American Fisheries Society Symposium 73:651–664, 2010
© 2010 by the American Fisheries Society
Community Ecology of Stream Fishes:
Synthesis and Future Direction
Keith B. Gido*
Division of Biology, Kansas State University, Manhattan, Kansas 66506, USA
Donald A. Jackson
Department of Ecology and Evolutionary Biology, University of Toronto
Toronto, Ontario M5S3G5, Canada
Synthesis
The broad range of subjects represented in this
symposium highlight exciting new advances
in the field of stream fish community ecology.
Some of the chapters expand on themes (e.g.,
community stability, species interactions) that
were represented in the 1987 publication Community and Evolutionary Ecology of North American Stream Fishes edited by W. J. Matthews and
D. C. Heins. Others present research that was
just developing in the late 1980s or represents
new lines of research (e.g., landscape ecology,
molecular ecology, ecological stoichiometry,
and riverscape perspectives). Many contemporary conservation challenges are similar to
those facing authors of the 1987 symposium,
but some of these challenges have escalated
because of increasing human population sizes
and associated demands on water resources.
The extent to which stream fishes have been affected by these issues is well documented in recent literature (e.g., Dudgeon et al. 2006; Jelks
et al. 2008), and it is clear that quality research
and innovative conservation actions are necessary to halt the rapid deterioration of stream
ecosystems worldwide (Angermeier 2010, this
volume). In light of these major conservation
challenges, the goal of this symposium was to
* Corresponding author: kgido@ksu.edu
bring together leaders in the field of stream fish
community ecology to highlight current advances in the field and pave a way for future researchers. Prefaces for the five main themes of
the book provide excellent syntheses of those
focal areas. Our review of the chapters and
those theme prefaces yield some consensus on
important advances and future directions for
the field of stream fish community ecology. In
particular, the continuing importance of linking patterns and processes was highlighted as
a critical objective for understanding and conserving stream fishes. In this synthesis, we will
use the framework of describing patterns and
process and make recommendations for future
direction. We start with a synthesis of advances in understanding patterns and process in
stream fish community ecology over the past
20+ years and follow this overview with recommendations for future research.
Revealing Pattern and Process in
Stream Fish Community Ecology
Describing patterns in the distribution and
abundance of organisms is central to the field
of ecology and is important in developing
and testing hypothesized factors structuring
stream fish communities. Large-scale patterns
of fish species distributions over space and
time described in the 1987 symposium were
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instrumental in identifying important linkages
between fish community structure and human
perturbations to the landscape (Cross and Moss
1987; Li et al. 1987; Pflieger and Grace 1987).
Papers in the 1987 symposium also linked species distributions to historical biogeography
(e.g., Boschung 1987; Mayden 1987). Over the
past 20+ years, survey information has accumulated, computing power has increased, and analytical methods for processing those data have
been developed and refined (Matthews 2010,
this volume). As a result, we are more aware of
the complexity of factors that regulate stream
fish communities and how scale of observation
influences our interpretation of those factors
(Hugueny et al. 2010, this volume; Matthews
2010). Some analytical methods present in
the 1980s remain valid and have new modifications to improve our understanding of stream
fish communities ( Jackson et al. 2010, this volume). New methods related to ecological niche
modeling (Buisson et al. 2010, this volume), autoregressive regression models (Grossman and
Sabo 2010, this volume), and structural equation modeling (Infante and Allen 2010; Zorn
and Wiley 2010; both this volume) illustrate recent advances in analytical approaches. A major
advancement associated with more extensive
data and analytical tools has been the generation and evaluation of conceptual models describing drivers of community structure, many
of which are based to a large extent on empirical
data collected across larger spatial and temporal
scales (e.g., Hugueny et al. 2010; Roberts and
Hitt 2010, this volume). Because of the breadth
of spatial and temporal scales, these large data
sets are also critical for developing predictive
models that forecast potential changes in stream
fish communities in response to global change
(e.g., Buisson et al. 2010).
Authors in the current symposium generally placed less emphasis on describing species
compositional changes and more emphasis on
describing functional organization of assemblages (e.g., Frimpong and Angermeier 2010;
Olden and Kennard 2010; Rahel 2010; all
this volume). Matching patterns of functional
organization or species composition with environmental variables (e.g., flows) has been
particularly helpful in testing main drivers of
fish community composition (Grossman et al.
1982; Grossman and Sabo 2010; Taylor 2010,
this volume). Indeed, papers published in the
past two decades by Poff and Allan (1995) and
more recently by Bunn and Arthington (2002)
have provided important conceptual frameworks to test linkages between hydrology and
dynamics of stream fish communities and to
develop effective approaches for managing flow
regimes to benefit native species (e.g., Tharme
2003; Annear et al. 2004). Although tracking
changes in species over space and time, particularly over long periods of time, remains
critical to truly understanding community dynamics or drivers thereof (e.g., Grossman et al.
2010, this volume; Taylor 2010; Turner et al.
2010, this volume), this additional focus on
functional organization has helped extrapolate
across regions.
In the 1987 symposium, there were a number of small-scale mechanistic studies characterizing resource use, physiology, and responses
of stream fishes to disturbance. Many of those
papers evaluated resource use of species at relatively small spatial scales (e.g., habitat selection
or resource partitioning studies by Angermeier
1987; Felley and Felley 1987; Fisher and Pearson 1987; Gorman 1987; Ross et al. 1987) and
invoked species interactions as a mechanism
driving those patterns of resource use. Within
these studies, the focus on patterns of resource
use among stream fishes was aimed at identifying potential competitive interactions. Studies
of physiological tolerances (Hlohowskyj and
Wissing 1987; Matthews 1987) and responses
to disturbance identified abiotic factors that
synthesis and future direction
constrained species distributions in space and
time. Small-scale mechanistic studies continue
today and can provide important insight into
processes that elicit a response to harsh conditions such as drought (Marsh-Matthews and
Matthews 2010, this volume) and interactions
among species (Copp et al. 2010, this volume).
These studies are critical for understanding
factors limiting species occurrence within and
across streams.
Linking Patterns with Process
The same year the 1987 symposium was published, Ricklefs (1987) noted that biological
communities failed to converge in similar physical environments (but see Winemiller 1991;
Winemiller et al. 1995) and put forth that both
regional and historical processes were important drivers of community structure. The field
of landscape ecology also has developed rapidly since 1987, providing a framework in which
to link local and regional processes (Ward et
al. 2002a; Wiens 2002). Fausch (2010, this
volume) notes the importance of this “renaissance” in the field of stream fish community
ecology, and chapters by Hugueny et al. (2010,
this volume), Infante and Allen (2010, this volume), and Zorn et al. (2010) provide examples
of research linking local and regional processes.
Winemiller (2010, this volume) points out that
metapopulation and metacommunity theories
provide a framework to link local and regional
processes through dispersal (Falke and Fausch
2010; Peres-Neto and Cumming 2010; both
this volume). However, quantifying dispersal
of individuals across the landscape is still a
major challenge for stream fish ecologists, particularly for early life stages or smaller-bodied
taxa. Douglas and Douglas (2010, this volume) show how modern molecular techniques
can be used to infer dispersal across large spatial and temporal scales, and Rodriguez (2010,
this volume) developed a model to account for
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variable movement behaviors of fish within a
population.
A major step towards linking pattern and
process in stream fish community ecology
stemmed from the work by Winemiller and
Rose (1992), who emphasized how life history
traits affect population response to patterns of
environmental variation, which paves the way
for studying the functional organization of
communities. By analyzing communities based
on functional traits, we can more easily generalize results across systems and link changes in
community structure to environmental factors
driving those patterns. For example, the widely
held view that stream flow is a “master” variable influencing stream fish community structure was influenced early on by Grossman et al.
(1982) and later by Poff and Allan (1995), who
have linked flows to functional traits of stream
fishes. Frimpong and Angermeier (2010) evaluate this trait-based approach and others provide examples of how functional compositions
of communities relate to abiotic conditions
( Jones et al. 2010; Olden and Kennard 2010;
both this volume).
Linking biodiversity to ecosystem function has emerged as a major subdiscipline of
ecology and has been central to the study and
conservation of stream fishes over the past two
decades. In regards to stream fishes, authors
have illustrated the importance of fishes from
a diverse array of functional groups (e.g., grazers, detritivores, invertivores, and piscivores)
on stream ecosystem structure and function
(Power and Matthews 1983; Dahl and Greenburg 1987; Power 1990; Gelwick and Matthews 1992; Flecker 1996; Gido and Matthews
2001). Stream fishes also can elicit trophic
cascades (Power et al. 1985), alter nutrient
dynamics (e.g., Vanni 2002), or stream ecosystem function (Taylor et al. 2006). Papers
in this symposium synthesize the important
roles fishes play in stream ecosystems (Gido
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gido and jackson
et al. 2010; Hoeinghaus and Pelicice 2010;
both this volume), while also outlining novel
approaches for identifying those roles (e.g.,
through altering element ratios [McIntyre and
Flecker 2010, this volume] or the flow of energy within and across habitats [Flecker et al.
2010, this volume]).
A Path for the Future
Over the past two decades, the amount of information has increased exponentially and
new concepts, approaches, and techniques
have been applied to stream fish community
ecology. The rapid expansion of this field has
increased our basic understanding of these
systems and informed managers on effective
conservation strategies. Unfortunately, the
conservation challenges facing stream fishes
has expanded and surely will accelerate in the
foreseeable future—human population is predicted to reach 9 billion by 2050 (Population
Reference Bureau, www.prb.org). Major shifts
in climate, land use, and distributions of biota, including exotic species, are projected. To
confront these challenges, stream fish ecologists will need to focus their research in areas
that will maximize information necessary to
appropriately conserve this diverse, beautiful,
and functionally important group of animals.
At the same time, there are still many species
and systems for which we have little basic
ecological understanding. Thus, there is still a
critical need for descriptive information on the
resource use, physiology, reproduction, and
population dynamics of many species.
This symposium provided an opportunity
to look at the current trajectories of research
and speculate on potentially productive areas
for future research. Below, we outline several
research foci that have provided useful insights for understanding processes structuring stream fish assemblages and make recommendations for future research in the fields.
We compiled this list from discussions with
participants from this symposium, colleagues,
and students. Our intention is not to present
an exhaustive list of potentially useful areas of
research—that list would be too large. Rather,
we present major areas that could be developed to address pressing questions that have
not been adequately answered in the past. We
refer to theme prefaces and individual chapters
of this book for more focused descriptions of
major advances and future directions in those
areas of research.
Macroecology: Comparative Studies
In his synthesis of the 1987 symposium, Clark
Hubbs (1987) identified the need to generalize results from studies conducted at limited
spatial and temporal scales. This continues
to be an important area of research and there
are new approaches to address this issue. The
future will likely make more data available for
such comparative studies through programs
like the U.S. Geological Survey (USGS) hydrologic gauging stations, gap analysis programs of the USGS and Department of Fisheries and Oceans Canada, national and local
monitoring programs (e.g., U.S. Environmental Protection Agency Environmental Monitoring and Assessment Program, Environment
Canada’s Ecological Monitoring and Assessment Network), and museum databases. The
use of meta-analysis (Rosenberg et al. 2000)
has been refined as a tool to uncover generalities from multiple studies. Given the breadth
and quantity of data that will be available, we
suggest several key actions that would facilitate
comparative studies among stream fish ecologists: (1) making data available to multiple investigators, (2) building collaborations among
scientists, (3) measuring community metrics
(e.g., functional traits such as trophic or reproductive guilds or functional groups [Matthews
1998:69]) that enable comparisons, and (4)
synthesis and future direction
developing tools and statistical approaches
to rigorously test generalities across regions.
Such studies will not only validate the generality of local experiments, but will help evaluate
the context dependency of those results across
major ecological gradients (see section on scaling below).
Methods of linking functional organization of fish assemblages with environmental
conditions (e.g., disturbance regimes) will
more explicitly link patterns and process. A
major limitation to this line of work is a lack of
information on the reproduction, feeding ecology, physiology, and behavior of many species.
If such data are available (e.g., Frimpong and
Angermeier 2009), promising new statistical
approaches are available to evaluate these linkages (e.g., RLQ analysis, Dolédec et al. 1996;
and fourth-corner analysis, Dray and Legendre
2008).
Technology: Major Advances in the Field
Will Be Facilitated by Technology
While accumulation of research over space
and time can incrementally advance the field
of community ecology, technological advancement can lead to punctuated steps towards answering key questions. For example, molecular
ecology has expanded at a rapid pace, and we
now have methods to quantify historical relationships among species using routine procedures (see Douglas and Douglas 2010, this
volume). Using this approach and new technologies to simultaneously evaluate genetic
differences among populations and the functions of genes responsible for those differences
(i.e., functional genomics, Turner et al. 2010,
this volume) should provide valuable insight
in to the evolutionary processes responsible
for structuring stream communities. An example of the use of molecular ecology in regards
to conservation is the ability to detect the presence of invasive species in a water body based
655
on the occurrence of its DNA extracted from
a water sample (e.g., Mahon et al. 2009); this
approach was recently used to detect the invasion of Asian silver carp Hypophthalmichthys
molitrix in Lake Michigan.
Rapid advances in molecular methods
and efficient cataloging of genetic information
have expanded our understanding of the phylogenetic relationship of fishes and has allowed
for stronger inferences in community ecology
(e.g., Felsenstein 1985). For example, DNA sequence data for many stream fishes are easily
accessed through the National Center for Biotechnology Information (GenBank), and these
data can be readily used to construct phylogentic hypotheses. Webb et al. (2002) pointed out
that improved phylogenetic hypotheses will
inform studies of community organization by
(1) examining the phylogenetic structure of
community assemblages, (2) exploring the
phylogenetic basis of community niche structure, and (3) adding a community context to
studies of trait evolution and biogeography.
Several studies have incorporated phylogenies
into stream fish community studies (e.g., Gotelli and Taylor 1999; Peres-Neto 2004; Alcaraz et al. 2005), but as data from new species
accumulates and methods of analysis improve,
inferences into the importance of evolutionary
constraints relative to other factors (e.g., biotic
interactions and abiotic forcing) will become
more robust.
Another example of technology that will
continue to improve is through ecoinformatic
approaches incorporating precise locality and
physiological measurements of fishes as well
as environmental sensors into geographic information systems. These databases will allow
synchronized linkages between environmental conditions and individual fishes, populations and communities. Such methods will
allow a higher resolution of data capture than
before, so it will be necessary to evaluate the
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consequence of these fine-resolution data and
potential for error propagation in uncovering
large-scale patterns and processes. Other new
technologies such as remote sensing, stable isotopes, fatty acids assays, and microchemistry
will all likely yield new information on movements and will allow us to integrate resource
use over larger spatial and temporal scales.
Some challenges still remain. For example, in
contrast to fine-resolution sensor technology,
will remotely sensed and biochemical data be
of sufficiently fine resolution to capture relevant ecological processes at smaller scales?
Connectivity: Spatial and Temporal
Connectivity of Stream Fish
Communities
Understanding spatial and temporal dependencies of stream fishes is critical in linking
local and regional processes. Although the
science and technology we use allows us to
measure dispersal of fishes over space and
time, the complexity of species life histories
has limited comprehensive investigations of
dispersal dynamics for many fish communities. We need innovative approaches to monitor movements of fishes across intermediate
spatial scales. Metacommunity theory (sensu
Leibold et al. 2004) has provided a framework
for understanding these dependencies, but the
theory is well beyond existing empirical research. A major hurdle, as mentioned above, is
quantifying dispersal patterns of fishes. Recent
advances in telemetry and biomarkers allow
us to tag very small fishes, but their numbers
are often too great and individuals too fragile
to mark enough fish to gain a comprehensive
assessment of movement across large spatial
and temporal scales. Understanding the consequences of connectivity across habitats will be
critical to future conservation of stream fishes.
Fausch et al. (2010) highlight the conundrum
of wanting to encourage movement of native
fishes while limiting the movement of nonnatives. Fragmentation of stream networks by
impoundments is central to this challenge, and
innovative methods of selectively allowing dispersal will greatly assist conservation issues.
Scaling: Linking Process to Pattern
A major obstacle in linking large-scale patterns to ecological processes is the scale at
which experiments are conducted (Hewitt et
al. 2007; Sandel and Smith 2009). The link
between small-scale mechanistic experiments
and observations and large-scale patterns still
presents a major challenge to the general field
of ecology (Ricklefs 2008). Several recent reviews (e.g., Jackson et al. 2001) have identified
approaches that would help link these processes. For example, mechanistic experiments
conducted along major environmental gradients (e.g., stream size and latitude) can be used
to infer the contingencies of local processes
on these larger gradients (Sandel and Smith
2009). In streams, some of the most consistent
patterns in diversity and community structure
are along gradients of stream size, productivity, and latitude (e.g., Oberdorff et al. 1995).
Experiments embedded along those gradients
are likely to provide insight into scaling local
processes.
Model Development and Validation:
Validation of Mathematical Models with
Empirical Data
The number of large-scale models (i.e., based
on satellite or climatic data) predicting distribution and abundance of stream fishes has expanded rapidly over the past few decades. This
work has been facilitated by the availability of
large data sets of both biological and environmental factors. In many cases, it is not clear if
these models are missing influential constraints
such as water chemistry or biotic interactions.
Advances in statistical modeling will occur if
synthesis and future direction
these models are able to incorporate processes.
To incorporate process, models must be based
on plausible mechanistic hypotheses and validated with empirical data. For example, a temperature model predicting species distribution
could be coupled with laboratory experiments
defining thermal tolerance of species or using
insight about habitat use derived from integrated telemetry biosensors. Or the observation of a parapatric distribution of species
could be coupled with a competition experiment. Although this sounds straightforward,
there are virtually an infinite number of experiments necessary to test all species interactions
or physiological responses of fish communities
across large spatial scales. A first step is to test
mechanisms in an appropriate venue. This may
require small mesocosms up to large, unreplicated experiments of the kind that have proven
valuable in furthering ecological understanding
and influencing policy (e.g., Schindler 1998).
Capitalizing on natural experiments also might
allow testing of mechanisms hypothesized to
be associated with large-scale processes, such
as species invasions (e.g., Smith et al. 2004).
What Role Can Stream Fish Community
Ecology Play in Advancing General
Ecological Theory?
We conclude this synthesis with a discussion
of how stream fish ecology can contribute to
general ecological theory. In particular, characteristics of streams and stream fishes make
them ideal for testing a number of general ecological theories (e.g., Ward et al. 2002b; Hugueny et al. 2010). At the largest scale, stream
catchments are “islands” and freshwater fishes
are separated from other catchments by land
or sea. These islands are often replicated across
the landscape and are useful in conducting
“natural” experiments that test patterns of fish
community structure along key environmental gradients. Thus, stream fishes continue to
657
be useful models for describing and testing
biogeographic theory (e.g., island biography
theory and community convergence of functional traits). At finer spatial scales, streams
are dendritic networks (Campbell-Grant et
al. 2007) with a hierarchical organization of
habitats (Frissell et al. 1986). Limited dispersal among network branches or unidirectional
movement of materials (Vannote et al. 1980)
also provides a backdrop for testing metapopulation and metacommunity theory (Fausch
2010). Metapopulation and metacommunity
theory has advanced rapidly, but often without empirical support with which to calibrate
models, let alone test the theory. Stream fish
ecologists are well positioned to advance this
area, especially when rigorous movement data
are able to be collected from fish communities.
Landscape ecology is often considered in a terrestrial setting with interest in the movement of
individuals among patches and across habitats,
often assuming simple but unrealistic straightline movements between patches. The sharp
aquatic-terrestrial boundary and strong longitudinal gradients make stream systems excellent models for measuring species movements
and their affinities to particular habitats. These
basic properties of streams are also useful in
testing genetic models (i.e., effective population size) because it is feasible to obtain information on the dimensions of fish movement,
dispersal barriers, and turnover rates of local
populations (Douglas and Douglas 2010).
Research on stream fishes and their prey
have advanced food-web theory (e.g., Power
1990; Nakano et al. 1999), due in part to compartmentalization of biota and resources in
streams, but also because the importance of
reciprocal connections between terrestrial and
aquatic systems has become increasingly apparent (e.g., Malison and Baxter 2010; Wesner,
in press). Moreover, rapid turnover rates of
aquatic taxa (primarily lower trophic levels)
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gido and jackson
allow us to conduct food-web manipulations
and observe responses over relatively short
intervals. Small-scale experiments of stream
organisms, but rarely with fish (e.g., Hargrave
2009), have provided information on the relationship between biodiversity and ecosystem
function (e.g., Cardinale and Palmer 2002).
However, we still have little understanding of
how stream fish diversity is associated with
ecosystem function and future research should
evaluate these associations (e.g., McIntyre et
al. 2008). A particular challenge, as highlighted in recent studies (Power et al. 2008; Murdock et al., in press), is that results are often
confounded by the complexity within major
food-web compartments (i.e., algal and macroinvertebrate taxonomic composition). Further
resolution of these interactions might be necessary to predict the context dependency of
fish effects in ecosystems.
Stream fish communities are ideal systems
for testing disturbance theory (e.g., Stanley et
al. 2010) because many streams are frequently
disturbed by flooding, drought or ice scouring
(e.g., Fisher et al. 1982; Townsend 1989; Poff
and Allan 1995; Mitchell and Cunjak 2007).
This nonequilibrium aspect of stream ecology
provides a platform to evaluate community assembly rules (i.e., priority effects, niche packing, e.g., Roberts and Hitt 2010, this volume).
Studies that couple disturbance with a landscape perspective (e.g., Fausch et al. 2002) at
intermediate scales should help to reveal key
trade-offs between species interactions and
spatial distribution of refugia. Moreover, continuing work in this area will help conservation
efforts by giving managers better understanding of how stream fishes will respond to changes in climate, hydrology, land use, and biota.
Research on stream fish communities over
the past two decades has shifted away from a
predominant focus on competition and resource partitioning (Matthews 2010). Nev-
ertheless, stream fish communities continue
to offer important opportunities to evaluate
various niche concepts. For example, stream
fishes have been used to test community
saturation and invasibility (e.g., Ross 1991;
Moyle and Light 1996) or for the importance
of density-dependent population regulation
(Grossman et al. 1998; Lobón-Cerviá 2009).
Despite previous experiments testing species
interactions (e.g., competition and predation),
we still only have a weak grasp on the role of
species interactions in regulating stream fish
community structure due to the complexity associated with factors such as ontogenetic
shifts, plasticity, spatial heterogeneity, indirect
food-web interactions, migration, and timelagged effects. Continuing long-term studies,
and studies conducted at larger spatial scales,
may reveal new insights into the importance of
biotic interactions and role of abiotic features.
In particular, small-scale mechanistic studies
combined with evaluation of large-scale patterns might be necessary to predict the causes
and consequences of species invasions.
In conclusion, we have noted that research highlighted in this symposium and in
the broader literature shows a considerable
change in its focus relative to the various foci in
the previous symposium. Ideally, this change
means that we have been addressing some of
the prominent issues from 20+ years ago and
that our field, and the broader field of ecology,
is seeing considerable progress. Alternatively,
it may represent the sometimes cyclical nature
of research questions presented in the literature (i.e., various themes repeatedly appearing at various times—sometimes under different names). However, the enthusiasm and
optimism expressed by the participants in this
symposium suggests that we are making considerable progress, especially given the many
new conceptual and technical approaches now
available. We both found that the symposium
synthesis and future direction
chapters published in Matthews and Heins
(1987) provided much stimulus for discussions and helped in the development of ideas
during our graduate work. We hope this current work also may serve to initiate and continue debate and discussion among existing and
future generations of stream fish ecologists.
Given the many challenges facing us, both in
basic and applied stream fish ecology, such
symposia and syntheses provide valuable insight into the progress made, our current state
of knowledge, and setting priorities and goals
for future studies.
Acknowledgments
We thank all participants of this symposium
and peer reviewers for thoughtful insights into
the past, present, and future challenges in the
field of stream fish community ecology. Many
of these ideas were generated from a formal
discussion following the symposium at the Ottawa meeting, as well as through informal communications with symposium participants and
other colleagues. We also thank Gary Grossman, Edie Marsh-Matthews, Erika Martin,
William Matthews, Josh Perkin, James Whitney, and Kirk Winemiller for valuable feedback
on earlier drafts of this synthesis.
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