SOIL BIOTA, SOIL

SYSTEMS, AND

PROCESSES

David C. Coleman
University of Georgia

I. Soils as Components of Ecosystems tivity in terrestrial ecosystems. See the legacy concept
II. Soils as Organizing Centers in Ecosystems in forest ecology.
III. Major Soil Processes
IV. Biodiversity in Soils
V. Conclusions
SOIL BIODIVERSITY is an intriguing, largely unap-
preciated facet of global biodiversity. There are many
GLOSSARY phyk, even "domains," widiin soils, which are largely
unseen, making use of the uniquely diverse physico-
domains The major divisions of the biota on earth, chemical complexity of soils, which is an intersection
namely: Bacteria, Archaea, and the Eucarya. of mineral, organic, aquatic, and aerial habitats. Organ-
ecosystem engineers The concept whereby members isms have evolved in soils literally since pre-Cambrian
of the macrofauna (e.g., termites and earthworms) times (more than 600 million years ago). They are still
are actually moving parts of the soil volume for their largely undescribed, and this is particularly true for the
own uses (e.g., making macropores, which permit prokaryotes, which have awaited die development of
flow of large amounts of water rapidly through the new techniques to characterize diem. By Unking several
soil). organismal groups to major processes in global bio-
immobilization The process wherein nutrients are geochemistry, it is proving possible to appreciate die
taken up or immobilized in litter and other organic wide array and diverse nature of soil organism functions
detritus until later .(usually weeks to months) in the in the biosphere.
decomposition process.
microfauna Animals that have high turnover rates, and
live in water films in soils. Small mesofauna, such I. SOILS AS COMPONENTS
as nematodes, are also water-film dwellers.
OF ECOSYSTEMS •
larger Mesofauna and macrofauna live in pores in
portions of soil profiles.
A. Soil-Forming Factors
mineralization The availability of inorganic nutrients
in the decomposition of organic detritus, occurring Soils are an intriguing, relatively diin (often <1 m.
after die immobilization phase; see above. depth) zone of physical-chemical and biological weadi-
organizing centers Soils are centers of history and ac- ering of the earths land surface. Soils are formed by

Encyclopedia of Biodiversity, Volume 5

Copyright © 2001 by Academic Press. All rights of reproduction in any form reserved. 305
306 SOIL B I O T A , SOIL SYSTEMS, AND PROCESSES

an array of factors, namely climate, organisms, parent due to the nature of the sandy surface material. As
material, the extent of slope, and aspect (relief) op- noted in the soil-forming factors diagram (Fig. 1), the
erating over time (Fig. 1). These factors affect major array of biota—namely microbiota, vegetation, and
ecosystem processes, such as primary production, de- consumers (herbivores, carnivores, detritivores)—is in-
composition, and nutrient cycling, which lead to the fluenced by soil processes and in turn has an impact
development of ecosystem properties unique to that on the soil system.
soil type, as a result of its previous history. For example,
a deep loess soil in Iowa, with a very fertile and deep B. Poly-Phasic Nature of Soils, Influence
surface or "A" horizon, containing considerable
on the Biota
amounts of organic matter, will be very different from
an "A" horizon developed in the Nebraska sandhills, Soils are perhaps the ultimate in interface media, located
with much greater porosity and lower water retention at the intersection of four principal entities: the atmo-

Controlling factors
Parent material Climate Vegetation Relief
•Man-

Rangeland Cropland Forests

Grazing Cultivation Seeding and planting

Species Fallow Site preparation
Nutrient input Crop selection Watershed
Rre Residue management management
Nutrient imputs Fire
Water management Harvest
Fire
Harvest (removal)
Processes
Energy inputs and transformations
Radiation
Preliminary production
Decomposition
Nutrient cycling
Immobilization
Mineralization
Weathering
Translocate
<D Transport
P Erosion
Gaseous
Leaching

Development of ecosystem properties

Vegetation
Consumers
Soil
Base status
Texture
Organic matter
Phosphorus
Sulfur
Nitrogen
Salinity
FIGURE 1 Soil-forming factors and processes, and the interaction over time. From Coleman et
d. (1983) and Coleman and Crossley (1996).
 SOIL BIOTA, SOIL SYSTEMS, AND PROCESSES 307

decomposition in the soil and litter horizons and trans-

ported to the photosynthetic layers of the ecosystem.
This is particularly true for less heavily managed, near-
natural ecosystems, many of which occur on soils of
relatively poor nutrient status. In these systems, mycor-
rhizas are often obligate partners in the obtaining of
adequate nutrients for the growing plants. Mycorrhizas
are known to be efficient at extracting nutrients from
both mineral and organic sources, enabling plants to
thrive in habitats that are considered poor in nutrients.
We need to be aware of these and other mutualistic
associations between microorganisms and roots, such
as rhizobia and actinorrhizas, various root-associated
symbiotic bacteria that facilitate the nitrogen fixation
process on which the entire ecosystem often depends.
These associations have arisen in soils over evolutionary
FIGURE 2 The pedosphere showing interactions of abiotic and biotic time and are key to an understanding of ecosystem
entities in the soil matrix. From Fitzpatrick (1984) and Coleman and
function.
Crossley (1996).

III. MAJOR SOIL PROCESSES

sphere, biosphere, lithosphere, and pedosphere (Fig.
2). Soils provide a wide range and variety of microhabi- A. Decomposition: Immobilization
tats, thus accommodating a very diverse biota. The mi- and Mineralization
crobes (bacteria and fungi) are found in numerous mi-
crosites, well-aerated or not; bacteria may thus respire A very large proportion (greater than 90%) of the terres-
either aerobically or anaerobically. There is an enor- trial net primary production is returned to the soil as
mous amount of surface area (hundreds of nr per gram dead organic litter. This litter, consisting of leaves,
of soil) on the soil particles, which range in size classes roots, and wood from trees and organic residues from
from clays (0.1-2 /j.m in diameter), to silts (2-50 yam agricultural fields, is decomposed on or in the soil, and
. in diameter), and sands (0.05-2 mm diameter). Numer- the nutrients contained within it recycled for further
ous microbes and micro- and meso-fauna (protozoa and use. The decomposition process drives complex food
nematodes) exist in water films on these particles and webs in the soil, with numerous interactions between
in or on the surfaces of microaggregates formed from the initial agents of decomposition, the bacteria and
the primary particles. In turn, the more mobile fauna, fungi, and the fauna which hi turn feed on them.
from collembola and mites (larger meso-fauna) to the Decomposition is the catabolism of organic com-
macrofauna (earthworms, millipedes, ants, termites, pounds in plant litter and other organic detritus. De-
and fossorial or earth-dwelling vertebrates), move composition is principally the result of microbial activi-
through macro- and micro-pores in the soil. The mac- ties; few soil animals have cellulases in their guts, which
rofauna play a role in moving parts of the soil profile allows them to hydrolyze the celluloses in plant resi-
around and form many sorts of burrows and pores; they dues. The decomposition of organic residues involves
are often termed "ecological engineers." the activities of a variety of soil biota, including both
microbes and fauna, which interact conjointly in the
process. For example, the initial breaking up of plant
litter usually is conducted by the chewing and macerat-
II. SOILS AS ORGANIZING CENTERS ing action of both large and small animals. This initial
IN ECOSYSTEMS breaking into smaller pieces, or "comminution," is a
process that benefits the fauna, which derive nutritional
Soils may be viewed as the organizing centers for terres- benefit from the litter or microbes initially colonizing
trial ecosystems. Major functions such as ecosystem the plant material. The increased surface area and fur-
production, respiration, and nutrient recycling are con- ther inoculation of the smaller pieces enhances the mi-
trolled by the rates at which nutrients are released by crobial access to, and breakdown of, these tissues.
308 SOIL BIOTA, SOIL SYSTEMS, AND PROCESSES

B. Nitrogen Cycle: Major Processes fixing rhizobia, in the genera Rhizobium and Bradyrhizo-
bium. The other principal nitrogen-fixing symbiont, the
Nitrogen enters the ecosystem via nitrogen fixation, in bacterium Frankia (Actinomycetaceae) forming the ac-
which the dinitrogen molecule (Nj) is separated into tinorrhiza (literally actinomycete-root), contains only
two nitrogen atoms, with considerable expenditure of a few species in the genus. However, approximately
energy and the assistance of the nitrogenase enzyme, 194 plant species in eight families and four different
to break the triple covalent bond. The atoms are am- subclasses of flowering plants have been identified as
monified and then used in the production of amino hosts. These plants share die general tendency to grow
acids and proteins in the plant. Another avenue for in marginal soils and play an important role as pioneer
nitrogen entry into soils is by lightning fixation, in species in early successional habitats.
which the extensive high-voltage energy in the lightning • In the nitrification steps, noted earlier, there are only
charge ruptures the dinitrogen molecule, hydrogens are a few genera and species of nitrifiers. Most of them are
attached, and then the ammonium is brought in by autotrophic and quite sensitive to changes in soil pH.
rainfall. As shown in Fig. 3, nitrogen is lost from the This means that diese organisms may be unusually
system via harvest and erosion of organic forms of N, prone to being diminished or"eliminated in regions
it can be ammonified in decomposition, and dien under- where diere is considerable acid rain.
goes nitrification to nitrate (NO3~), whereupon it can
be taken up by biota, eidier plant roots or into microbial
tissues. If there is adequate energy and low amounts of
IV. BIODIVERSITY IN SOILS
oxygen present, there can be denitrification, in which
the nitrogen is lost as either nitrogen gas (N2) or N2O,
A. Evolutionary History
nitrous oxide. For further details, consult textbooks on
ecology or ecosystem studies. Soils, as we know them, with well-differentiated pro-
The nitrogen cycle is of critical importance to biodi- files, probably developed concurrently with the origin
versity considerations, because key points in the cycle of a land flora in die early Devonian era, about 425
are dependent on relatively species-poor assemblages million years ago. The microorganisms that inhabit die
of microbes, including die nitrogen fixation and nitrifi- soils, particularly die prokaryotic microbes such as die
cation steps. There are only a few species of nitrogen cyanobacteria, originated perhaps 3 billion years ago.

N fixation
Stream water
losses
Harvest and
erosion losses
N 2 N20
Dentrification
losses

Atmospheric
deposition

FIGURE 3 Inputs and outputs of N, which make up the intersystera transfers in an ecosystem.
Numbers indicate groups of organisms active at a given stage of the N cycle. (1) Rhizobia, Frankia,
(2) ammonifiers, (3) nitrifying bacteria, and (4) denitrifying bacteria. Denitrifying bacteria must
compete with biotic uptake of NOj by other microbes and plants; thus the rate of nitrification
sets an upper limit on denitrification losses. Streamwater losses represent the excess of available
N over that taken up by biotic processes. Modified from Waring and Schlesinger (1985).
 . SOIL BIOTA, SOIL SYSTEMS, AND PROCESSES 309

B. Diversity of Biota that have a unit membrane-bound nucleus and other

organelles, such as mitochondria. Eukaryotic organisms
Biodiversity is an inclusive concept, including a wide are often multicellular. This scheme is based on an
range of functional attributes in ecosystems in addition increasing body of evidence from ribosomal RNA
to being concerned with numbers of species present in (rRNA) phylogenies, that the archaebacteria are worthy
the system. This differentiates it from the concept of of the same taxonomic status as eukaryotes and bacteria.
species diversity, which is concerned with the identity As shown in Fig. 4, the universal rRNA tree develops
and distribution of species in a given habitat or region. from a postulated "cenancestor," leading to the relative
Soil biodiversity is best considered by focusing on positions of the three great domains.
the groups of soil organisms that play key roles in
ecosystem functioning. Spheres of influence (SOI) of 1. Number of Species of Prokaryotes
soil biota are recognized, such as the root biota, the Recent estimates of the number of prokaryotic species
shredders of organic matter, and the soil bioturbators. range from 100,000 to 10 million. Interestingly, the
These organisms influence or control ecosystem pro- number of described species of bacteria in soil amount
cesses and have further influence via their interactions only to about 4000. This discrepancy is due largely to
with key soil biota (e.g., plants). What is the extent of the fact that only a small proportion, usually less than
redundancy within functional groups within these SOI? 1%, of the bacteria present in soil or any other medium
Some soil organisms, such as the fungus and litter- are amenable to culturing and subsequent micro-
consuming microarthropods, are very speciose. For ex- scopic observation.
ample, there are up to 170 species in one Order of It should be noted that, on the basis of the accepted
mites, Oribatida (members of the Arachnida, eight- criterion for separating taxa in microbial studies, which
legged arthropods), in the forest floor of one watershed is a greater than 70% DNA homology, a mouse and a
in western North Carolina. The soil biota considered human would be considered as being in the same spe-
at present to be most at risk are some of the species- cies. This leads to complications, as we shall see, in
poor functional groups, such as specialized bacteria, discussing the total amount of genetic diversity of all
that is, nitrifiers and nitrogen fixers (see diagram). Oth- organisms, including the as-yet largely unknown diver-
ers include fungi forming mycorrhiza (literally fungus- sity of Archaea and Eubacteria. The latter now are esti-
root), a symbiotic association that benefits both plant mated to have an array of 36 kingdoms, which are
and fungus, with the plant supplying high-quality car- genetically as diverse as the Kingdoms Animalia, Plan-
bon to the fungus, and the fungal hyphae exploring a tae, and Fungi in older classification systems.
greater volume of the soil, obtaining scarce mineral
nutrients, particularly phosphorus. Other species-poor 2. Biomass and Numbers of Bacterial
functional groups include macrofaunal shredders of or- Species on Earth.
ganic matter (e.g., millipedes) and bioturbators of soils, This figure is vastly underestimated. We are just now
which includes various types of earthworms and ter- delineating the overall genetic makeup of isolates taken
mites. from soils, which are determined by the use of molecu-
lar probes. The total numbers of bacteria on earth in
C. Three Great "Domains" of Organisms all habitats is truly staggering: 4-6 X 1030 cells, or 350
to 550 petagrams of Carbon. One petagram is 1013 g,
on Earth
or one billion metric tonnes. The amount of the total
All of life exists in three great "urkingdoms," or do- that is calculated to exist in soils is approximately
mains. These domains are (a) the Bacteria (eubacteria), 2.6 X 1029 cells, or about 5% of the total on earth.
which are the bacteria as generally considered; (b) Arch- A majority of bacteria exist in oceanic and terrestrial
aea (archaebacteria), which include the methanogens subsurfaces, especially in the deep mantle regions, ex-
(methane-producers), most extreme halophiles (ones tending several kilometers below the earth's surface.
living in hypersaline environments), and hyperther-
mophiles (ones living in volcanic hot springs, and in 3. Viruses as Quasi-Orgam'sms
mid-sea ocean hot-water vents); and (c) Eucarya (eu- Viruses are quasi-organisms, not included in the three
karyotes) (Fig. 4). The first two domains are prokary- domains. Viruses are RNA or DNA molecules contained
otes, which are unicellular organisms, lacking a unit within protein envelopes. Viral particles are metaboli-
membrane-bound nucleus and other organelles, usually cally inert, carrying out neither biosynthetic nor respira-
having their DNA in a single circular molecule. Eukary- tory functions. They multiply only within host cells,
otes, in comparison, consist of all of the organisms by inducing a living host cell to produce the necessary
310 .SOIL BIOTA, SOIL SYSTEMS, AND PROCESSES

Animals Fungi

Plants
Entamoeba
Euryarchaeota
Crenarchaeota Euglena
Kinetoplasta
(e.g. TVypanosoms)
Low G-'-C Gram
Positives Parabasalia
O/E - Purples (e.g. Trichomonas)
a - Purples and Mitochondria "Archezoa"
y/p - Purples Microsporidia [?]
Spirochaetes (e.g. Nosema)
Fusobacteria Metamonda
(e.g. Giardia)
Flexibacter / Bacteriodes
Cyanobacteria & Chloroplasts
Thermus
Aquifex

"Cenancestor"
FIGURE 4 Schematic drawing of a universal rRNA tree showing the relative positions of evolutionary pivotal groups in the
domains Bacteria, Archaeo, and Eucarya. The location of the root (the cenancestor) corresponds to that proposed by reciprocally
rooted gene phytogenies. The question mark beside the Archezoa group Microsporidia denotes recent suggestions that it might
branch higher in the eukaryotic portion of the tree. (Branch lengths have no meaning in this tree.) From Brown and Doolittle (1997).

viral components. Once assembled, the replicated vi- of enzymes, such as cellulases, and even lignases in
ruses escape from the cells. Viruses infect all sorts of some specialized forms, decomposing substrates in situ,
animals, plants, and microbes. Viruses parasitizing bac- imbibing the decomposed subunits and translocating
terial cells are commonly called bacteriophages, or sim- them back through the hyphal network. Fungi are very
ply phages. Although little is known about the ecology abundant, particularly hi undisturbed forest floors in
of viruses, they can persist in soils for many years and which literally thousands of kilometers of hyphal fila-
decades. Some research on viruses in deserts showed ments will occur per gram of leaf litter.
that they were inactivated in soils at acid pH levels Fungi are still little-described, with possibly less than
between 4.5 and 6. There is little information on the 5% of them known to Science (69,000 described; per-
overall species diversity of viruses in soils. Current esti- haps 1,500,000 in existence (Table I)). This is largely
mates are 5000 species known and perhaps 130,000 because of the fact that so many fungi are associated
in existence. with tropical plants and animals, and these in rum have
not been described.
As noted earlier, the roles of mycorrhizas in soil
4. Numbers and Biodiversity of Eukaryotes
systems are being increasingly viewed as central to
a. Fungal Diversity much of terrestrial ecosystem function. The total num-
Fungi are multicellular eukaryotes that are found in ber of mycorrhizal species may be just 1000 or 2000,
many habitats worldwide. They have long, ramifying but they are essential to the growth and reproduction
strands (hyphae), which can grow into and explore of numerous families of plants. Recent experimental
many microhabitats, and are used for obtaining water studies have noted that species richness, namely with
and nutrients. The hyphae secrete a considerable array large versus small numbers of species of Arbuscular
 SOIL BIOTA, SOIL SYSTEMS, AND PROCESSES 311

TABLE I 70 to 80% of all soil ciliates may yet be unknown. This

Comparison of the Numbers of Known and Estimated Total high proportion may hold true for the other protozoan
Species Globally of Selected Groups or Organisms groups as well.
Known Estimated Percentage c. Mesofauna
Group species total species known
I Nematodes Nematodes feed on a wide range of
Vascular plants 220,000 270,000 81 foods. A general trophic grouping is bacterial feeders,
Bryophytes 17,000 25,000 68 fungal feeders, plant feeders, and predators and omni-
Algae 40,000 60,000 67 vores. Anterior (stomal or mouth) structures can be
Fungi 69,000 1,500,000 5 used to differentiate general feeding or trophic groups.
Bacteria 3,000 30,000 10 The feeding categories are a good introduction, but
Viruses 5,000 130,000 4 feeding habits of many genera are complex or poorly
known. For example, some genera in immature phases
Source: Hawksworth (1991).
will feed on bacteria and dien become predators on
other fauna once diey have matured. Because of the
wide range of feeding types and the fact that nematodes
mycorrhiza, has a positive impact on plant primary seem to reflect ages of the systems in which they occur
production in macrocosms of North American old fields (e.g., annual versus perennial crops, or old fields and
(fields undergoing succession and not intensively pastures and more mature forests), diey have been used
managed). as indicators of overall ecosystem condition. This is a
growing area of research in soil ecology, and one in
b. Microfauna which the intersection of community analysis and eco-
The unicellular eukaryotes, or Protoctista, include a system function could prove very fruitful. Current spe-
wide range of organisms, which are more often called cies described total some 5000, and upward of 20,000
protozoans. These include the flagellates, naked amoe- may exist.
bae, testacea, and ciliates (Fig. 5). These organisms ii. Collembola Collembolans, or "springtails," are
range in size from a few cubic micrometers in volume primitive Apterygote (wingless) insects. They are called
to larger ciliates, which may be up to 500 micrometers "springtails" because many of diem have a spring-like
in length and 20 to 30 micrometers in width. Protozoa lever, or furcula, which enables them to move many
are quite numerous, reaching densities of from 100,000 body lengths away' from predators by use of it, in a
to 200,000 per gram of soil. Bacteria, dieir principal springing fashion. Collembolans are ubiquitous mem-
prey, often exist in numbers up to 1 billion per gram bers of die soil fauna, often reaching abundances on
of soil. All of these organisms are true water-film dwell- 100,000 or more per square meter. They occur through-
ers and become dormant or inactive during episodes of out the soil profile, where dieir major diet is decaying
drying in the soil. They can exist in inactive or resting vegetation and associated microbes (usually fungi).
stages for literally decades at a time hi very exeric envi- However, like many members of die soil fauna, collem-
ornments. bolans defy placement in exact trophic groups. Many
About 40,000 extant protozoan species have been collembolan species will eat nematodes when those are
described, but many more undoubtedly are awaiting abundant. Some feed on live plants or dieir roots. One
scientific discovery. Foissner (1997) notes that about family (Onychiuridae) may feed in die rhizosphere and
360 protozoan species per year are being discovered. ingest mycorrhizae or even plant padiogenic fungi.
In an extensive survey of soils from Africa, Australia, Eight families of Collembolans occur in soils. Many
and Antarctica, in some cases nearly half of the total Collembolans are opportunistic species, capable of rapid
species described were new to science. This was particu- population growth under suitable conditions. Eggs are
larly true in Africa, where of 507 species identified, 240 laid in groups. Collembolans become sexually mature
of diem, or 47%, were previously undescribed. Even in widi die fifth or sixth-instar, but diey continue to molt
a more extensively investigated region, Australia, 43% throughout life. Although many species are bisexual,
of the total of 361 species were new to science. In some of die common species are parthenogenic, con-
Antarctica, 95 species were described, with only 14, or sisting of females only. Collembolan "blooms" are a
15%, being unknown. phenomenon of late winter or early spring, when some
Because many habitats have been uninvestigated yet, species may appear in large numbers on the surface of
and the isolation procedures are still imperfect, from snow banks, on die surface ice of pond water, or on
312 , SOIL B I O T A , SOIL SYSTEMS, AND PROCESSES

Contractile vacuole
Nucleus

B
Test

Nucleus

Contractile vacuole

Food vacuole

Pseudopod

Frontal Adoral zone of

membranelles

Undulating membrane

Contractile vacuole
Cirri /
Macronucleus
Anal or
transverse

Caudal
\
FIGURE 5 Morphology of four types of soil Protozoa: (a) flagellate (Bodo), (b) naked amoeba
(Naegleria), (c) testacean (Hyalosphenia), and (d) dilate (Oxytridia). From Coleman and Cros-
sley (1996).

lichen-covered granite outcrops. There are some 6500 with varied strategies for feeding, reproduction, and dis-
described species and possibly more than 10,000 in persal.
existence. Four suborders of mites occur frequently in soils:
iii. Mites (Acari) The soil mites, Acari, are chelic- the Oribatei, Prostigmata, Mesostigmata, and Astig-
erate arthropods related to the spiders. They are often mata. Occasionally, mites from other habitats are ex-
the most abundant microarthropods in many types of tracted from soil samples. These include, for example,
soils. A 100-g sample may contain as many as 500 plant mites (also called spider mites), predaceous mites,
mites representing nearly 100 genera. This diverse array normally found on green vegetation, and parasites of
includes participants in three or more trophic levels, vertebrates or invertebrates. The most numerous ones
 SOIL B I O T A , SOIL SYSTEMS, AND PROCESSES 313

are the true soil mites. The oribatid mites (Oribatei) soil processes comes from agroecosystems and involves
are the characteristic mites of the soil and are usually a small group of European lumbricids (family Lumbrici-
fungivorous or detritivorous. Mesostigmatid mites are dae in the order Oligochaeta). In North America, south
nearly all predators on other small fauna, although a of the southern limit of the Wisconsinan glaciation,
few species are fungivores and may become numerous several native genera exist. However, exotic (often pere-
at times. Astigmatid mites are associated with rich, de- grine European lumbricids) earthworm species have
composing nitrogen sources and are rare except in ag- been introduced into much of this area following hu-
ricultural soils. The Prostigmata contains a broad diver- man population changes and colonizations. Impacts of
sity of mites with several feeding habits. Very little is exotic earthworms on native species are not well under-
known of the niches or ecological requirements of most stood, although there is evidence that when native habi-
soil mite species, but some interesting information is tat is destroyed and native earthworm species extir-
emerging. For further details on the life-history charac- pated, exotic earthworms colonize the newly empty
teristics of these interesting animals, refer to Coleman habitat. As more extensive studies are carried out, it is
and Crossley (1996). About 20,000 species have been becoming clear that earthworms are present in a wide
described and possibly in excess of 80,000 exist. variety of tropical as well as temperate ecosystems.
Earthworms have important rolessn the fragmenta-
d. Macrofauna tion, breakdown, and incorporation of soil organic mat-
i. Termites Termites (Isoptera) are one of the ma- ter (SOM). This affects the distribution of SOM and
jor ecosystem "engineers" particularly in tropical re- also its chemical and physical characteristics. Changes
gions. Termites are social insects with a well-developed in any of these soil parameters may have significant
caste system. By their ability to digest wood, they have effects on other soil biota, by changing their resource
become economic pests of major importance in some base (e.g., distribution and quality of SOM, microbes,
regions of the world. Termites are arranged in five dif- or microarthropods) or by changing the physical struc-
ferent families. The termites in a more primitive family, ture of the soil. Recent evidence indicates that earth-
the Kalotermitidae, possess a gut flora of protozoans, worm activities impact the communities of other soil
which enables them to digest cellulose. Their normal biota through their effects on the chemical and physical
food is wood that has come into contact with soil. Many characteristics of SOM, causing changes in oribatid spe-
species of termites construct runways of soil, or along cies richness and microarthropod abundances. It is
root channels, and some are builders of large, spectacu- probable that earthworm-induced changes in the micro-
lar mounds. Members of the phylogenetically advanced bial and microarthropod communities will also have
family Termitidae possess a formidable array of .micro- impacts both higher and lower in the soil food web.
bial symbionts (bacteria and fungi, but not protozoa), Some 3650 species of earthworms have been described
which enable them to process and digest the humified and possibly as many as 8000 exist.
organic matter in tropical soils and to grow and thrive
on such a diet.
Although termites are mainly tropical in distribution, V. CONCLUSIONS
they occur in temperate zones and deserts as well Ter-
mites are often considered the tropical analogs of earth- It is apparent that a large proportion of the biota associ-
worms since they reach large abundances in the tropics ated with soils are as yet undescribed, with the most
and process large amounts of litter. Termites parallel extreme cases being the bacteria and fungi. However,
earthworms in ingestive and soil turnover functions. of even somewhat more extensively studied groups,
The principal difference is that earthworms egest much such as Oribatid mites, more than half remain unknown
of what they ingest in altered form (that enriches micro- to science. Therefore, it is premature to give even a
bial action), whereas termites can transfer large rough estimate of the total numbers of species that
amounts of soil/organic material into building nests occur in many of these taxa, as such large percentages
and mounds (carbon sinks). More than 2000 species of the total number of organisms are skill unknown. It
of termites have been described, and probably up to is incumbent on the rising generation of ecologists and
10,000 exist. biologists to develop more innovative ways to describe,
ii. Earthworms The earthworm fauna of North catalog, and understand the myriad patterns and pro-
America is surprisingly poorly known, given the impor- cesses in the biosphere, which are due in large part to
tance of these animals to soil processes and soil struc- the actions of the biota. It is hoped that some of the
ture. Much of the evidence for earthworm effects on observations in this chapter, plus the insights offered
314 SOIL BIOTA, S O I L SYSTEMS, A N D PROCESSES

by the references cited in the bibliography, will encour- sity: A probability-based approach using large sample collections
age this effort. from Africa, Australia and Antarctica. Biodiversity and Conserva-
tion 6, 1627-1638.
Hammond, P. M. (1994). Described and estimated species numbers:
An objective assessment of current knowledge. In Microbial Diver-
See Also the Following Articles sity and Ecosystem Function (D. Allsopp, R. R. Colwell, and D. L
Hawksworth, Eds.), pp. 29-71. CAB International, Wall-
ARCHAEA, ORIGIN OF • BACTERIAL BIODIVERSITY • ingford, UK.
EUKARYOTES, ORIGIN OF • FOOD WEBS • FUNGI • NITROGEN Hansen, R. A., and Coleman, D. C. (1998). Litter complexity and
AND NITROGEN CYCLE • SOIL CONSERVATION composition are determinants of the diversity and species compo-
sition of oribatid mites (Acari: Oribatida) in litterbags. Applied
Soil Ecology 9, 17-23.
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