Stream and River Introductory article

Community Structure . Introduction

Article Contents

Laurie C Alexander, University of Maryland, College Park, Maryland, USA . Stream Biota

Margaret A Palmer, University of Maryland, College Park, Maryland, USA . Factors Controlling Community Structure
. Summary

Streams and rivers are inhabited by diverse communities of fish, insects, crustaceans and
other organisms that have adapted to the physical and chemical conditions found in
running water environments. The assemblage of species found in any particular stream is
not random: it is determined by many current and historical factors, including regional
environmental conditions, species interactions, evolutionary processes and human
activities. All of these factors interact in complex ways to create the observed species
diversity, abundance and composition in stream and river communities.

Introduction
Communities are not haphazard groupings of species.
Many large-and small-scale factors affect the species
composition of stream communities, and no single factor
provides a complete explanation for the observed patterns
of community structure in nature. Patterns of community
structure provide insights into the origin and maintenance
of biological diversity in freshwater streams, and into
stream ecological function. We describe the major groups
of stream biota, the key factors that shape stream
community structure, and the scales at which these factors
interact. We also discuss some effects of human activities
on stream communities, and provide examples to illustrate
how physical (abiotic) or biological (biotic) factors interact
to change the species composition of a stream.
Stream Biota
There are over a hundred thousand species of stream
organisms described to date and probably many more
undescribed. A convenient way to discuss stream biota is
by grouping them according to their size: microbes (40
micrometres or less in size), meiofauna (up to a millimetre
in size), and the larger macrofauna. These groupings do not
imply homogeneity in lifestyles, feeding modes, or even
locomotion – indeed diversity within each of these three
groups is as great as among the groups.
Microbes and meiofauna
Microbes and meiofauna are the most abundant and
diverse groups of organisms living in freshwater ecosys-
tems. One cubic centimetre of stream water contains up to
109 bacteria alone. Compared with larger organisms, little
is known about the species composition of microbial
communities because of their small size. Traditional
methods of classification and identification depend on
morphological features that are difficult to see in single-
celled organisms. However, new molecular methods of
genetic identification and characterization are contribut-
ing to our understanding of microbial diversity, speciation
and community structure.
An easily observed part of the stream microbial
community is the ‘slime’ or biofilm that forms on the
surfaces of submerged rocks, wood, soils, leaves and
plants. Although it appears amorphous, this biofilm is a
highly structured community of aerobic and anaerobic
bacteria, fungi, algae (primarily diatoms and cyanobacter-
ia), and small protists, bound together in a polysaccharide
matrix secreted by individual organisms.
The meiofauna are defined as those organisms that will
pass through a 1 mm mesh size but are retained by a 40 mm
mesh size. They may reach abundances as high as 106 per
square metre of streambed and include representatives of a
large number of invertebrate classes as well as the larger
protists. Common stream meiofauna include: rotifers,
microcrustaceans (cladocerans, copepods, ostracods),
larval insects, oligochaetes and nematodes. Permanent
meiofauna spend their entire lives within the size range of
meiofauna (e.g. copepods) while temporary meiofauna
spend only their early life stages as members of the
meiofauna (e.g. early instar midges). Stream meiofauna are
an important part of the food web – they consume
microbes and other meiofauna, and act as a food source for
larger stream predators, including fish, freshwater crusta-
ceans, and insects.
Members of both groups may live in the water column
(planktonic), the substrate surface (benthic), substrates
beneath the streambed (hyporheic), or in or on sediments
and vegetation along the stream margins (riparian).
Stream microbes include the bacteria, fungi, algae and
cyanobacteria (‘blue-green algae’). Single-celled, silica-

ENCYCLOPEDIA OF LIFE SCIENCES © 2002, John Wiley & Sons, Ltd. www.els.net 1
 Stream and River Community Structure

encased diatoms and dinoflagellates are usually grouped Table 1 Local species richness of freshwater stream biota.
with the algae because of their plant-like attributes, Values are estimates of the maximum number of species
especially their ability to conduct photosynthesis. expected in a single reach based on published data for
temperate streams

Vascular plants and bryophytes Taxonomic group No. of species

A relatively small number of vascular plants and bryo- Meiofauna:

phytes are found in streams, primarily because most are Protozoa 600
unable to withstand strong currents. Their growth is also Flatworms 20
often restricted by lack of fine sediments suitable for Gastrotrichs 10
rooting and by limited light levels. Those that are found Rotifers 120
include some flowering plants, mosses and liverworts, Tardigrades 15
collectively referred to as macrophytes. Vascular plants are Nematodes 25
completely absent from many streams, or limited to small Oligochaetes 30
patches in shallow pools and other areas of slower flow. Microcrustacea (copepod- 100
Even where they are limited, macrophytes can have s, ostracods, cladocerans)
significant impacts on the microbial and invertebrate Macrofauna:
community structure because they greatly enhance the Bivalves 40
stream habitat for benthic organisms by offering a large Crustacea (e.g. crayfish, am- 8
and complex growing surface for biofilms. Diatoms, phipods)
bacteria and invertebrates are abundant on the stems and Insects
leaves of submerged vegetation. Aquatic plants offer a Mayflies (Ephemeroptera) 30
much more varied and complex three-dimensional habitat Stoneflies (Plecoptera) 15
for microorganisms and macroinvertebrates than the bare Dragonflies (Odonata) 35
rock surface, and studies have shown that the microbial Caddis flies (Trichoptera) 40
community structure of the biofilm is quite different on Aquatic beetles (Coleoptera) 70
aquatic plants than on rock substrates on the streambed. True bugs (Hemiptera) 30
Although not useful as a food source for most inverte- Megaloptera 3
brates, macrophytes offer refuge from predators and Midges (Chironomidae) 125
scouring floods. Other true flies (Diptera) 300

Macroinvertebrates
The major macroinvertebrate groups living in freshwater
streams are insects (mayflies, stoneflies, dragonflies, caddis
flies, aquatic beetles, true flies – including midges,
mosquitoes and black flies – true bugs, dobsonflies and
alderflies), freshwater crustaceans (e.g. crayfish, freshwater
shrimp, amphipods and isopods), molluscs (snails and
bivalves – freshwater clams and mussels), oligochaetes
(segmented worms), leeches, water mites, and turbellaria
(flatworms).
The aquatic insects are a diverse group (Table 1) and are
often the most abundant and conspicuous species found in
streams. In most cases, aquatic insect larvae and nymphs
develop underwater, then emerge from the stream for a
terrestrial adult stage. The terrestrial adult stage is often
shorter than the aquatic immature stage, and is important
for dispersal and finding suitable mates. Insects are an
important food source for other animals both underwater
(e.g. fish) and on land (e.g. birds, spiders). Insects also
contribute to the processing of allochthonous inputs
(leaves, wood and other plant materials that fall into
streams) by mechanically breaking them into smaller
particles that can be more efficiently processed by stream
microbes. Thus aquatic insects form an important link
between aquatic and terrestrial ecosystems, as well as an
important link between the microbes and the larger
predators.
In recent years, aquatic macroinvertebrates, especially
insects, have become important as bioindicators of water
quality and stream health. The morphology, physiology
and behaviour of different taxa are sufficiently diverse that
macroinvertebrate community structure reveals much
about environmental conditions. Aquatic insect species
are adapted to a wide range of flow regimes, water
conditions, nutrient levels, temperature regimes, and in-
stream habitats, so changes in their community structure
can help us detect changes in environmental conditions.
For example, the mayflies (order Ephemeroptera), stone-
flies (Plecoptera) and caddis flies (Trichoptera), collectively
referred to ‘EPT taxa’, are known to be intolerant to
nutrient enrichment like that caused by release of sewage,
fertilizers, or farm wastes into a stream. Members of other
groups, including oligochaetes (worms) and some flies (e.g.
family Chironomidae or midges), are more tolerant of
these conditions. So a shift in community structure such

2

that EPT taxa are replaced by midges might indicate
elevated levels of organic pollution. New research into the
relationships between environmental conditions and
stream invertebrate community structure is seeking ways
to improve the use of these organisms in stream monitoring
programmes.
Vertebrates
Stream communities include representatives from all
vertebrate classes, including amphibians (e.g. salaman-
ders, frogs), reptiles (e.g. turtles, snakes, crocodiles), birds
(e.g. ducks, dippers), mammals (e.g. beavers, otters), and
the principal vertebrate member of lotic food webs, fish.
Many stream vertebrates are predators, and as such exert a
powerful structuring force on the communities of streams
they inhabit. Detritivorous and grazing vertebrates,
including many fish species, also play an important role
in shaping stream community structure by reducing algal
abundance, thereby limiting an important food source for
other grazers. Stream invertebrates, which are consumed
by predaceous fish and compete for food resources with
grazing fish, are especially affected by the presence of this
group of aquatic vertebrates.
There are approximately 8500 species of freshwater fish,
most of which live in rivers and connected floodplains. The
life cycles of some freshwater fish include a marine stage.
Anadromous fish such as salmon and lamprey spawn in
freshwater but migrate to saltwater for part of their
growth; catadromous fish such as eels spawn in the sea but
migrate into freshwater streams for development. The
natural connectivity of stream networks has been inter-
rupted in hundreds of thousands of streams by the building
of impoundments for human purposes (e.g. drinking water
reservoirs, hydroelectric power, flood control, recreation).
The effect of impoundments on fish community structure
and dispersal is a major concern of conservationists and
environmental agencies.
Factors Controlling Community
Structure
Life in streams and rivers is affected by the characteristics
of the landscapes through which they flow. Over long
periods of time, landforms built up by sedimentation and
geological activity are eroded by processes of weathering
and the movement of glaciers and water. The complex
topography and underlying geology of the resulting
landscapes determine how precipitation falling on Earth
will accumulate to form the channelized flows of water in
which stream organisms live. Climate, water chemistry,
local geology (e.g. soils, channel size, type of bottom
substrates), and flow act as first-order constraints to
determine which taxa we can expect to find in a catchment.
Stream and River Community Structure
Many species we may never expect to find in a given stream
system simply because they cannot survive the rigours of
the environment. However, even if an organism fully
adapted to these conditions could occur in a catchment, we
may not find it there, because final composition of stream
communities is determined by a whole host of second-
order factors including local habitat types, food avail-
ability, predation and competition (Figure 1).
Climatic and chemical factors
The distribution of a species is constrained by the
temperature regime, and thus tightly linked to geographic
latitude and altitude. The correlation of temperature to
species distribution is based in part on the evolutionary
adaptation of metabolism and life cycle to the ranges and
cycles of temperature in which the species lineage
originated and diversified. Stream temperature is a
function of seasonal weather cycles, geographic location,
type and amount of riparian vegetation, and contact of
stream channels with groundwater sources. Unlike lake
ecosystems, which experience thermal stratification and
diurnal cycling of water temperature, stream water is well
mixed and therefore temperature is relatively constant
throughout a channel. In cases where groundwater
Flow
Abiotic
magnitude, frequency
°C, O2, pH
extremes
Substrate
size, heterogeneity
Habitat
availability, diversity
Biotic
Food
availability, diversity
Competition, predation
intensity, frequency
Figure 1 Factors controlling the community structure of stream and
riverine communities. Factors are shown hierarchically in terms of general
importance; however, factors do interact and take on different levels of
importance depending on the regional and local context. For example,
flood or drought magnitude may determine which species colonize an
area, but if temperatures exceed their tolerances or the bottom substrate is
too silty for them to feed, they may be absent.
3
 Stream and River Community Structure
constitutes a large percentage of flow, stream water
temperatures may vary as little as 1–28C year round,
making them almost completely independent of air
temperature.
Stability and predictability of water temperature is
important to stream biota because the life cycles of some
organisms are linked to seasonal cycles, and even very
small changes in stream temperature can trigger events
such as initiation of mating behaviours, hatching of eggs,
or emergence of terrestrial adults. Like all biota, stream
organisms have thermal and chemical tolerances. Some
(e.g. the midge family, Chironomidae) are tolerant of wide
ranges while others can only tolerate very narrow ranges
(trout will generally not be found if the temperature
exceeds 208C, and cannot survive above 258C). Some
insects have larger body size and fewer generations per year
in colder streams compared with warmer streams. In one
mayfly species (Ephemerella ignita), the percentage of eggs
that hatch drops from 90% at 108C to about 40% at 208C.
Streams vary immensely in the chemical composition of
their waters, which reflect the environmental conditions of
the surrounding landscapes. Variables of greatest impor-
tance to stream biota include (a) pH; (b) water hardness; (c)
nutrient level, measured as several nitrogen-and phos-
phorus-based compounds; (d) dissolved gases, especially
carbon dioxide and oxygen; (e) dissolved major ions,
including Ca2 1 , Na 1 , Mg 1 ; (f) suspended inorganic
elements, including iron, aluminium and silicon. In
general, these factors are most influential at their extremes.
For example, when waters are extremely low in oxygen or
very acidic, species may be unable to survive. Below the
tolerance limits, there are, however, gradients in species
composition and diversity in response to some chemical
factors. For example, species richness and diversity have
been shown to decrease with pH. Species composition may
also be affected – certain invertebrates are absent from
lower pH streams (mayflies, molluscs) whereas others may
be well represented (black flies, stoneflies). Fish are very
sensitive to low pH, especially in combination with
elevated aluminium levels. Acid mining drainage streams
– where the pH is extremely low – are largely depauparate
of fauna, and may have a community consisting of just a
few acid-tolerant algal species.
Flow and substrate characteristics
In undisturbed areas, climatic factors (cycles of precipita-
tion, sunlight and temperature) interacting with the
physical properties of the catchment (altitude, stream
gradient, underlying geology, hydrologic soil type, and
vegetation cover) dominate the large-scale factors that
affect stream community structure. Flow parameters of
greatest importance to stream organisms are the levels of
stream discharge (volume of water and its rate of flow
through a channel), flood frequency and predictability,
4
which together define a stream’s flow regime. These
parameters are largely determined by regional differences
in seasonal patterns of precipitation, connectivity of
stream channels to sources of groundwater sources, basin
topography, and terrestrial vegetation in the riparian zone.
Discharge is of great importance to stream plants and
animals, which must adapt to the forces of current. Current
velocity is perhaps the single most important physical
factor in shaping the structure of stream communities. In
addition to the obvious effect of dislodgement, forces
associated with velocity affect the amount and type of
stream habitat available for colonization by stream
communities by determining (a) the size and arrangement
of substrate particles; (b) the rate at which nutrients and
particulate carbon sources are transported though a
stream; (c) in-stream movement of individual animals,
including dispersal, migration, and drift; (d) rate of bank
erosion and channel shape; and (e) the availability of
dissolved oxygen (DO). Organisms that depend on passive
diffusion of DO across gills (e.g. fish, mayflies) or through
their body walls (e.g. chironomids, aquatic worms), and
organisms that filter-feed (e.g. black flies, net-building
caddis flies), depend on fast flow for survival. Other
organisms that are not well adapted to resist the forces of
flow (e.g. adult beetles, meiofauna) require slower flow or
protective refuge on plants, wood, or within the streambed
and may be washed downstream in periods of flood.
Discharge is the product of baseflow from groundwater
sources (which moves underground into stream channels)
and overland flow from precipitation or snowmelt. Over-
land flow and sedimentation into streams are reduced by
vegetation, which changes stream conditions in four
important ways: (a) plants slow runoff by intercepting
precipitation and physically obstructing overland flow, so
the physical impact of floods on stream organisms is
reduced; (b) the soil bacteria and fungi associated with
terrestrial plants alter the chemical composition of runoff
in some cases, significantly cleansing the water (e.g. of
excess nitrogen); (c) vegetation reduces the rate at which
weathering of terrestrial rocks occurs or soil erosion
occurs; and (d) by slowing runoff, riparian vegetation
reduces the temperature of water that enters the stream.
It is also important to emphasize that some level of
flooding is normal in streams and rivers and without this,
species composition shifts towards more lentic (low or no
flow) adapted species. Some ecologists have found
evidence that species richness of stream macroinverte-
brates may be highest when there is moderate but not
extreme or zero flow variability (the intermediate dis-
turbance hypothesis). This hypothesis states that species
richness is increased by intermediate levels of flooding
because when flow is constant, those species best adapted
(i.e. with highest reproductive and survival rates) to the
particular flow level will out compete other species,
whereas when flow is extremely variable, few species will
be able to survive the extremes. Furthermore, research has
 Stream and River Community Structure

shown that the organisms have adapted over evolutionary
time to particular flood frequencies and predictabilities.
The life cycles of many stream organisms are synchronized
with annual flood or drought cycles, so that hatching eggs
and small fry or larvae can develop during periods of
slower flow or in moist streambeds. One of the major
impacts of human activities on streams is alteration of
water flow in streams and rivers (e.g. by dams or other land
use changes) and thus there is a move around the world to
restore natural flow regimes.
Local habitat and faunal movements
Some of the most important factors controlling commu-
nity structure in streams are those relating to local habitat –
not only habitat quality but also the number of habitat
types (habitat diversity or heterogeneity), their stability,
and their arrangement on the streambed. In general, for
both fish and stream invertebrates, species abundance and
diversity increase as habitat heterogeneity increases. The
presence of woody snags, overhanging banks, macro-
phytes, and many different sizes of bottom substrates (e.g.
cobble, boulders, sand) increase the number of habitat
types and food niches available to fauna. The presence of
wood from adjacent riparian vegetation seems particularly
important to fish and insect diversity. Wood along with
large boulders helps stabilize the channel and provides
refugia for fish and invertebrates during floods. Wood and
the decaying leaves often trapped on the wood (‘woody
debris dams’) provide rich resources for biofilm develop-
ment, which in turn supports a large number of meiofauna
and macrofauna, including fish. For this reason, debris
dams are considered biological ‘hotspots’ in streams – total
production is high, community structure is complex with
many feeding groups present (algal and microbial grazers,
leaf shredders, predators) (Figure 2).
Dispersal abilities of fauna often determine whether or
not they will be present in a particular stream. If the stream
has frequent, unpredictable floods (as in urban settings,
where impervious surfaces produce flood-levels of runoff
with each heavy rain), only those taxa that broadly disperse
and rapidly colonize will be found. This includes insects

Figure 2 Wood that falls into streams and traps leaves and other organic material supports biologically productive and diverse biotic assemblages. The
fauna live both on and in the wood, as well as on the biofilm that covers submerged pieces.
(a) Microbes and meiofauna: http://www.microscopy-uk.org.uk/mag/wimsmall/smal3.html
(b) Elmidae (adult riffle beetle): http://entweb.clemson.edu/museum/misc/aqua/aqua22.htm
(c) Aeshnidae (dragonfly nymph): http://twri.tamu.edu/
(d) Hydropsychidea (caddisfly larvae): http://waterknowledge.colostate.edu/arctopsy.htm
(e) Simuliidae (blackfly larvae): http://www.fish.washington.edu/naturemapping/water/1invert.html
(f) Pteronarcyidae (stonefly nymph): http://waterknowledge.colostate.edu/pteronar.htm
(g) Peltoperlidae (stonefly nymph): http://www.dec.state.ny.us/website/dow/stream/plefamilypageone.htm
(h) Tipulidae (crane fly larva): http://www.chebucto.ns.ca/Science/SWCS/ZOOBENTH/tipulida.html
(i) Ephemerellidae (mayfly nymph): http://www.wlu.ca/  wwwbiol/bio305/Database/Ephemerella.htm

5
 Stream and River Community Structure
with strongly dispersing aerial adults, particularly those
that feed as adults and spend longer times in flight. Those
fauna with a strictly benthic lifestyle (e.g. many crustacea)
will have a tough time surviving in such settings. At smaller
local scales (e.g. a particular pool or riffle) the community
composition at a point in time is greatly influenced by local
movements. While fish may move around to select
preferred habitats, many stream organisms are at the
whims of currents and may be swept away when they move
to the tops of stones to feed. For this reason, communities
dominated by strong swimmers with habitat preferences or
immobile organisms that can remain in their preferred
habitat will exhibit more predictable community structure.
In these cases, we can reasonably predict what species will
be found, based on habitat characteristics (Figure 3),
particularly if the flow is low or highly predictable (e.g.
has regular spring floods and summer baseflows). Fauna
that have high dispersal rates but poor control of where
they move (e.g. passive dispersers that drift with the flow)
may exhibit quite unpredictable community structure,
particularly in streams that have extremely flashy or
sporadic flow regimes.
Food availability
All ecosystems require a continual influx of energy to
function. Lotic ecosystems are sustained by energy from
two sources: solar radiation, which fuels in-stream primary
production; and stored energy in the form of nonliving
organic matter (e.g. leaves, wood, fruit, soil particulates,
dissolved nutrients) transported into the stream from other
ecosystems by wind, runoff, gravity and groundwater.
The relative contribution of each energy source affects
the food base and structure of the stream’s food web. In
streams with abundant sunlight, primary producers such
as algae, cyanobacteria, diatoms and dinoflagellates form
the lowest layer of the stream food web. Biofilm microbes
and meiofauna discussed earlier also live in close associa-
Flashy,
unpredictable Stochastic, temporally
variable communities
Flow regime
Stable, predictable
Constant
or highly
communities
predictable
Low High
Extent of dispersal
Figure 3 Community structure of stream fauna may be highly predictable
over time or space depending on the frequency and magnitude (‘extent’)
of dispersal and the constancy and predictability of flow. For communities
with a high degree of predictability, the factors in Figure 1 may be useful in
predicting the diversity and composition of communities.
6
tion with the complex green algal communities observed in
autochthonous streambeds. The biofilm community in-
corporates algae, trapped detritus and other organic
materials into the polysaccharide matrix, which is grazed
and fed upon by many larger stream animals. Together
these two interrelated communities form an important
food source called periphyton or aufwuchs. The dominant
feeding guilds in such streams would be ‘scrapers’ and
‘grazers’, which feed on attached periphyton. These guilds
include several families of Ephemeroptera, Trichoptera,
Diptera and Coleoptera, and scraping algivorous fish such
as the stoneroller Campostoma.
In many other streams, however, primary production is
limited by low sunlight or by fish and invertebrate grazing.
In these streams the food web is supported by external
(allochthonous) carbon sources, especially the annual leaf-
fall from deciduous trees. For example, a small, forested
stream shaded by a closed canopy of deciduous trees would
probably have a detritus-based food web dominated by
‘shredders’, which break down coarse particulate organic
matter (CPOM), and ‘collectors’, which process fine
particulate organic matter (FPOM), as well as by their
predators. The shredders include families of Plecoptera,
Trichoptera, Crustacea, Diptera and Gastropoda; and the
collectors include families of Diptera and Ephemeroptera.
Predators might include invertebrate-feeding fish, such as
trout, as well as predaceous Plecoptera, Trichoptera,
Diptera and Crustacea. Fungi and bacteria that break
down the leaf material also act as food for higher trophic
levels, including insects, molluscs and grazing fish.
Biotic interactions
Predation and competition are known to influence com-
munity structure in virtually all ecosystems, and streams
are no exception. While biotic interactions may be less
important in structuring communities if physical condi-
tions are extremely harsh (e.g. if there are unpredictable
flash floods), they are considered important determinants
of species composition and relative abundance in most
running-water systems. For example, fish predators may
be highly selective, foraging on a single species or suite of
species. This may reduce species diversity locally unless the
predator is removing prey that would otherwise dominate
(outcompete other species). The complexity of ways in
which predators may influence entire communities in
streams is nicely illustrated by research in a California
stream. Here, fish (steelhead trout, roach) consume
damselfly nymphs that in turn prey on algivorous midge
(chironomid) larvae. When damselfly densities are reduced
by fish feeding, the midge larvae graze the stream’s
filamentous algae to a low mat. When fish are removed,
damselfly nymph feeding reduces the number of midges
feeding on algae, and the algae grow into a tall turf. This
example of a ‘trophic cascade’ (here, a top predator

influencing the structure of the entire community by a
series of trophic interactions) has become a classic study
among ecologists.
Community structure in streams may also be influenced
by direct competitive interactions such as when larger trout
actively displace smaller species from the best feeding
areas, or when caddis flies battle for the best rock upon
which to live and commence filter-feeding. A few field
studies in which individuals have been removed from a
stream have provided evidence that competition for food
may be strong among grazing insects. Declines in stream
insect populations have also been shown to be related to
large-scale outbreaks of parasites (e.g. a host-specific
protozoan that infects caddis flies). Such parasites may
act as important regulators of community structure if they
alter the population cycles of abundant or trophically
important community members.
Summary
In stream communities, species composition and distribu-
tion are the result of many interacting physical and
biological factors, including climate, flow regime, water
chemistry, local habitat heterogeneity and stability, faunal
movement, food base, trophic function, and competition.
Historical environmental and evolutionary processes
Stream and River Community Structure
worked together to create a regional species pool of
potential stream community members, from which local
habitat conditions, species interactions, and dispersal
interact to determine which of those species will be able
to colonize a given stream.
Some factors regulating stream community structure
can be predicted given adequate knowledge of environ-
mental conditions (e.g. loss of species with stream
acidification), biotic factors (e.g. prey population explo-
sion and possible trophic cascade following elimination of
a predator), or their interaction (e.g. seasonal cycles of
species replacement). Other important regulators of
community structure are not predictable including sto-
chastic events (e.g. flash floods caused by hurricanes) or
series of events (e.g. genetic changes interacting with
environmental factors), and the side effects of capricious
human behaviour (e.g. introduction of invasive species,
fragmentation and destruction of essential habitats).
Further Reading
Allan JD (1995) Stream Ecology: Structure and Function of Running
Waters. London: Chapman and Hall.
Cummins KW, Cushing CE and Minshall GW (eds) (1995) River and
Stream Ecosystems. New York: Elsevier.
Giller PS and Malmqvist B (1999) The Biology of Streams and Rivers.
Oxford: Oxford University Press.
Morin PJ (1999) Community Ecology. Massachusetts: Blackwell Science.