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Ocean Transport Paths for the Early Life History Stages of Offshore-Spawning flatfishes: a Case Study in the Gulf of Alaska

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Ocean Transport Paths for the Early Life History Stages of Offshore-Spawning flatfishes: a Case Study in the Gulf of Alaska F I S H and F I S H E R I E S , 2008, 9, 44–66 Ocean transport paths for the early life history stages of offshore-spawning flatfishes: a case study in the Gulf of Alaska Kevin M Bailey1, Alisa A Abookire2 & Janet T Duffy-Anderson1 1Alaska Fisheries Science Center, 7600 Sand Point Way NE, Seattle, WA 98115, USA; 2Alaska Fisheries Science Center, Kodiak Facility, 301 Research Court, Kodiak AK 99615 USA Abstract Correspondence: Offshore- and deepwater-spawning flatfish species face the problem of transport of Kevin M Bailey, Alaska Fisheries their planktonic stages to shallow juvenile nursery grounds that are often far Science Center, 7600 shoreward in bays or estuaries. We compare life history attributes of four offshore- Sand Point Way NE, spawning flatfish species in the Gulf of Alaska: Pacific halibut (Hippoglossus stenolepis), Seattle, WA 98115, arrowtooth flounder (Atheresthes stomias), rex sole (Glyptocephalus zachirus) and Dover USA sole (Microstomus pacificus) to examine how their larvae get from a spawning location Tel.: +1 206 526 4243 at the edge or beyond the continental shelf to specific inshore nursery zones. We Fax: +1 206 526 utilize historical records of survey catches of different life stages to characterize the 6723 stage-specific changes in distribution of spawning, planktonic stages and juvenile E-mail: kevin. nursery areas. We infer transport mechanisms based on the shifts in distribution of [email protected] the life stages and in comparison with local physical oceanography. This comparison Received 24 May 2007 provides insight into the different mechanisms marine species may use to solve the Accepted 6 Nov 2007 common ‘problem’ of planktonic drift and juvenile settlement. Keywords Fish larvae, juvenile nursery, larval distribution, larval transport, ocean currents, spawning strategy Introduction 45 Data 46 The Gulf of Alaska setting 47 Shelf features 47 Circulation 48 Life history attributes of offshore-spawning flatfishes 49 Spawning 49 Patterns in egg and larval distributions 55 Juvenile nurseries 56 Discussion of transport mechanisms 58 Spawning location and season 58 Transport onto the shelf 59 Transport along the shelf 60 Larval behaviours 62 Settlement and recruitment issues 62 Future studies 63 Acknowledgements 64 References 64 Ó 2008 The Authors 44 Journal compilation Ó 2008 Blackwell Publishing Ltd Transport of offshore-spawning flatfish larvae K M Bailey et al. Some flatfish species are known to utilize behavio- Introduction ural adaptations to modify their transport as Organisms with planktonic stages release their plankton, or passively drifting particles. Among offspring into an environment where they not the best known and most sophisticated of these only face the immediate issue of survival, but they behaviours is selective tidal stream transport must also ‘anticipate’ survival at later stages, (STST), by which larvae alter their position in the which often have different and sometimes highly water column to take advantage of vertically specific requirements. If the release area of eggs is stratified tidal currents (Creutzberg et al. 1978; within suitable habitat territory for juveniles, then Rijnsdorp et al. 1985; Tanaka et al. 1989; Jager it is advantageous to be retained there, and if 1999; Forward and Tankersley 2001). Using STST, distant from suitable habitat, then eggs and larvae larvae that are spawned near to suitable nursery need to be delivered there. This problem is acute habitat can maintain themselves there by vertically for marine fishes, where planktonic stages have migrating into counter-currents, and larvae that high mortality rates (commonly measured at need directed transport can migrate into tidal 5–40% per day) and whose larval stages may currents flowing in the preferred direction. This metamorphose into quite different juvenile forms. behaviour is sophisticated in the sense that larvae Flatfishes share the same planktonic problems need to have an innate sense of which direction to with many other species, but they represent an move in after receiving specific cues that reveal the extreme in the process of metamorphosis between orientation of water movement. The complex larvae and juveniles. Juvenile flatfishes exhibit processes by which a larva recognizes the cues markedly different morphological forms and and clues which control its vertical migratory and behaviours and have different environmental settlement behaviour are not well understood requirements than their pelagic precursors. (Metcalfe et al. 2006). Most flatfish species have their juvenile nurseries Besides STST, there are other ways flatfishes have inshore of the spawning grounds (Minami and developed their strategies to maximize delivery to Tanaka 1992), and often these nurseries have nearshore nursery areas (Bailey et al. 2005). Like specific qualities characterized by depth, tempera- many other animals with planktonic stages, they ture, salinity, sediment and predator/prey consider- locate their offspring where and when food abun- ations, which define it as suitable habitat (Gibson dance is high and predator abundance is low. They 1994). For species that spawn offshore in deep may broadcast their offspring where the probability water, finding suitable nursery habitat poses a of their transport in an advantageous direction is particularly difficult and interesting problem for optimized (Gibson 1999; Rooper et al. 2006). Thus, larvae: how to hit a target whose direction is often there may be local adaptation to regional currents, orthogonal to prevailing currents? Is hitching a or adaptations to take advantage of predictable planktonic ride on the right mass of water a random seasonal events (Boehlert and Mundy 1987). For process in which only a few lucky passengers arrive example, some flatfish species have extremely long at their destination? Or can discriminating behav- planktonic periods of up to 2 years to ride seasonal iour among adults or larvae influence the outcome downwelling currents to their inshore juvenile of the transport process? Success in this process of nurseries (Pearcy et al. 1977; Markle et al. 1992). arriving at suitable inshore nurseries could play a In such circumstances, larvae may settle faculta- role in recruitment variability. The concept that tively when suitable habitat is encountered (Tanaka transport plays an important role in population et al. 1989; Abookire and Bailey 2007). However, regulation is supported by numerous correlative when larvae are spawned near or within suitable studies that associate planktonic transport to juvenile habitat or there are specific transport variability in year-class strength (Parker 1989; mechanisms to deliver them there, planktonic life Wilderbuer et al. 2002). Detailed field research may be as short as 10 days (Minami and Tanaka (Van der Veer and Witte 1999), modelling studies 1992). Even within a species, there may be local (Van der Veer et al. 1998) and comparative adaptations reflecting quite different strategies. For approaches (Van der Veer et al. 1990; Rijnsdorp example, plaice (Pleuronectes platessa, Pleuronecti- et al. 1992) also support the importance of larval dae) larvae in the North Sea utilize bottom currents flatfish transport in the recruitment process. for transport to near-nursery areas and then use Ó 2008 The Authors Journal compilation Ó 2008 Blackwell Publishing Ltd, F I S H and F I S H E R I E S , 9, 44–66 45 Transport of offshore-spawning flatfish larvae K M Bailey et al. STST to enter nurseries (Van der Veer et al. 1998); Table 1 Total length (cm) and weight values used to in the Kattegat/Belt Sea, spawning occurs offshore estimate mature flatfishes and the range of their spawning and larvae ride wind-driven currents inshore season. Based on these parameters, adult flatfish data (Nielsen et al. 1998); and in the Irish Sea, plaice were isolated from the NMFS Bottom Trawl Survey spawn nearshore and larvae utilize retention fea- database (1976–2006). tures to maintain themselves there (Nash and Geffen 1999). Total length Weight Spawning In this paper, we compare life history attributes Species Sex (cm) (kg) season of four offshore-spawning flatfish species in the Gulf of Alaska (GOA): Pacific halibut (Hippoglossus Pacific Males and ‡125(1) 19.5(2) Dec–Mar(3) stenolepis, Pleuronectidae), arrowtooth flounder halibut females (4) (5) (6) (Atheresthes stomias, Pleuronectidae), rex sole Arrowtooth Females ‡53 1.4 Jan–Mar flounder (Glyptocephalus zachirus, Pleuronectidae) and Dover (7) (7) (7) Rex sole Females >34 0.37 Oct–May sole (Microstomus pacificus, Pleuronectidae), to Dover Females >43(8) 0.61(8) Feb–May(8) examine how they solve a common ontogenetic sole problem of getting from a spawning location at the edge or beyond the continental shelf to specific (1)Clark et al. (1999). (2) inshore nursery zones. We utilize time series of Clark (1992). (3)St. Pierre (1984). survey catches of different life stages to characterize (4)Zimmermann (1997). the horizontal distribution of spawning adults, (5)Turnock et al. (2005). planktonic egg and larval stages, and settled juve- (6)Blood et al. (2007). niles. Historical data are also used to characterize (7)Abookire (2006). (8) vertical distributions of eggs and larvae. We then Abookire and Macewicz (2003). infer species-specific transport mechanisms based on the shift in spatial distribution between successive life stages. These inferences are discussed in relation were measured for length in each haul and then to cross-shelf circulation. New detailed information assigned a spawning season according to parame- on circulation is
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