186 
Fishery Bulletin 117(3) 
atypically warm years, when upwelling and forage condi¬ 
tions were above average, the frequency of occurrence of 
multiple brooding in chilipepper (S. goodei) was common 
and nearly equal in both Southern and Central Califor¬ 
nia, whereas previous studies had not detected multiple 
broods in chilipepper of Central California (Wyllie Eeh- 
everria, 1987; Stafford et al., 2014). This finding might 
indicate that species capable of producing multiple 
broods do so more frequently under warmer ocean con¬ 
ditions when food resources are sufficient, potentially 
providing some advantage over single-brooding species 
in the face of warming trends. 
Fisheries managers are beginning to prepare for the 
effects of climate change on fisheries by incorporating 
climate variables (such as ocean temperature trends) 
into harvest policies (Hill et ah, 2017) and by conducting 
climate vulnerability assessments (Morrison et ah, 2015; 
Hare et ah, 2016). Climate vulnerability assessments 
evaluate which species will be at the greatest risk (or 
most vulnerable) to the effects of climate change, and 
these assessments consequently are intended to inform 
and prioritize research and management actions. Spawn¬ 
ing frequency is 1 of 12 biological attributes used in the 
current criteria for evaluating vulnerability to climate 
change, with species that spawn in one single event per 
year considered to be more vulnerable than those that 
spawn several times per year (Hare et al., 2016). There¬ 
fore, multiple-brooding species may have an evolution¬ 
ary advantage relative to single-brooding species in the 
face of either warming or more variable ocean conditions, 
necessitating an accurate assessment of the reproductive 
ecology of all Sebastes species in evaluating the vulnera¬ 
bility of different species to climate change. 
Accounting for multiple brooding as a maternal effect 
in population models can substantially affect stock 
assessments, particularly if the likelihood of producing 
multiple broods (increasing overall reproductive output) 
is size dependent, as demonstrated for chilipepper (Lefe- 
bvre et ah, 2018). Currently, stock assessment models of 
multiple-brooding species do not account for such fac¬ 
tors in estimating reproductive output and stock status, 
potentially biasing assessment results if the effect is to 
nominally increase reproductive output of larger, older 
individuals. Taking multiple brooding into account in 
general, as well as in the context of climate change, will 
improve stock assessments and help managers develop 
appropriate management responses to climate variabil¬ 
ity and change now and into the future (Barneche et ah, 
2018). Our results provide robust predictions of this like¬ 
lihood and provide context to the observed distribution 
of multiple-brooding species relative to single-brooding 
shelf rockfish species. However, our insights are not 
strictly mechanistic and are only the first step toward 
understanding the mechanisms and evolution of multi¬ 
ple brooding in rockfish species of the continental shelf. 
Future research should explicitly explore the mecha¬ 
nisms that drive multiple versus single brooding to aid 
in dosing the knowledge gap for multiple-brooding rock¬ 
fish species. 
Acknowledgments 
We v/ould like to thank S. Sogard, S. Beyer, L. Lefebvre, 
N. Kashef, and D. Stafford at the Fisheries Ecology Divi¬ 
sion of the Southwest Fisheries Science Center and the 
University of California’s Cooperative Institute for Marine 
Ecosystems and Climate because their combined efforts 
were critical to inform and guide this effort. Additionally, 
we are deeply appreciative of the detailed comments pro¬ 
vided by 3 anonymous reviewers and feel that incorpo¬ 
rating their recommendations has greatly improved this 
manuscript. 
Literature cited 
Barneche, D. K., D. R. Robertson, C. R. White, and D. J. Marshall. 
2018. Fish reproductive-energy output increases 
disproportionately with body size. Science 360:642-645. 
Berkeley, S. A., M. A. Hixon, R. J. Larson, and M. S. Love. 
2004. Fisheries sustainability via protection of age structure 
and spatial distribution of fish populations. Fisheries 
29:23-32. 
Beyer, S. G., S. M. Sogard, C. J. Harvey, and J. C. Field. 
2015. Variability in rockfish (Sebastes spp.) fecundity: species 
contrasts, maternal size effects, and spatial differences. 
Environ. Biol. Fish. 98:81-100. 
Boehlert, G. W., and M. M. Yoklavich. 
1983. Effects of temperature, ration, and fish size on growth 
of juvenile black rockfish, Sebastes melanops. Environ. 
Biol. Fish. 8:17-28. 
Boehlert, G. W., M. Kusakari, and J. Yamada. 
1991. Oxygen consumption of gestating female Sebastes 
schlegeli: estimating the reproductive costs of livebearing. 
Environ. Biol. Fish. 30:81-90. 
Burnham, K. P., and D. R. Anderson. 
2002. Model selection and multimodel inference: a practical 
information theoretic approach, 2 nd ed., 488 p. Springer, 
New York. 
Checkley, D. M., and J. A. Barth. 
2009. Patterns and processes in the California Current Sys¬ 
tem. Prog. Oceanogr. 83:49-64. 
Conrath, C. L. 
2017. Maturity, spawning omission, and reproductive com¬ 
plexity of deepwater rockfish. Trans. Am. Fish. Soc. 
146:495-507. 
Cope, J. M., J. BeVore, E. J. Dick, K. Ames, J. Budrick, D. L. Erick¬ 
son, J. Grebel, G. Hanshew, R. Jones, L. Mattes, et al. 
2011. An approach to defining stock complexes for US West 
Coast groundfishes using vulnerabilities and ecological 
distributions. N. Am. J. Fish. Manage. 31:589-604. 
Craney, T. A., and J. G. Surles. 
2002. Model-dependent variance inflation factor cutoff val¬ 
ues. Qual. Eng. 14:391-403. 
Cushing, D. H. 
1990. Plankton production and year-class strength in fish 
populations: an update of the match/mismatch hypothesis. 
Adv. Mar. Biol. 26:249-293. 
Dick, E. J., and A. D. MacCall. 
2010. Estimates of sustainable yield for 50 data-poor 
stocks in the Pacific Coast groundfish fishery manage¬ 
ment plan. NOAA Tech. Memo. NMFS-SWFSC-46G, 
208 p. 
