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Rev Fish Biol Fisheries
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Review of trap-and-haul for managing Pacific salmonids

(Oncorhynchus spp.) in impounded river systems
Tobias J. Kock . John W. Ferguson . Matthew L. Keefer . Carl B. Schreck
Received: 29 June 2020 / Accepted: 31 October 2020

The Author(s) 2020
Abstract High-head dams are migration barriers for reducing exposure to dangerous in-river conditions,
Pacific salmon Oncorhynchus spp. in many river and reintroducing ecological processes upstream from
systems and recovery measures for impacted stocks dams. Information gathered from decades of operation
are limited. Trap-and-haul has been widely used in on facility design criteria and fish handling protocols,
attempts to facilitate recovery but information from and robust literature on fish collection and passage are
existing programs has not been synthesized to inform available. While many aspects of trap-and-haul have
improvements to aid recovery of salmonids in systems been evaluated, effects on population productivity and
with high-head dams. We reviewed 17 trap-and-haul sustainability remain poorly understood. Long-term
programs regarding Pacific salmon to: (1) summarize and systematic studies of trap-and-haul outcomes are
information about facility design, operation and rare, and assessments can be confounded by concur-
biological effects; (2) identify critical knowledge rent management actions and broad ecological and
gaps; and (3) evaluate trap-and-haul as a current and climatic effects. Existing data suggest that perfor-
future management tool. Existing programs are oper- mance and effectiveness vary among programs and
ated to address a range of management goals including over various time scales within programs. Although
restoring access to historical habitats, temporarily critical information gaps exist, trap-and-haul is an
important management and conservation tool for
T. J. Kock (&)
providing Pacific salmonids access to historical habi-
U.S. Geological Survey, Western Fisheries Research tats. Successful application of trap-and-haul programs
Center, 5501A Cook-Underwood Road, Cook, requires long-term commitment and an adaptive
WA 98605, USA management approach by dam owners and stakehold-
e-mail: [email protected]
ers, and careful planning of new programs.
J. W. Ferguson
Anchor QEA, LLC, 1201 3rd Avenue #2600, Seattle, Keywords Pacific salmon Trap-and-haul
WA 98101, USA Oncorhynchus Dams Passage Impoundments
M. L. Keefer
Department of Fish and Wildlife Sciences, College of
Natural Resources, University of Idaho, 875 Perimeter
Drive, Moscow, ID 83844-1136, USA
C. B. Schreck
Department of Fisheries and Wildlife, Oregon State,
University, Corvallis, OR 97331, USA
123
Introduction Trap-and-haul is being used to restore connectivity

to reaches both downstream and upstream of dams.
Dams provide many benefits worldwide including Programs typically rely on collection facilities located
flood control, municipal and agricultural water sup- in a dam forebay (reviewed by Kock et al. 2019a) to
plies, and hydropower generation. However, these collect juvenile migrants for downstream transport
structures and the reservoirs they create are often whereas adult migrants are captured for upstream
barriers to movement for migratory fishes and other transport in trapping facilities located in a dam tailrace
aquatic species. Fish mortality has been directly (NMFS 2011). In some cases, O. mykiss kelts, adult
attributed to dam passage (Čada 2001; Muir et al. steelhead that have successfully spawned and are
2001; Skalski et al. 2002; Schilt 2007; Larinier 2008) returning to the ocean, are also collected and trans-
and to indirect effects from factors such as predation, ported downstream (Trammell et al. 2016). Transport
migration delay, pathogen transmission, and thermal of juveniles and adults to in-river release sites is
perturbations (Poe et al. 1991; Clarkson and Childs generally accomplished using fish-hauling tanker
2000; Schreck et al. 2006; Keefer et al. 2012; Colvin trucks with the capability to provide supplemental
et al. 2015). Substantial efforts have focused on oxygen for fish during transport (Sigourney et al.
developing safe and effective fish passage options for 2015; DeWeber et al. 2017; Colvin et al. 2018).
upstream (Clay 1995; Roscoe and Hinch 2010; Bunt However, at some Columbia River dams, juveniles
et al. 2012; Katopodis and Williams 2012; Pompeu (Ward et al. 1997; McMichael et al. 2011) or
et al. 2012; Williams et al. 2012) and downstream downstream-migrating adults (Evans et al. 2008) are
migrants at dams (Ferguson et al. 2007; Schilt 2007; transported in barges with flow-through or recirculat-
Adams et al. 2014; Shi et al. 2015; Skalski et al. 2016; ing river water. For all transport methods, protocols
Fjelstad et al. 2018). Unfortunately, fish passage have been developed to protect fish during transit. A
facilities have often failed to fully mitigate negative main goal of these protocols is to minimize fish stress.
effects of dams and their impoundments (Nicola et al. For example, maximum fish density thresholds are
1996; Caudill et al. 2007; Fukushima et al. 2007; used to ensure that fish are not overcrowded while
Ferguson et al. 2011). inside holding and transport tanks. Water temperatures
In some river systems, resource managers have and dissolved oxygen levels are monitored and
elected to collect migrants and transport them (here- maintained during transport and fish are loaded and
after trap-and-haul; Fig. 1) past dams, impoundments, released using water-to-water transfer methods. Addi-
and other migration barriers to facilitate migration and tionally, fish are sometimes released into ‘‘stress-relief
dispersal. Trap-and-haul has been used as a routine ponds’’ after transport which has been shown to
fisheries management tool (Sigourney et al. 2015) and attenuate the effects of stress incurred during the
as a conservation strategy to maintain or restore fish holding and transport (NMFS 2011).
populations upstream from dams without fish passage Existing trap-and-haul programs are operated to
(Sard et al. 2016; Lusardi and Moyle 2017). Addi- provide passage for either juveniles or adults (here-
tionally, trap-and-haul has been employed as an after one-way trap-and-haul), or for multiple life
emergency procedure to protect endangered Snake stages (juveniles downstream and adults upstream,
River sockeye salmon (Oncorhynchus nerka) when in- hereafter two-way trap-and-haul; Lusardi and Moyle
river migratory conditions become hazardous (Kozf- 2017). Some trap-and-haul programs have been oper-
kay et al. 2017). While volitional passage of Pacific ating for decades (Table 1), especially in the U.S.
salmon (Oncorhynchus spp.) at barriers is preferred Pacific Northwest where hydropower development
over passage that requires collection and transporta- has contributed to widespread Pacific salmonid pop-
tion (e.g., NMFS 2011), volitional passage at many ulation declines and extirpations (Nehlsen et al. 1991;
locations is constrained by economic or engineering Kareiva et al. 2000; McClure et al. 2003).
considerations. At many high-head dams ([ 30 m Although some programs have operated for dec-
tall), for example, fishway construction may be ades, efforts to review and synthesize the effectiveness
unrealistic, leaving trap-and-haul or dam removal as of trap-and-haul as a fish conservation and manage-
the only potentially viable options for sustaining or ment strategy are lacking. In some cases, individual
reintroducing populations upstream of the dam. elements of the trap-and-haul process (i.e., juvenile
123
Fig. 1 Conceptual diagram of a two-way trap-and-haul program and key elements considered in program development
collection efficiency, adult prespawn mortality) have valuable insights into several aspects of trap-and-haul
been evaluated. However, a comprehensive assess- but provided limited information on what has been
ment is needed to evaluate if the strategy should learned about developing and operating trap-and-haul
continue to be used at existing and new locations. In a facilities, handling and transporting fish, or how fish
recent review, Lusardi and Moyle (2017) assessed respond biologically to trap-and-haul.
two-way trap-and-haul as a conservation tool, with an Given that trap-and-haul operations have occurred
emphasis on potential application for threatened for nearly 100 years in the Pacific Northwest and that
Chinook salmon (O. tshawytscha) in central Califor- there is continued interest in using trap-and-haul at
nia. The authors reviewed several existing two-way existing and new locations (Clancey et al. 2017;
(i.e., both downstream juvenile and upstream adult Hardiman et al. 2017; Herbold et al. 2018; Upper
transport) trap-and-haul programs in the Pacific Columbia United Tribes 2019), we conducted this
Northwest, highlighted potential risks of fish transport, review to: (1) summarize information about facility
and provided recommendations for effective charac- design and operation and the biological effects of trap-
teristics of two-way trap-and-haul programs. These and-haul in the U.S. Pacific Northwest; (2) identify
included establishing clearly defined success metrics, critical knowledge gaps; and (3) evaluate trap-and-
confirming that adequate spawning, incubation, and haul as a current and future management tool. To
rearing habitats exist in recipient rivers, and that two- address these objectives, we focused on trap-and-haul
way trap-and-haul be conducted as part of a more operations for Pacific salmonids at dams without
comprehensive program that includes other manage- volitional fish passage facilities. We also include
ment actions such as habitat restoration and dam examples of where trap-and-haul is used to circumvent
removal (Lusardi and Moyle 2017). They concluded other impediments to illustrate the breadth of trap-and-
that no two-way trap-and-haul program was an haul applications. To address how fish respond to trap-
unequivocal success and that new programs should and-haul we reviewed literature from existing trap-
proceed with extreme caution. The authors provided and-haul sites and other established salmonid transport
123
Table 1 Trap-and-haul locations, program type, fish species, and transport distance
Location Year program began Species transported Approximate number Approximate
of fish transported transport time
(min)
Two-way trap-and-haul programs

Baker River, Adult program began in 1925 SO, CO, CT Adults = 10,000 s Adults = 60 min
Washington Juvenile program began in 1958 Juveniles = 100,000 s Juveniles = 20,
60 min
Cowlitz River, Adult program began in 1994 ST, CK, CO, CT Adults = 10,000 s Adults = 30, 45,
Washington Juvenile program began in 1995 Juveniles = 100,000 s 90 min
Juveniles = 45 min
Deschutes River, Adult program began in 2012 SO, CK, ST, BT Adults = 10 s Adults = 45 min
Oregon Juvenile program began in 2009 Juveniles = 100,000 s Juveniles = 45 min
Lewis River, Adult program began in 2005 ST, CK, CO, CT Adults = 10,000 s Adults = 90 min
Washington Juvenile program began in 2012 Juveniles = 10,000 s Juveniles = 45 min
North Fork Skokomish Juvenile program began in 2015 SO, CO Juveniles = 10,000 s Juveniles = 20 min
River, Washington
Upstream transport trap-and-haul programs
Fall Creek, Oregon Adult program began 1965 CK 100 s 15 min
McKenzie River, Adult program began in 1993 CK, BT, RT, CT, 100 s 60 min, 120 min
Oregon WF
Middle Fork Willamette Adult program began in 1994 CK 1000 s 90 min, 150 min,
River, Oregon 210 min
North Santiam River, Adult program began in 2000 CK 1000 s 25 min, 30 min,
Oregon 40 min
Snake River, Adult program began in 2001 SO 100 s 210 min
Washingtona
South Fork Skykomish Adult program began in 1958 ST, CO, SO, CK, 10,000 s 20 min
River, Washington CT, PS, CH, BT
South Santiam River, Adult program began in 1967 ST, CK 1000 s 10 min, 20 min,
Oregon 45 min
Toutle River, Adult program began in 1988 ST, CO, CT 100 s 10 min, 20 min,
Washington 40 min
Wynoochee River, Adult program began in 1972 ST, CO, CK 1000 s 20 min
Washington
White River, Adult program began in 1941 CK, ST, CO, PS, 100,000 s 30 min
Washington BT, CH
Yakima River, Adult program began in 2009 SO 1000 s 120 min
Washington
Elwha River, Adult program began in 2012 CO 100 s 45 min
Washington
ST steelhead, SO sockeye salmon, CO coho salmon, CK Chinook salmon, PS pink salmon, CS chum salmon, CT cutthroat trout, RT
rainbow trout, BT bull trout
a
Adult sockeye salmon are transported from the Stanley Basin to a conservation hatchery near Boise, Idaho for holding, then back to
the Stanley Basin for spawning. In 2015 fish were captured at Lower Granite Dam to save them from dangerous in-river conditions
programs with directly relevant research results. The existing programs and inform discussions among
summaries and recommendations herein are intended resource managers and stakeholders at sites where
to improve management and monitoring strategies in trap-and-haul is being considered.
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Upstream migrant trap-and-haul programs completed but was then abandoned until the program
was re-started in 1991. Returning Chinook salmon are
Upstream migrant trap-and-haul is the most common collected from the tailrace of the dam via an adult
type of trap-and-haul program currently used in the fishway and trap and transported upstream by truck *
Pacific Northwest. Most of these programs are oper- 15 min to a release site upstream of the reservoir. The
ated to move returning adult salmon and steelhead average annual number of adults transported in
around high-head dams where they are released to 1991–2018 was 470 fish with a peak of 2805 adults
spawn naturally. In some cases, these programs are in 2004 (data provided by Greg Taylor, USACE). The
used to move fish around natural barriers, areas program goal is population recovery upstream of Fall
affected by natural disaster, or to remove endangered Creek Dam (Sharpe et al. 2016). Progeny of trans-
fish from dangerous in-river conditions below dams. ported adults move downstream and enter Fall Creek
These programs rely on fish traps to collect upstream Reservoir. Passage options are limited at the dam so
migrants and transport is by fish-hauling trucks. most juvenile Chinook salmon rear in the reservoir
during their first year (Monzyk et al. 2015a). In recent
Willamette River basin, Oregon years, Fall Creek Reservoir has been drained in
November to flush juvenile salmon downstream and
Numerous upstream migrant trap-and-haul programs to reduce populations of non-native piscivores that
are operated in the Willamette River basin, Oregon. prey on salmon in the reservoir (Murphy et al. 2019).
The U.S. Army Corps of Engineers (USACE) operates Although the Fall Creek salmon population was nearly
the Willamette Valley Project, a group of 13 dams that extirpated, it is currently self-sustaining and is one of
provide flood risk management, power generation, and the few natural-origin-only Chinook salmon popula-
other societal benefits. Eleven of the dams are high- tions in the Willamette basin.
head projects that were built during the 1950s–1960s The McKenzie River supports the most abundant
and block spring-run Chinook salmon and winter-run population of spring-run Chinook salmon in the
steelhead from historical upstream habitats (NMFS Willamette River basin (Myers 2017). Access to
2008). In the early 1990s, the Oregon Department of headwater tributaries in the McKenzie River is
Fish and Wildlife (ODFW) began transporting and blocked by Blue River Dam (rkm 3 of the Blue River;
releasing surplus hatchery Chinook salmon adults height = 82 m), Cougar Dam (rkm 7 of the South Fork
upstream of several Willamette River dams to evaluate McKenzie River; height = 158 m), and Trail Bridge
the feasibility of re-establishing anadromous popula- Dam (rkm 124 of the McKenzie River; height = 31
tions (NMFS 2008). These early releases resulted in m), which were all constructed in the 1960s. Trap-and-
the successful production of offspring that moved haul efforts in the McKenzie River were initiated in
downstream and attempted to pass dams (Keefer et al. 1993 using adult Chinook salmon collected at hatch-
2012, 2013), which led to the integration of adult trap- ery traps. The McKenzie River program moves adults
and-haul into the fish recovery program for the basin upstream of Trail Bridge and Cougar dams. In 2010, a
(NMFS 2008). Upstream transport of adults in the new adult collection facility was completed in the
Willamette basin occurs in Fall Creek, and the Cougar Dam tailrace. This facility serves as the
McKenzie, Middle Fork Willamette, North Santiam, collection point for unmarked (i.e., presumed natural-
and South Santiam rivers (Table 1). The goals of origin) Chinook salmon that return to the South Fork
individual tributary programs differ (Sharpe et al. McKenzie River. Both natural-origin and hatchery-
2016), and safe downstream fish passage through origin Chinook salmon are transported upstream of
reservoirs and past dams are key limitations to Cougar Reservoir but are segregated by release site
population expansion in several tributaries (Keefer with hatchery-origin fish released farther upstream
et al. 2012, 2013; Monzyk et al. 2015a, b; Romer et al. than natural-origin fish. The goal of the natural-origin
2016). releases is to recover Chinook salmon populations in
Trap-and-haul of adult spring-run Chinook salmon the upper South Fork McKenzie River while the goal
at Fall Creek Dam (rkm 12; height = 64 m) began in of the hatchery-origin releases is to produce prey for
1965 during dam construction. Adult transport bull trout (Salvelinus confluentus) residing in the
occurred for several years after the dam was upper system. The average annual number of adult
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Chinook salmon transported around McKenzie River upper North Santiam basin, steelhead reintroduction
dams was 1382 fish in 1993–2018. In 2014, the efforts are on hold until downstream fish passage is
USACE constructed an experimental floating collector available at Detroit Dam (Mapes et al. 2017). Trap-
designed to collect downstream migrants in the and-haul operations in the North Santiam River began
Cougar Dam forebay. That device was operated for in 2000 (Table 1) using an adult collection and holding
two collection seasons but collected few fish (Beeman facility located at Minto Dam (rkm 67; 4 m), down-
et al. 2016a, b) which led to a decision to abandon its stream of Detroit and Big Cliff (rkm 74; height = 58
use. A new, larger surface collector is being designed m) dams; a rebuilt Minto Fish Facility began operating
to provide downstream passage at Cougar Dam. in 2013. The North Santiam reintroduction effort relies
Improvement to downstream fish passage and survival on transporting hatchery-origin Chinook salmon *
at Cougar Dam is considered critically important for 25–40 min upstream from Detroit Reservoir where
establishing viable Chinook salmon populations in the they are released at three tributary locations. The mean
upper South Fork McKenzie River basin. Reintroduc- annual number of adult Chinook salmon transported
tion efforts have not been initiated upstream of Trail upstream of Detroit Dam was 1307 fish during
Bridge Dam, so there are currently no plans for 2000–2018 (data provided by Greg Grenbemer,
constructing downstream fish passage facilities at this ODFW). Studies have shown that juvenile outmi-
dam, and natural-origin Chinook salmon are not grants can pass Detroit Dam through turbines or over
transported upstream of the dam. Adult hatchery- spillways (Beeman and Adams 2015; Kock et al.
origin Chinook salmon are transported upstream of 2015) where substantial mortality can occur (Nor-
Trail Bridge Dam for the sole purpose of providing mandeau 2010). As a result, Myers (2017) recom-
juvenile salmon as prey for bull trout. mended that reintroduction plans for Chinook salmon
Trap-and-haul in the Middle Fork Willamette River and steelhead upstream of Detroit Dam be synchro-
is a complex process with multiple purposes (Sharpe nized with the development of juvenile passage
et al. 2016). Adult trapping occurs at Dexter Dam (rkm facilities at the dam. The USACE is currently design-
29; height = 36 m), the downstream-most dam in the ing selective water withdrawal and floating screen
Middle Fork Willamette River, and at Willamette structures that will be hydraulically connected and
Hatchery (rkm 2) with releases conducted at multiple work together to collect downstream migrants in the
sites in various upstream reaches and tributaries. The dam forebay and allow the temperature of water
intent of the adult releases includes restoring biolog- passing the dam to be controlled. This temperature
ical contributions of salmon to the ecosystem, provid- control is critical for supporting spawning, rearing,
ing forage for native bull trout that were recently and migration of salmonids downstream of the dam.
reintroduced, and supporting research on Chinook Anadromous fish management objectives are
salmon prespawn mortality (Sharpe et al. 2016; Myers unique in the South Santiam River relative to other
2017). Middle Fork Willamette River reservoirs have Willamette River tributaries. Natural-origin winter-
high growth opportunity for juvenile salmonids but run steelhead adults have been transported upstream of
contain large populations of predatory fishes and Foster Dam (rkm 62; height = 38 m) since it began
consequently juvenile mortality rates can be high operating in 1968 and trap-and-haul of adult Chinook
(Brandt et al. 2016; Kock et al. 2019b). Juvenile salmon began in 1996. Currently, these programs
Chinook salmon primarily pass dams during fall and transport only natural-origin fish of both species with
winter months in the Middle Fork Willamette River the objective of increasing natural production and
when reservoir elevations are relatively low and deep- reducing the influence of hatchery fish on the basin’s
water passage routes are available at the dams, though populations (Evans et al. 2016; Sharpe et al. 2016;
passage mortality through these routes can be quite Wiegel et al. 2019). Adult collection initially relied on
high (Keefer et al. 2012, 2013). a fish trap that was included in the original construc-
On the North Santiam River, trap-and-haul has been tion of Foster Dam in 1968. That structure was
used to reintroduce Chinook salmon upstream from replaced with a new adult collection facility consisting
Detroit Dam (rkm 79; height = 141 m), which cur- of a fishway and trap located adjacent to the Foster
rently lacks downstream fish passage facilities. While Dam tailrace in 2014. The mean annual number of
winter-run steelhead also historically occupied the adults transported upstream during 1996–2018 was
123
979 Chinook salmon (1996–2018) and 580 steelhead mountain whitefish Prosopium williamsoni, rainbow
(1967–2018). Collected adults are transported trout (resident O. mykiss), and cutthroat trout around
upstream to several release locations, including the the dam (Fig. 2). On average, 147,307 fish were
head of Foster Reservoir (Sharpe et al. 2016; DeWeber transported annually during 2000–2017, which
et al. 2017; Naughton et al. 2018). Downstream included a peak count of 650,336 fish in 2011. Several
migrants at Foster Dam pass through turbines, spill- factors have contributed to adult returns of this
bays, or through a weir designed to facilitate down- magnitude. In the mid-1980s, hatchery production of
stream passage; juvenile fish passage efficiencies and spring-run Chinook salmon began on the White River,
survival rates have generally been higher at Foster minimum instream flow requirements were estab-
Dam than at other Willamette Project dams (Hughes lished, and fish screens were added to water diversions
et al. 2014, 2016). The overall success of the South in 1997 (personal communication, Fred Goetz,
Santiam program can be attributed to the efficacy of USACE). Additionally, pink salmon began returning
the adult trap-and-haul program and relatively suc- to the fish trap at Mud Mountain Dam in 1995.
cessful juvenile passage, though several challenges Regionally, pink salmon predominantly return in odd
remain (Myers 2017). Evans et al. (2016) reported numbered years. The average number of pink salmon
Chinook salmon cohort replacement rate for fish transported and released upstream of Mud Mountain
transported upstream exceeded 1.0 during three con- Dam in odd numbered years during 2000–2017 was
secutive years (2007–2009). Cohort replacement rate 293,681 fish. The existing fish facility was designed to
refers to the number of adults that return to spawn in a handle approximately 20,000 fish each year, so a new
given site compared to the number of adults that were facility designed to collect up to 60,000 fish/d is
transported upstream to produce those fish. currently under construction that will be the largest
Although trap-and-haul is used throughout the adult fish collection and transport facility in the United
Willamette River basin to move adult Chinook salmon States. Overall, the number of adults and species
and steelhead upstream of project dams, most of these collected and transported annually at Mud Mountain
programs are not ready to be assessed for effectiveness Dam has increased substantially since the 1980s
in population recovery because downstream fish (Fig. 2).
passage facilities are not present at most dams. Most
programs are currently operated to maintain ecolog- Skykomish River, Washington
ical functions provided by anadromous fish presence
(i.e., nutrient enhancement, forage for other fish On the South Fork Skykomish River, trap-and-haul is
populations) and support research aimed at improving used to transport tens of thousands of adult salmonids
trap-and-haul methods but cannot fully address the around a series of impassable waterfalls (Sunset,
fragmentation caused by dams across the entire life Canyon and Eagle Falls; 7.6–31.7 m tall). The
cycle. It is anticipated that these programs will program began in 1958 to provide access to quality
eventually move towards the goal of recovering spawning and rearing habitat that would otherwise be
anadromous fish populations upstream of project dams unavailable to anadromous fish and expand fishery
when downstream fish passage facilities are developed opportunities on salmonid stocks in the Snohomish
and deployed. River, a large tributary to Puget Sound. To implement
the program, a fish ladder and sorting facility were
White River, Washington constructed downstream of Sunset Falls (rkm 84). At
the sorting facility, hatchery fish can be removed when
Numerically, the trap-and-haul program on the White required by protocols (personal communication, Ed
River, Washington, is the largest adult transport Eleazer, Washington Department of Fish and Wildlife
program operating in the Pacific Northwest. The [WDFW]). Upstream transport (* 20 min) is con-
program began in 1941 while Mud Mountain Dam ducted for coho salmon, Chinook salmon, pink
(rkm 45; height = 132 m) was under construction and salmon, chum salmon, sockeye salmon, steelhead,
trap-and-haul is used to transport Chinook salmon, bull trout and cutthroat trout (Table 1). The mean
coho salmon, steelhead, sockeye salmon, chum annual number of fish transported from 1958 to 2018
salmon O. keta, pink salmon O. gorbuscha, bull trout, was 25,447, and the peak number transported was
123
Fig. 2 Seventy-eight years of adult trap-and-haul: Mud Mountain Dam on the White River, Washington
124,228 fish in 2009. Juvenile passage occurs voli- and pass through a spillway located on the side of the
tionally as the waterfalls do not prevent outmigration. SRS. Overall, the number of adult fish collected and
transported under this program is small and averaged
Toutle River, Washington 198 winter-run steelhead and 202 coho salmon each
year from 1989 to 2018 (WDFW, unpublished data).
The 1980 eruption of Mount St. Helens had catas-
trophic effects on aquatic communities in the North Wynoochee River, Washington
Fork Toutle River, Washington (Major et al. 2000).
The upper 25 km of the river were buried in a layer of Trap-and-haul is used to transport adult steelhead,
mud, ash and debris estimated to be 45 m deep (Voigt coho salmon and Chinook salmon around Wynoochee
et al. 1981; Glicken 1998). In 1987, the USACE began Dam (rkm 80; height = 53 m) and reservoir, located
construction of a sediment retention structure (SRS; on a major tributary to the Chehalis River. The
height = 56 m) designed to trap sediments upstream program began operating in 1972 when Wynoochee
of the structure and minimize downstream sediment Dam was constructed. Fish collection occurs two
accumulation. The SRS was not designed with fish miles downstream of the dam where fish are prevented
passage capabilities, so a fish collection structure was from moving upstream by a low-head barrier dam. At
also constructed 3.2 km downstream from the SRS the barrier dam fish are diverted into a series of pools
(Loch and Downing 1990). Since 1989, WDFW has that terminate in a large holding pool. On hauling days
been using the Toutle River fish collection facility to fish are removed from the holding pool and trucked *
trap adult coho salmon, winter-run steelhead, and 20 min upstream to a release site located upstream of
cutthroat trout for transport upstream of the SRS. Wynoochee Reservoir. We were unable to access data
Transported fish are released directly into three describing the number of fish transported annually on
tributaries of the North Fork Toutle River with the Wynoochee River. Juvenile passage is facilitated
transport times ranging from 10–40 min. Progeny of by not operating the dam’s turbines for 77 consecutive
transported adults can move downstream volitionally days each year during the primary outmigration period
123
which allows fish to pass through downstream passage yielded interesting results. Matala et al. (2019) found
outlets rather than through the turbines. that fish from the two populations exhibited spatial and
temporal separation during the spawning period, with
Yakima River, Washington adults originating from Lake Wenatchee spawning
earlier and farther upstream than adults from Osoyoos
Adult trap-and-haul has been used to reintroduce Lake. Genetic analysis of samples taken from smolts
sockeye salmon in the Yakima River basin. Four and returning adults found that 5% or less of the
natural glacial lakes in the upper Yakima River basin sampled fish were hybrids produced from mixed-stock
historically supported sockeye salmon populations matings. The authors also found that smolts produced
that were extirpated in the early 1900s due to dam in Cle Elum Reservoir were significantly larger at
construction (Bureau of Reclamation [BOR] 2007). In emigration (average of 140 mm and 129 mm for
2009, the Confederated Tribes and Bands of the Osoyoos-origin and Wenatchee-origin, respectively)
Yakama Nation (Yakama Nation) began reintroducing than smolts emigrating from either Osoyoos Lake or
adult sockeye salmon into Cle Elum Reservoir, a lake Lake Wenatchee (84 mm average; Matala et al. 2019).
historically used by sockeye salmon in the basin that Differences in reproductive success were also evident
was modified by the construction of Cle Elum Dam as 70% of the smolts sampled were produced from
(rkm 345; height = 150 m) at the natural lake outlet parents from Lake Wenatchee. This transferred to
(Matala et al. 2019). The reintroduction is using donor cohort replacement rates as well, as the Lake
stocks of sockeye salmon from Osoyoos Lake (British Wenatchee stock had a replacement rate of 0.80
Columbia–Washington) and Lake Wenatchee (Wash- compared to 0.17 for Osoyoos Lake stock (Matala
ington). Adults from the two populations are collected et al. 2019). Over the course of the study (2013–2016),
using an adult trap at Priest Rapids Dam (rkm 639; 53% of the adults that were outplanted into Cle Elum
height = 54 m) on the Columbia River in eastern Reservoir were of Lake Wenatchee-origin, 43% were
Washington (Matala et al. 2019). Trapped sockeye of Osoyoos Lake-origin, and the remaining fish were
salmon adults are trucked from Priest Rapids Dam to hybrids. These results show that genetic studies can be
Cle Elum Reservoir with a transport time of * 2 h. useful for assessing how fish from different source
The number of fish available for transport varies populations adapt to a novel environment after trap-
annually based on the total run size of sockeye salmon and-haul.
in the Columbia River. The average annual number of
transported sockeye salmon was 5760 fish during
2009–2018. Outplanted adults spawn in Cle Elum Two-way trap-and-haul programs
Reservoir and upstream in the Cle Elum River (Matala
et al. 2019). A single downstream passage route, Two-way trap-and-haul is generally defined as down-
located in one of the dam’s spillbays, is seasonally stream trap and transport of juvenile fish and upstream
available for juvenile outmigrants at Cle Elum Dam trap and transport of adult fish around a dam or other
when reservoir water levels are sufficiently high. passage obstacle (Lusardi and Moyle 2017). Two-way
Sockeye salmon adults began returning to the Yakima trap-and-haul programs are operated on four rivers in
River in 2013 as a result of reintroduction efforts with the northwestern United States including the Baker,
an average of 1160 fish returning annually during Cowlitz, and Lewis rivers in Washington and the
2013–2019 (data courtesy of Yakama Nation Fish- Deschutes River in Oregon (Fig. 3). At these locations
eries). The BOR, who owns and operates Cle Elum trap-and-haul is used to move fish around multiple
Dam, is currently working to construct on-site juvenile high-head dams ([ 30 m) and reservoirs. In addition
and adult collection facilities to support this effort to the existing programs, two-way trap-and-haul
(BOR 2011). The long-term plan is to reintroduce programs are planned for the North Fork Skokomish
sockeye salmon to all lakes historically used by River (Washington), on several Willamette River
sockeye salmon in the Yakima River Basin (BOR tributaries (Oregon), and are being considered on the
2012). upper Columbia River (Washington, British Colum-
The strategy of using two separate stocks of adult bia), upper Sacramento River (California), and other
sockeye salmon in the reintroduction effort has river systems in California.
123
efficiencies over time and were eventually replaced

with state-of-the-art floating surface collectors in 2008
(Upper Baker Dam) and 2013 (Lower Baker Dam).
Studies conducted to estimate collection efficiencies
of the Baker River collectors have shown that * 85%
of juvenile sockeye salmon and * 91% of coho
salmon O. kisutch are collected and transported
downstream of Lower Baker Dam (Kock et al.
2019a). The adult collection facility, located down-
stream of Lower Baker Dam, has also evolved over
time, with the latest facility completed in 2010.
Collected adults are transported and released upstream
of Upper Baker Dam or taken to hatcheries for
spawning. Numerous fish species are collected at
juvenile and adult collectors in the Baker River
system, but transport is limited to sockeye salmon,
coho salmon and cutthroat trout O. clarkii. Collec-
tively, improvements to trap-and-haul facilities and
hatchery supplementation have resulted in large
increases in adult returns over time (personal commu-
nication, Nick Verretto, Puget Sound Energy).
Cowlitz River, Washington

Fig. 3 Map of Washington, Oregon, Idaho and Northern
California showing locations where trap-and-haul are currently
operated (closed black circles), operated intermittently (closed In the Cowlitz River basin, upstream passage of
grey circles), or being considered for future operations (open anadromous fish was blocked in 1968 by the con-
circles). Locations in Washington include the Baker River (A), struction of Mossyrock Dam (rkm 105; height = 185
Elwha River (B), South Fork Skykomish River (C), Upper
m), that lacks fish passage. Resource managers
Columbia River (D), North Fork Skokomish River (E), White
River (F), Keechelus (G), Kachess (H), and Cle Elum implemented a trap-and-haul program in the mid-
(I) reservoirs on the Yakima River, Wynoochee River (J), 1990s (Table 1) to reintroduce Chinook salmon,
Cowlitz River (K), North Fork Toutle River (L), Bumping River steelhead, coho salmon, and cutthroat trout to the
(M), Lower Granite Dam (O), and Lewis River (N). Locations in upper basin. This timing was based, in part, on the
Oregon include the North Santiam River (P), Deschutes River
(Q), South Santiam River (R), McKenzie River (S), Fall Creek construction of Cowlitz Falls Dam (rkm 142; height =
(T), and Middle Fork Willamette River (U). Locations in Idaho 43 m) in the early 1990s, the uppermost dam in the
include Eagle Hatchery (V), and Redfish Lake (W). Locations in system. Cowlitz Falls Dam was constructed with a
California include the Upper Sacramento River (X), Yuba River surface collection system designed for juvenile
(Y), Tuolumne River (Z), and Carmel River (AA)
salmonids. The original system collected downstream
Baker River, Washington migrants during 1996–2016 and a new, larger system
was added in 2017. Collected individuals are trans-
The Baker River trap-and-haul program, which began ported downstream * 45 min and released below the
transporting adult salmonids in 1925 after Lower lowest dam in the complex, Mayfield Dam (height =
Baker Dam (river kilometer [rkm] 2; height = 87 m) 76 m), near the adult collection facility. At this site
was constructed, is operated to maintain anadromous adult salmonids are collected and transported
fish populations upstream of impassable dams in the upstream to one of four locations: the Tilton River
system. Juvenile collectors began operating at Lower (* 30 min transport time), Lake Scanewa, the
Baker Dam in 1958 and at Upper Baker Dam (rkm 15; impoundment created by Cowlitz Falls Dam
height = 95 m) in 1960 (Table 1). The juvenile (* 45 min), the Cowlitz River near the town of
facilities were modified to improve collection Packwood, Washington (* 90 min), and the Cispus
River upstream of Lake Scanewa (* 90 min). The
123
trap-and-haul program has successfully re-established ineffective at providing fish passage, primarily
anadromous fish populations in the upper Cowlitz because fish failed to locate the collection facility,
River basin, and current efforts are focused on presumably due to complex water currents in the
increasing populations to a level where stocks are reservoir that failed to promote high discovery rates by
healthy and harvestable (personal communication, outmigrants; facility use was abandoned in 1968.
John Serl, WDFW). Thus, anadromous fish were excluded from the upper
Deschutes River basin for nearly 4 decades until
Lewis River, Washington reintroduction efforts began in 2007 with releases of
summer-run steelhead fry upstream of Round Butte
Three dams were constructed on the North Fork Lewis Dam (rkm 166). In 2009, a new downstream collection
River during 1932–1958, blocking anadromous fish facility began operating at the dam. Collected fish are
from accessing historical spawning habitat in the transported downstream of Pelton Dam (rkm 155;
upper watershed. Efforts to reintroduce winter steel- height = 62 m) and released into the Deschutes River.
head, spring-run Chinook salmon, and coho salmon In 2012, returning adults were collected in the adult
upstream of the Lewis River dams began with an trap downstream of Pelton Dam and the two-way trap-
upstream (one-way) trap-and-haul program in 2012. and-haul program became fully functional. Transport
Adult salmon and steelhead were collected in an time for upstream and downstream migrants is * 45
existing fish trap at Merwin Dam (rkm 31; height = min. Juvenile collection at Round Butte Dam peaked
98 m), the downstream-most dam on the North Fork recently with a total of 933,896 fish in 2017 (PGE
Lewis River. The fish trap was replaced with a new 2018). However, the number of adults transported
upstream collection and sorting facility in 2013. Once upstream has remained low. For example, a total of 20
collected, anadromous adults are loaded onto fish adult Chinook salmon and 30 adult steelhead were
hauling trucks and transported * 60 min to a release transported upstream in 2017 (PGE 2018). Although
site upstream of Swift Reservoir, the upstream-most self-sustaining anadromous fish populations have not
reservoir on the North Fork Lewis River. Juveniles, yet established upstream of the Pelton Round Butte
produced by transported adults, move downstream Hydroelectric Project, efforts continue in the basin to
through Swift Reservoir and arrive at Swift Dam (rkm reach this goal.
77; height = 156 m) where they can be collected in a
floating surface collector that began operating in 2012. North Fork Skokomish River, Washington
Collected juveniles are trucked * 45 min to a release
site located downstream of Merwin Dam. Recent adult On the North Fork Skokomish River, anadromous
returns to the Merwin trap suggest that many of these salmonid runs were severely affected by the construc-
juveniles survive, enter the ocean, and eventually tion of Cushman Dam No. 1 (rkm 32; height = 84 m)
return to the Lewis River: 54% of the early-run coho and No. 2 (rkm 28; height = 72 m) in the 1920s and
salmon collected in 2017 were of natural-origin, up 1930s. These dams did not include fish passage until
from 35% in 2016 and 7% in 2015 (PacifiCorp 2018). 2014 and 2015 when adult collection and juvenile
Natural-origin fish collected at Merwin Dam origi- surface collection systems were installed at Cushman
nated upstream of Swift Dam and thus are products of Dam No. 2 and No. 1, respectively. These installations
the trap-and-haul program (PacifiCorp 2018). A total facilitated plans to reintroduce and restore anadro-
of 17,551 adult fish were captured in the trap in 2017, mous fish upstream. A floating surface collector began
and 8569 of these were transported upstream while the operating in 2015 with assessment of collection
others were taken to the hatchery (PacifiCorp 2018). performance using experimental releases of juvenile
coho salmon and sockeye salmon (from the Baker
Deschutes River, Oregon River project) upstream of Cushman Dam No. 1
(Tacoma Power 2016a, 2017). Collected juveniles are
The Pelton Round Butte Hydroelectric Project was loaded onto trucks and transported * 20 min down-
constructed on the Deschutes River during 1957–1964 stream where they are released below Cushman Dam
and included both upstream and downstream passage No. 2. Reintroduction efforts have not proceeded to
facilities. However, the downstream facilities were the point where adults are transported and released
123
upstream of Cushman Dam No. 1, although testing of response to high temperatures, fishery managers
collection efficiency at the adult facility at Cushman implemented trap-and-haul operations and collected
No. 2 is ongoing (Tacoma Power 2016b). 51 adult sockeye salmon at an adult trap located in the
Lower Granite Dam (rkm 173; height = 30 m; Fig. 1)
fish ladder on the Snake River, Washington. The fish
Intermittent use of trap-and-haul were transported around warm water conditions in the
free-flowing portions of the Snake and Salmon rivers
In some situations, trap-and-haul is used intermittently directly to Eagle Fish Hatchery in southern Idaho
to rescue fish from perilous conditions or to jumpstart (average transport time = 8.3 h), with no mortalities
recovery efforts. Three examples are provided from occurring during trapping and transport (NOAA
the Snake River, Washington–Idaho, the Elwha River, 2015). Fish were transported by truck in two 946 L
Washington, and the Carmel River, California. transport tanks with continuous oxygen flow and a
water recirculating pump (Kozfkay et al. 2017). Cubed
Snake River–Salmon River, Washington–Idaho ice was placed in the tank partway through transport to
temper the water from * 17 C at Lower Granite
Snake River sockeye salmon no longer encounter Dam to * 13 C at Eagle Fish Hatchery (NOAA
impassable dams during their migration (Waples et al. 2015). That same year an additional 24 PIT-tagged fish
1991), but adult trap-and-haul is an important conser- passed Lower Granite Dam and moved upstream
vation strategy that has been used to protect the volitionally, but only 7 (29%) of those fish survived
population from extinction. By the late 1980s, the and arrived in the Sawtooth Valley (NOAA 2015). The
Snake River sockeye salmon population had declined Snake River sockeye salmon example suggests that
to a perilous level and were listed as endangered under applying trap-and-haul intermittently can be an
the U.S. Endangered Species Act (NMFS 1991). This important conservation strategy to protect at-risk
population exists solely within the Sawtooth Valley populations from extreme environmental conditions
basin, Idaho, and is supported by a conservation or to support broodstock collection, even when it
broodstock program (Kozfkay et al. 2019). Anadro- requires long transport distances and times during
mous adults that return to the Sawtooth Valley are summer.
trapped at one of two locations. In years of low returns
all trapped adult fish are transported by truck to the Elwha River, Washington
conservation facility, Eagle Fish Hatchery, approxi-
mately 3.5 h away (Fig. 3; Baker et al. 2017). In years Two dams were breached on the Elwha River in 2012
when the run size is larger (several hundred fish) a to restore anadromous salmonid populations in the
portion of the fish are transported to Eagle Fish system. The river historically supported runs of
Hatchery and others are moved upstream of the traps Chinook salmon, coho salmon, sockeye salmon, pink
and allowed to naturally spawn. In relatively high salmon, chum salmon and steelhead, and researchers
abundance years, the number of fish transported to the have been monitoring how fish recolonize the newly
hatchery often exceeds the brood requirements at the accessible habitat since the dams were removed.
hatchery, and any excess fish at the hatchery are Resource managers implemented trap-and-haul of
transported back to the Sawtooth Valley and released excess hatchery-origin coho salmon during
for spawning (personal communication, Dan Baker, 2011–2017 (Liermann et al. 2017; McHenry et al.
Idaho Fish and Game). Trap-and-haul has also been 2018) because coho salmon exhibit low straying rates
used as an emergency procedure in years when (Pess 2009), potentially limiting the rate at which they
conditions along the migration corridor are poor and could recolonize areas upstream of the former dams.
there is potential for high mortality of adult sockeye Researchers found that transported hatchery-origin
salmon (Keefer et al. 2008a; Kozfkay et al. 2017). For coho salmon successfully spawned in novel habitats
example, Columbia, Snake, and Salmon river water they were released into and produced outmigrating
temperatures in 2015 were exceptionally high in June progeny at comparable rates to other populations in the
and July, coincident with adult sockeye salmon region (Liermann et al. 2017). The authors also
migration through the system (NOAA 2015). In reported that more than half of the spawning adults
123
observed during later study years were not fish that had natural disaster, and to limit exposure by endangered
been transported and concluded that these were fish to dangerous in-river conditions.
progeny from adults transported in previous years
that were returning to their natal sites (Liermann et al.
2017). The Elwha example suggests trap-and-haul can Design and operation of trap-and-haul facilities
be used to kickstart reintroduction efforts in rivers
affected by impoundments. In this section, we summarize the substantial amount
of knowledge that exists for design and operation of
Carmel River, California trap-and-haul facilities to provide a general under-
standing for readers outside the community of trap-
The steelhead population in the Carmel River has and-haul practitioners. We present information on
experienced dramatic declines over several decades adult facilities followed by juvenile facilities in cases
due to several factors including dam construction and where both are discussed. In several cases we provide
extended drought (Arriaza et al. 2017). Efforts to information about various performance metrics of
recover the population have included dam removal trap-and-haul programs (e.g., trap collection effi-
(San Clemente Dam in 2015), habitat restoration, and ciency and prespawn mortality rate). There are many
habitat and fish interventions to minimize mortality similarities between volitional passage facilities and
and increase steelhead dispersal in the system. Trap- trap-and-haul facilities, such as post-passage effects,
and-haul has been a key element in these efforts. Since fallback downstream over a dam after release, stress,
1989, juvenile steelhead have been collected and low trap effectiveness, and migration delay. While
removed from stream reaches where water levels these issues exist for both volitional passage and trap-
become critically low during May–October and are and-haul facilities, providing a direct comparison
reared in a hatchery for later release or transferred to between the two approaches to passing salmonids at
reaches where water levels are sufficiently high to migration barriers was beyond the scope of our review.
allow volitional upstream and downstream movement.
Additionally, trap-and-haul has been used to rescue Fish traps and collection facilities
upstream- and downstream-migrating adult steelhead
that become stranded due to low water levels. These Several studies have described fish traps or collection
fish are typically transported to the ocean and released facilities for adult salmonids (Harmon 2003; Keefer
(Arriaza et al. 2017). et al. 2004, 2005; Morrisett et al. 2018) and existing
guidelines are available for trap construction and
Summary of existing trap-and-haul programs operation (e.g., NMFS 2011). Furthermore, extensive
information useful for designing these facilities is
Our review of 17 existing trap-and-haul programs available from studies of fishways and other passage
(Table 1) indicates these are being operated to achieve devices (e.g., Clay 1995; Haro et al. 1998, 2004;
a broad range of management goals. Many are being Noonan et al. 2012; Williams et al. 2012; Fjelstad et al.
operated experimentally as one of several manage- 2018). Several of the cited studies have emphasized
ment actions aimed at maintaining or improving the importance of siting and design of fishway
salmon and steelhead populations in a given basin entrances. Clay (1995), for example, described the
and some are operated intermittently as conditions entrance as the ‘‘most important single part of any
dictate. Several programs are providing fish access to fishway’’, and the same is likely true for adult
areas located upstream of impassable dams as an collection facilities. To ensure that discovery rates
interim measure, prior to completion of downstream are high, the collector entrance should either have a
passage facilities that may eventually allow full substantial volume of flow (relative to other discharge
implementation of population recovery efforts. In sources) or be placed at a location where fish naturally
some cases, trap-and-haul is being used to provide congregate in response to hydraulic or other naviga-
access to areas located upstream of a natural passage tion cues (Clay 1995; Williams et al. 2012; Fjelstad
barrier, to provide passage to areas blocked due to a et al. 2018). NOAA (2014) recommended that attrac-
tion flow be directed perpendicular to the flow of the
123
river if the attraction volume is low relative to the river forebays fluctuate over a relatively narrow range,
volume, and that attraction flow be directed parallel to water is typically passed through multiple routes (e.g.,
the river if attraction volume is relatively large. The turbines, spillways, sluiceways, fish bypasses) during
source of attraction flow can be extremely important seasons when juvenile salmonids are outmigrating.
because some sources may contain different olfactory When multiple routes are available for downstream
cues or have a different thermal profile than water in a fish passage, fish residence time in a forebay may be
dam tailrace (e.g., Caudill et al. 2013). Potentially reduced (Venditti et al. 2000). However, fish injury
confusing cues or physiochemical variation among and mortality rates can vary considerably among
water sources may cause fish to delay or reject entry passage routes (e.g., Coutant and Whitney 2000;
into a collector or fishway. Bickford and Skalski 2000; Muir et al. 2001; Pracheil
In some cases, the collector entrance is located at a et al. 2016); risk tradeoffs (i.e., between passage delay
river level where fish enter a trap directly. For and mortality risk) are therefore common at such
example, on the North Fork Skokomish River, the facilities. At water storage and flood-control projects,
adult fish trap is immediately downstream from large fluctuations in reservoir water elevations present
Cushman Dam No. 2, and fish can move directly from several challenges to downstream passage, including
the dam’s tailrace into the trap’s holding pool. changes in the availability of downstream passage
Collected fish are crowded into a hopper and raised routes and potentially forced passage via more
to the top of the dam via a tram and jib crane, where hazardous routes such as turbines (e.g., Coutant and
they can be processed at the fish sorting facility Whitney 2000; Schilt 2007; Keefer et al. 2012, 2013).
(Tacoma Power 2016b). However, many adult collec- Downstream passage and collection systems also
tors require fish to enter and ascend a fishway prior to must account for behavioral differences between
entering a trap (e.g., collection facilities in the species and life history types. For example, yearling
Cowlitz, Lewis, and South Santiam rivers). At these Chinook salmon and steelhead undergoing smoltifi-
locations, conditions must promote discovery of the cation generally exhibit directed outmigration behav-
entrance and entry into the fishway opening(s), ior and move downstream quickly, typically in spring
upstream movement through the fishway, and trap when river flows peak, and water temperatures are
entry. cool (Achord et al. 2007; Scheuerell et al. 2009;
Trap designs include features to help retain fish Bourret et al. 2016). Conversely, subyearling Chinook
after entry such as finger weirs (North Fork Toutle salmon often spend time rearing in the migration
River), false weirs (McCutcheon et al. 1994), or Fyke- corridor and therefore move downstream more slowly
style entrances (Tacoma Power 2016b; PacifiCorp than yearlings (e.g., Tiffan et al. 2000, 2012; Venditti
2018). The latter are vertical V-shaped entrances with et al. 2000; Schroeder et al. 2016). The often-
the widest part of the opening located at the down- protracted migration window of subyearlings may
stream end of the entrance, which allows fish to guide include periods when reservoir surface water temper-
along the narrowing entrance as they move upstream. atures are warm and downstream passage options at
Fyke-style openings are wide enough to allow trap dams are limited. An example of this life history
entry but narrow enough that fish have a low occurs in the Willamette River basin, where juvenile
probability of rediscovering the entrance and moving Chinook salmon exhibit a continuum of life histories
out of the holding pool and downstream. At other that include groups of fish that outmigrate as fry, as
locations, such as the barrier dam on the Cowlitz subyearling smolts, and as yearling smolts (Schroeder
River, the ladder exit is located at a higher elevation et al. 2016). Consequently, outmigration occurs during
than the holding pool (i.e., a false weir), so passing fish most months of the year, and migrants can differ
drop into the holding pool and are retained. substantively in size and behavior. Behavioral differ-
Creating effective downstream fish passage at dams ences are also common among populations and among
is a challenging task that can be influenced by a variety life history types of Pacific salmon species (Quinn
of factors including type of dam, reservoir size and 2005; Hillgruber and Zimmerman 2009). For example,
thermal regimes, fish behavior, and differences in chum and pink salmon juveniles outmigrate within
behavior among species. At run-of-river projects weeks of hatching, coho salmon and sockeye salmon
where reservoir storage capacities are limited and primarily outmigrate during their second year of life,
123
Chinook salmon outmigrate in their first or second

year, and steelhead can rear in freshwater for
1–7 years before outmigrating (Quinn 2005).
At water storage projects where fish collection is
needed, surface-oriented systems are used to capture
juvenile salmonids in dam forebays for downstream
transport in several existing trap-and-haul programs
(Fig. 1). These devices were first used in the 1950s and
1960s in attempts to collect outmigrants in large water
storage reservoirs in the Pacific Northwest (Fish
Commission of the State of Oregon 1960; Collins Fig. 4 Collection efficiency estimates for juvenile Chinook
and Elling 1964; Sweeney et al. 2007). The original salmon (filled circles), coho salmon (open circles), sockeye
salmon (filled triangles), and steelhead (open triangles) at
surface collectors, often termed ‘‘gulpers’’, were
surface collectors in Washington and Oregon, 2008–2018. Data
operated using inflows of * 4.2 m3/s or less and by obtained from annual reports from various locations described
most accounts the systems collected few fish. How- in this paper
ever, the concept continued to be developed, and
newer versions in the 2000s used inflows as high as head-of-reservoir collection systems is an emerging
170 m3/s (Kock et al. 2019a). Several of the high- research topic in trap-and-haul science (Liedtke et al.
volume devices collect a high percentage of tagged 2010; Clancey et al. 2017). The survival benefits for
outmigrants ([ 90%; Kock et al. 2019a) while others juveniles transported past reservoirs and dams are
have been far less effective (\ 40%; Fig. 4). The potentially substantial, but there are not currently any
development of surface collectors has continued, and operational head-of-reservoir collection systems and
new facilities are being designed at various locations no data are available to assess the efficacy of this
(e.g., BOR 2016; USACE 2018). Surface collectors approach.
have been designed to capitalize on the tendency for
many juvenile salmonids to migrate near the surface Guidance systems
and follow the bulk flow as they attempt to pass a dam
(Johnson and Dauble 2006; Sweeney et al. 2007). Systems that guide fish towards collection entrances
Supplement 2 in Kock et al. (2019a) provides a (Scruton et al. 2003; Nestler et al. 2008;) or away from
detailed description of the various components of dangerous passage routes or undesirable locations
current state-of-the-art surface collection systems. (Schilt 2007; Kock et al. 2009; Perry et al. 2014;
Head-of-reservoir collection (Fig. 1) is currently Hansen et al. 2019) have received considerable
being considered at several sites where in-reservoir attention in the primary literature. Effective guidance
conditions are thought to limit the number of juvenile systems have enhanced collection at many juvenile
outmigrants that survive, move downstream, and enter and adult collection facilities, including at several of
a dam forebay for potential collection (Liedtke et al. the trap-and-haul case study sites (e.g., on the Baker,
2010; Clancey et al. 2017; Kock et al. 2019b). Head- Lewis, and North Fork Skokomish rivers). Commonly
of-reservoir systems may be effective in reservoirs used physical guidance structures such as nets or walls
that are very large, have complex bathymetry or direct fish towards collection entrances to facilitate
hydrology, or lack velocity cues needed for juvenile discovery rates and minimize passage delay (Adams
salmonids to successfully orient and move down- et al. 2001; Johnson et al. 2005; Kock et al. 2019a). At
stream (e.g., Shasta Reservoir on the Sacramento several locations, suspended vertical net guidance
River, California; Clancey et al. 2017). Such systems systems are used to restrict downstream migrants from
may also be appropriate in reservoirs with large passing through alternate routes and to guide fish to
populations of piscivorous fishes that can substantially collector entrances (Kock et al. 2019a). Net systems
reduce survival of downstream migrants, as in Look- are used on some forebay collectors to funnel fish to
out Point Reservoir on the Middle Fork Willamette the collector entrance (‘‘guide nets’’), decrease the
River (Kock et al. 2019b) and in Fall Creek Reservoir likelihood of fish turning around inside the collector
(Murphy et al. 2019). Development and testing of (‘‘lead nets’’), and prevent fish from passing the dam
123
via alternate routes (‘‘exclusion nets’’; Kock et al. was not available for repairs or maintenance. Liedtke
2019a). Non-physical guidance systems have used et al. (2013) radio-tagged 9 winter-run steelhead and
sound, light, electricity, or aeration (i.e., bubble 11 coho salmon collected in the adult trap and then
screens) to repel fish from locations where passage released the tagged fish * 6 rkm downstream to
conditions are dangerous, such as through turbines or evaluate collection efficiency. None of the coho
into water diversion intakes (Feist and Anderson 1991; salmon and 3 (33%) of the steelhead were eventually
Schilt 2007; Perry et al. 2014). Non-physical systems recollected at the trap (Liedtke et al. 2013). Given the
are attractive because they have the potential to affect small sample sizes in this evaluation, the results are
fish movements without obstructing water flow, but anecdotal, but they do indicate that trapping efficiency
their effectiveness is questionable. In some cases, was apparently very low at the site. Despite the
these systems have produced unintended results such apparent low collection success at the facility, several
as increasing passage rates through dangerous passage hundred winter-run steelhead, coho salmon, and
routes (Schilt et al. 2007; Kock et al. 2009) and may be cutthroat trout have been collected each year and
most effective if used in combination (e.g., strobe transported upstream, maintaining anadromous fish
lights, sound, and bubble screens; Perry et al. 2014). populations in the river.
Adult trap collection efficiency was estimated at
Collection efficiency of fish facilities Foster Dam on the South Santiam River using a
similarly small sample (n = 17) of hatchery-origin
Collection efficiency of adult fish traps has been Chinook salmon in 2017 (M. L. Keefer, unpublished
empirically evaluated at some locations and observa- data). The fish were collected and tagged at a
tions from other sites are useful for understanding temporary trap * 27 km downstream from Foster
factors that may reduce collection success. At Merwin Dam (i.e., they were naı̈ve to the Foster adult fish
Dam on the Lewis River, the adult trap was evaluated collection facility). All 17 salmon moved upstream
in 2015–2017 and two metrics were calculated: trap and were detected in the Foster Dam tailrace, 16 (94%)
entrance efficiency, defined as the proportion of entered the Foster fishway one or more times, but only
tagged fish that entered the trap of all tagged fish that 7 (44%) were eventually collected in the trap. The low
entered the dam tailrace; and trap collection effi- collection efficiency in the radiotelemetry study
ciency, defined as the proportion of tagged fish affirmed observations by fish managers and research-
collected of all tagged fish that entered the dam ers of low adult collection at the Foster facility. The
tailrace (PacifiCorp 2018). Mean annual trap entrance behaviors did not appear to be related to hydraulic or
efficiency estimates were 87.5% for winter-run steel- structural features of the fishway or trap, which were
head (total n = 446 tagged, 3 years), 46.5% for coho recently rebuilt using established specifications
salmon (total n = 184, 2 years), and 90.0% for Chi- (NMFS 2011), but instead may have been related to
nook salmon (total n = 40, 1 year); mean annual trap confusing olfactory or temperature cues. Water for the
collection efficiencies were 70.1% (winter-run steel- facility is drawn from deep within Foster Reservoir
head), 36.2% (coho salmon), and 38.0% (Chinook and large (4–8 C) temperature gradients can occur
salmon) (PacifiCorp 2018). The results from the between the South Santiam River, the Foster Dam
Merwin Dam study illustrate the variability in adult tailrace, and the collection facility (M. L. Keefer,
trapping efficiency among species and that low unpublished data). The circumstances surrounding
collection rates may be an impediment to achieving these apparently low collection rates at the Foster adult
some management goals. The data are being used to fish collection facility highlight the need for post-
inform physical and operational modifications to the construction evaluations and adaptive management at
Merwin adult fish trap to increase future trap collec- these types of facilities.
tion performance. Where reservoir surface collectors are used to
The adult fish collection facility on the North Fork collect downstream migrants, fish collection effi-
Toutle River was in a general state of disrepair after ciency estimates have been obtained in numerous
years of operating under high sediment loads in the telemetry studies and results have varied substantially
system. Features of the collection facility such as the by site, year, species, and life history stage (Fig. 4;
crowder and fish lift were not operational, and funding also see Al-Chokhachy et al. 2018; Kock et al. 2019a).
123
Surface collectors at dams on the Baker and Clacka- physiologically prepared to migrate downstream,
mas rivers have collected 75–98% of the tagged fish may not be susceptible to surface collection during
released during collection efficiency studies (Kock periods of thermal stratification (Fig. 5; also see
et al. 2019a). These facilities are operated using high Beeman et al. 2016a).
inflow (14.2–35.4 m3/s) in relatively small forebays
(7–49 ha). At other locations (e.g., Cushman Dam No. Sorting and holding
1 on the Skokomish River and Swift Dam on the Lewis
River) collection efficiencies have been much lower, Fish sorting is an important element in most adult trap-
ranging from 3 to 30% (PacifiCorp 2019; Tacoma and-haul programs. Sorting can be used to remove
Power 2017). The Cushman and Swift facilities are non-target fish or species (e.g., hatchery-origin fish or
operated using lower inflow (7.1–17.0 m3/s) in large non-native species) from the transported population,
forebays (81–149 ha), compared to surface collectors collect scale or genetic samples to assess age structure
on the Baker and Clackamas rivers. Other recent or monitor reproductive success (Sard et al.
studies have shown that surface collector modifica- 2015, 2016), mark or tag fish to monitor behavior
tions (e.g., reducing noise produced by pumps at Swift and survival after fish are released (Keefer et al. 2010;
Dam on the Lewis River; PacifiCorp 2019) or DeWeber et al. 2017; Naughton et al. 2018; Kock et al.
operational alterations (e.g., extended periods of 2018a), and distribute fish in accordance with man-
constant inflow at Round Butte Dam on the Deschutes agement strategies and regulatory requirements. Most
River; PGE 2018) can substantially improve collec- adult fish collection facilities were designed with
tion efficiency. Several existing surface collectors (see sampling stations integrated into the infrastructure of
previous section ‘‘Downstream trap-and-haul’’) have the collection facility. Various methods of anesthesia
been operated for short periods (\ 5 years), and (carbon dioxide, electro-anesthesia, isoeugenol, etc.)
collection efficiency improvements will likely be have been used to sedate fish for handling at these
realized in coming years as operators become familiar facilities. Many of the guidelines for adult trapping
with each system and modify structures or operations systems and fish handling and sampling include
to adapt to the local environmental and biological considerations for factors such as fish density, fish
circumstances. size, and water temperature (NMFS 2011). Holding
Results from a recent meta-analysis of surface pools at trap-and-haul facilities should be designed to
collectors by Kock et al. (2019a) showed that inflow provide safe conditions where fish can be held for
volume, fish guidance net presence, and size of the short periods (24–96 h) prior to transport. NMFS
collector entrance positively affected collection effi- (2011) provides criteria for factors such as holding
ciency, and that effective forebay area (i.e., the size of
the forebay that fish can access) was inversely
associated with collection. There is also emerging
information that suggests factors such as excessive
noise caused by surface collector pumps may limit
collection success (PacifiCorp 2019). Kock et al.
(2019a) reported steelhead had the highest collection
efficiencies, on average, across projects and Chinook
salmon had the lowest collection efficiencies among
species studied (steelhead, coho salmon, sockeye
salmon, and Chinook salmon). These differences
may be due, in part, to differences in migration timing,
vertical distribution, or physiological readiness to
migrate. For example, subyearling Chinook salmon Fig. 5 Graph showing mean daily depths (black dots) and
move downstream in summer in some rivers when ranges (error bars) of acoustic–tagged subyearling Chinook
salmon within 20 m (horizontal distance) of a surface collector
reservoir surface water temperatures are high, and then
entrance (3.7 m deep) in the forebay of Cougar Dam, South Fork
reside deep in the water column where cooler water is McKenzie River, Oregon, 2014. Legend at top shows water
available. These populations, even when temperature. Figure reprinted from Beeman et al. (2016a)
123
pool capacity under a range of water temperatures, North Fork Skokomish River, a fish lift/hopper system
holding pool inflow, and minimum water depth of the raises fish from the holding pool to an elevation where
holding pool. The document also includes suggestions fish can be released into a tank on a transport truck
for structures that can be used to provide shade and (NMFS 2011). Similarly, a fish lock like the one at
minimize adult jumping, which can lead to fish injury Merwin Dam on the Lewis River, is a mechanical-
or mortality during holding. hydraulic system that raises fish in a water chamber or
Following collection in a surface-oriented system, tower to an elevation where they can be processed or
juvenile salmonids are often conveyed to a fish loaded onto a transport truck. At other locations (e.g.,
holding facility where they can be sampled or held. Baker River, Cowlitz River) holding pools are in large
Most systems pass juveniles through a series of overhead structures under which transport trucks are
separator bars that sort fish by size. Size separation driven and gravity-driven water-to-water transfer
is typically justified by a need to prevent predation of occurs. Some facilities require additional handling to
small fish by larger fish during holding. Smolt-sized move fish to transport trucks. For example, at Roza
fish are often retained for downstream transport while Dam on the Yakima River, fish are crowded from a
fry- and parr-sized fish (i.e., non-migrants such as holding pool into a distribution flume that leads to a
sockeye salmon fry that are incidentally captured) may sampling station where each fish is anesthetized and
be either transported downstream (e.g., Upper Baker processed. After sampling, fish are inserted into a
Dam on the Baker River, Swift Dam on the Lewis Whooshh Fish Transport System (Mesa et al. 2013;
River) or returned to the reservoir to continue rearing Fast et al. 2016; Geist et al. 2016; Kock et al. 2018a)
(e.g., Cowlitz Falls Dam on the Cowlitz River, Round that uses differential air pressure to move adults
Butte Dam on the Deschutes River). Sampling of 12.2 m from the sampling station to the transport
collected juveniles varies among projects. At some truck. On the North Fork Toutle River, transfer
locations, biological data are collected for all fish or all facilities are no longer operational, so fish are hand-
are marked or tagged. At other locations, a portion of netted and carried to a sampling station where they are
the catch is sampled, and non-sampled fish are directly processed and then hand-loaded into transport trucks.
transferred to holding tanks or raceways. NMFS The transport of juvenile and adult salmonids from
(2011) criteria for juvenile holding tanks includes a collection facilities to in-river release sites is relatively
minimum depth of 0.76–0.91 m (30–36 in) and an consistent among trap-and-haul programs. Fish are
inflow of * 19 l/min/kg (* 5 gal/min/lb) of fish. transported in trucks that can maintain dissolved
Juvenile salmonids are typically crowded into a oxygen levels in the transport tank at 6–7 ppm and
hopper from holding tanks on the surface collection maintain water temperature at the ambient level
system as the first step in downstream transport. The (NMFS 2011). It is recommended that transport tanks
hopper is lifted directly to a transport truck or is placed for hauling adult salmonids be ‘‘closed’’ systems with
on a small barge or tramway that leads to a location tanks filled to the maximum water level to reduce
where the hopper can be loaded onto the truck. Fish are water sloshing that could kill or injure fish. Some
transferred to a tank on the transport truck using water- trucks have baffles in the tank to reduce sloshing. As
to-water transfer. an example, a typical transport truck used in the
Willamette River basin has a 5678 l (1500 gal)
Transport and release uninsulated stainless-steel transport tank that contains
two aerators, supplemental oxygen, and temperature
A variety of approaches have been used to move adult and oxygen monitors (personal communication, Doug
salmonids from holding tanks or pools to vehicles for Garletts, USACE). For most of the trap-and-haul
upstream transport. Several of the case study facilities programs in Table 1, transport times range from 10 to
described above were designed to limit handling 60 min, but in some cases, fish are hauled for longer
during this process and to transfer fish using auto- periods. Release sites used by the various trap-and-
mated features such as crowders or fish lifts. Various haul programs vary widely in terms of the number
lifting devices are used to load fish onto transport used and their locations within a basin. However, the
trucks including fish lift/hopper systems, and fish fish release process is generally consistent. Transport
locks (NMFS 2011). At Cushman Dam No. 2 on the trucks are positioned near the release site, a gravity-
123
fed pipe is attached to the egress opening on the maintaining oxygen and water temperature at safe
transport tank, the tank is opened, and fish pass levels during transport, recommended the addition of
through the pipe into the receiving water body. NMFS salt (5–10% solution; Moyle and Cech 1988; Mazic
(2011) recommended that fish should not drop more et al. 1991) to the transport water to minimize
than 1.8 m (6 ft) from the pipe to the water surface, the osmoregulatory stress for juveniles and adults, and
impact velocity should not exceed 7.6 m/s (25 ft/s), summarized available information on water condi-
and the receiving water should be at least 0.9 m (3 ft) tioners, anesthetics, acclimation, and carrying
deep. capacity.
Factors such as loading time, hauling density,
transport timing, and disease transfer have been
evaluated at several locations with varying results. Biological responses to trap-and-haul
Colvin et al. (2018) evaluated factors affecting hauling
mortality of adult spring-run Chinook salmon in the Trap-and-haul programs and volitional fish passage
South Santiam River and Middle Fork of the Wil- facilities both have the potential to affect individual
lamette River in 2006–2013. They found that fish fish and fish populations through a variety of biolog-
loading time and hauling density were positively ical mechanisms. Many fish experience migration
associated with hauling mortality but observed that delays and stress during handling and passage events,
these relationships varied between rivers and among which in turn can affect swimming performance,
years. The authors noted that trip-specific factors such increase disease transmission rates and infection risks,
as water quality, dissolved oxygen levels, and and even lead to premature death. Adult fish that pass
pathogen loads in the transport tank likely affected dams volitionally or are transported upstream may fall
hauling mortality on a trip-by-trip basis. back past the dams (Boggs et al. 2004; Naughton et al.
Downstream transport of juveniles occurs daily 2018) or fail to locate spawning habitat, and not
during most of the fish collection season at most contribute to population productivity. Similarly, juve-
facilities. At some locations, daily collection numbers nile fish that use fish bypass systems or are transported
require multiple transport events each day during peak may be injured or infected in these systems. Trans-
outmigration periods. Following transport, fish are ported fish have an additional risk of elevated stray
typically released into holding ponds where they rates as adults compared to fish that remain in-river
remain for at least 24 h to recover from handling stress during juvenile outmigration, resulting in decreased
associated with collection and transport. The holding reproductive potential of the transported population
ponds are often referred to as ‘‘stress relief ponds’’ and (Keefer and Caudill 2014). We consider these factors
are typically located adjacent to the free-flowing river. and present information about what is known for trap-
NMFS (2011) requires all transported juvenile fish to and-haul in the following sections.
be transferred to stress release ponds where they Migration delay and survival.
remain for at least 24 h, and then be allowed to There is limited information on how trapping
volitionally enter the river and resume downstream affects migration timing and survival in existing
migration. If fish remain in the stress relief pond trap-and-haul programs, but several studies (e.g.,
for [ 48 h, NMFS (2011) requires fish to be forced Bromaghin et al. 2007; Marauskas et al. 2014;
from the pond by crowding or draining of the pond. Morrisett et al. 2018) have evaluated these factors in
Fish-handling that occurs during loading, transport, other settings and provide useful insights. Murauskas
and release is a critical period in the trap-and-haul et al. (2014) evaluated passage delay and blockage of
process because of the relationship between stress and adult sockeye salmon at a combined passage and
fish health. Harmon (2009) conducted a review of trapping facility on the Wenatchee River, Washington,
transport methods to identify best practices for reduc- in 2008–2012. Trapping operations occurred 7 days
ing stress and maintaining water quality in fish per week in 2008–2010 and median passage delay
transport tanks. He reported that it was important to ranged from 0.4 to 8.7 days while an estimated 8–38%
minimize potential stressors and the duration of of the returning adults were prevented from migrating
exposure to those stressors to maximize transport upstream past the trap (Murauskas et al. 2014). After
survival rates. The review included methods for reducing trap operations to \ 24 h per week in
123
2011–2012 (fish were allowed to volitionally pass severity of the stress varies seasonally; (2) is associ-
when the trap was not operating), the authors reported ated with recovery times of several days; (3) can lower
that median passage delay decreased to 6 min and swimming ability; and (4) result in cumulative stress
nearly all adults migrated to the spawning grounds. due to exposure to various elements encountered
Morrisett et al. (2018) evaluated how trapping at sequentially in a collection facility. The trapping and
Lower Granite Dam affected in-ladder transit time and holding environment affects stress levels of trapped
upstream homing success for Chinook salmon, sock- fish. For example, juvenile Chinook salmon trapped
eye salmon and steelhead in 2012–2016. The authors during the day under darkened conditions appeared
found that trapping delayed mean transit time through less stressed than those exposed to the sun, and
the fish ladder by 4–18 h and noted that fish delayed by exposure to high densities of other fish species was
10 h or more had lower homing success than fish that stressful during collection (Schreck and Li 1985).
freely passed through the fish ladder and were not Thus, providing shade or creating darkened conditions
routed to the trap. These examples illustrate the need in trapping and holding facilities, and minimizing
to carefully consider how trap operations may affect holding densities, appear to be reasonable measures
fish behaviors and collection rates as well as post- for minimizing stress. Schreck and Li (1985) found
transport effects in a trap-and-haul program. that loading juvenile fish into a transport vehicle
appeared to be the most stressful aspect of the actual
Handling effects transport process and reported it took fish several days
to recover. Stewart et al. (2017) found it could take
Data from the Yakima River basin are useful for juvenile salmon weeks to recover from the cumulative
assessing the safety of transporting adult salmon. stress of capture and transport. Transporting juvenile
Spring-run Chinook salmon broodstock for the Cle salmon can have other effects such as lowered disease
Elum Supplementation and Research Facility resistance, reduced ability for conditioning to an
(CESRF), operated by the Yakama Nation, are odorant, and failure to complete smoltification fol-
collected at Roza Dam each year. Adult Chinook lowing transport and release (i.e., residualization;
salmon destined for the CESRF are PIT-tagged prior Schreck 1980). However, if conditions are good (i.e.,
to transport, which allows researchers to monitor post- good water quality, low fish density) and trip duration
transport survival of adults to the time of spawning, is appropriate, fish can partially recover from stress
and to track fecundity, fertilization success, and while a vehicle is en route to the release site (Schreck
embryo survival rates after spawning occurs. Data 1980, 1985). Results from these studies illustrate the
collected in 2001–2018 showed 96.3% of the [ need to carefully consider how fish are collected, held,
10,000 adult Chinook salmon transported from Roza and transported to minimize stress and its associated
Dam to CESRF (about 60 min transport time) sur- effects on fish health and performance.
vived to spawning (data courtesy of Yakama Nation In summary, the design and operation of trap-and-
Fisheries). The high survival rates, over nearly 2 haul systems is critical for determining the amount of
decades, indicate collection, handling, and transport of stress fish will incur during collection, transport and
adult Chinook salmon can be safely accomplished release. Stress endured during trap-and-haul may
under suitable conditions and have minimal effects on substantially affect fish performance and survival
survival to the time of spawning. following release. Therefore, facility operations and
Numerous in situ and laboratory studies have handling procedures should aim to minimize severity
shown that fish trapping and handling can negatively and duration of stress. Here, we provide recommen-
affect physiological responses (e.g., those associated dations for measures to minimize stress based on
with the stress response) and whole organism perfor- personal experience with trap-and-haul operations and
mance (e.g., disease resistance, escape behavior, supported by a large body of research (Wedemeyer
ability to enter saltwater, ability to imprint on olfac- 1976; Schreck 1980; Schreck and Li 1985; Maule et al.
tory cues, etc.) of both juvenile and adult salmonids 1988; Schreck et al. 1989, 1995, 1997, 2006; Benda
(Specker and Schreck 1980; Schreck and Li 1985; et al. 2015; Schreck and Tort 2016; Cogliati et al.
Maule et al. 1988; Congleton et al. 2000). Schreck and 2019):
Li (1985) found trapping of fish: (1) is stressful and
123
• Trap-and-haul operations should be designed to be disruption of the integrity of the skin caused by
fish-friendly and to minimize stressor exposure crowding and handling.
duration. Fish collection, transfer, holding and • Trap-and-haul programs often incorporate a fish
transport structures should not produce turbulent tagging element into the process. From a stress
water conditions and should be free of sharp angles management perspective, tagging should be done
and edges that can cause injuries. Providing shade in such a way that the fish perceived it as part of
or creating darkened conditions, particularly in some other stressful event (i.e., so the fish receives
shallow water, can be beneficial. Water-to-water a single rather than two, sequential stressors). For
transfers are highly recommended because netting example, fish could be tagged as part of the loading
and direct handling induce high stress responses, process, rather than separately before loading
especially if fish are dewatered. If netting is occurs.
necessary, soft mesh should be used and nets with • The effects of stress experienced during trap-and-
knotted twine should be avoided because the knots haul is less severe if fish are released in locations
can cause skin injuries. Further, mesh should be where they can regain orientation without exposure
appropriately sized for the fish being handled. If to high-velocity currents that are energetically
mesh size is too large for a particular size class of demanding. The release site should not contain
fish eye injuries and blindness can occur during substantial predator concentrations to allow suffi-
netting because a fish’s snout can pass through the cient time for fish to recover without facing the
mesh. Furthermore, overloading nets should be challenges of predation avoidance.
avoided because fish in the bottom of a full net can
be crushed or injured. This is particularly prob-
Fallback
lematic for small juveniles and sexually mature
female adults.
The term ‘‘fallback’’ refers to the downstream move-
• Overcrowding that negatively affects water quality
ment, generally at a dam, by fish that would be
in the holding and transport phases should be
expected to actively move upstream to locate potential
avoided. The severity of stressors in trap-and-haul
spawning areas. Fallback has been observed and
can be reduced by minimizing the risk of infection
documented for a variety of Pacific salmon species
with pathogenic organisms to optimize post release
(Reischel and Bjornn 2003; Boggs et al. 2004; Richins
performance of fish. Minimizing cross infection
and Skalski 2018), Atlantic salmon (Salmo salar;
can be accomplished by lowering fish density,
Hagelin et al. 2016; Frechette et al. 2019), and
elevating flow, and for many pathogens, maintain-
alewives (Alosa pseudoharengus; Frank and Mather,
ing cool water temperature. Treating fish with
2009) among others. Fallback rates for Pacific salmon
antibiotics to increase their survival after release is
at mainstem dams on the Snake and Columbia rivers
an option for consideration. Holding fish in a
have been well documented (Reischel and Bjornn
system that does not harbor necessary hosts of
2003; Boggs et al. 2004; Naughton et al. 2006), with
certain pathogens would eliminate the risk of
annual run-specific estimates at individual dams
further infection of these pathogens.
ranging from 1.9 to 22.0% for adult Chinook salmon,
• The timing of specific operations should be
sockeye salmon, and steelhead. Although high num-
considered as a measure that could limit stress.
bers of upstream migrants can fall back at dams with
For example, juvenile releases that occur during
fishways, many of the fish eventually reascended the
dark evening hours when fish are prone to migra-
dam where fallback occurred and resumed upstream
tion and less vulnerable to predators could be used
migration (Reischel and Bjornn 2003; Boggs et al.
to reduce stress and increase survival.
2004; Naughton et al. 2006). Other fish remained
• Adding salt to transport water to approximate the
downstream from the fallback location (e.g., Keefer
physiological level of saline can help fish over-
et al. 2008b; Richins and Skalski 2018). Reascension
come osmoregulatory difficulties associated with
is not an option for adult salmonids that fall back via
stress and be beneficial in reducing the severity of
spillways or turbines at most high-head dams. Most
the transport experience. Commercially available
fallback fish at these sites are lost to the anadromous
additives for transport water also help with
123
population upstream of the dam, although those that prespawn mortality in trap-and-haul programs, but
escape serious injury may potentially re-enter adult readers should be aware that prespawn mortality
traps or spawn at downstream sites (Naughton et al. occurs in systems where volitional passage is possible
2018). (Bowerman et al. 2016), as well as in undammed river
In the Cowlitz River, Kock et al. (2016) reported systems with thriving salmon populations (Quinn et al.
fallback rates for transported adults ranging from 7 to 2007).
22% for natural-origin and hatchery-origin steelhead Evaluation of prespawn mortality related to trap-
and Chinook salmon following released upstream (3.9 and-haul has received considerable attention in the
rkm) of Cowlitz Falls Dam during 2005–2009 and Willamette River basin (Keefer et al. 2010; Benda
2012. Fallback by hatchery-origin fish was lower for et al. 2015; DeWeber et al. 2017; Bowerman et al.
fish released at alternate release sites located farther 2018; Colvin et al. 2018; Naughton et al. 2018). In the
upstream than the primary release location (Kock et al. Middle Fork Willamette River, mean annual mortality
2016). Similarly, on the South Santiam River, of * 30–60% was reported for several transported
Naughton et al. (2018) reported 14% of tagged groups of tagged Chinook salmon, and some release
Chinook salmon released into Foster Reservoir fell groups experienced [ 90% mortality (Keefer et al.
back downstream of Foster Dam compared to 1% of 2010; Naughton et al. 2018). In the South Santiam
tagged Chinook salmon released in the river upstream River, DeWeber et al. (2017) reported prespawn
of Foster Reservoir. In 2017, Kock et al. (2018b) mortality rates were low (B 13%) in 2009–2012 but
monitored behavior and movements of adult sockeye were very high (74%) in 2013. Outside the Willamette
salmon after release upstream of Cle Elum Dam and River basin, Kock et al. (2018a) reported 8% of
found 20% of tagged fish moved downstream of the sockeye salmon that were transported upstream and
dam prior to spawning. On the Elwha River, fallback released directly into Cle Elum Reservoir died prior to
over a dam was not an option because the dams had spawning in 2017.
been removed but McHenry et al. (2018) reported Survival to spawning of trap-and-hauled adult
hatchery-origin coho salmon outplanted into the salmonids can be highly variable among watersheds
mainstem Elwha River fell back to a hatchery at a and among years (Keefer et al. 2010; DeWeber et al.
higher rate than fish outplanted into tributaries. Key 2017; Bowerman et al. 2018; Kock et al. 2019b).
takeaways from these findings are: (1) fallback rates Variation in prespawn mortality may be in part
can be substantial at locations where fish volitionally explained by seasonal, annual, and watershed-scale
pass dams and at locations where trap-and-haul is used variation in environmental conditions affecting infec-
to move fish around dams; (2) fallback seems to occur tion and/or disease resistance rates, as prespawn
for similar proportions (up to 25%) of fish, regardless mortality has frequently been associated with patho-
of how they pass dams; and (3) increasing the distance gens (Benda et al. 2015; Bowerman et al. 2016).
between release sites and the dam is a potential Holding early-run fish in water of good quality and
strategy for reducing fallback rates in trap-and-haul free of pathogens prior to transport may enhance
programs. survival (Benda et al. 2015). Survival of transported
adults also appears to be negatively affected by the
Prespawn mortality time it takes to load fish into trucks, fish density in the
tank, timing of hauling trips, thermal exposure, and
Adult salmon can experience significant mortality low instream flows at release sites (Colvin et al. 2018).
during freshwater migration or when holding on the Warm water temperature, in particular, appears to be
spawning grounds prior to spawning (Keefer et al. an important driver of prespawn mortality (Keefer
2010; Hinch et al. 2012; Bowerman et al. 2016). The et al. 2010; Benda et al. 2015; Bowerman et al. 2018;
latter is termed ‘‘prespawn mortality’’ and it can be Naughton et al. 2018). Spring-run Chinook salmon
exacerbated by trap-and-haul programs due to stress may be especially vulnerable due to their extended
during collection and transport (Benda et al. 2015; holding prior to spawning during the warmest (sum-
Colvin et al. 2018), and the fact that fish are released mertime) part of the year. This observation was
into an environment they did not volitionally enter. supported by Benda et al. (2015) who showed that
This section focuses on factors known to affect outplanted Chinook salmon were 12.6 times more
123
likely to experience prespawn mortality than groups of (Baumsteiger et al. 2008). Evans et al. (2016) evalu-
Chinook salmon held at a hatchery facility in 13 C ated reproductive outcomes of transported hatchery-
water. Bowerman et al. (2018) observed that prespawn origin and natural-origin Chinook salmon on the South
mortality rates were highly variable when average Santiam River and found individuals had highly
water temperature was in the 18–20 C range, but that variable reproductive success with some fish produc-
nearly all estimates of prespawn mortality exceeded ing as many as 40 adult progeny while others produced
80% when mean water temperature was [ 20 C. none. The authors also noted a possible trend towards
Other factors such as sex, fish origin, and transport reduced fitness in mate pairs from hatchery-origin and
protocols (i.e., timing, fish density) also affect pres- natural-origin fish which led them to conclude that use
pawn mortality rates. Keefer et al. (2010) found that of natural-origin fish may improve population pro-
females had lower physical condition and higher ductivity in such programs. Weigel et al. (2019)
prespawn mortality than male counterparts, while evaluated steelhead spawning success in the South
Bowerman et al. (2018) found prespawn mortality Santiam River and found that more than half of the fish
rates were positively correlated with the proportion of outplanted in 2012–2016 produced either juvenile or
hatchery-origin fish on the spawning grounds. The adult offspring. Sard et al. (2015) found that factors
adult Chinook salmon transport study by Colvin et al. such as release location and release date had incon-
(2018) indicated fish density and transport truck sistent effects on the reproductive success of Chinook
loading time were predictive of transport mortality. salmon in the South Fork McKenzie River and
DeWeber et al. (2017) reported prespawn mortality observed that origin had little effect. Reproductive
decreased for groups of salmon hauled later in the success has been high enough to achieve cohort
season, but noted that other studies (e.g., Evans et al. replacement in some locations (South Santiam River;
2015; Sard et al. 2015) have shown early-transport Evans et al. 2016) but not others (South Fork
groups have higher rates of reproductive success. The McKenzie River; Sard et al. 2016).
mentioned studies, and others, have generally con-
cluded that prespawn mortality is likely influenced by Operations that minimize stress for fish are critical
multiple additional factors, such as previous dam
passage, fishing pressure, instream habitat, energetic The design and operation of trap-and-haul systems is
status, and pathogen loads. Collectively, research on critical for determining the amount of stress fish will
prespawn mortality suggests risks can be substantial in incur during collection, transport and release. Stress
some circumstances, can vary widely even at single endured during trap-and-haul can substantially affect
study sites, and causation can be difficult to conclu- fish performance and survival following release;
sively establish. facility operations and handling procedures should
therefore aim to minimize the severity and duration of
Reproductive success stress. In this section we provide recommendations for
measures to minimize stress based on personal expe-
Research has shown that many transported adult rience with trap-and-haul operations and supported by
salmonids survive and are reproductively successful a large body of research (Wedemeyer 1976; Schreck
upstream of impassable dams when trap-and-haul is 1980; Schreck and Li 1985; Maule et al. 1988; Schreck
used (Baumsteiger et al. 2008; Sard et al. 2015, 2016; et al. 1989, 1995, 1997, 2006; Benda et al. 2015;
Evans et al. 2016; Weigel et al. 2019). Baumsteiger Schreck and Tort 2016; Cogliati et al. 2019).
et al. (2008) used parentage analysis to estimate the Trap-and-haul operations should be designed to be
reproductive success of outplanted hatchery-origin fish-friendly and to minimize stressor exposure dura-
Chinook salmon in Shitike Creek, Oregon in 2002 and tion. Fish collection, transfer, holding and transport
2003. They found hatchery-origin outplants produced structures should not produce turbulent water condi-
juveniles from matings between outplanted males and tions, be free of sharp angles and edges that can cause
females as well as from matings with natural-origin injuries, and provide shade or darkened conditions,
fish present in the stream. They also observed some particularly in shallow water conditions. Water-to-
outplanted males mated with up to seven females and water transfers are highly recommended because
some outplanted females mated with up to four males netting and direct handling induce high stress
123
responses, especially if fish are dewatered. If netting is currents that are energetic demanding. The release site
necessary, soft mesh should be used and nets with should not contain substantial predator concentrations
knotted twine should be avoided because the knots can to allow sufficient time for fish to recover without
cause skin injuries. Further, mesh should be appropri- facing the challenges of avoiding predation.
ately sized for the fish being handled; the use of mesh Each trap-and-haul system is unique and within a
sizes too large for a particular size class of fish can lead given system day-to-day variation in stress experi-
to eye injuries and blindness because a fish’s snout can enced occurs (Schreck et al. 2006). The severity of
pass through the mesh. Furthermore, overloading nets stress experienced is a function of numerous variables
should be avoided because fish in the bottom of a full including: (1) the general wellbeing of the fish as they
net can be crushed or injured. This is particularly enter a trapping system (e.g., are they already stressed
problematic for small juveniles and sexually mature or infected?); (2) how close other environmental
female adults. variables are to being optimal for the species involved
Overcrowding that negatively affects water quality (e.g., water that is warmer than optimal can have an
in the holding and transport phases should be avoided. exponentially-increasing negative effect as it
The severity of stressors in trap-and-haul can be approaches tolerance levels); (3) the duration of each
reduced by minimizing the risk of infection with stressful element of trap-and-haul experienced (in
pathogenic organisms to optimize post-release fish general, the longer the exposure to a stressor, the
performance. Minimizing cross infection can be exponentially greater the stress experienced); and (4)
accomplished by lowering fish density, elevating flow, the cumulative effects of stressors of each element of
and for many pathogens, maintaining cool water trap-and-haul (e.g., traps can be injurious to fish, the
temperature. Treating fish with antibiotics to increase loading of a transportation vehicle, and the release
their survival after release is an option for consider- system and process can each be very stressful). There
ation. Holding fish in a system that does not harbor are critical unknowns in predicting success of trap-
necessary hosts of certain pathogens would eliminate and-haul programs, but minimizing stress experienced
the risk of further infection by these pathogens. by fish throughout the process is clearly an important
The timing of specific operations should be con- element affecting performance of fish subjected to
sidered as a measure that could limit stress. For trap-and-haul.
example, juvenile releases that occur during evening
hours when fish are prone to migration and less
vulnerable to predators could be used to reduce stress Critical gaps in trap-and-haul knowledge
and increase survival. The addition of additives, such
as salt, to transport water can be beneficial in reducing The trap-and-haul case studies described above illus-
the severity of the experience. Adding salt to approx- trate how oftentimes the programs evolve substan-
imately the level of physiological saline can help fish tively over time. Technological advances, regulatory
overcome osmoregulatory difficulties associated with changes, improved biological understanding, and
stress. Commercially available additives for transport changing stakeholder priorities are common and these
water help with disruption of the integrity of the skin all precipitate new programmatic goals and objectives
caused by crowding and handling. Trap-and-haul (Fig. 1). As part of our review, we identified several
programs often incorporate a fish tagging element key information needs that are common across many
into the process. From a stress management perspec- trap-and-haul programs. The examples in this section
tive, it would be desirable if the tagging process was can be used by fisheries managers to both assess how
done in such a way that the fish perceived it as part of trap-and-haul processes can be improved at current
some other stressful event (i.e., so the fish receives a facilities and to inform the planning and implemen-
single rather than two, sequential stressors). For tation of future programs.
example, fish could be tagged as part of the loading
process, rather than separately, before loading occurs. Juveniles: post-transport survival
The effects of stress experienced during trap-and-haul
is less severe if fish are released in locations where Few studies have evaluated post-transport survival of
they can regain orientation without exposure to heavy juvenile salmon and steelhead for the case study
123
projects we reviewed. In general, evaluations on Downstream transport of juveniles in the two-way

juvenile downstream migrants tend to focus on factors trap-and-haul programs described above currently
such as collection efficiency because effective collec- occurs without segregation, although juveniles gener-
tion of downstream migrants is a bottleneck for many ally are separated from large-bodied fish including
programs (Lusardi and Moyle 2017; Kock et al. adult salmonids. Studies that compare post-release
2019a). These studies provide insights into the physiology and survival of groups of fish hauled with
proportion of downstream migrants that are success- and without species separation and under various
fully collected and transported but typically provide densities and water temperatures would be straight-
little information on survival following release. Post- forward and would provide valuable information to
transport survival studies of downstream migrants inform whether segregation benefits outweigh addi-
would help managers understand how factors such as tional handling stress.
handling and transport methods contribute to delayed
mortality of transported fish. There are several good Juveniles: head-of-reservoir collection
research templates for evaluating post-transport sur-
vival questions, with many developed as part of the The development of effective head-of-reservoir juve-
large-scale juvenile transportation program in the nile collection systems is an important emerging
Columbia and Snake rivers (e.g., Muir et al. 2006; research and development need. Such systems could
Rechisky et al. 2014; Gosselin and Anderson 2017). facilitate two-way trap-and-haul programs at locations
Examples include experimental post-transport holding where reservoirs are large, have complex bathymetry,
evaluations and a variety of tagging studies to evaluate warm surface water temperatures, lack downstream
short- and long-term survival. While the Columbia and velocity cues, or contain large populations of pisciv-
Snake river transportation program uses different orous predators. Juvenile salmon and steelhead mor-
transport methods (barges) with longer transport times tality in these types of impoundments can be
([ 24 h) and distances (up to * 470 rkm) than the substantial (Rieman et al. 1991; NMFS 2008). The
case study projects we described, their study designs ability to collect a high proportion of downstream
could be readily adapted to assess post-transport migrants at head-of-reservoir sites and transport them
survival of juvenile salmonids associated with other around the reservoir may improve the likelihood of
juvenile or adult trap-and-haul programs. program success at these locations. However, head-of-
reservoir collection has rarely been attempted and
Juveniles: separation by species for transport successful systems have not been developed to date.
Clancey et al. (2017) described a reservoir screen
Research has shown that co-transport of juveniles system that may eventually be used to modify water
from two or more fish species can result in increased temperature near a potential head-of-reservoir fish
stress levels and may have implications for post- collection system in Shasta Lake (California). The
release survival (Congleton et al. 2000; Kelsey et al. head-of-reservoir juvenile collection system in Shasta
2002). For example, Congleton et al. (2000) found Lake would consist of an inclined plane collector with
stress indices of juvenile Chinook salmon were highest guidance nets and a temperature curtain located within
during the peak of the outmigration period when approximately 1 km of where juveniles enter the
transport loading densities were high, and salmon reservoir from a tributary (BOR 2015). Currently, it is
stress levels were correlated with steelhead loading unclear whether downstream migrants can be suc-
density. Kelsey et al. (2002) reported cortisol levels cessfully collected in reservoirs, and feasibility testing
were significantly higher for juvenile Chinook salmon of prototype head-of-reservoir designs is needed.
held in tanks with juvenile steelhead than for salmon
held without steelhead, and salmon behaviors differed Adults: prespawn mortality
among treatments. While these co-transport results
suggest sorting collected fish by species and hauling As described previously, prespawn mortality of trans-
them separately may be a good practice, it is possible ported adults is a major management issue for some
additional handling offsets potential benefits of sep- trap-and-haul programs. The mortality has been
aration, particularly when transport times are short. associated with a number of covariates, including fish
123
traits (e.g., pathogen burden, injury status, energetic spatially segregate groups of hatchery-origin and
reserves, origin), handling and transport effects (e.g., natural-origin fish.
collection and holding methods, transport density, One of the more promising genetic monitoring
transport distance), and river environment encoun- approaches is the use of parentage analysis to estimate
tered before and after transport. Several research adult-to-adult cohort replacement rates (Botsford and
groups have addressed portions of the complex suite of Brittnacher 1998; Sard et al. 2016). Cohort replace-
factors that affect mortality in transported adult ment can be estimated as the number of adults that
populations, but considerable uncertainty remains. It return in future years (i.e., across all adult age classes)
is likely the most influential factors will vary among divided by the number of adults transported and
species, trap-and-haul programs, locations, environ- released in a given brood year. Replacement greater
mental conditions and years. Research that may help than 1.0 would indicate that fish in the trap-and-haul
untangle the mechanisms affecting prespawn mortal- program are self-sustaining or population growth is
ity include: studies addressing effects of specific trap- increasing. In contrast, a sequence of replacement
and-haul protocols (e.g., Colvin et al. 2018) on adult estimates less than 1.0 would indicate population
stress and pathogen transmission; studies that evaluate decline and potential sink dynamics. Comparing
alternatives to direct adult transport to spawning sites, cohort replacement rates among transported and non-
such as temporary holding in facilities with high water transported populations in a multi-year study would be
quality (e.g., Benda et al. 2015) or release into useful for understanding whether trap-and-haul
thermally-stratified reservoirs where adults can select upstream of migration barriers significantly improves
preferred thermal niches prior to spawning (e.g., overall salmonid population dynamics.
Naughton et al. 2018); and studies that integrate
behavior, experiences at collection facilities, and Populations: productivity
presumptive mortality risk factors.
A main goal of trap-and-haul programs is to expand
Adults: post-transport reproductive success population abundance and spatial distribution in a
and cohort replacement given basin by providing access for adult spawners to
additional, and in many situations, more suitable habi-
In an earlier section we discussed genetic studies by tat than if they were not provided upstream transport.
Baumsteiger et al. (2008), Sard et al. (2015, 2016), and Population expansion is most likely to occur if
Evans et al. (2016) that provided information on the conditions such as water temperature and spawning
reproductive success of transported adults. These habitat quality upstream from a dam can support
studies showed that success varied substantially biological outcomes (e.g., growth, survival) that result
among individual adults and among years in trap- in a net increase in basin-wide productivity. Hypo-
and-haul populations in the Deschutes and Willamette thetically, the combined juvenile productivity from
river basins. Additional research on this topic would adults that spawn upstream and downstream of a dam
be valuable for several reasons. The use of genetics to would be higher than productivity of the downstream
track the reproductive success of individual fish can group alone, unless transported adults that were culled
reveal important insights into factors such as adult from the downstream population are not contributing
transport timing, release location, and the effects of to overall reproductive success.
origin on the success of producing viable offspring. In addition to increased productivity and positive
Such information can be useful for optimizing trap- population growth rates, McElhany et al. (2000)
and-haul methods to maximize the number of juvenile identified three additional parameters that are key to
fish produced by transported adults. Additionally, evaluating and establishing the viability (i.e., having a
genetic or fish-marking studies may provide insights negligible risk of extinction) of a salmonid population:
into factors such as how adults disperse from release overall population abundance, population spatial
locations and whether fish released in different structure, and diversity. Trap-and-haul programs can
locations or at different times have a reproductive influence population viability through each of these
advantage. In the South Fork McKenzie River, for parameters. Generating productivity estimates for
example, fishery managers use separate release sites to entire watersheds is expensive and logistically
123
challenging. This is true even for a straightforward life history types. Chinook salmon in some impounded
scenario such as with coho salmon, which have a non- rivers, for example, move downstream as fry, sub-
complex life cycle. The abundance of female spawners yearling smolts, yearling smolts, and age 1 ? smolts
requires information on the number of smolts pro- (Keefer et al. 2012; Schroeder et al. 2016). Collection
duced, marine survival and harvest exploitation rates, efforts tend to target yearling smolts because these fish
and the proportion of smolts that return as adult are larger, are physiologically prepared for seaward
females (Bradford et al. 2000). Estimating smolt migration, and perhaps are more likely to seek
abundance requires either direct smolt assessments at downstream passage routes at dams. Active smolt
migrant traps or information on the number of outmigration for several species occurs in spring when
spawners, fecundity, egg deposition rates, and egg- reservoir surface temperatures are cool and surface
to-fry, fry-to-parr and parr-to-smolt survival (Ward collection is more likely to be feasible. In contrast,
and Slaney 1993). Not surprisingly, there have been younger life stages tend to move downstream in a less
few sustained efforts to calculate such estimates in directed fashion that includes rearing behavior (e.g.,
existing trap-and-haul programs. However, collecting Connor et al. 2013; Bourret et al. 2016) and often
these data would allow potential impacts (e.g., John- occurs in summer when reservoir surface collection
son et al. 2012) or benefits to population productivity strategies are less effective. Thus, these fish are more
from trap-and-haul to be evaluated, similar to evalu- challenging to collect, and the collection period often
ations of other conservation actions. For example, must occur over an extended period. Facilitating
spatially explicit, multiple life-stage population mod- collection of all juvenile life history types should be
els have been used to evaluate habitat variables prioritized at all sites as this is a key parameter of
(Honea et al. 2009) and climate effects on habitat population viability (McElhany et al. 2000).
restoration (Battin et al. 2007). Statistical analyses can More generally, understanding existing—and ide-
assess changes in productivity (expressed as the per ally historical—life history diversity in a trap-and-
capita growth rate in the absence of harvest) associated haul population is a prerequisite to effectively pre-
with specific actions, such as hatchery production serving or restoring populations. Such traits are valued
(Buhle et al. 2009). Quantifying spawner-recruit and needed for a population to persist and maintain
relationships, variability in productivity among spe- long-term viability under future environmental con-
cies, watersheds and programs, and understanding ditions and intra- and inter-year variability. Conser-
potential limiting factors would help managers assess vation actions that address human impacts on
the effectiveness of trap-and-haul as a recovery or phenotypic variation may be necessary to preserve
population expansion tool. evolutionarily significant biodiversity, such as the
premature migrating phenotypes also known as
Populations: life history diversity ‘‘spring-run’’ (Thompson et al. 2019). Trap-and-haul
is a potential conservation action to address the loss of
Simplifying the life-history portfolio of a population phenotypic variation and life history diversity in target
through anthropogenic activities has negative conse- populations. However, when considering trap-and-
quences on long-term population viability, and prac- haul as part of a reintroduction effort, the adaptive
tices that help diversify population structure may be suitability of potential source stocks is needed, for
critical for populations subject to disturbance regimes example, due to the loss of spring-run alleles from
(Greene et al. 2010) such as climate change (Battin selective pressures associated with dams (e.g.,
et al. 2007). These activities can have major effects on Thompson et al. 2019).
phenotypic variation across a broad array of species
and traits, and maintaining intraspecific phenotypic Populations: genetic, phenotypic,
variation is an important component of conserving and demographic effects
biodiversity (Thompson et al. 2019).
Trap-and-haul programs generally focus on col- The presence of migration barriers and the process of
lecting downstream migrants during the yearling collecting and transporting fish around these barriers
smolt life stage, but this management approach may are likely to have a variety of population-level impacts
be insufficient for populations that express a mix of (e.g., Angilletta et al. 2008; Thompson et al. 2019) that
123
are currently not well understood. Migration delays at upstream population (e.g., Araki et al. 2008). To date,
collection facilities can affect the phenology of evaluations of source-sink dynamics have received
upstream and downstream migrations and may also little attention in trap-and-haul programs, despite the
have important delayed effects on survival and fitness. potential importance of these effects on programmatic
For example, early migrating adult phenotypes that and regional management and conservation goals.
arrive late on spawning grounds may have reduced
reproductive opportunity and decreased lifetime fit- Populations: effects on conspecifics and other
ness (e.g., Dickerson et al. 2005). Similarly, ocean endemic species
arrival timing for juveniles can be a critical determi-
nant of growth opportunity and survival to adulthood Adult source-sink dynamics are just one example of
Scheuerell et al. 2009; Satterthwaite et al. 2014; how trap-and-haul populations may affect con-
Weitkamp et al. 2015). Trap-and-haul operations specifics. The timing and location of post-transport
should clearly strive to collect and transport fish at juvenile releases may also affect juveniles that rear
times when fish would be volitionally migrating, downstream via resource competition, especially for
though this may initially be impractical in some fish that may still be rearing (as opposed to active-
systems. Managers and researchers should also migrant smolts). Juvenile salmonid release sites have
attempt to understand the degree to which collection also been associated with predator aggregations (Col-
facilities select for specific phenotypes or demo- lis et al. 1995; Shively et al. 1996; Scheel and Hough
graphic groups, as these types of selection biases can 1997) that may increase mortality risk for both
rapidly and significantly reduce the productivity and transported and resident fish. These effects, and
resilience of populations (e.g., Hard et al. 2008; Naish perhaps others like pathogen transmission, can poten-
et al. 2008; Ohlberger et al. 2018). tially occur at multiple spatial and temporal scales
from site-specific effects within watersheds resulting
Populations: source/sink dynamics in broader impacts on regional populations. Very few
studies have examined potential effects of trap-and-
In some impounded rivers, adult salmonids success- haul on other native species (Hardiman et al. 2017;
fully reproduce and may be self-sustaining down- Upper Columbia United Tribes 2019), but such
stream from dams. These fish may be natural-origin impacts should be considered. Transported salmonids
individuals using historical spawning sites in tribu- may compete with or prey upon other species and have
taries or main stem sites, or they may be the progeny of the potential to introduce pathogens or parasites. Trap-
hatchery stocks or of hatchery-wild hybrids. In situa- and-haul effects on both conspecifics and other
tions where origin is mixed or uncertain, it may be endemics may be of particular concern when large
important for managers to limit the collection and numbers of fish are transported relative to the abun-
transport of adult fish that originated downstream from dance of natural-origin fish, and potential impacts
the dam for trap-and-haul programs. This may be should be carefully considered early in the decision-
especially true when the downstream population has making process.
legal status (i.e., threatened or endangered, or species
of concern) and low abundance. Trap-and-haul oper- Operations: pathogen and parasite transmission
ations that capture natural-origin adults from down-
stream populations will likely result in lost production The potential for infected fish to infect other individ-
from the source populations (e.g., Schtickzelle and uals during high-density holding and transport can be
Quinn 2007; Johnson et al. 2012). Conversely, man- high (Ogut et al. 2005; Van Gaest et al. 2011), but
agers may seek to preserve a natural-origin population pathogen infection prevalence and transmission rates
upstream from a dam by using selective trap-and-haul have rarely been studied in trap-and-haul programs.
(e.g., Evans et al. 2016). In such cases, incidental For adults, horizontal transmission could have signif-
collection of hatchery-origin or hybrid fish from below icant effects on the reproductive potential of trans-
the dam may result in the transfer of undesirable ported fish if infectivity rates are high and result in
genetic or phenotypic traits from the downstream prespawn mortality, as hypothesized by Benda et al.
population and reduce the productivity or fitness of the (2015) and Colvin et al. (2018). Juvenile migrants are
123
also at risk of infection in collection facilities and so it is not surprising that trap-and-haul use has
during transport, while both juveniles and adults have expanded in the Pacific Northwest in recent decades.
the potential to infect non-transported conspecifics or Implementation of trap-and-haul began as early as
congenerics after release. Research needs include: 1925, but more than half of the programs reviewed
studies of pathogen and parasite prevalence in trap- were implemented since 2000 (9 of 17 programs;
and-haul facilities and populations; identification of Table 1). Older facilities have been modified if needed
optimal fish transport densities to minimize infection (e.g., Baker and Deschutes rivers) and the following
rates; effectiveness tests of antibiotics or other programs are now being considered:
prophylactic treatments on transported fish, where
• On the upper Columbia River (Washington), the
permitted (i.e., with hatchery-origin fish); and studies
Upper Columbia United Tribes are moving for-
of the interactions among infectivity rates, collection
ward with reintroduction efforts aimed at restoring
facilities and transport tanks, environmental risk
salmon and steelhead to portions of the more than
factors like warm water temperatures, and biological
1770 km of river and stream habitat blocked by
risk co-factors such as elevated stress and
Grand Coulee Dam (rkm 597; UCUT 2019).
immunosuppression.
• On the Yakima River (Washington), trap-and-haul
will support efforts to provide passage upstream of
six headwater dams, including Cle Elum Dam
Discussion
(BOR and DOE 2012), as described previously.
• In California, reintroduction may be necessary to
Trap-and-haul: a tool for management
achieve delisting criteria for ESA-listed salmonids
and conservation
in the Central Valley (NMFS 2014). Unoccupied
habitats that historically supported winter-run
Trap-and-haul is one of the only options currently
Chinook salmon, spring-run Chinook salmon, or
available to resource managers for reestablishing
steelhead have been prioritized, including in the
connectivity in impounded river systems. While dam
McCloud River, Battle Creek, the Yuba River, and
removal is often proposed and discussed, it has rarely
the San Joaquin River (NMFS 2014); NMFS has
occurred in large rivers to date. Therefore, trap-and-
been discussing the feasibility of reintroducing
haul is an important tool that provides access for non-
salmon and steelhead above high-head dams on the
listed, threatened, and endangered salmon and steel-
Yuba and Tuolumne rivers with stakeholders
head to spawning and rearing habitats located
(personal communication, Steve Edmundson,
upstream of impassable dams. Areas above dams
National Oceanic and Atmospheric
often contain habitats that were occupied historically
Administration).
and are of high cultural importance, and reestablishing
• On the Chehalis River (Washington), local author-
connectivity among river reaches can support fish
ities have incorporated trap-and-haul into a pro-
population viability through increased abundance,
posed flood retention dam designed to operate
productivity, diversity and spatial structure.
intermittently for short periods (up to 32 days)
The 17 programs reviewed here were implemented
when major and catastrophic floods are retained
to achieve a broad range of management objectives. In
(https://chehalisbasinstrategy.com/eis/proposed-
some cases, trap-and-haul is used simply to provide
project/).
passage at a natural barrier (e.g., South Fork Sky-
komish River). At other locations, it has been used in Critics of trap-and-haul point to costs associated
attempts to reestablish self-sustaining populations of with developing, installing, operating, and maintain-
anadromous fish upstream of a dam (e.g., Deschutes ing facilities and the potential for negative biological
and South Santiam rivers). Options for addressing fish effects, such as those described above. These are valid
passage at high-head dams are limited, and trap-and- concerns that also apply to volitional fish passage
haul can be used to address multiple management facilities. Given the limited options for providing fish
objectives. Furthermore, ESA-listed salmon and steel- passage at high-head dams, there remains an imminent
head populations on the U.S. west coast have not need to advance the state-of-knowledge on trap-and-
recovered (Northwest Fisheries Science Center 2015), haul programs and their operations. Research on trap-
123
and-haul continues and new studies are likely to hatchery supplementation, water management scenar-
provide additional insights that will improve our ios, ocean productivity, and climate. The characteri-
understanding of how this tool affects survival, adult zation of a program as unsuccessful has little merit if
spawning, reproductive success, juvenile life history neighboring populations that lack trap-and-haul are
diversity and overall population productivity. Data also in decline due to factors not related to trap and
collected to date have primarily focused on quantify- haul (i.e., poor ocean conditions, climate regime,
ing the number of fish transported. At some sites hatchery effects). Conversely, characterizing a pro-
trapping efficiency has also been evaluated. However, gram as successful has little merit if cohort replace-
research techniques have advanced and can provide ment is persistently less than 1.0 (e.g., Sard et al.
valuable information on individual- and population- 2015). Also, results from the evaluation of individual
level responses to trap-and-haul techniques. For aspects of a trap-and-haul program should be viewed
example, in the South Santiam River, genetic studies in the context of other passage options such as how
have addressed productivity and population replace- volitional fishways perform. For example, up to 20%
ment questions (Evans et al. 2016; Weigel et al. 2019) of the transported adults fall back at dams in some
and telemetry studies have evaluated post-release adult trap-and-haul programs (Kock et al.
adult behaviors and survival (Keefer et al. 2010; 2016, 2018b; Naughton et al. 2018), but this percent-
Naughton et al. 2018). Similarly, genetic studies in the age is nearly identical to fallback documented at some
Cle Elum River have been used to identify stock- run-of-river dams on the Snake and Columbia rivers
specific differences in spawning times, spawning where volitional passage occurs (Reischel and Bjornn
location, and productivity (Matala et al. 2019), and 2003; Boggs et al. 2004; Naughton et al. 2006). Data to
telemetry has been used to estimate losses due to inform biological metrics such as cohort replacement
fallback and prespawn mortality (Kock et al. 2018b). rates need to be collected in basins with and without
These examples illustrate the type of detailed infor- trap-and-transport to place overall trends in context,
mation that can be obtained and the importance of although this is challenging from a study design and
conducting a suite of studies to assess biological cost standpoint. As noted above in the knowledge gaps
benefits and consequences of trap-and-haul and to section, information on multiple performance metrics
collect data that can be used to optimize methods and is needed to fully evaluate and quantify the effects of
protocols. trap-and-haul on fish populations. While identifying
Context is critical when evaluating overall program clearly defined metrics of success is an important first
effectiveness. Given this review, it seems appropriate step in the development of a trap-and-haul program
that effectiveness be judged by stakeholders involved (Lusardi and Moyle 2017), assessing success is
in each basin and program because they understand its complex and its definition may substantively differ
management objectives, and their judgements should among stakeholders, river basins, and regions.
be informed by results of scientific studies and
analyses. Lusardi and Moyle (2017) concluded that Tools are needed that address climate change
no two-way trap-and-haul program is an unequivocal
success. However, given the diverse views by stake- The development of trap-and-haul programs to
holders of what constitutes program success and the reestablish salmonids upstream of impassable dams
different management objectives trap-and-haul pro- may be critical for population persistence if climate
grams are designed to meet, this conclusion is change predictions are realized in the coming decades
premature and unsubstantiated in our view, and does (Mantua et al. 2010; Shanley and Albert 2014;
not align with evidence from many one-way trap-and- Herbold et al. 2018; Crozier et al. 2019). Crozier
haul programs. Evaluating the success of a trap-and- et al. (2019) conducted a climate vulnerability assess-
haul program requires that multiple factors be consid- ment for salmon and steelhead populations in Wash-
ered. These include program-specific goals identified ington, Oregon, and California and found that 23 of the
by stakeholders and resource managers, potential 33 distinct population segments (DPS) evaluated had
effects of parallel management actions in the basin, high or very high vulnerability ratings. Several of
in-river conditions that fish face prior to collection and these DPSs were vulnerable to warm stream temper-
could face in the future, and population-level effects of atures during both juvenile and adult freshwater life
123
stages. In many rivers, volitional fish passage is such as this one, are needed to inform resource
available at main stem run-of-river and low-head management and conservation actions.
diversion dams, but high-head storage dams block
access to headwater, tributary streams that can contain Integrated, adaptive, and sustained management
extensive suitable spawning and rearing habitat. These
areas are likely to be more resistant to climate change Trap-and-haul is often implemented as one of several
compared to areas located lower in watersheds. High- management actions in a watershed that includes—for
elevation tributaries can serve as climate refuges, example—habitat restoration, streamflow regulation,
support increased diversity, and provide for larger water diversion screening, and hatchery supplemen-
populations if fish are provided access (Crozier et al. tation. To optimize success, these activities should be
2019). In California, improving access to upstream implemented as an integrated suite of actions. Recent
habitats is likely to improve the ability of salmon to studies have identified trap-and-haul as an important
persist in a changing climate (Herbold et al. 2018). element in the management toolbox (Lusardi and
While access to these areas is blocked in many basins, Moyle 2017; Herbold et al. 2018; Crozier et al. 2019)
interest in developing reintroduction programs to and there are specific aspects of the river environment
restore salmonid populations upstream of impassable that can sometimes be managed to enhance these
dams is ongoing. Short of dam removal, trap-and-haul activities. For example, while high-head dams restrict
is one of few, and may be the only economical and passage to upstream areas, they also store cold water
culturally acceptable option at these dams to restore deep in their reservoirs. At some locations, such as
watershed connectivity and expand population Cougar and Detroit dams in the Willamette River
productivity. Basin, this cold water is used to manage water
Developing new technologies and refining best temperatures in downstream river reaches to support
management practices are critical steps needed to resident and anadromous salmonid spawning and
support trap-and-haul efforts in the future and help rearing. In 2004, Cougar Dam was retrofitted with a
address effects of climate change. Fish collection is water temperature control tower that allows water to
often a bottleneck (Lusardi and Moyle 2017) that can be released from varying depths in the forebay. A
result in migration delay, increase stress levels in fish, similar project is under development at Detroit Dam in
and in extreme cases prevent passage for a significant conjunction with a new juvenile fish collection
portion of a juvenile or adult population. Technolog- facility. The integrated fish transport and river tem-
ical advances aimed at increasing attraction, entrance, perature management facilities should help address
and retention characteristics at existing fish traps and multiple fishery objectives upstream and downstream
collection facilities will significantly improve the of dams in both river systems.
process of trapping and transporting fish. Also, These Willamette River Basin examples point to a
development of new collection systems capable of growing trend where salmonid productivity and
collecting fish at novel locations, such as head-of- anadromy is supported throughout a watershed by a
reservoir sites, may significantly improve the ability to broad suite of management actions. This includes
maximize fish collection rates, reduce losses that occur managing environmental conditions downstream of
during migration through the reservoir, and support dams, re-establishing connectivity to headwater habi-
maintenance of life history diversity and overall tats upstream of dams through adult trap-and-haul and
productivity of salmon and steelhead populations. supporting life history diversity and life-cycle pro-
Optimized fish holding and handling procedures are ductivity by collecting juveniles across the entire
likely to be important in the future to minimize stress calendar year and transporting them downstream. This
and its associated effects on the physiologic and expanded capability to manage a broad spectrum of
performance characteristics of transported fish, the river will be useful to resource managers. For
because transported individuals are likely to face example, downstream water temperatures and flow
increasingly challenging conditions in other portions could be managed to increase the effectiveness of
of their migratory pathways (Herbold et al. 2018; habitat restoration, improve conditions for adult
Crozier et al. 2019). Periodic reviews to summarize salmon and steelhead migrating upstream to reach a
the state-of-knowledge on trap-and-haul programs, collection facility, reduce stress while holding prior to
123
collection, minimize stress during collection, holding phase of a trap-and-haul program. The efficacy of the
and transport, and reduce prespawn mortality after program relies on collection of information about
release. where juvenile and adult fish congregate near potential
collection locations under different operational sce-
Implementation needs to be adaptively managed narios, how juvenile fish distribute in the water column
and supported long-term of a reservoir seasonally, and during which times of
the year fish are expected to be near the collection
Trap-and-haul requires a long-term commitment of location and available for capture. Similarly, the
time and resources and the resulting studies and physical nature of the collection system needs to be
operations being implemented must be adaptively evaluated carefully to optimize the design of the
managed and supported to ensure they are effective various system elements and ascertain not only how
(Fig. 1). This is because design criteria have been well the system will collect fish but also the effect of
developed to maximize the likelihood of collection the collection on fish wellbeing. All elements includ-
success and ensure fish safety at juvenile and adult ing discovery, entry, passage to holding facilities,
collection facilities, but variability in performance separation and handling, transport vehicle loading, in-
occurs due to localized environmental conditions, route specifics, vehicle release system, release timing,
project operations, the configuration of the reservoir and release location need to be considered.
relative to the dam, and by fish species and life stage. This review highlights the importance of starting
Performance can also vary within a given year and with design guidelines and then assessing the site-
among years as environmental conditions and water specific criteria required for a successful passage
levels in a reservoir change, and with basin hydrology program through rigorous observation of local condi-
and climate. tions. Implementation of trap-and-haul programs
Each trap-and-haul system is unique, and within a requires years, if not decades, to design, install, and
given system day-to-day variation in the level of stress test facility performance, discuss test results with
experienced occurs (Schreck et al. 2006). The severity stakeholders, and make decisions on adaptive changes
of stress experienced is a function of numerous to facilities or operations. The changes will then
variables including: (1) the general wellbeing of the require additional design and testing. In short, the
fish as they enter a trapping system (e.g., are they development of a successful trap-and-haul program
already stressed or infected); (2) how close the other requires a commitment by stakeholders, regulators,
environmental variables being experienced by the fish and project owners to a long-term process of continued
are to being optimal for the species involved (e.g., evaluation that ideally follows an adaptive manage-
water that is above optimal can have an exponentially- ment strategy developed and agreed upon before
increasing negative effect as it approaches tolerance implementation, and a financial commitment to the
levels); (3) the duration of each stressful element of approach.
trap-and-haul that is experienced (in general, the
longer the exposure to a stressor, the exponentially
greater the stress experienced); and (4) the cumulative Summary
effects of stressors of each element of trap-and-haul
(e.g., traps can be injurious to fish, and the loading of a The goal of this review was to provide a comprehen-
transportation vehicle, and the release system and sive synthesis of the current state-of-knowledge on
process can each be very stressful). There are critical trap-and-haul for management of Pacific salmonid
unknowns in predicting the success of trap-and-haul populations in impounded river systems. Numerous
programs, but minimizing stress experienced by fish trap-and-haul programs exist throughout the Pacific
throughout the process is clearly an important element Northwest, with operating histories ranging from
affecting the performance of fish subjected to trap- years to decades and with a broad range of manage-
and-haul. ment goals and objectives. The development and
The considerable variation in effectiveness among operation of trap-and-haul infrastructure is guided by
systems discussed above underscores the importance established criteria and a solid research base on many
of conducting robust evaluations during the pre-design aspects of fish behavior. However, fish handling, an
123
intrinsic requirement of trap-and-haul, can increase the permitted use, you will need to obtain permission directly
fish stress which is negatively associated with fish from the copyright holder. To view a copy of this licence, visit
http://creativecommons.org/licenses/by/4.0/.
health and survival. Thus, improvements to infras-
tructure and fish handling protocols should strive to
minimize stress effects.
Existing programs have experienced a range of References
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Rev Fish Biol Fisheries

Review of trap-and-haul for managing Pacific salmonids

Received: 29 June 2020 / Accepted: 31 October 2020

Introduction Trap-and-haul is being used to restore connectivity

Two-way trap-and-haul programs

efficiencies over time and were eventually replaced

Cowlitz River, Washington

Chinook salmon outmigrate in their first or second

projects we reviewed. In general, evaluations on Downstream transport of juveniles in the two-way

J Fish Manag 28:1472–1485. https://doi.org/10.1577/M07- https://odfw.forestry.oregonstate.edu/willamettesalmonidrme/

behavior. Hydrobiologia 635:237–249. https://doi.org/10. https://doi.org/10.1577/1548-8659(1998)127%3c0118:

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