1. Iteroparity and Steelhead: what we know and don’t know John R. McMillan Oregon State University

2. Introduction to Reproduction Reproductive strategies (big bang v. bet- hedge) –Annual strategy in plants and semelparous strategy in animals Reproduce one time –Perennial strategy in plants and iteroparous strategy in animals Reproduce more than one time Salmonid species –Pacific salmon – semelparous (Altukhov et al. 2000) –All other species – iteroparous (Wilson 1997)

3. Introduction to Reproduction Reproductive strategies (big bang v. bet- hedge) –Annual strategy in plants and semelparous strategy in animals Reproduce one time –Perennial strategy in plants and iteroparous strategy in animals Reproduce more than one time Salmonid species –Pacific salmon – semelparous (Altukhov et al. 2000) –All other species – iteroparous (Wilson 1997)

4. Selection and Traits Life history theory (Stearns 1976; Charlesworth 1994; Crespi and Teo 2002) –Semelparity Select for higher juvenile survival – Increased egg size Select for no adult survival –Increased investment in egg weight, female nest guarding, secondary sexual characteristics, breed under higher densities –Iteroparity Low or inconsistent juvenile survival –Smaller egg size High adult survival –Reduced investment in eggs, secondary sexual characteristics, lower breeding density, and no female nest guarding

5. Selection and Traits Life history theory (Stearns 1976; Charlesworth 1994; Crespi and Teo 2002) –Semelparity Select for higher juvenile survival – Increased egg size Select for no adult survival –Increased investment in egg weight, female nest guarding, secondary sexual characteristics, breed under higher densities –Iteroparity Select for lower juvenile survival –Smaller egg size Select for higher adult survival –Reduced investment in egg weight, secondary sexual characteristics, no female nest guarding, breed under lower densities

6. Steelhead Patterns Steelhead –One-time reproduction generally most common Highly variable rates of repeat spawning (0 – 79 %) –Iteroparous individuals Typically female (Burgner et al. 1992; Wertheimer and Evans 2005) Typically smaller sized (Teo and Crespi 2002; Hendry and Stearns 2004) More common in ocean-maturing life history (Busby et al. 1996) Latitude and distance from sea –Highest levels at latitudinal extremes (e.g., Russia, Savvaitova et al. 1999; South America, Riva-Rossi 2007) –Similar observations for Atlantic salmon (Jonsson and Jonsson 2004) –Perhaps because energy consumption increases with water temperature –Lowest levels appear to be in furthest inland populations (Meehan and Bjornn 1991; Narum et al. 2008)

7. Steelhead Patterns Steelhead –One-time reproduction generally most common Highly variable rates of repeat spawning (0 – 79 %) –Iteroparous individuals Typically female (Burgner et al. 1992; Wertheimer and Evans 2005) Typically smaller sized (Teo and Crespi 2002; Hendry and Stearns 2004) More common in ocean-maturing life history (Busby et al. 1996) Latitude and distance from sea –Highest levels at latitudinal extremes (e.g., Russia, Savvaitova et al. 1999; South America, Riva-Rossi 2007) –Similar observations for Atlantic salmon (Jonsson and Jonsson 2004) –Lowest levels appear to be in furthest inland populations (Meehan and Bjornn 1991; Busby et al. 1996; Narum et al. 2008)

8. NorthSouth RussiaWashington - OregonAK - Canada Latitudinal Distribution of Repeat Spawn Rates for Steelhead Derived largely from Busby et al. 1996

9. Recruitment First-time spawners don’t always replace themselves (e.g., Waddell Creek, Keogh River; Hal Michael, personal communication) Repeat spawners in Atlantic salmon can produce disproportionate share of recruitment (Chadwick 1987; Mills 1989) Why is Iteroparity Important?

10. Fitness –Increased lifetime fitness Greater fecundity Repeat spawning female steelhead produced twice as many offspring as one- time spawners (Seamons et al., in prep) –Spread risk over multiple generations (Fleming and Reynolds 2004; Hendry and Stearns 2004)

11. What Human Factors Influence Iteroparity? Environment –Habitat conditions experienced by adults during return to ocean Dams Climate Oversummering habitat in some cases Sport and commercial fisheries –Level of adult exploitation and encounter rates during return to ocean Selection against larger fish could select against iteroparous individuals –This has been suggested to have occurred in Atlantic salmon (see Jonsson and Jonsson 2004)

12. Fisheries –Sport Rate of adult exploitation & encounter by sport anglers during return to ocean? –Energy expenditure –Commercial Selection against larger fish could select against iteroparous individuals –This has been suggested to have occurred in Atlantic salmon (see Jonsson and Jonsson 2004)

13. Patterns, Processes, Uncertainties Lots of hypotheses, little data –Does iteroparity matter? –What causes iteroparity? Natural v. human influences –How does this factor into our management regimes? Abundance v. diversity