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Steelhead Overshoot and Fallback Rates in the Columbia–Snake River Basin and the Influence of Hatchery and Hydrosystem Operations

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Steelhead Overshoot and Fallback Rates in the Columbia–Snake River Basin and the Influence of Hatchery and Hydrosystem Operations North American Journal of Fisheries Management © 2018 American Fisheries Society ISSN: 0275-5947 print / 1548-8675 online DOI: 10.1002/nafm.10219 ARTICLE Steelhead Overshoot and Fallback Rates in the Columbia–Snake River Basin and the Influence of Hatchery and Hydrosystem Operations Shelby M. Richins and John R. Skalski* Columbia River Basin Research, School of Aquatic and Fishery Sciences, University of Washington, 1325 4th Avenue, Suite 1515, Seattle, Washington 98101, USA Abstract Tributary overshoot occurs when adult salmonids homing to natal sites continue upstream past the mouth of their natal stream. Although overshooting is a common behavior by steelhead Oncorhynchus mykiss in the Columbia River basin, it has not been adequately quantified or explained. Using multistate release–recapture models, we examined the prevalence of overshooting and fallback to natal tributaries by 37,806 PIT-tagged steelhead from 14 tributaries of the Columbia River basin during 2005–2015. Eight populations had overshooting rates exceeding 50% in at least 1 year. Source of hatchery stock, rearing location, and release practices were found to have appreciable effects on overshoot rates. Overshooting was elevated in hatchery stocks reared upstream of release sites, but this effect may be lessened by utilizing endemic broodstocks and acclimating juveniles within the release basin. For one population of hatchery steelhead, acclimation within the release basin was found to decrease overshooting from 81% to 40%. Across both hatchery and wild populations, successful homing was found to decline 4 percentage points for every 5-percentage- point increase in overshoot rate. Average annual fallback probabilities ranged from 0.18 for Walla Walla River hatch- ery steelhead to 0.75 for Umatilla River wild steelhead. Fish stocks with the greatest fallback probabilities also had the greatest interannual variability in fallback rates. For John Day River wild steelhead and Tucannon River hatchery steelhead, the interannual range in fallback probabilities exceeded 0.50. We found evidence that spill at dams during March may enhance the fallback of overshooting steelhead and contribute to increased homing to natal tributaries. Therefore, additional attention should be paid to facilitating downstream dam passage of adult salmon. Homing is a general pattern in many migratory animals Tributary overshoot occurs when adult fish homing to in which reproductive adults return to natal sites or the natal sites continue upstream past the mouth of their natal site of previous reproduction. Homing in rivers presents a stream (Ricker 1972). Subsequent return downstream is special challenge compared to lakes or marine environ- called “overshoot fallback” (Boggs et al. 2004; Naughton ments; decisions must be made at each branch of the river, et al. 2006). Overshooting has been documented in steel- and homeward migrations may include movement in head (Keefer et al. 2008a; Copeland et al. 2015), Chinook many compass directions. Pacific salmon Oncorhynchus Salmon O. tshawytscha (Boggs et al. 2004; Keefer et al. spp. and steelhead O. mykiss are believed to imprint using 2008b; Gallinat and Ross 2009), Sockeye Salmon O. nerka multiple “landmarks” along their out-migration route as (Ricker and Robertson 1935), and Atlantic Salmon Salmo juveniles—a process called sequential imprinting (Ueda salar (Økland et al. 2001). Observed overshoot distances 2012; Bett and Hinch 2016). As adults, salmon and steel- range from less than 1 river kilometer (rkm; Ricker and head navigate complex river systems by identifying waters Robertson 1935) to 200 rkm or more (Boggs et al. 2004). that contain these memorized olfactory stimuli (Johnsen There is growing awareness that tributary overshoot is and Hasler 1980; Quinn 2005). a common behavior by steelhead in the Columbia River *Corresponding author: [email protected] Received December 20, 2017; accepted July 27, 2018 1 2 RICHINS AND SKALSKI basin, with as many as half the fish in some populations Khan et al. 2013). Survival of postspawn steelhead mov- passing their natal river and ascending upstream dams ing through dams is lowest through turbines and juvenile (Bumgarner and Dedloff 2011; Copeland et al. 2015; Kee- bypass systems (Harnish et al. 2015). When available, fer et al. 2016). Unless steelhead successfully fall back at both prespawn steelhead (Khan et al. 2009) and steelhead dams to return downstream to natal sites, high levels of kelts (Wertheimer and Evans 2005; Wertheimer 2007; overshooting may significantly deplete naturally spawning Khan et al. 2013; Rayamajhi et al. 2013; Harnish et al. populations. Although tributaries may also receive over- 2015) exhibit a very strong preference for safer surface shooting steelhead from other rivers, large influxes of routes, such as spillways and sluiceways. The study by strays—particularly hatchery strays—can negatively Khan et al. (2009), who strongly recommended sluiceway impact population fitness (Chilcote 2003; Chilcote et al. operation during winter months to prevent turbine fall- 2011). back by adult steelhead at The Dalles Dam (rkm 309 on The act of homing may require exploration of multiple the Columbia River), supports the hypothesis that surface pathways to determine the correct route (Ricker 1972; flow at dams may benefit steelhead that overshoot their Keefer et al. 2008b). Therefore, it is unclear whether tribu- natal rivers. tary overshoot is part of the normal homing process or Many populations of salmon and steelhead in the whether it is exacerbated by human-induced changes to Columbia River basin are at risk of extinction (NMFS river systems and steelhead populations. Disruption of 2012); therefore, reliable estimates of overshoot and fall- memory development during imprinting or the decay back rates, as well as a greater understanding of the fac- of memories over time may inhibit the homing abilities of tors influencing these behaviors, are important for the salmonids and lead to higher rates of straying (reviewed management and conservation of steelhead in the Colum- by Keefer and Caudill 2014). Hatchery practices (Nish- bia River basin. Unfortunately, previous studies may have ioka et al. 1985; Pascual et al. 1995; Marchetti and Nevitt underreported overshooting rates because they used fish 2003), juvenile barging (Bugert and Mendel 1997; Keefer with unknown sources (Boggs et al. 2004; Keefer et al. et al. 2008c; Bond et al. 2017), temperature modification 2008a) or utilized a tally-based approach that ignored (Isaak et al. 2018), migration pathway obstruction (Khan detection efficiencies (Bumgarner and Dedloff 2011; Mur- et al. 2009), or extended ocean residency (Labelle 1992) doch et al. 2012; Keefer et al. 2016). To address these can have unexpected consequences for homing by adult knowledge gaps, our objectives were to (1) quantify over- salmonids and may also influence overshoot behavior. shooting and fallback to natal tributaries for multiple pop- In addition to correctly navigating to natal sites, many ulations of steelhead in the Columbia River basin using salmonids must also locate temporary holding areas that multistate release–recapture methods; (2) assess the extent are both thermally appropriate and energetically efficient. to which tributary overshoot and fallback are associated Interior Columbia River basin steelhead are stream matur- with hatchery rearing, juvenile barging, and ocean resi- ing, meaning that they (1) enter freshwater before they are dence time; and (3) investigate the effect of winter spill fully mature, (2) overwinter in rivers, and (3) spawn the and dam outflow on fallback to natal tributaries. following spring (Robards and Quinn 2002). A review by Quinn et al. (2016) found that this “premature migration” occurs in many anadromous fishes, potentially because of METHODS mortality risks at sea or physical factors in freshwater sys- This study used tagging and detection data from the tems (e.g., temperature or flow) that seasonally limit Columbia River basin PIT Tag Information System (PTA- access to spawning sites. The early return of adult steel- GIS), operated by the Pacific States Marine Fisheries head to freshwater may prompt overshooting to reach Commission (PSMFC 2015). Passive integrated transpon- suitable holding areas prior to their final movements to der tags are small, internal tags (10–14 mm long) with dis- spawning grounds. Overshooting by Chinook Salmon tinct identification codes that can be detected when a (Keefer et al. 2008b) and steelhead (Richins 2017) in the tagged fish passes through a magnetic field (McCutcheon Columbia River basin was found to increase as water tem- et al. 1994; Gibbons and Andrews 2004). Known-source peratures rose in late summer, indicating that the behavior steelhead from nine tributaries of the Columbia River and may serve thermoregulatory purposes. five tributaries of the Snake River were selected based on Overshooting within the Columbia River hydrosystem adequate sample sizes of PIT-tagged steelhead and the is of concern because downstream dam passage can be presence of instream detection sites in the natal tributaries hazardous for large adult fish (Ferguson et al. 2008) and from 2005 to 2015. All hatchery and wild populations can add significant migration delays (Wertheimer and included at least 250 tagged adults distributed across Evans 2005; Rayamajhi et al. 2013; Harnish et al. 2015). 8 years or more (Table 1). All steelhead were tagged as General options for downstream dam passage include tur- juveniles, and natal tributaries
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