1. Art and Diseño

American Fisheries Society Symposium 69:557–575, 2009
© 2009 by the American Fisheries Society
Fishway Evaluations for Better Bioengineering:
An Integrative Approach
Theodore Castro-Santos*
S.O. Conte Anadromous Fish Research Center, U.S. Geological Survey, Leetown Science Center
Post Office Box 796, One Migratory Way, Turners Falls, Massachusetts 01376, USA
Aline Cotel
Department of Civil and Environmental Engineering, University of Michigan
Ann Arbor, Michigan 48109, USA
Paul Webb
School of Natural Resources and the Environment, University of Michigan
Ann Arbor, Michigan 48109, USA
Abstract.—Effective fishway design requires extensive integration of biological and hydraulic data. Many relevant biological parameters remain poorly characterized, however, and
the lack of adequate biological data has long been recognized as a central weakness in fish
passage technology. This is of particular concern given the growing recognition of the importance of passing a broad diversity of species. Part of the reason for this weakness is the difficulty of identifying relevant biological, hydraulic, and other physical parameters. We propose
that by both exploring questions suggested by current knowledge, and also by increasing the
frequency and refining the methods with which fishways are evaluated, two results can be
achieved: our understanding of design effectiveness can be improved, and research questions
can be prioritized through adaptive management. We describe a framework and rationale
for fishway evaluations that identifies several promising avenues of research. Understanding
correlates of passage performance is increasingly important as fish passage needs expand on
a global scale.
Background
Fishway engineers have long recognized the importance of fish swimming capacity to fishway design.
Indeed, the recognized need to pass fish upstream
of dams and other obstacles inspired many seminal
studies on fish swimming performance, energetics,
and biomechanics (Denil 1909, 1937; Stringham
1924; McLeod and Nemenyi 1940; Bainbridge
1960; Weaver 1963, 1965; Brett 1964, 1967;
Beamish 1978). Close collaboration between biologists and fishway engineers reached a zenith in the
1950s and 1960s at the Columbia River Fisheries
Research Laboratory (CRFRL). The CRFRL enabled researchers to gather valuable information on
* Corresponding author: [email protected]
hydraulics, swimming performance, and physiological limits to passage on a scale that permitted testing
of full-scale prototype fishways with live, actively
migrating salmon, (Collins 1952, 1962; Collins
and Elling 1960; Collins et al. 1963; Weaver 1963;
1965). Many of today’s most common fishway designs were developed or modified at this facility.
By the late 1960s, however, close collaborations between fishway engineers and biologists
became less frequent. Bioengineers continued to
grapple with improving fishway performance (Haro
et al. 1998, 2004; Bunt et al. 1999; Larinier and
Travade 1999, 2002), and work on fish swimming
performance and biomechanics proliferated (Lighthill 1971, 1977; Webb 1971a, 1971b, 1975; Brett
1972, 1973, 1995; Brett and Glass 1973; Wardle
1975; Brett and Groves 1979; Alexander 1983; Vi-
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deler and Wardle 1991), but the two areas of research progressed independently, having lost the
core of integration that had existed previously.
At the time, there was a sense that sufficient
information had been gathered to provide adequate
passage (Powers et al. 1985; Bell 1991). Salmon
passed structures in large numbers, and although
fishway engineers recognized that knowledge gaps
remained, it was unclear how important these factors were (Powers et al. 1985; Orsborn and Powers
1986; Orsborn 1987); specific recommendations for
how to gather the needed data and how to prioritize
research questions were lacking. Further, both managers and engineers were primarily concerned with
salmonid passage. This taxonomic bias had deep
cultural and economic roots (Clay 1995). Although
the need for passage of other taxa has long been acknowledged, there is a growing consensus that failure to provide passage for a broad range of both diadromous and riverine species poses a serious threat
to both freshwater and terrestrial ecosystems (Meyer
et al. 2007). Part of this recognition comes from a
growing awareness of the mobility of riverine organisms, and part comes from a body of literature that
suggests that fishways often perform poorly. This is
especially true for nonsalmonid species (Slatick and
Basham 1985; Bunt et al. 1999, 2000, 2001; Bunt
2001; Boggs et al. 2004; Cooke et al. 2005), but recent data suggest that even salmon may not pass as
well as previously thought (Naughton et al. 2005).
Why some fishways work better than others
and why some species perform better than others in
fishways is poorly understood. Although some efforts have been made to identify conditions conducive to passage (Monk et al. 1989; Bunt et al. 1999,
2001; Haro et al. 1999; Bunt 2001), a coherent
framework and rationale for evaluations is lacking.
Below, we propose a framework for evaluating fishways and explore how such a framework might lead
to better integration of biology and engineering toward improved fish passage technology.
A Framework for Fishway
Evaluations
Comprehensive evaluations of fishways are only
possible if their purpose and expected product are
clearly defined. Powers et al. (1985) described the
objective of a fishway as follows:
If an artificial barrier to upstream migration is created … an alternate passage
route must be provided for the anadromous and/or resident fish. The alternate
route should not add to the level of stress
(or even total energy expenditure) compared with the level of energy expenditure
experienced by the fish under pre-[barrier]
conditions.
This description, though reasonable, does not readily lend itself to quantification. Also, it misses the
main point: persistence of populations above and
below barriers depends on protecting fitness (i.e.,
spawning success). The “predammed condition” is
almost always impossible to document, and cumulative effects of the structure itself, compromised
riverine and marine habitats, and interests of commercial and recreational fisheries may direct managers to attempt to provide superior passage than
existed before human alteration. Although such
management policies can be controversial, passage
is sometimes even provided in locations that contain natural obstacles.
The Ideal Fishway Dichotomy
A more generalized approach is to define the ideal
fishway. This is more a conceptual than a realistic
goal, but identifying ideal characteristics can serve
as a heuristic exercise for establishing a rationale for
evaluating fishway performance. The objective of an
ideal fishway, we propose, is to make the dammed
reach transparent to the movement of native species, allowing unfettered access to free-flowing
reaches above and below the obstacle (note that this
may include passage through reservoirs and other
features associated with the obstacle). If this objective is accepted, then the ideal fishway must have
the following characteristics: (1) any individual of
any native species wishing to move upstream or
downstream must be able to enter the fishway without experiencing any delay; (2) entry is immediately
followed by successful passage, with (3) no temporal
or energetic costs and (4) no stress, disease, injury,
predation, or other fitness-relevant costs associated
with passage.
This describes the ideal condition; it is also useful to contemplate the worst-case scenario. From
the biological perspective, this would be immediate
extinction of the local population. This extinction
fishway evaluations for better bioengineering
could result in several ways, for example, (1) by denying access to essential habitat, (2) by capturing and
removing all individuals, or (3) by efficiently passing
all individuals upstream (in the case of anadromous
migrants) but then failing to provide adequate downstream passage for adults and/or juveniles.
A similar continuum can be defined from an
operational perspective. Here, the objective is to design, build, and operate a structure with a minimum
of cost. An ideally designed structure from this perspective would cost nothing to construct; would require no maintenance; would use no water, power,
or other resources to operate; and would be free of
licensing restrictions. In the absence of performance
criteria, the ideal operational solution can thus be
described as doing nothing. The worst-case scenario
is essentially indeterminate, however, because it is
always possible to commit more resources to solving a particular problem, usually with diminishing
marginal gains as costs increase.
There is clearly an underlying dichotomy between biological and operational goals. However, it
is important to recognize that success in one goal
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does not necessarily imply failure in the other. Three
hypothetical scenarios illustrate some potential bioengineered outcomes (Figure 1). Consider the “obligate philopatriot,” species A: an example might be
a salmon population excluded from its spawning
habitat by a dam. In the absence of fish passage, this
population will quickly go extinct. On the other
hand, modest efforts at providing passage can yield
substantial benefits, and a biologically ideal situation might be approached without excessive costs
from the operational perspective.
A second example (“opportunistic invader,”
species B) also suggests that good passage success
may be attainable at reasonable cost. It is worth
noting here, though, that doing nothing does not
result in local extinction, merely a suboptimal condition for that species. An example of this might be
the sea lamprey Petromyzon marinus, which persists
in altered habitat but ascends certain fishways designed for salmon with seeming ease. The lamprey
example is also interesting given the experience of
the Laurentian Great Lakes. Here, the species is
invasive and massive efforts to exterminate it have
Worse
C
B
Costs
A
Best
Worst
Biological Ideal
Best
Figure 1.—A heuristic description of the intersection of operational (“Costs”) and biological ideals. Three
species are presented: species A is an obligate philopatriot (i.e., must return to specific spawning grounds to
reproduce). Failure to provide fish passage quickly drives this population to extinction. Species B is an opportunistic invader. Passage provisions help this species, but populations persist without them, and extirpation may
be difficult or impossible. Species C is a facultative philopatriot that benefits from fish passage but can also be
harmed if passage provisions are not adequate.
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failed. In this context, engineered solutions for extirpation, including construction of passage barriers, have proved unattainable, and emphasis has
shifted to control (Madenjian et al. 2002; Sullivan
et al. 2003).
A third example (“facultative philopatriot,”
species C) shows the dangers of unintended consequences; an example might be the American shad
Alosa sapidissima in its native range. Although they
often migrate long distances upriver, American shad
are able to spawn in lower reaches. Thus, the widespread extirpations of Atlantic salmon Salmo salar
that resulted from damming rivers in eastern North
America merely restricted the available spawning
habitat of American shad, and populations were
maintained at stable but reduced levels. Provisions of upstream passage, however, have proved
difficult (Foote 1976; Rideout et al. 1985; Slatick
and Basham 1985; Monk et al. 1989; Larinier and
Travade 2002; Sullivan 2004), and it appears that
potential benefits of providing passage have not always been realized. Indeed, there is some concern
that fishways, once constructed, can have negative
effects on individuals and perhaps even populations
(Foote 1976; Sullivan 2004). Although we know
of no examples of extinctions of any species that
have occurred as a result of mitigation measures, the
potential for negative results is real and should be
openly considered when evaluating performance of
these structures.
Clearly, the goal of managers and fish passage
engineers is to optimize designs with respect to both
biological and operational ideals. Intrinsic incentives exist to identify and minimize factors associated with the costs of constructing and operating
fishways. Optimization with respect to biological
goals, however, is harder to characterize. Although
fitness maximization is the desired outcome, it is
unclear how this should be measured. The only biological parameter consistently included in fishway
design is swimming performance. But is this an adequate predictor of passage success? How important
is delay? What are the behavioral variables associated with passage success, and how might they be
quantified? How do turbulence and other hydraulic
complexities influence passage performance? How
do energetics, stress, and disease associated with fish
passage affect spawning success and subsequent survival? How do all of these translate into successful
downstream migrations of juveniles and iteroparous
adults? Hidden in these questions is the threat of
uninformed compromise: how can a manager argue
for more costly design criteria if s/he is unable to
quantify benefits?
Rationale and Criteria for Fishway
Evaluations
The proliferation of questions surrounding biological components of passage success quickly becomes
overwhelming. This suggests that passage predictions from fundamental biological principles are
likely to be imprecise. Instead, this complexity of
potentially important and interacting biological
factors argues for the importance of coordinated
efforts to catalog performance of existing fishways.
Further, there is a pressing need to characterize passage performance for a range of species and through
a diversity of fishway types.
In the United States alone, there are thousands
of engineered fishways; if we consider culverts as
de facto fish passage structures, the number reaches into the millions. Despite the ubiquity of these
structures, fishway evaluations are surprisingly
rare and tend to focus on a small number of highprofile projects (Reback et al. 2004; Bernhardt
et al. 2005; Palmer et al. 2005). This is a missed
opportunity: collectively, these structures can be
considered a functional experiment; the more information becomes available on how various taxa
pass them, the easier it will become to identify and
prioritize hypotheses as to what governs passage
performance.
Information from evaluations is most useful if
performance parameters are reported in consistent
and biologically relevant terms. The more information provided the better. Thus, documentation of
spawning fish upstream, or even window counts, indicate some level of success. This type of information
can be misleading, however, and can mask significant
problems if large numbers of individuals fail to enter,
fail to pass, or otherwise incur fitness costs associated
with the structure. We propose a suite of variables that
might be quantified in fishway evaluations (Table 1).
The list is not intended to be exhaustive, nor is it reasonable to expect that all identified variables will be
quantified in each evaluation. Instead, it is intended
as a framework from which managers can draw and
assess how well fishways are being evaluated.
Note that many of the parameters identified
fishway evaluations for better bioengineering
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Table 1.—Biological indicators of fish passage success.
Biological variables
Units
Goal
Dependent variables
Proportion guided to fishway entrance Pg
Maximize Pg and dPg/dt
Proportion entering fishway (Pe)
Pe
Maximize Pe and dPe/dt
Proportion passing (Pp)
Maximize PP per units
Height (H)
Pp/m Height (dPp/dH) and
Distance (D)
Pp/m Distance (dPp/dD)
Passage time (“delay,” t)
h
Minimize t
Guidance
Pg/h Maximize dPg/dt
Entry
Pe/h Maximize dPe/dt
Passage
Pp/h Maximize dPp/dt
Failure (PF)
PF/h Context-dependenta
Condition/health
Minimize fishway effects on condition
Energy content (E)
J/kg Minimize ∆E
Change in E
Per time (dE/dt)
J/kg/h Minimize dE/Dt
Per height (dE/dH)
J/kg/m Minimize dE/dH
Per distance (dE/dD)
J/kg/m Minimize dE/dD
Stress indicators
[Cortisol ] Minimize fishway-induced stress
[Plasma ions]
[Glucose]
Sex steroids Injury
Wounds
Minimize injuries
Scale loss
Proportion surviving PS
Maximize long-term survival, including spawning
success of adults and migration success of
juveniles (survival and postmigratory
performance)
Covariates
Species
Category
Identify species-specific differences; understand forcing mechanisms
Body length
m, cm, mm
Relate size to overall passage, hydraulic and other physical effects
Temperature (T)/water quality
°C, d°C, Effect of temperature and water quality on passage
integral performance and bioenergetics Time (t)
h
Effects of residence time and seasonality on passage
success, bioenergetics
Hydraulic, physical characteristics
See Table 2
Evaluate effects on all biological dependent
variables
Depending on importance of upstream habitat and risks of prolonged residence, it might be better to abandon fishway quickly (increase dPF/dt) or retain fish to improve PP (minimize dPF/dt).
a
in Table 1 scale along biologically and structurally
relevant dimensions. Thus proportion passing is
useful information, but if it is presented as a continuous distribution with respect to fishway height
and length, a much clearer understanding of where
failure occurs emerges. Once the location of failure
has been characterized, it becomes much easier to
formulate hypotheses as to why.
Quantification along these axes will also allow for least-biased estimates of covariate effects on
failure rates. Just as mortality rates act only on the
surviving portion of a population, factors affecting
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failure rate in a fishway act on an ever-decreasing
population of ascending fish (Castro-Santos and
Haro 2006). The structural similarity between fish
passage and mortality data means that well-established methods of survival analysis can be readily
applied to fish passage studies. Successful ascenders
constitute censored observations and will bias results unless explicitly accounted for. Survival analysis methods minimize this bias; they also allow failure rate to be modeled as either constant or varying
with distance, enabling researchers to estimate both
the magnitude and location of covariate effects.
(Hosmer and Lemeshow 1999; Castro-Santos and
Haro 2003; Haro et al. 2004; Castro-Santos 2005).
Note that in these applications, the term “survival
analysis” can be misleading, given that the dependent variables may not be directly linked to survival
at all. Less ambiguous terms include “event-time
analysis,” “failure analysis,” or perhaps most generally “analysis of censored data.”
In addition to rates of passage and failure,
some effort should be made to characterize delay
incurred at the structure. The proportion of available fish entering a fishway changes over time;
because the objective is to pass fish expeditiously
through the desired route, rather than simply excluding them from less desirable routes, it is appropriate to report these data as probability functions that are continuous with respect to time,
rather than multinomial responses (Castro-Santos
and Haro 2003). Furthermore, fishway entry is
actually a two-step process, each step having a discrete zone: (1) the guidance zone, where fish are
guided to the fishway entrance; and (2) the entry
zone, where fish are able to detect the structure
and must choose to enter it. Performance within
these two zones is best measured as time-based
functions (i.e., guidance rate, the proportion of
available fish arriving at the detection zone per
unit time, and entry rate, the proportion entering
per unit time. Once again, these data are prone to
censoring, with individuals leaving the respective
zones, entering alternate routes, and so forth. Having once entered, additional delays are incurred as
fish ascend or descend the fishway, whether or not
they ultimately pass the structure.
The amount of delay incurred at fishways is
likely to be associated with fitness costs. These include energetic costs of passage, rates of injury, exposure to disease, and so forth. These factors, along
with their consequent effects on survival and recruitment, remain poorly understood. We propose
some variables to measure in Table 1 but encourage
researchers to further explore the relationship between postpassage condition and spawning success
(for adults) and successful seawater entry or adult
return rates (for juveniles). For iteroparous species,
postspawning survival is another feature that can
have serious implications for population viability
(Charnov and Schaffer 1973; Glebe and Leggett
1981). Migratory delay, both up-and downstream,
holds considerable potential to affect this important
life history stage (see also McCormick et al. 2009,
this volume), and the cumulative effects of multiple
barriers should be considered wherever they occur.
Many of the issues laid out in the preceding
paragraphs have long been recognized (British Institution of Civil Engineers 1942; Powers et al. 1985;
Orsborn 1987). Recent advances in animal tracking technology and analytical techniques, however,
have now made it possible to address these questions directly. Passive integrated transponder telemetry promises to supplant visible external and coded
wire tags for many mark–recapture experiments
(Prentice et al. 1990; Skalski et al. 1998; Axel et al.
2005) and has also been used to provide fine-scale
spatiotemporal data on behaviors associated with
passage performance (Castro-Santos et al. 1996;
Lucas et al. 1999; Sullivan 2004). Both radio and
acoustic telemetry technology are being used to collect movement data over scales ranging from submeter to intercontinental. Telemetry of physiological
and biomechanical processes (e.g., McKinley and
Power 1992; Hinch et al. 1996; Lowe 1996; Lowe
et al. 1998) also holds the potential to improve our
understanding of fishway energetics. More work is
needed in this area, though, especially with regard
to contributions of anaerobic metabolism during
passage through zones of high velocity.
In some cases, these technological advances
have outstripped the statistical methodology needed
to fully exploit them. There is an ongoing need for
new and adapted techniques for gleaning as much
information as possible from the data these technologies provide (e.g., Turchin 1998; Skalski et al.
2001; Castro-Santos and Haro 2003; Johnson et
al. 2004; Castro-Santos 2006). This is particularly
important with respect to guiding fish toward desirable passage routes and away from undesirable
ones. Simply excluding fish from hazardous routes
fishway evaluations for better bioengineering
may incur added delays that offset the benefits of
safer passage. Telemetry and hydroacoustic systems
can provide a wealth of fine-scale data on threedimensional movements, but effective methods for
using these data to quantify attraction have yet to
be demonstrated.
In addition to biological indicators of passage
performance, concurrent reporting of engineering
and hydraulic design specifications of the fishway
under evaluation is also important. Here again,
because we have limited information on hydraulic
correlates of passage success, it makes sense to seek
the most detailed descriptions possible. Current
standard criteria for fish passage include maximum
and mean flow velocity, total kinetic energy (or the
related energy dissipation factor [EDF]), and various dimensions describing pools, chutes, weirs, and
so forth. These, however, have proven to be poor
predictors of passage performance, and more detail
is needed. Recent work shows that eddy structure,
vorticity, and circulation all affect fish behavior and
may be important determinants of fishway performance (Crowder and Diplas 2002; Dabiri 2005;
Tritico et al. 2007). We propose a suite of structural
and hydraulic variables to quantify in Table 2. These
criteria can ultimately be used as covariates to help
explain biological performance measures.
Here again, technological advances hold great
potential for improving our understanding of hydraulics, both in engineered fishways and in natural
rivers. In particular, flow visualization using digital
particle image velocimetry and computational fluid
dynamics modeling can provide much more detailed
descriptions of fishway hydraulics than were previously available (e.g., Khan 2006; Cea et al. 2007).
These same technologies have also allowed significant advances in fish locomotion theory (Fish and
Lauder 2006; Schultz and Webb 2002), with the
intriguing possibility that hydraulic complexity and
biomechanics could be linked to produce improved
fishway designs.
Below, we pose several questions, based on current and developing knowledge, to serve as a starting point for future fish passage research. As more
data become available, it will be possible, and indeed necessary, to refine these questions. In this way,
understanding of bioengineering factors underlying
fishway performance can progress. Ultimately, this
should lead toward development of generalized predictive models to help direct management choices
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for site-specific fishway design. Developing questions may still function on the level of prototype
fishways (e.g., distance between resting pools as
predictors of passage performance). However, as hydraulic and biological details of fish passage emerge,
the moment-to-moment experience of ascending
fish will become more apparent. As these experiences become linked to performance measures, more
basic biological questions can be prioritized and
tested at the organismal level.
Some Questions Suggested by the
Current Literature
Question 1: How Does Turbulence
Structure Influence Swimming
Performance?
Technical fishways are structures engineered using
hydraulic principles and (sometimes) swimming
performance estimates—generally, these are built of
standard construction materials and make no effort
to mimic a natural river. These fishways typically use
turbulence to dissipate head, with design prescriptions including maximum EDF values. Turbulence
may be defined in various ways, and it is likely that
these affect conclusions regarding turbulence effects
on fish swimming performance or other biologically
relevant parameters. Most studies find turbulence
increases costs of swimming performance, irrespective of how it is defined or induced in experimental
systems (Powers et al. 1985; Orsborn 1987). Recent work by Enders et al. (2003, 2005) also seems
to support the practice of limiting turbulence in
fishways. On the other hand, Nikora et al. (2003)
found negligible effects of turbulence created by
wavy walls on swimming ability of inanga Galaxius maculatus. Other studies show that turbulence
may be beneficial, certainly promoting resting opportunities and probably facilitating translocation
through fishways. Thus Liao et al. (2003a, 2003b;
Liao 2004) found that rainbow trout Oncorhynchus
mykiss swimming in the presence of a controlled
von Karman vortex street appeared to surf on the
vortices, experiencing substantial reductions in activity of swimming muscles and presumably saving
energy in the process. Webb (1998a) also described
reduced swimming activity by river chub Nocomis
micropogon and smallmouth bass Micropterus dolo-
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Table 2.—Engineering and hydraulic descriptors of fishways. These variables should be considered when
evaluating design effects on passage performance.
Variable
Design
Design type
Dimensions
Chute
Pool
Weir
Total length
Number of pools by pool type
Slope Resting/turn pools
Dimensions
Elevation
Distance
Hydraulics
Reynolds number
Froude number
Energy dissipation factor
Total kinetic energy
Flow velocity
Minimum
Maximum
Mean
Turbulence
SD of velocity (“RMS”; three-dimensional)
Vortices
Size
Shedding frequencya Rotation period
Vorticity
Impulse: linear
Impulse: angular
Force or thrust
Circulation
a
Units
Category
m, m2, m3
m, m2, m3
m, m2
m
Integer
m rise: m run or degrees from horizontal
m, m2, m3
m, m2, m3
m
m
–
–
J/m3
J/m3
m/s
m/s
m/s
m/s
m
Hz
s
rad/s
kg m/s
kg m2/s
kg m/s2
m2/s
Shedding frequency can also be presented as period (seconds elapsed between vortices) or Strouhal number.
mieu swimming behind cylinders, and similar use of
refuges is well known among stream species.
Evidently turbulence is not intrinsically costly.
This fact raises the tantalizing possibility that turbulence structure might be controlled in such a way
that hydraulic head is dissipated without reducing
passage performance. Indeed, it may be possible to
enhance passage through judicious control of turbulence structure. Before such applications could be
realized, though, much more detail is needed to understand the correlations between turbulence struc-
ture and swimming performance, from mechanical,
energetic, and behavioral perspectives.
One approach that might elucidate interactions of kinematics and energetics with respect to
swimming in turbulent flow is to explore turbulence structure in natural environments (Crowder
and Diplas 2002; Cotel et al. 2006; Tritico et al.
2007). Using developing technology like the portable small-scale Particle Image Velocimetry system
(Tritico et al. 2007), researchers can obtain instantaneous measurements of velocity and vorticity at
fishway evaluations for better bioengineering
high spatial resolution. The detailed information on
flow structure that this provides should help answer
questions arising from the work described above:
What ranges of size, period, and vorticity of eddies
occur in the natural environment? How readily do
fish pass flows with these various characteristics?
How do fishways compare with natural rivers in
this respect? Are there limits to design parameters
for technical fishways? Depending on the answers,
these questions could provide feedback for prioritization of design schemes.
Question 2: How Does Morphology
Affect Fish Passage?
A common concern regarding fish passage is that
fishways are often size-selective (Powers et al. 1985).
This does not necessarily reflect swimming ability
(Bunt et al. 2000), especially when it is the smaller
individuals that pass (Sullivan 2004). Perhaps hydraulic structure somehow interacts with the morphology of the fish to produce selective passage.
Several morphological characters besides body
length affect swimming functions and performance
and hence could be implicated. These include body
depth, general shape (gobbose versus fusiform versus elongate), number and structure of fins, and so
forth. Morphological form may vary to some extent
within species, and variation is large among species.
Aspects of such interspecific variation also should
include consideration of musculoskeletal systems
(Johnston 2001), sensory apparatus (Coombs and
Montgomery 1999), and life history (Northcote et
al. 1970). Advances in this area could result from
comparative data on passage performance, but also
from smaller scale information relating swimming
capacity in controlled environments to controlled
hydraulic conditions.
Question 3: Does Sensory Physiology
Affect Passage Performance?
The lateral line system comprises an important
sensory modality with which fish detect and process certain hydraulic stimuli. Lateral line morphology and associated neural processing vary
with life history and habitat type, with fish inhabiting lentic and lotic environments having
structures that are relatively more and less sensitive to high-frequency disturbances, respectively
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(Coombs et al. 1988; Mogdans and Bleckmann
2001; Engelmann et al. 2003). Could this factor
be implicated in passage performance? Data from
the Connecticut River (Sullivan 2004 and personal
observation) show a stark contrast between passage
performance of Atlantic salmon Salmo salar and
American shad Alosa sapidissima. Both species are
strong swimmers (Beamish 1978; Peake et al. 1997;
Castro-Santos 2005), but fishways and turbulent
zones that salmon traverse with ease pose barriers to
shad movements (Sullivan 2004; Castro-Santos and
Haro 2005). Moreover, many shad do not abandon
the fishways, but hold, sometimes for weeks, in the
relatively quiescent waters of resting pools, where
they gradually sicken and die. Why?
Engelmann et al. (2003) found that goldfish
Carassius auratus were much more sensitive than
trout to stimuli in moving water. Like the goldfish,
clupeids possess a lateralis structure that appears to
be designed to be sensitive to absolute velocity and
small perturbations in flow. Could this heightened
sensitivity explain the avoidance behaviors these
fish appear to exhibit in the presence of turbulent
flow? Perhaps the hydraulic environments common
to salmon fishways are too noisy for American shad
to pass. Larinier and Travade (2002) suggest that
fishways of even moderate height seem to be problematic for shad. Given their swimming ability, it is
unlikely that this is a result of fatigue. Could it be
a stress response that triggers these fish to abandon
their migrations?
Question 4: How Does Behavior
Affect Fish Passage?
Connections between sensory physiology and behavior are also important in influencing entry rates.
Flow velocity, acceleration, turbulence, and discharge, as well as temperature, pH, and CO2 concentrations, have all been shown to influence rates
of entry by fish into open-channel flumes (Collins
1952; Weaver 1963; Haro et al. 1998; CastroSantos 2002, 2004). How and why these behaviors
vary among species remains nearly unexplored, even
though optimizing conditions for entry into fishways remains one of the most important and challenging problems of fish passage (Bunt 2001).
Other aspects of behavior may influence passage
performance as well. In one fishway, passage rates of
alewife Alosa pseudoharengus declined as density in-
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creased (Dominy 1973). How do social interactions
like schooling behavior affect passage performance?
What role might vision or other sensory modalities
play in this? Similarly, the presence of conspecifics
upstream has been shown to improve attraction and
passage of sea lamprey (Sorensen and Vrieze 2003).
Other chemical cues also influence homing ability
and migratory behavior (Hasler 1971; Carruth et
al. 2002; Sorensen and Stacey 2004), suggesting the
potential exists to influence passage performance,
especially among depleted populations.
The well-known biotic factors competition and
predation might also influence passage performance.
Fish passageways tend to concentrate fishes at greater
densities than in adjacent natural habitats. In addition, preferred tracks through a fishway may be limited, such that fish compete for them. Predation both
above and below dams remains a major concern surrounding fish passage and is one reason that passage
delays should be minimized (Venditti et al. 2000; Agostinho et al. 2002; Laake et al. 2002; Aarestrup and
Koed 2003). If fishway entrances act efficiently to
attract target species, they might also prove attractive
to predators, which could then intercept the fish or
otherwise exclude them from the fishway. We know
of no studies that evaluate this question, but the potential is troubling and worthy of examination.
One obvious way in which behavior can influence swimming performance is through volitional
gait selection and swim speed optimization. Existing
designs tend to be overly conservative with respect to
flow velocity, in part because methods typically used
to quantify swimming capacity are themselves intrinsically conservative. Most studies of fish locomotion
have used enclosed chambers or respirometers (Brett
1964). By restricting the movement of study animals,
these devices allow researchers to closely monitor and
control flow velocity and minimize turbulence while
offering a fixed frame of reference for the researcher.
Such enclosed spaces reduce swimming performance,
however, and fish that are allowed to swim volitionally in open-channel flow exhibit gait shifts and related behaviors that substantially expand their ability
to traverse velocity barriers (Peake 2004; Peake and
Farrell 2004, 2005; Tudorache et al. 2007). Furthermore, the speeds at which unrestrained fish actually
swim and the ability to stage multiple attempts can
dramatically affect distance of ascent, depending on
how closely they approximate the distance-maximizing optimum (Castro-Santos 2004, 2005, 2006).
Data from fish swimming in open-channel flow cast
doubt on the one biological performance component
that is consistently incorporated into fishway design.
Characterizing the magnitude of error associated
with the wealth of literature describing swimming capacity of constrained fish would be very helpful and
could yield considerable benefits in improved design
efficiencies.
Question 5: Do Fishways Affect Disease
Transmission?
Because diadromous species are often under stress
when they enter freshwater, they may be more
susceptible to disease or other factors that reduce
fitness. The example cited above (Dominy 1973),
where alewife pass more slowly at higher densities,
suggests that there may be greater risk of disease
transmission under these conditions. Also, migrating fish often suffer injuries and scale loss, either
within or outside the fishway. This fact, along with
elevated densities and increased stress, will likely act
to increase rates of disease transmission. Although
the potential consequences seem significant (Foote
1976; Powers et al. 1985; Tierney and Farrell 2004)
and extensive studies exist with respect to rates of
injury associated with downstream passage (e.g.,
Reischel and Bjornn 2003; Boggs et al. 2004; Cada
et al. 2006), we know of no studies examining fishway-induced injury and disease transmission.
Question 6: What Are the Energetic
Consequences of Fish Passage?
One of the more exciting technologies to arise in recent years is in the field of physiological telemetry.
Using radio tags that transmit rectified and integrated
electromyograms (EMG tags), researchers have been
able to identify zones of difficult passage within rivers
and even to project the consequences of the energetic
costs incurred there (Hinch et al. 1996; Hinch and
Rand 1998; Rand and Hinch 1998; Hinch and Bratty 2000). These data have far-reaching implications,
including the likely consequences of climate change
(Rand et al. 2006). Although some concern remains
regarding the reliability of the data this technology
provides (Geist et al. 2002), its potential for fishway
research is obvious.
Concerns over the energetic costs of fish passage
and migration motivated several seminal studies (e.g.,
fishway evaluations for better bioengineering
Idler and Clemens 1959; Brett 1962). These concerns are particularly acute for those species that do
not feed during their migrations: a primary measure
of a successful migration is that the animals arrive at
their habitat with sufficient energy reserves to spawn
successfully and, in the case of iteroparous migrants,
to return to their feeding habitat. Indirect observations of fishway energetics have been performed by
collecting fish above and below structures (e.g., Leonard and McCormick 1999), but direct observations
are lacking. With improved data on activity levels of
fish ascending fishways, it will become possible to
measure their effect on energy depletion.
Energetics, while affecting fitness, is itself affected by environmental and behavioral factors. As
mentioned above, turbulent flow affects swimming
energetics, but mechanisms and magnitude of this
effect are still ambiguous. Delay or residence time
clearly can be expected to have an effect, but this
will depend on where the fish spend most of their
time. If fish exhibit anaerobically fuelled bursting
behavior, this will have much higher costs than if
they are able to swim at aerobically powered speeds
(Boisclair and Tang 1993). It is worth noting that
the EMG tags mentioned above do not accurately
transmit data from anaerobic swimming behaviors,
a limitation that may be overcome using alternate
technologies (e.g., Lowe et al. 1998). Likewise, temperature will influence swimming energetics. This
has implications both for climate change and for
more immediate anthropogenic thermal impacts
such as power plant effluents (Mustard et al. 1999).
Energetics may also be associated with disease risk
and reproductive physiology.
Question 7: How Do Hormonal Levels
Influence Fish Passage?
Hormonal ontogeny is closely linked to migratory
behavior throughout the Animalia (Dingle 1996).
Among fish, this effect is particularly well documented for out-migrating salmonids (Hoar 1988;
McCormick 1994). Although hormonal correlates
of adult salmonid migrations have also been documented (Ueda et al. 1984; Leonard et al. 2001),
we are aware of only one investigation that linked
changes in circulating levels of hormones to upstream migratory behavior (Sato et al. 1997).
How and whether the physical environment
within and around fishways influences hormonal lev-
567
els of migratory fish is an open question. Stress in
fish is known to affect the timing of reproduction,
behavior during spawning, and the survival of offspring (Schreck et al. 2001). Some data exist to show
that stress levels increase and sex steroids decrease in
association with fishway residence and that this varies
by fishway type (D. T. Lerner and S. D. McCormick,
U.S. Geological Survey, S.O. Conte Anadromous
Fish Research Center, personal communication)
Given growing concerns regarding endocrine
disrupting compounds in the environment (McCormick 2009), the potential influence of these
compounds on migratory motivation and performance is worthy of examination.
Despite the scope of topics covered in the
preceding questions, this list is neither exhaustive
nor comprehensive. As stated above, improved information on performance of existing fishways will
at once expand and focus the list; and this result
will be best attained if evaluations are conducted in
ways that provide the greatest possible scope and detail on biologically relevant performance measures.
Such data will improve measures of biological effectiveness (or lack thereof ). This in turn will allow
for better cost–benefit analysis and ultimately optimized bioengineered fishway designs.
It is also worth noting that the preceding sections have a pronounced bias toward improving upstream passage. Many of the same principles apply
to downstream passage, though, and we in no way
wish to diminish the importance of this issue. To
the contrary, the concept of transparency requires
that passage be facilitated in both directions, and
improved downstream passage technology remains
one of the most pressing needs in diadromous
fishery management. Often, solutions for up- and
downstream passage are considered separately. This
is a practical matter that has to do with different flow
requirements for up- and downstream fishways. In
the same way, it is likely that improved passage for a
range of species will only be achieved by providing
more than one type of passage structure at a given
location.
The Qualified Promise of
Nature-Like Fishways
Inevitably, fish passage structures are designed on a
site-specific basis, and engineers must rely on the
568
castro-santos et al.
best available knowledge in order to adapt the structures to local conditions. Insufficient information
on behavior can lead to a dilemma: should the design be conservative and maintain low flow velocities? If so, there is a risk that the conditions will not
stimulate the fish to move through. Conservative
designs are often recommended by management
agencies, despite the possibility of a reduced cost–
benefit ratio. This trend troubled Orsborn (1987),
and solutions remain elusive.
One approach to the one-size-fits-all problem
of fishway technology is to design fishways that provide diverse hydraulic conditions. This is the rationale most often proposed for constructing so-called
nature-like fishways. The theory underlying these
designs is that if the fishway can adequately mimic
its surrounding environment, including both dimensional diversity of substrate and spatially diverse
hydraulic characteristics (called “stream simulation”
in the culvert literature; Bates et al. 2003), then all
native fauna (including invertebrates) should be able
to pass it or even occupy it as habitat. This notion
is intuitively appealing, and nature-like fishways are
being widely constructed in Europe, with increasing
support elsewhere (Jungwirth 1996; Eberstaller et
al. 1998; Santos et al. 2005).
As with technical fishways, the enthusiasm to
construct nature-like fishways has preceded and exceeded the willingness or ability to evaluate them
(Elvira et al. 1998; FAO 2002). Available data indicate that they do sometimes meet their objectives of
passing a diversity of species (Eberstaller et al. 1998;
Santos et al. 2005). This success is not universal,
however, even for salmonids (Aarestrup et al. 2003),
and passage for some species has proved ineffective
(Schmutz et al. 1998).
A further criticism of the nature-like approach
is that these fishways tend to have very low slopes
and so require extensive resources in land and materials to construct. They also can require a great deal
of flow to operate (after all, it is difficult to simulate
a natural river without passing a significant amount
of the total discharge through the structure). Moreover, even by attempting to simulate the natural
features of rivers questions remain as to the requirements of various fish species; the fact that it might
look natural to a human does not eliminate this
concern. Nevertheless, the nature-like approach explicitly recognizes the lack of supporting knowledge
for effective technical fishway design and so holds
real promise in the absence of reliable technical
solutions. Perhaps the added costs associated with
these structures will create an economic incentive to
gather the information needed for more economical technical solutions (e.g., Stuart and Berghuis
2002).
Surrogate Species and the
Big Picture
Perhaps the single greatest challenge in fish passage technology is the development of structures
and design concepts that will pass a broad range of
species. Of particular concern is the development
of hydropower in regions with diverse and poorly
understood fish stocks. Tremendous hydropower
resources exist in countries like Brazil and China
(Zhong and Power 1996; Anderson et al. 2009, this
volume). Often, these same systems provide habitat for economically, sociologically, and ecologically
important species, and planned and ongoing development may obstruct vital migratory corridors (Agostinho et al. 2002, 2005; Fernandes et al. 2004).
Biological data relevant to passage performance
are insufficient even among the comparatively sparse
native fauna in developed countries; this problem is
even more acute in those regions currently undergoing the most rapid economic development. Because
it is unrealistic to expect that all of the necessary data
be acquired for the full diversity of taxa, managers
are faced with the difficult task of identifying surrogate species that can be expected to be representative of local fauna. Although broad generalizations
have been proposed linking body form to function
and behavior (e.g., Webb 1998b; Webb and Gerstner 2000), this link is largely lacking with respect
to fish passage. Even more troubling is the extent
of intraspecific diversity of swimming performance;
differences exist among individuals and among populations, leading some to question the reliability of
even species-specific models (Northcote et al. 1970;
Taylor and Mcphail 1986; Castro-Santos 2006;
Castro-Santos and Haro 2006). Certainly, it is unrealistic to expect existing salmonid-based models
to be of much value on a global scale (Santos et al.
2007).
Identification of surrogate species or trait characteristics associated with passage would be a major step toward improving fish passage worldwide.
fishway evaluations for better bioengineering
In this paper, we have attempted to identify the
biological variables most likely to lead to this goal.
Further work on relations between hydraulics and
morphology, sensory biology, life history, behavior,
disease, energetics, and endocrinology can all help
in the development of improved fishway technology. The most pressing question is how to prioritize
these areas of research. By starting from the natural
experiments that comprise existing structures while
focusing on both biological and design objectives,
practical solutions are likely to emerge.
References
Aarestrup, K., and A. Koed. 2003. Survival of migrating
sea trout (Salmo trutta) and Atlantic salmon (Salmo
salar) smolts negotiating weirs in small Danish rivers. Ecology of Freshwater Fish 12:169–176.
Aarestrup, K., M. C. Lucas, and J. A. Hansen. 2003. Efficiency of a nature-like bypass channel for sea trout
(Salmo trutta) ascending a small Danish stream studied by PIT telemetry. Ecology of Freshwater Fish
12:160–168.
Agostinho, A. A., L. C. Gomes, D. R. Fernandez, and H.
I. Suzuki. 2002. Efficiency of fish ladders for neotropical ichthyofauna. River Research & Applications
18:299–306.
Agostinho, A. A., S. M. Thomaz, and L. C. Gomes. 2005.
Conservation of the biodiversity of Brazil’s inland
waters. Conservation Biology 19:646–652.
Alexander, R. M. 1983. The history of fish mechanics.
Pages 1–35 in P. W. Webb and D. Weihs, editors.
Fish biomechanics. Praeger, New York.
Anderson, E., M. Montoya, A. Soto, H. Flores, and M.
McClain. 2009. Challenges and opportunities for
co-management of a migratory fish (Prochilodus nigricans) in the Peruvian Amazon. Pages 741–756 in
A. J. Haro, K. L. Smith, R. A. Rulifson, C. M. Moffitt, R. J. Klauda, M. J. Dadswell, R. A. Cunjak, J.
E. Cooper, K. L. Beal, and T. S. Avery, editors. Challenges for diadromous fishes in a dynamic global environment. American Fisheries Society, Symposium
69, Bethesda, Maryland.
Axel, G. A., E. F. Prentice, and B. P. Sandford. 2005. PITtag detection system for large-diameter juvenile fish
bypass pipes at Columbia River basin hydroelectric
dams. North American Journal of Fisheries Management 25:646–651.
Bainbridge, R. 1960. Speed and stamina in three fish.
Journal of Experimental Biology 37:129–153.
Bates, K., B. Barnard, B. Heiner, J. P. Klavas, and P. D.
Powers. 2003. Design of road culverts for fish passage. Washington Department of Fish and Widlife,
Olympia.
569
Beamish, F. W. H. 1978. Swimming capacity. Pages 101–
187 in W. S. Hoar and D. J. Randall, editors. Fish
physiology, volume VII. Locomotion. Academic
Press, London.
Bell, M. C. 1991. Fisheries handbook of engineering requirements and biological criteria. U.S. Army Corps
of Engineers, Portland, Oregon.
Bernhardt, E. S., M. A. Palmer, J. D. Allan, G. Alexander,
K. Barnas, S. Brooks, J. Carr, S. Clayton, C. Dahm,
J. Follstad-Shah, D. Galat, S. Gloss, P. Goodwin,
D. Hart, B. Hassett, R. Jenkinson, S. Katz, G. M.
Kondolf, P. S. Lake, R. Lave, J. L. Meyer, T. K.
O’Donnell, L. Pagano, B. Powell, and E. Sudduth.
2005. Synthesizing US river restoration efforts. Science 308:636–637.
Boggs, C. T., M. L. Keefer, C. A. Peery, T. C. Bjornn,
and L. C. Stuehrenberg. 2004. Fallback, reascension, and adjusted fishway escapement estimates for
adult Chinook salmon and steelhead at Columbia
and Snake River dams. Transactions of the American
Fisheries Society 133:932–949.
Boisclair, D., and M. Tang. 1993. Empirical-analysis of
the influence of swimming pattern on the net energetic cost of swimming in fishes. Journal of Fish
Biology 42:169–183.
Brett, J. R. 1962. Some considerations in the study of
respiratory metabolism in fish, particularly salmon.
Journal of the Fisheries Research Board of Canada
19:1025–1038.
Brett, J. R. 1964. The respiratory metabolism and swimming performance of young sockeye salmon. Journal of the Fisheries Research Board of Canada
21:1183–1226.
Brett, J. R. 1967. Swimming performance of sockeye
salmon in relation to fatigue time and temperature.
Journal of the Fisheries Research Board of Canada
24:1731–1741.
Brett, J. R. 1972. Metabolic demand for oxygen in fish,
particularly salmonids, and a comparison with other
vertebrates. Respiration Physiology 14:151–170.
Brett, J. R. 1973. Energy expenditure of sockeye salmon
(Oncorhynchus nerka) during sustained performance.
Journal of the Fisheries Research Board of Canada
30:1799–1809.
Brett, J. R. 1995. Energetics. Pages 3–68 in C. Groot, L.
Margolis, and W. C. Clarke, editors. Physiological
ecology of Pacific salmon. UBC Press, Vancouver.
Brett, J. R., and N. R. Glass. 1973. Metabolic rates and
critical swimming speeds of sockeye salmon (Oncorhynchus nerka) in relation to size and temperature.
Journal of the Fisheries Research Board of Canada
30:379–387.
Brett, J. R., and T. D. D. Groves. 1979. Physiological
energetics. Pages 279–352 in W. S. Hoar, D. J. Randall, and J. R. Brett, editors. Fish physiology. Vol-
570
castro-santos et al.
ume 8: bioenergetics and growth. Academic Press,
New York.
British Institution of Civil Engineers 1942. Report of the
committee on fish-passes. William Clowes and Sons,
London (UK) (Reprinted 1948).
Bunt, C. M. 2001. Fishway entrance modifications enhance fish attraction. Fisheries Management and
Ecology 8:95–105.
Bunt, C. M., S. J. Cooke, and R. S. McKinley. 2000.
Assessment of the Dunnville fishway for passage of
walleyes from Lake Erie to the Grand River, Ontario. Journal of Great Lakes Research 26:482–488.
Bunt, C. M., C. Katopodis, and R. S. McKinley. 1999.
Attraction and passage efficiency of white suckers
and smallmouth bass by two Denil fishways. North
American Journal of Fisheries Management 19:793–
803.
Bunt, C. M., B. T. van Poorten, and L. Wong. 2001. Denil fishway utilization patterns and passage of several warmwater species relative to seasonal, thermal
and hydraulic dynamics. Ecology of Freshwater Fish
10:212–219.
Cada, G., J. Loar, L. Garrison, R. Fisher, and D. Neitzel.
2006. Efforts to reduce mortality to hydroelectric
turbine-passed fish: locating and quantifying damaging shear stresses. Environmental Management
37:898–906.
Carruth, L. L., R. E. Jones, and D. O. Norris. 2002.
Cortisol and pacific salmon: a new look at the role
of stress hormones in olfaction and home-stream
migration. Integrative & Comparative Biology
42:574–581.
Castro-Santos, T. 2002. Swimming performance of upstream migrant fishes: new methods, new perspectives. Doctoral dissertation. University of Massachusetts, Amherst.
Castro-Santos, T. 2004. Quantifying the combined effects of attempt rate and swimming capacity on passage through velocity barriers. Canadian Journal of
Fisheries and Aquatic Sciences 61:1602–1615.
Castro-Santos, T. 2005. Optimal swim speeds for traversing velocity barriers: an analysis of volitional highspeed swimming behavior of migratory fishes. Journal of Experimental Biology 208:421–432.
Castro-Santos, T. 2006. Modeling the effect of varying
swim speeds on fish passage through velocity barriers. Transactions of the American Fisheries Society
135:1230–1237.
Castro-Santos, T., and A. Haro. 2003. Quantifying migratory delay: a new application of survival analysis
methods. Canadian Journal of Fisheries and Aquatic
Sciences 60:986–996.
Castro-Santos, T., and A. Haro. 2005. Turners Falls fish
passage studies 2005: results from PIT and radio telemetry studies. U.S. Geological Survey, Biological
Resource Discipline, S.O. Conte Anadromous Fish
Research Center, Internal Report # 2005-04, Turner
Falls, Massachusetts.
Castro-Santos, T., and A. Haro. 2006. Biomechanics
and fisheries conservation. Pages 469–523 in R. E.
Shadwick and G. V. Lauder, editors. Fish physiology. Volume 23: Fish biomechanics. Academic Press,
New York.
Castro-Santos, T., A. Haro, and S. Walk. 1996. A passive integrated transponder (PIT) tagging system for
monitoring fishways. Fisheries Research 28:253–
261.
Cea, L., L. Pena, J. Puertas, M. E. Vazquez-Cendon, and
E. Pena. 2007. Application of several depth-averaged turbulence models to simulate flow in vertical slot fishways. Journal of Hydraulic Engineering
133:160–172.
Charnov, E. L., and W. M. Schaffer. 1973. Life-history
consequences of natural selection: Cole’s result revisited. American Naturalist 107:791–793.
Clay, C. H. 1995. Design of fishways and other fish facilities. Lewis Publishers, Boca Raton, Florida.
Collins, G. B. 1952. Factors influencing the orientation
of migrating anadromous fishes. Fisheries Bulletin
52:375–396.
Collins, G. B. 1962. Ability of salmonids to ascend high
fishways. Transactions of the American Fisheries Society 91:1–7.
Collins, G. B., and C. H. Elling. 1960. Fishway research
at the Fisheries-Engineering Research Laboratory.
U.S. Fish and Wildlife Service Circular 98.
Collins, G. B., C. H. Elling, J. R. Gauley, and C. S.
Thompson. 1963. Effect of fishway slope on performance and biochemistry of salmonids. Fisheries
Bulletin 63:221–253.
Cooke, S. J., C. M. Bunt, S. J. Hamilton, C. A. Jennings,
M. P. Pearson, M. S. Cooperman, and D. F. Markle.
2005. Threats, conservation strategies, and prognosis for suckers (Catostomidae) in North America:
insights from regional case studies of a diverse family of non-game fishes. Biological Conservation
121:317–331.
Coombs, S., J. Janssen, and J. F. Webb. 1988. Diversity of
lateral line systems: evolutionary and functional considerations. Pages 553–593 in J. Atema, R. R. Fay, A.
N. Popper, and W. N. Tovolga, editors. Sensory biology of aquatic animals. Springer-Verlag, New York.
Coombs, S., and J. C. Montgomery. 1999. The enigmatic
lateral line system. Pages 319–361 in R. R. Fay and
A. N. Popper, editors. Comparative hearing: fish
and amphibians. Springer, New York.
Cotel, A. J., P. W. Webb, and H. Tritico. 2006. Do brown
trout choose locations with reduced turbulence?
Transactions of the American Fisheries Society
135:610–619.
fishway evaluations for better bioengineering
Crowder, D. W., and P. Diplas. 2002. Vorticity and circulation: spatial metrics for evaluating flow complexity in stream habitats. Canadian Journal of Fisheries
and Aquatic Sciences 59:633–645.
Dabiri, J. O. 2005. On the estimation of swimming and
flying forces from wake measurements. Journal of
Experimental Biology 208:3519–3532.
Denil, G. 1909. Les eschelles a poissons et leru application aux barrages des Meuse et d’Ourthe. Annales
des Travaux Publics de Belgique 2:1–152.
Denil, G. 1937. La mecanique du poisson de riviere: les
capacites mecaniques de la truite et du saumon. Annales des Travaux Publics de Belgique 38:412–423.
Dingle, H. 1996. Migration: the biology of life on the
move. Oxford University Press, New York.
Dominy, C. L. 1973. Effect of entrance-pool weir elevation and fish density on passage of alewives (Alosa
pseudoharengus) in a pool and weir fishway. Transactions of the American Fisheries Society 102:398–
404.
Eberstaller, J., M. Hinterhofer, and P. Parasiewicz. 1998.
The effectiveness of two nature-like bypass channels
in an upland Austrian river. Pages 363–383 in M.
Jungwirth, S. Schmutz, and S. E. Weiss, editors. Fish
migration and fish bypasses. Fishing News Books,
Oxford, UK.
Elvira, B., G. G. Nicola, and A. Almodovar. 1998. A catalogue of fish passes at dams in Spain. Pages 203–207
in M. Jungwith, S. Schmutz, and S. Weiss, editors.
Fish migration and fish bypasses. Fishing News
Books, Oxford, UK.
Enders, E. C., Boisclair, D., and Roy, A.G. 2003. The effect of turbulence on the cost of swimming for juvenile Atlantic salmon (Salmo salar). Canadian Journal
of Fisheries and Aquatic Sciences 60:1149–1160.
Enders, E. C., D. Boisclair, and A. G. Roy. 2005. A model
of total swimming costs in turbulent flow for juvenile Atlantic salmon (Salmo salar). Canadian Journal
of Fisheries and Aquatic Sciences 62:1079–1089.
Engelmann, J., S. Krother, H. Bleckmann, and J. Mogdans. 2003. Effects of running water on lateral line
responses to moving objects. Brain, Behavior &
Evolution 61:195–212.
FAO (Food and Agriculture Organization of the United
Nations). 2002. Fish passes. Food and Agriculture
Organization of the United Nations, Rome.
Fernandes, C. C., J. Podos, and J. G. Lundberg. 2004.
Amazonian ecology: tributaries enhance the diversity of electric fishes. Science 305:1960–1962.
Fish, F. E., and G. V. Lauder. 2006. Passive and active
flow control by swimming fishes and mammals. Annual Review of Fluid Mechanics 38:193–224.
Foote, P. S. 1976. Blood lactic acid levels and mortality of American shad (Alosa sapidissima) utilizing
the Holyoke dam fishlift Massachusetts, 1974 and
571
1975. Master’s thesis. University of Massachusetts,
Amherst.
Geist, D. R., R. S. Brown, K. Lepla, and J. Chandler.
2002. Practical application of electromyogram radiotelemetry: the suitability of applying laboratoryacquired calibration data to field data. North American Journal of Fisheries Management 22:474–479.
Glebe, B. D., and W. C. Leggett. 1981. Latitudinal differences in energy allocation and use during the
freshwater migrations of American shad (Alosa sapidissima) and their life history consequences. Canadian Journal of Fisheries and Aquatic Sciences
38:806–820.
Haro, A., T. Castro-Santos, J. Noreika, and M. Odeh.
2004. Swimming performance of upstream migrant
fishes in open-channel flow: a new approach to predicting passage through velocity barriers. Canadian
Journal of Fisheries and Aquatic Sciences 61:1590–
1601.
Haro, A., M. Odeh, T. Castro-Santos, and J. Noreika.
1999. Effect of slope and headpond on passage of
American shad and blueback herring through simple Denil and deepened Alaska steeppass fishways.
North American Journal of Fisheries Management
19:51–58.
Haro, A., M. Odeh, J. Noreika, and T. Castro-Santos.
1998. Effect of water acceleration on downstream
migratory behavior and passage of Atlantic salmon
smolts and juvenile American shad at surface bypasses. Transactions of the American Fisheries Society 127:118–127.
Hasler, A. D. 1971. Orientation and fish migration. Pages
429–510 in W. S. Hoar and D. J. Randall, editors.
Environmental relations and behavior. Academic
Press, New York.
Hinch, S. G., and J. Bratty. 2000. Effects of swim speed
and activity pattern on success of adult sockeye
salmon migration through an area of difficult passage. Transactions of the American Fisheries Society
129:598–606.
Hinch, S. G., R. E. Diewert, T. J. Lissimore, A. M. J.
Prince, M. C. Healey, and M. A. Henderson. 1996.
Use of electromyogram telemetry to assess difficult
passage areas for river-migrating adult sockeye salmon. Transactions of the American Fisheries Society
125:253–260.
Hinch, S. G., and P. S. Rand. 1998. Swim speeds and energy use of up-river migrating sockeye salmon (Oncorhynchus nerka): role of local environment and fish
characteristics. Canadian Journal of Fisheries and
Aquatic Sciences 55:1821–1831.
Hoar, W. S. 1988. The physiology of smolting salmonids.
Pages 275–343 in W. S. Hoar and D. Randall, editors. Fish physiology. Volume XIB: The physiology
of developing fish. Academic Press, New York.
572
castro-santos et al.
Hosmer, D. W., and S. Lemeshow. 1999. Applied survival
analysis. Wiley, New York.
Idler, D. R., and W. A. Clemens. 1959. The energy expenditures of Fraser River sockeye salmon during
the spawning migration to Chilko and Stuart lakes.
International Pacific Salmon Fisheries Commission,
Vancouver.
Johnson, G. E., J. B. Hedgepeth, J. R. Skalski, and A. E.
Giorgi. 2004. A Markov chain analysis of fish movements to determine entrainment zones. Fisheries
Research 69:349–358.
Johnston, I. A. 2001. Genetic and environmental determinants of muscle growth patterns. Pages 141–186
in D. Randall and A. P. Farrell, editors. Fish physiology, volume 18: Muscle development and growth.
Academic Press, London.
Jungwirth, M. 1996. Bypass channels at weirs as appropriate aids for fish migration in rhithral rivers. Regulated Rivers Research & Management 12:483–492.
Khan, L. A. 2006. A three-dimensional computational
fluid dynamics (CFD) model analysis of free surface
hydrodynamics and fish passage energetics in a vertical-slot fishway. North American Journal of Fisheries
Management 26:255–267.
Laake, J. L., P. Browne, R. L. DeLong, and H. R. Huber.
2002. Pinniped diet composition: a comparison of
estimation models. Fisheries Bulletin 100:434–447.
Larinier, M., and F. Travade. 1999. Downstream migration:
problems and facilities [In French]. Bulletin Francais
de la Peche et de la Pisciculture 364(Supplement):181–
210.
Larinier, M., and F. Travade. 2002. The design of fishways
for shad. Bulletin Francais de la Peche et de la Pisciculture 364(Supplement):135–146.
Leonard, J. B. K., M. Iwata, and Ueda, H. 2001. Seasonal
changes of hormones and muscle enzymes in adult
lacustrine masu (Oncorhynchus masou) and sockeye
salmon (O. nerka). Fish Physiology and Biochemistry 25:153–163.
Leonard, J. B. K., and S. D. McCormick. 1999. Effects of
migration distance on whole-body and tissue-specific energy use in American shad (Alosa sapidissima).
Canadian Journal of Fisheries and Aquatic Sciences
56:1159–1171.
Liao, J. C. 2004. Neuromuscular control of trout swimming in a vortex street: implications for energy
economy during the Karman gait. Journal of Experimental Biology 207:3495–3506.
Liao, J. C., D. N. Beal, G. V. Lauder, and M. S. Triantafyllou. 2003a. Fish exploiting vortices decrease
muscle activity. Science 302:1566–1569.
Liao, J. C., D. N. Beal, G. V. Lauder, and M. S. Triantafyllou. 2003b. The Karman gait: novel body kinematics of rainbow trout swimming in a vortex street.
Journal of Experimental Biology 206:1059–1073.
Lighthill, M. J. 1971. Large amplitude elongate body
theory of fish locomotion. Proceedings of the Royal
Society of London Series B 179:125–138.
Lighthill, M. J. 1977. Mathematical theories of fish swimming. Pages 131–144 in J. H. Steele, editor. Fisheries mathematics. Academic Press, London.
Lowe, C. G. 1996. Kinematics and critical swimming
speed of juvenile scalloped hammerhead sharks.
Journal of Experimental Biology 199:2605–2610.
Lowe, C. G., K. N. Holland, and T. G. Wolcott. 1998. A
new acoustic tailbeat transmitter for fishes. Fisheries
Research 36:275–283.
Lucas, M. C., T. Mercer, J. D. Armstrong, S. McGinty,
and P. Rycroft. 1999. Use of a flat-bed passive integrated transponder antenna array to study the migration and behaviour of lowland river fishes at a fish
pass. Fisheries Research 44:183–191.
Madenjian, C. P., G. L. Fahnenstiel, T. H. Johengen,
T. F. Nalepa, H. A. Vanderploeg, G. W. Fleischer,
P. J. Schneeberger, D. M. Benjamin, E. B. Smith,
J. R. Bence, E. S. Rutherford, D. S. Lavis, D. M.
Robertson, D. J. Jude, and M. P. Ebener. 2002. Dynamics of the Lake Michigan food web, 1970–2000.
Canadian Journal of Fisheries and Aquatic Sciences
59:736–753.
McCormick, S. D. 1994. Ontogeny and evolution of salinity tolerance in anadromous salmonids: hormones
and heterochrony. Estuaries 17:26–33.
McCormick, S. D., D. T. Lerner, M. Y. Monette, K.
Nieves-Puigdoller, J. T. Kelly, and B. T. Björnsson.
2009. Taking it with you when you go: how perturbations to the freshwater environment, including
temperature, dams, and contaminants, affect marine
survival of salmon. Pages xxx–xxx in A. J. Haro,
K. L. Smith, R. A. Rulifson, C. M. Moffitt, R. J.
Klauda, M. J. Dadswell, R. A. Cunjak, J. E. Cooper, K. L. Beal, and T. S. Avery, editors. Challenges
for diadromous fishes in a dynamic global environment. American Fisheries Society, Symposium 69,
Bethesda, Maryland.
McKinley, R. S., and G. Power. 1992. Measurement of
activity and oxygen consumption for adult lake sturgeon (Acipenser fulvescens) in the wild using radiotransmitted EMG signals. Pages 307–318 in I. G.
Priede and S. M. Swift, editors. Wildlife telemetry:
remote monitoring and tracking of animals. Horwood, New York.
McLeod, A. M., and P. Nemenyi. 1940. An investigation
of fishways. State University of Iowa, Iowa City.
Meyer, J. L., D. L. Strayer, J. B. Wallace, S. L. Eggert, G.
S. Helfman, and N. E. Leonard. 2007. The contribution of headwater streams to biodiversity in river
networks. Journal of the American Water Resources
Association 43:86–103.
Mogdans, J., and H. Bleckmann. 2001. The mechanosen-
fishway evaluations for better bioengineering
sory lateral line of jawed fishes. Pages 181–213 in
B. G. Kapoor and T. J. Hara, editors. Sensory biology of jawed fishes: new insights. Science Publishers,
Plymouth, UK.
Monk, B., D. Weaver, C. Thompson, and F. Ossiander.
1989. Effects of flow and weir design on the passage behavior of American shad and salmonids in an
experimental fish ladder. North American Journal of
Fisheries Management 9:60–67.
Mustard, J. F., M. A. Carney, and A. Sen. 1999. The use
of satellite data to quantify thermal effluent impacts.
Estuarine Coastal and Shelf Science 49:509–524.
Naughton, G. P., C. C. Caudill, M. L. Keefer, T. C.
Bjornn, L. C. Stuehrenberg, and C. A. Peery. 2005.
Late-season mortality during migration of radiotagged adult sockeye salmon (Oncorhynchus nerka)
in the Columbia River. Canadian Journal of Fisheries and Aquatic Sciences 62:30–47.
Nikora, V. I., J. Aberle, B. J. F. Biggs, I. G. Jowett, and
J. R. E. Sykes. 2003. Effects of fish size, time-tofatigue and turbulence on swimming performance:
a case study of Galaxias maculatus. Journal of Fish
Biology 63:1365–1382.
Northcote, T. G., S. N. Williscr, and H. Tsuyuki. 1970.
Meristic and lactate dehydrogenase genotype differences in stream populations of rainbow trout below
and above a waterfall. Journal of the Fisheries Research Board of Canada 27:1987–1995.
Orsborn, J. F. 1987. Fishways: historical assessment of design practices. Pages 122–130 in M. J. Dadswell, R.
J. Klauda, C. M. Moffitt, R. L. Saunders, R. A. Rulifson, and J. E. Cooper, editors. Common strategies of
anadromous and catadromous fishes. American Fisheries Society, Symposium 1, Bethesda, Maryland.
Orsborn, J. F., and P. D. Powers. 1986. Fishways: an assessment of their development and design. Part 3 of
4. Final project report. Bonneville Power Administration, BPA Report DOE/BP-36523-2, Portland,
Oregon.
Palmer, M. A., E. S. Bernhardt, J. D. Allan, P. S. Lake,
G. Alexander, S. Brooks, J. Carr, S. Clayton, C. N.
Dahm, J. F. Shah, D. L. Galat, S. G. Loss, P. Goodwin, D. D. Hart, B. Hassett, R. Jenkinson, G. M.
Kondolf, R. Lave, J. L. Meyer, T. K. O’Donnell, L.
Pagano, and E. Sudduth. 2005. Standards for ecologically successful river restoration. Journal of Applied Ecology 42:208–217.
Peake, S. 2004. An evaluation of the use of critical swimming speed for determination of culvert water velocity criteria for smallmouth bass. Transactions of the
American Fisheries Society 133:1472–1479.
Peake, S., R. S. McKinley, and D. A. Scruton. 1997. Swimming performance of various freshwater Newfoundland salmonids relative to habitat selection and fishway design. Journal of Fish Biology 51:710–723.
573
Peake, S. J., and A. P. Farrell. 2004. Locomotory behaviour and post-exercise physiology in relation to
swimming speed, gait transition and metabolism in
free-swimming smallmouth bass (Micropterus dolomieu). Journal of Experimental Biology 207:1563–
1575.
Peake, S. J., and A. P. Farrell. 2005. Postexercise physiology and repeat performance behaviour of freeswimming smallmouth bass in an experimental
raceway. Physiological and Biochemical Zoology
78:801–807.
Powers, P. D., J. F. Orsborn, T. W. Bumstead, S. KlingerKingsley, and W. C. Mih. 1985. Fishways: an assessment of their development and design. Bonneville
Power Administration, U.S. Department of Energy
No. DOE/BP-36523–4, Portland, Oregon..
Prentice, E. F., T. A. Flagg, and S. McCutcheon. 1990.
Feasibility of using implantable passive integrated
transponder (PIT) tags in salmonids. Pages 317–322
in N. C. Parker, A. E. Giorgi, R. C. Heidinger, D. B.
Jester, Jr., E. D. Prince, and G. A. Winans, editors.
Fish marking techniques. American Fisheries Society, Bethesda, Maryland
Rand, P. S., and S. G. Hinch. 1998. Swim speeds and energy use of upriver-migrating sockeye salmon (Oncorhynchus nerka): simulating metabolic power and
assessing risk of energy depletion. Canadian Journal
of Fisheries and Aquatic Sciences 55:1832–1841.
Rand, P. S., S. G. Hinch, J. Morrison, M. G. G. Foreman,
M. J. MacNutt, J. S. Macdonald, M. C. Healey, A.
P. Farrell, and D. A. Higgs. 2006. Effects of river
discharge, temperature, and future climates on energetics and mortality of adult migrating Fraser River
sockeye salmon. Transactions of the American Fisheries Society 135:655–667.
Reback, K. E., P. D. Brady, K. D. McLaughlin, and C. G.
Milliken. 2004. A survey of anadromous fish passage
in coastal Massachusetts (parts 1–4). Massachusetts
Division of Marine Fisheries, Technical Report TR15, Boston.
Reischel, T. S., and T. C. Bjornn. 2003. Influence of
fishway placement on fallback of adult salmon
at the Bonneville Dam on the Columbia River.
North American Journal of Fisheries Management
23:1215–1224.
Rideout, S. G., L. M. Thorpe, and L. M. Cameron. 1985.
Passage of American shad in an Ice Harbor style
fishladder after flow pattern modifications. Pages
251–256 in F. W. Olson, R. G. White, and R. H.
Hamre. Proceedings of the Symposium on Small
Hydropower and Fisheries. American Fisheries Society, Bethesda, Maryland.
Santos, H. D. E., P. D. Pompeu, and C. B. Martinez. 2007. Swimming performance of the migratory neotropical fish Leporinus reinhardti
574
castro-santos et al.
(Characiformes:Anostomidae). Neotropical Ichthyology 5:139–146.
Santos, J. M., M. T. Ferreira, F. N. Godinho, and J. Bochechas. 2005. Efficacy of a nature-like bypass channel in a Portuguese lowland river. Journal of Applied
Ichthyology 21:381–388.
Sato, A., H. Ueda, M. Fukaya, M. Kaeriayama, Y. Zohar,
A. Urano, and K. Yamauchi. 1997. Sexual differences in homing profiles and shortening of homing duration by gonadotropin-releasing hormone
analog implantation in lacustrine sockeye salmon
(Oncorhynchus nerka) in Lake Shikotsu. Zoological
Science 14:1009–1014.
Schmutz, S., C. Giefing, and C. Wiesner. 1998. The efficiency of a nature-like bypass channel for pike-perch
(Stizostedion lucioperca) in the Marchfeldkanalsystem. Hydrobiologia 372: 355–360.
Schreck, C. B., W. Contreras-Sanchez, and M. S. Fitzpatrick. 2001. Effects of stress on fish reproduction,
gamete quality, and progeny. Aquaculture 197:3–
24.
Schultz, W. W., and P. W. Webb. 2002. Power requirements of swimming: do new methods resolve old
questions? Integrative & Comparative Biology
42:1018–1025.
Skalski, J. R., J. Lady, R. Townsend, A. E. Giorgi, J. R. Stevenson, C. M. Peven, and R. D. McDonald. 2001.
Estimating in-river survival of migrating salmonid
smolts using radiotelemetry. Canadian Journal of
Fisheries and Aquatic Sciences 58:1987–1997.
Skalski, J. R., S. G. Smith, R. N. Iwamoto, J. G. Williams,
and A. Hoffmann. 1998. Use of passive integrated
transponder tags to estimate survival of migrant juvenile salmonids in the Snake and Columbia rivers.
Canadian Journal of Fisheries and Aquatic Sciences
55:1484–1493.
Slatick, E., and L. R. Basham. 1985. The effect of Denil fishway length on passage of some nonsalmonid
fishes. Marine Fisheries Review 47:83–85.
Sorensen, P. W., and N. E. Stacey. 2004. Brief review of
fish pheromones and discussion of their possible uses
in the control of non-indigenous teleost fishes. New
Zealand Journal of Marine and Freshwater Research
38:399–417.
Sorensen, P. W., and L. A. Vrieze. 2003. The chemical
ecology and potential application of the sea lamprey
migratory pheromone. Journal of Great Lakes Research 29:66–84.
Stringham, E. 1924. The maximum speed of fresh-water
fishes. American Naturalist 18:156–161.
Stuart, I. G., and A. P. Berghuis. 2002. Upstream passage
of fish through a vertical-slot fishway in an Australian subtropical river. Fisheries Management and
Ecology 9:111–122.
Sullivan, T. J. 2004. Evaluation of the Turners Falls fish-
way complex and potential improvements for passing adult American shad. Master’s thesis. University
of Massachusetts, Amherst.
Sullivan, W. P., G. C. Christie, F. C. Cornelius, M. F.
Fodale, D. A. Johnson, J. F. Koonces, G. L. Larson, R. B. McDonald, K. M. Mullett, C. K. Murray, and P. A. Ryan. 2003. The sea lamprey in Lake
Erie: a case history. Journal of Great Lakes Research
29:615–636.
Taylor, E. B., and J. D. Mcphail. 1986. Prolonged and
burst swimming in anadromous and fresh-water
threespine stickleback, Gasterosteus aculeatus. Canadian Journal of Zoology-Revue Canadienne de Zoologie 64:416–420.
Tierney, K. B., and A. P. Farrell. 2004. The relationships
between fish health, metabolic rate, swimming performance and recovery in return-run sockeye salmon, Oncorhynchus nerka (Walbaum). Journal of Fish
Diseases 27:663–671.
Tritico, H. M., A. J. Cotel, and J. N. Clarke. 2007. Development, testing and demonstration of a portable
submersible miniature particle imaging velocimetry device. Measurement Science & Technology
18:2555–2562.
Tudorache, C., P. Viaenen, R. Blust, and G. De Boeck.
2007. Longer flumes increase critical swimming
speeds by increasing burst and glide swimming duration in carp (Cyprinus carpio, L.). Journal of Fish
Biology 71:1630–1638.
Turchin, P. 1998. Quantitative analysis of animal movement. Sinauer, Sunderland, Massachusetts.
Ueda, H., O. Hiroi, A. Hara, K. Yamauchi, and Y. Nagahama. 1984. Changes in serum concentrations
of steroid-hormones, Thyroxine, and Vitellogenin
during spawning migration of the chum salmon,
Oncorhynchus keta. General and Comparative Endocrinology 53:203–211.
Venditti, D. A., D. W. Rondorf, and J. M. Kraut. 2000.
Migratory behavior and forebay delay of radio-tagged
juvenile fall Chinook salmon in a lower Snake River
impoundment. North American Journal of Fisheries
Management 20:41–52.
Videler, J. J., and C. S. Wardle. 1991. Fish swimming
stride by stride: speed limits and endurance. Reviews
in Fish Biology and Fisheries 1:23–40.
Wardle, C. S. 1975. Limit of fish swimming speed. Nature 255:725–727.
Weaver, C. R. 1963. Influence of water velocity upon orientation and performance of adult migrating salmonids. Fisheries Bulletin 63:97–121.
Weaver, C. R. 1965. Observations on the swimming
ability of adult American shad (Alosa sapidissima).
Transactions of the American Fisheries Society
94:382–385.
Webb, P. W. 1971a. The swimming energetics of trout: I.
fishway evaluations for better bioengineering
Thrust and power output at cruising speeds. Journal
of Experimental Biology 55:189–520.
Webb, P. W. 1971b. The swimming energetics of trout: II.
Oxygen consumption and swimming capacity. Journal of Experimental Biology 55:521–540.
Webb, P. W. 1975. Hydrodynamics and energetics of fish
propulsion. Bulletin of the Fisheries Research Board
of Canada 190:1–158.
Webb, P. W. 1998a. Entrainment by river chub Nocomis
micropogon and smallmouth bass Micropterus dolomieu on cylinders. Journal of Experimental Biology
201:2403–2412.
575
Webb, P. W. 1998b. Swimming. Pages 3–24 in D. H.
Evans, editor. The physiology of fishes. CRC Press,
Boca Raton, Florida.
Webb, P. W., and C. L. Gerstner. 2000. Fish swimming
behaviour: predictions from physical principles. Pages 59–77 in P. Domenici and R. W. Blake, editors.
Biomechanics in animal behaviour. BIOS Scientific
Publishing, Oxford, UK.
Zhong, Y. G., and G. Power. 1996. Environmental impacts of hydroelectric projects on fish resources in
China. Regulated Rivers Research & Management
12:81–98.