crusts. U.S. Department of

Agriculture, Natural Resources Conservation Service, Soil Quality Institute and Grazing
Lands Technology Institute. This is a non-technical information bulletin gear toward the
public land user.
Ladyman, J.A.R. and E. Muldavin. 1996. Terrestrial cryptogams of the Pinyon-
Juniper woodlands in the Southwestern United States: a review. USDA, USFS
General Technical Report RM-GTR-280. Fort Collins, CO. 33 pp.

The Great Basin Naturalist Volume 53 No. 1, March 1993 (Proceedings of the
Symposium on Soil Crust Communities). Contains multiple technical papers on
biological soil crusts.

West, N.E. 1990. Structure and function of soil microphytic crusts in wildland
ecosystems of arid and semi arid regions. Advances in Ecological Research 20: 179-
223.

Examples of
Biological Crusts

Moss-dominated Lichen-dominated Biological crust dominated by

cyanobacteria and lichens
biological crust biological crust

Soil particles
Physical soil crust being held together by
filamentous cyanobacteria
Photo credit:
Jayne Belnap, USGS, BRD

Healthy
sagebrush-brunchgrass
community
 Mycorrhizal Fungi
 

What They Are and A Few Interesting Facts

 Mycorrhizal fungi colonize the roots of many plants. Mycorrhizal fungi don’t harm
the plant; on the contrary, they develop a "symbiotic" relationship that helps the
plant be more efficient at obtaining nutrients and water. In return, the plant
provides energy to the fungus in the form of sugars.

 Here’s how that symbiotic relationship works. The fungus is actually a network of
filaments that grow in and around the plant root cells, forming a mass that
extends considerably beyond the plant’s root system. This essentially extends the
plant’s reach to water and nutrients, allowing it to utilize more of the soil’s
resources.

 There are two main categories of mycorrhizae common to western rangelands in

the United States.
o Vesicular-arbuscular mycorrhizae or VAM. VAM is a type of
endomycorrhizae (endo = inside), and is the most widespread of the
mycorrhizae. These fungi actually reside inside the cells of the plant root.
They’re typically found associated with most grasses, forbs, shrubs, and a
few trees such as juniper. They are generalists, have only a few species,
and are slow to disperse.
o Ectomycorrhizae (ecto = outside) grow around the root and between the
root cells, but unlike VAM, the fungus doesn’t actually penetrate the root
cells. The fungus also forms a considerable mass in the soil surrounding
the plant roots. The fruiting, or reproductive bodies, of these fungi are
sometimes visible as something we all recognize— mushrooms!
Ectomycorrhizae are commonly associated with forest trees of temperate
regions. On rangelands, they are found in riparian areas (the places next
to water), open woodlands, and shrub oak communities. They are host-
specific, have many species, and can disperse far and quickly.

 Not all fungi are mycorrhizal. There are also fungi that help decompose the
organic matter in litter and soil. However, they play a lesser role than bacteria in
this important process in semi-arid and arid rangeland soils.

 Why They Are Important

 Some plants are "mycorrhizal-obligate," meaning that they can’t survive to

maturity without their fungal associate. Important mycorrhizal-obligate plants in
western North America are sagebrush, bitterbrush, and some native
bunchgrasses.
  Mycorrhizae are particularly important in assisting the host plant with the uptake
of phosphorus and nitrogen, two nutrients vital to plant growth.

 Mycorrhizae actually increase the surface area associated with the plant root,
which allows the plant to reach nutrients and water that might not be available
otherwise. Put simply, mycorrhizae extends the plant’s reach, allowing it to get to
more of what it needs to survive. That makes the plant stronger, especially during
drought periods. Stronger individuals means that the community is more resilient
to disturbance. Some mycorrhizae may even protect their host plant against
unwanted pathogens.

Learn More!

Allen, M.F. 1991. The ecology of mycorrhizae. Cambridge Univ. Press. New York. 184
pp.

Ingham, Elaine R. 1998. The soil biology primer, soil fungi. USDA, Natural Resources
Conservation Service, Soil Quality Institute.

Read, D.J. 1991. Mycorrhiza in ecosystems. Experientia 47: 376-391.

Wicklow-Howard, M. 1994. Mycorrhizal ecology of shrub-steppe habitat in

Proceedings-Ecology and Management of Annual Rangelands. USDA, Forest Service.
Inter-mountain Research Station. General Technical Report INT-GTR-313. pp. 207-210.

Wicklow-Howard, M. 1998. The role of mycorrhizal fungi in rangelands. pp. 23-25 in

Rosentreter, R. and A. DeBolt, editors. The Ellen Trueblood Symposium. Technical
Bulletin No. 98-1, Bureau of Land Management, Boise, Idaho

Soil Bacteria and Actinomycetes

 

What They Are and A Few Interesting Facts

Bacteria are minuscule, one-celled organisms that can only be seen with a powerful light
(1000X) or electron microscope (we're talking TINY). They can be so numerous that a
pinch of soil can contain millions of organisms. Bacteria are tough—they occur
everywhere on earth and have even been found over a mile down into the core of the
earth.

Bacteria can be classified into five functional groups. Autotrophic (literally, self-feeding)
bacteria are photosynthetic. They are the primary producers. Decomposers consume
soil organic matter, plant litter, and simple carbon compounds, releasing the nutrients in
these substances for use by living plants. Mutualists, such as nitrogen-fixing bacteria,
form associations with plants and help them absorb nutrients. Pathogens are the bad
guys— they cause disease in plants. The last group, the chemolithotrophs (literally,
chemical and rock-eating) obtain energy from minerals rather than from carbon
compounds.

Bacteria are common throughout the soil, but tend to be most abundant in or adjacent to
plant roots, an important food source.

Actinomycetes are a broad group of bacteria that form thread-like filaments in the soil.
They are responsible for the distinctive scent of freshly exposed, moist soil.

Why They Are Important

Bacteria are important in the carbon cycle. They contribute carbon to the system by
fixation (photosynthesis) and decomposition. Bacteria are important decomposers in
grassland environments. Actinomycetes are particularly effective at breaking down
tough substances like cellulose (which makes up the cell walls of plants) and chitin
(which makes up the cell walls of fungi) even under harsh conditions, such as high soil
pH. Some management activities, particularly those that change nutrient levels in the
soil, can shift the dominance of decomposers from bacterial to fungal. When one group
becomes dominant where it shouldn't be, there is also a shift in the rest of the system.
The shift from bacterial to fungal dominance, for instance, can enhance the conditions
favoring weed invasions on rangelands.

Bacteria are particularly important in nitrogen cycling. Free-living bacteria fix

atmospheric nitrogen, adding it to the soil nitrogen pool. Other nitrogen-fixing bacteria
form associations with the roots of leguminous plants such as lupine, clover, alfalfa, and
milkvetches. Actinomycetes form associations with some non-leguminous plants
(important species are bitterbrush, mountain mahogany, cliffrose, and ceanothus) and
fix nitrogen, which is then available to both the host and other plants in the near vicinity.
Some soil nitrogen is unusable by plants until bacteria convert it to forms that can be
easily assimilated.

Some bacteria exude a sticky substance that helps bind soil particles into small
aggregates. So despite their small size, they help improve water infiltration, water-
holding capacity, soil stability, and aeration.

Wait! Aren't there also "bad" bacteria? Yes, there are, but some soil bacteria suppress
root-disease in plants by competing with pathenogenic organisms. The key is in
maintaining a healthy system so that the good guys can do their work.

Bacteria are becoming increasingly important in bioremediation, meaning that we

(people) can use bacteria to help us clean up our messes. Bacteria are capable of
filtering and degrading a large variety of human-made pollutants in the soil and
groundwater so that they are no longer toxic. The list of materials they can detoxify
includes herbicides, heavy metals, and petroleum products.
Learn More!

Ingham, Elaine. 1998. The soil biology primer, soil bacteria. USDA, Natural
Resources Conservation Service, Soil Quality Institute.

Kennedy, A.C. and R.I Papendick. 1995. Microbial characteristics of soil quality.
Journal of Soil and Water Conservation 50 (3) 243-248.

Vollmer, A.T., A. Au, and S.A. Bamberg. 1977. Observations on the distribution of
microorganisms in desert soil. Great Basin Naturalist 37 (1) 81-86.

Soil Protozoa
What They Are and A Few Interesting Facts

Protozoa are tiny single-celled animals that mainly feed on bacteria (think of them as
little grazers), although some eat other protozoa and organic matter. While protozoa are
many times larger than bacteria, you still need a microscope to see them. A pinch of soil
can contain thousands.

Protozoa are classified into three groups based on structural peculiarities. Ciliates are
the biggest and use many hair-like cilia to scoot through soil and water. Amoebae are
small (relatively), amorphous, and use temporary feet to move around. Flagellates are
the smallest and swim using a few whip-like flagella. All protozoa need water to move
through soil, however, they only need a thin film surrounding the soil particles to get
around. Believe it or not, they can be quite active even in very parched desert soils.

Protozoa are found in soils everywhere: even in very dry desert. However, they are most
abundant near plant roots, because that's where both bacteria and organic matter (i.e.,
food) are concentrated in the soil.

Why They Are Important

Protozoa play an important role in the structure of the soil food web. Protozoa eat
bacteria and release nitrogen and other nutrients in their waste. Since protozoa are
concentrated near plant roots, the plant can benefit from this supply of nutrients.
Protozoa can stimulate the rate of decomposition by maximizing bacterial activity.

Protozoa are in turn consumed by nematodes and microarthropods. Remember what we

said about missing parts? Think of what would happen if there were no protozoa.

Okay, so they aren't all good guys. Some protozoa attack roots and cause disease in
rangeland plants. However, other protozoa feed on root pathogens, thus reducing plant
disease.

Learn More!
Clarholm, Marianne. 1985. Interactions of bacteria, protozoa and plants leading to
mineralization of soil nitrogen. Soil Biol. Biochem. 17(2)181-187.

Ingham, Elaine R. 1998. The soil biology primer, soil protozoa. USDA, Natural
Resources Conservation Service, Soil Quality Institute.

Whitford, Walter G. 1996. The importance of the biodiversity of soil biota in arid
ecosystems. Biodiversity and Conservation 5: 185-195.

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Ciliate Protozoa
Photo credit:
Cells Alive

Naked amoeba soil protozoa eat thousands of bacteria daily.

Photo credit:
Wilhelm Foissner, Institute of Zoology, University of Salzburg

 
Testate amoeba soil protozoa make a protective shell of silica, soil particles or calcium.
Photo credit:
Wilhelm Foissner, Institute of Zoology, University of Salzburg

Flagellate soil protozoa.

Photo credit:
Wilhelm Foissner, Institute of Zoology, University of Salzburg

 
 Ciliate soil protozoa eat tens of thousands of bacteria daily.
Photo credit:
Wilhelm Foissner, Institute of Zoology, University of Salzburg

Soil Nematodes

What They Are and A Few Interesting Facts

Nematodes are tiny roundworms that are common in soils everywhere, from the freezing
Arctic to dry, hot deserts. They are particularly abundant in grassland ecosystems. To
give you an idea of exactly how common nematodes are, consider this: one cubic foot of
soil can contain millions.

Nematodes can be most easily classified according to their feeding habits. Some graze
on bacteria and fungi. Some like plant roots; others prey on other tiny animals. Some
aren't fussy at all and will eat any of the above mentioned food items.

Nematodes can't move through the soil unless a film of moisture surrounds the soil
particles. Under hot, dry conditions, nematodes can become dormant, allowing them to
survive long periods of drought. As soon as water becomes available, they quickly
spring back to life.

Why They Are Important

Among the thousands of species that have been identified, many are considered
beneficial because they boost the nutritional status of the soil. Nematodes feed on
decaying plant material, along with organisms that assist in the decomposition of organic
matter (bacteria and fungi). This helps disperse both the organic matter and the
decomposers in the soil. Increased organic matter concentration and decomposition
boost nitrogen and phosphorus levels.
Because some nematodes prey on other animals, they can be useful for control of pest
insects. Nematodes are also being investigated for their potential as biological controls
for noxious weeds.

Nematodes aren't all good guys. Some damage the roots of domestic crops, costing
U.S. farmers an estimated $8 billion a year. Nematode infestations can be identified by
yellowing, stunted plants that grow in sparse stands. Research is underway to develop
plants that can resist nematode predation.

Learn More!

Freckman, Diana W., ed. 1982. Nematodes in Soil Ecosystems. Austin, Tx., University
of Texas Press. 206 pp.

Freckman, Diana W. 1988. Bacterivorous nematodes and organic-matter

decomposition. Agric. Ecosystems Environ., 24: 195-217.

Ingham, Elaine. 1998. The soil biology primer, soil nematodes. USDA, Natural
Resources Conservation Service, Soil Quality Institute.

Zak, John C. and Diana W. Freckman. 1991. Soil communities in deserts:

microarthopods and nematodes. in Polis, Gary A., ed. The Ecology of Desert
Communities. Desert Ecology Series. pp. 55-88.

oil Arthropods
What They Are and A Few Interesting Facts

 Soil arthropods are invertebrates, meaning they don't have a spinal column. Their
legs are jointed. If you saw one, you'd probably call them something very
scientific, such as "bugs."

 Soil arthropods can be microscopic or quite large (i.e., large enough to cause a
moment of panic if you find one crawling up your pant leg).

 There are lots of different arthropods:

o insects, such as ants and springtails
o crustaceans, such as sowbugs
o arachnids, such as spiders and mites
o myriapods, such as centipedes and millipedes
o scorpions
 Arthropods perform many different functions in the soil community. Some are
shredders, others predators. Some arthropods eat plants, while others feed
strictly on fungus.

Why They Are Important

  They stir up and churn the soil, mixing in air (aerating), which is needed by other
organisms in the soil community.

 They shred organic matter into small pieces, assisting other soil organisms in the
decomposition process.

 They regulate populations of other soil organisms, like protozoa, which help
maintain a healthy soil food web and control disease-causing organisms. In turn,
soil arthropods make a nice dinner for larger animals such as burrowing
mammals, birds, and lizards.

 They help distribute beneficial microbes in the soil.

 Through consumption, digestion, and excretion of soil organic matter, soil

arthropods help improve soil structure and change nutrients into forms available
to plants.

Learn More!

Moldenke, Andrew R. 1998. The soil biology primer, soil arthropods. USDA, Natural
Resources Conservation Service, Soil Quality Institute.

Moldenke, Andy. 1993. Denizens of the soil: small, but critical. Natural Resources
News-Special Edition-August, 1993.

Nash, M. H. and W.G. Whitford. 1995. Subterranean termites: regulators of soil

organic matter in the Chihuahuan desert. Biolog. Fertil. Soils 19: 15-18.

Whitford, Walter G. 1991. Subterranean termites and long-term productivity of

desert rangelands. Sociobiology 19: (1 ) 235-243.

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Microscopic view of Springtail

Photo credit:  SSSA

Male Ologamasid mite

Photo credit:  Dr. David Evans Walter, Department of Zoology & Entomology, The University of
Queensland, Australia

Basket mite
Photo credit:  Dr. David Evans Walter, Department of Zoology & Entomology, The University of
Queensland, Australia

 
 Ant lion waiting for prey
Photo credit:  Bill Chissoe, SR Noble Electron Microscopy Lab of the Univerity of Oklahoma

Harvester ant mound

Photo credit: BLM

Harvester ants are granivores (they eat seeds)

Photo credit: Jim Munger,
Boise State University

 
  

Burrowing Mammals
What They Are and A Few Interesting Facts

 Burrowing mammals dig burrows and tunnels in rangeland that can be quite
extensive. These creatures can be as large as a badger or as small as a shrew.
They include ground squirrels, pocket gophers, prairie dogs, kangaroo rats,
kangaroo mice, pocket mice, and many other mammals.

 Burrowing mammals are very common on semi-arid and arid rangelands. Their
underground pathways and dwellings provide protection from predators and
weather extremes in open landscapes.

 Some burrowing mammals, such as badgers, dig many burrows that are not
maintained. Others, such as prairie dogs, build vast burrow and tunnel systems
that can be occupied and maintained for hundreds of years.
Why They Are Important

 Their digging mixes subsurface materials with surface soils, litter, and feces. This
helps fertilize the soil and buries carbon, which benefits many plants and soil
microorganisms.

 Their burrows and tunnels allow water from high intensity storms to rapidly
infiltrate into the soil instead of running off. Burrows carry oxygen deep into the
soil, helping to aerate the soil around plant roots.

 Some mammals, such as kangaroo rats, kangaroo mice, and pocket mice, bury
seeds in caches that serve as a valuable seed source for plant establishment.
The burying of organic matter with the seed provides a supply of nutrients for
seedling survival.

 Small mammals, including shrews and moles, consume and help control soil
arthropod populations.

 Their burrowing activities help transport mycorrhizal and other fungus spores.

 They contribute to the faunal diversity of the landscape. Their burrows provide
homes for burrowing owls, snakes, lizards, and other small mammals. In
Montana, up to 163 different animal species live in prairie dog burrows.

Learn More!

Butler, David. 1995. Zoogeomorphology: Animals as geomorphic agents.

Cambridge Univ. Press. New York.

Parmenter and VanDevender, Thomas R. 1995. Vertebrates in the desert grassland.

pp. 194-223 in The Desert Grassland. McClaran, M.P. and T.R. VanDevender, eds. The
University of Arizona Press. Tucson.

Steinberger, Y. and W.G. Whitford. 1983. The contribution of rodents to

decomposition processes in a desert ecosystem. Journal of Arid Environments (6):
177-181.

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