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Early Life History and Fisheries Oceanography: New Questions in a Changing World

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Early Life History and Fisheries Oceanography: New Questions in a Changing World Early Life History and Fisheries Oceanography: New Questions in a Changing World Llopiz, J. K., Cowen, R. K., Hauff, M. J., Ji, R., Munday, P. L., Muhling, B. A., ... & Sponaugle, S. (2014). Early Life History and Fisheries Oceanography: New Questions in a Changing World. Oceanography, 27(4), 26-41. doi:10.5670/oceanog.2014.84 10.5670/oceanog.2014.84 Oceanography Society Version of Record http://cdss.library.oregonstate.edu/sa-termsofuse SPECIAL ISSUE ON FISHERIES OCEANOGRAPHY Early Life History and Fisheries Oceanography New Questions in a Changing World By Joel K. Llopiz, Robert K. Cowen, Martha J. Hauff, Rubao Ji, Philip L. Munday, Barbara A. Muhling, Myron A. Peck, David E. Richardson, Susan Sogard, and Su Sponaugle CAPTION. Larval stage of selected ABSTRACT. In the past 100 years since the birth of fisheries oceanography, research fish (from left to right): armored on the early life history of fishes, particularly the larval stage, has been extensive, and searobin, largetooth flounder, byth- itid brotula, and little tunny. Photos much progress has been made in identifying the mechanisms by which factors such by Cedric Guigand as feeding success, predation, or dispersal can influence larval survival. However, in recent years, the study of fish early life history has undergone a major and, arguably, necessary shift, resulting in a growing body of research aimed at understanding the consequences of climate change and other anthropogenically induced stressors. Here, we review these efforts, focusing on the ways in which fish early life stages are directly and indirectly affected by increasing temperature; increasing CO2 concentrations, and ocean acidification; spatial, temporal, and magnitude changes in secondary production and spawning; and the synergistic effects of fishing and climate change. We highlight how these and other factors affect not only larval survivorship, but also the dispersal of planktonic eggs and larvae, and thus the connectivity and replenishment of fish subpopulations. While much of this work is in its infancy and many consequences are speculative or entirely unknown, new modeling approaches are proving to be insightful by predicting how early life stage survival may change in the future and how such changes will impact economically and ecologically important fish populations. 26 Oceanography | Vol.27, No.4 BACKGROUND aberrant drift hypothesis (1914, 1926), a governing early life survival (Houde, Understanding the early life stages of groundbreaking theory that highlighted 2008; Hare, 2014), yielded important dis- fishes, and the processes influencing their the importance of variability in the trans- coveries in fish biology (e.g., daily incre- survival, is at the heart of fisheries ocean- port and dispersal of eggs and larvae by ments in fish otoliths, or “ear stones”), ography, along with the field’s historical ocean currents in shaping recruitment drove advances in technology and roots. In the early twentieth century, the patterns. Thus, Hjort touched upon two techniques (e.g., coupled biological- founding father of fisheries oceanogra- of the three major challenges faced by fish physical models), and underpinned phy, Johan Hjort of Norway, led interna- larvae in the plankton: finding food and large-scale interdisciplinary programs tional efforts to understand why import- reaching suitable juvenile habitat. Real (e.g., California Cooperative Oceanic ant fish stocks yield such remarkably progress in understanding the third crit- Fisheries Investigations [CalCOFI], large variations in catch. While most ical hurdle—avoiding predators—did not Fisheries Oceanography Coordinated researchers were focused on the move- come for another 75 years (Bailey and Investigations [FOCI], Global Ocean ment of adults in and out of fishing Houde, 1989). Ecosystem Dynamics [GLOBEC]) that grounds as a cause of recruitment fluc- With Hjort laying the foundation for all contributed greatly to our ability to ask tuations (Sinclair, 1997), Norway’s enor- early life history research in the twentieth and answer interesting questions about mous 1904 year class of herring inspired century, the race to solve the “recruitment survivorship in the early life stages of Hjort to think instead about the recruit- problem” picked up considerable steam fishes. Some of the most important find- ment process—that is, the survival of in the last quarter of the century (Houde, ings have elucidated the ways in which the early life stages. This year, we cel- 2008), especially after David Cushing larval growth rates interact with overall ebrate the centennial of the publica- extended Hjort’s critical period hypoth- mortality rates (growth-mortality hypoth- tion of Hjort’s (1914) seminal work enti- esis to the entire larval period with his esis), largely due to reduced susceptibil- tled “Fluctuations in the great fisheries match-mismatch hypothesis (Cushing, ity to predation for a larva growing more of northern Europe, viewed in the light 1974, 1990). This hypothesis emphasized quickly through its vulnerable planktonic of biological research” (see Browman, the degree to which fish larvae and their period (Cushing, 1975; Houde, 1987, 2014). This is where Hjort first laid out planktonic prey overlap temporally (as 1997; Anderson, 1988). his most enduring contribution to the determined by the timing of productivity Building upon thorough reviews of the field of fisheries oceanography, the criti- blooms and fish spawning) as a key factor history and hypotheses of early life his- cal period hypothesis, which posited that in determining larval survival and recruit- tory and fisheries oceanography (Houde, recruitment levels could be greatly influ- ment. The match-mismatch hypothe- 2008; Peck et al., 2012; Hare, 2014), we enced by the degree to which sufficient sis has had a remarkable impact on the focus here on a pressing contemporary types and abundances of planktonic prey direction of larval fish research, and it question in early life history research: the were available to millimeter-scale fish lar- laid the cornerstones for modern fisheries implications of global change and other vae as they first transitioned to exogenous oceanography. A wave of early life history anthropogenic impacts on fish early life feeding. In addition to the critical period research led to the development of sev- survival. As in Hjort’s time, planktonic hypothesis, Hjort also put forth his eral other hypotheses on the mechanisms fish larvae today still have to find food, Oceanography | December 2014 27 avoid predation, and end up in suit- population connectivity? Perhaps most of change will likely vary regionally; tem- able juvenile habitat, yet a whole new importantly, how will all of these ongoing peratures near the equator, for example, suite of issues has emerged (Figure 1), and impending global changes interact to should increase more rapidly than those and we must now consider how anthro- affect the early life stages of fishes? Our at higher latitudes (Collins et al., 2013; pogenically induced stressors affect nat- goal here is to review some of the prog- Kirtman et al., 2013). Tropical larvae ural processes. Specifically, how will ress made in addressing these important therefore may be subject to more dramatic climate-driven warming and ocean acid- questions, and to call attention to critical temperature shifts, and they may also be ification directly and indirectly affect the knowledge gaps yet to be filled. particularly sensitive to increases, given life histories of planktonic larvae in their that they are adapted to relatively narrow journey from spawning sites to juvenile TEMPERATURE EFFECTS and stable thermal regimes (Tewksbury nursery areas? How will ocean produc- The increase in ocean temperatures et al., 2008). Polar populations, too, are tivity change in magnitude as well as in throughout the next century will have generally stenothermal, and they might time and space, and how will dynamics major implications for the structure and be similarly sensitive to warming. Because of productivity, zooplankton composi- functioning of marine ecosystems. With mid-latitude fish populations are already tion and abundance, and spawning out- average sea surface temperatures poten- subject to a wider range of tempera- put (in light of fishing pressure) influence tially rising 3°C or more by the year 2100 tures, partly due to interdecadal fluctu- early life survival and recruitment? How (Collins et al., 2013), effects on fish early ations in circulation and surface heating will ocean currents be impacted by cli- life stages inhabiting those surface waters in those regions (Wang et al., 2010), they mate change to affect larval transport and could be profound. Rates and magnitudes could be more tolerant of temperature variation than their tropi- cal and polar counterparts. " "!! *!" ! Nevertheless, fish early life " !! ! ' " *!! "& ".!*!" '!# #. 0& "!, " " # stages across latitudes will "! * 2 !" # #1 !" ! be affected by changing temperatures, whether by !*!" " " acute or chronic (i.e., sub- "! & & !" &# #.! lethal) mechanisms. !" &# In both embryos (fer- tilized eggs and yolk-sac ' (" /!" /! larvae) and exogenously !& '' !!! & #"* " feeding larvae, lethal tem- & "* peratures exist at which &' survival is simply not pos- (# " " # sible. Surprisingly, pro- 0 '- )""1 tocols used to determine the critical limits that !"/!%" define the thermal niches !& '' of juvenile and adult fishes (Pörtner and Peck, 2010) have not
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