Fishway Evaluations for Better Bioengineering: An ...

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

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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

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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-

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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

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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.

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