Larvae. Juvenile. Adult. Bipartite life cycle of benthic marine fishes with pelagic larvae. Pelagic Environment. settlement.

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13 - Marine Ecology

Bipartite life cycle of benthic marine fishes with pelagic larvae Larvae survive, grow, develop, disperse reproduce Pelagic Environment Benthic Environment settlement Adult Juvenile survive, grow, mature

Bipartite life cycle of benthic marine fishes with pelagic larvae

Closed Populations Open Populations Production Supply Production Supply Little or no exchange among populations Significant exchange among populations Production Supply Supply Production Decouples local production and supply

Pelagic duration - a proxy for dispersal potential Dispersal Distance (km) 10000 1000 100 10 1 0.1 0.01 invertebrates fish Predicted by passive dispersal 0.001 0.0001 0.01 0.1 1 10 100 1000 10000 Propagule Duration (hr) Shanks et al. 2003 Ecological Applications

Larval duration of 24 coastal fish species from western North America SPECIES Larval duration midpoint (range) Shanks et al. 2003 Ecological Applications MEAN = 94 days

Indirect Estimates of Larval Fish Dispersal Distances Pelagic-duration Isolation by distance 10000 Dispersal Distance (km) 10 1000 100 10 1 0.1 0.01 0.001 = 30 days 0.0001 0.01 0.1 1 10 100 1000 10000 Fish 5 0 1 m 10 m 100 m 1 km 10 km 100 km 1000 km Estimated dispersal scale Kinlan & Gaines 2003 Ecology Palumbi 2003 Ecological Applications Propagule Duration (hr) Shanks et al. 2003 Ecological Applications 10 s to 100 s of km

The paradigm shift Generally assumed that larvae disperse away from natal population

Anemonefish recruitment study - Moorea, French Polynesia

1. Surveys Manta Tows Scuba Diving

2. Characteriza;on

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A material

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4. Return the fish

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Location of all Amphiprion chrysopterus collected around Moorea (French Polynesia) 1-10 11-20 >21 individuals

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Moorea, PGEM

Recruitment studies demonstrate spillover effect Harrison et al. 2012

Potential consequences of larval dispersal: local recruitment (replenishment) not necessarily tied to local production recruitment critical to replenishment of local populations local replenishment reliant on recruitment of larvae produced elsewhere open spatial structure of local and regional populations openness decreases as likelihood that larvae return to adult population increases

Recruitment is important and fascinating Temporal scale 100 yrs 1 decade 1 year 1 month 1 week 1 day 1 hour 1 min Langmuir cells Seagrass beds Turbulent eddies Surface waves Plankton migration Kelp forests Internal waves Seasonal current shifts Small-scale fronts, plumes, runoff Seasonal upwelling Mesoscale eddies Surface tides Internal tides Coastally trapped waves PDO ENSO Coastal filaments, Upwelling / relaxation 1 cm 1 dm 1 m 10 m 100 m 1 km 10 km 100 km 1000 km 10000 km After T. Dickey, unpublished Linear spatial scales but its very complex!

Recruitment is important and fascinating Temporal scale 100 yrs 1 decade 1 year 1 month 1 week 1 day 1 hour 1 min Larval production Langmuir cells Seagrass beds Turbulent eddies Surface waves Plankton migration Kelp forests Internal waves Seasonal current shifts Small-scale fronts, plumes, runoff Seasonal upwelling Mesoscale eddies Surface tides Internal tides Coastally trapped waves PDO ENSO Coastal filaments, Upwelling / relaxation 1 cm 1 dm 1 m 10 m 100 m 1 km 10 km 100 km 1000 km 10000 km After T. Dickey, unpublished Linear spatial scales but its very complex!

Recruitment is important and fascinating Temporal scale 100 yrs 1 decade 1 year 1 month 1 week 1 day 1 hour 1 min Larval dispersal, mortality Langmuir cells Seagrass beds Turbulent eddies Surface waves Plankton migration Kelp forests Internal waves Seasonal current shifts Small-scale fronts, plumes, runoff Seasonal upwelling Mesoscale eddies Surface tides Internal tides Coastally trapped waves PDO ENSO Coastal filaments, Upwelling / relaxation 1 cm 1 dm 1 m 10 m 100 m 1 km 10 km 100 km 1000 km 10000 km After T. Dickey, unpublished Linear spatial scales but its very complex!

Recruitment is important and fascinating Temporal scale 100 yrs 1 decade 1 year 1 month 1 week 1 day 1 hour 1 min Larval settlement Langmuir cells Seagrass beds Turbulent eddies Surface waves Plankton migration Kelp forests Internal waves Seasonal current shifts Small-scale fronts, plumes, runoff Seasonal upwelling Mesoscale eddies Surface tides Internal tides Coastally trapped waves PDO ENSO Coastal filaments, Upwelling / relaxation 1 cm 1 dm 1 m 10 m 100 m 1 km 10 km 100 km 1000 km 10000 km After T. Dickey, unpublished Linear spatial scales but its very complex!

Sources of spatial and temporal variation in recruitment Larval dispersal Settlement Larval production Post-settlement

Sources of spatial and temporal variation in recruitment Larval production: - adult fecundity (eggs per female) size structure / sex ratio adult condition - adult abundance

Offspring production: climatic variability Holbrook et al. 1997 Ecological Applications Ocean climate change Surfperch production Population responses (4 species) Benthic productivity Perch recruitment (Love et al. 1998 Fishery Bulletin) Power plant studies: (Brooks et al. 2002 Mar. Freshwater Res.) - Bight-wide patterns of juvenile impingement - declines in recruitment for many spp. (1980-1991) - attributed to reduced production (but maybe larval survival) - reflecting large-scale decline in productivity

Sources of spatial and temporal variation in recruitment Larval dispersal (direction, distance, delivery): - larval duration - larval behavior - oceanographic features - interaction among these Larval production: - timing of reproduction - location of reproduction

Physical processes and larval behavior (1) Larval cues: (light, pressure, temperature, structure) e.g., Norris 1963, Ecological Monographs - Opaleye (Girella nigricans) - recruitment related to tide pool temp. - lab experiments: thermal preference - coast-wide patterns of recruitment - hypothesized mechanisms of larval delivery: - internal waves - thermal / structural cues - upwelling ONSHORE TRANSPORT Shanks 1983 Mar. Ecol. Prg. Ser.

Olive rockfish Structure - schooling Long larval duration (3-4 months) Yellowtail rockfish Black rockfish Kelp rockfish Benthic - solitary Short larval duration (1-2 months) Gopher rockfish Black-&-yellow rockfish

Physical processes and larval distribution (2) shifts in vertical distribution with ontogeny -- upwelling e.g., Larson et al. 1994, Lenarz et al. 1995, CalCOFI Rpt.s - vertical distribution of early and late larval rockfishes proportion 0 0.2 0.4 0.6 structure - schooling spp. benthic - solitary spp. depth (m) 13 37 depth late larvae pelagic juveniles early larvae kelp bed onshore 87-117 offshore

1.0 Lenarz et al. 1995 CalCOFI Mid-water complex Long larval duration (3-4 months) Olive, Yellowtail and Black rockfish Proportion 0.5 0 0.5 1.0 1986 1992 Kelp, Black-&-yellow, and Gopher rockfish Benthic complex Short larval duration (1-2 months) Relative Abundance 100 75 50 25 0 25 50 75 100 El Nino La Nina La Nada (1998) (1999) (2000)

(1998) (1999) (2000) Kelp rockfish Gopher rockfish Relaxation Black-&-yellow rockfish Benthic complex Short larval duration (1-2 months) Fish per 240 m 3 20 Kelp, Black-&-yellow, and Gopher rockfish 16 12 8 4 0 El Niño La Niña Normal

(1998) (1999) (2000) Mid-water complex Long larval duration (3-4 months) Upwelling Olive rockfish Yellowtail rockfish Fish per 240 m 3 70 50 30 10 Olive, Yellowtail and Black rockfish Black rockfish El Niño La Niña Normal

Sources of spatial and temporal variation in recruitment Settlement: - larval cues - habitat structure - priority effects conspecific cues predation competition

Settlement (post-settlement): habitat structure Kelp Bass Recruit Density 3 (Number / 60 m ) 2 (Number / 10 m ) 30 10 5 3 1 5 4 3 2 1 0 0 100 200 300 400 B B A 0 40 80 120 160 Blade Biomass Per Reef Area 2 (grams / 10 m ) Carr Ecology 1994 1,200 B B 800 A 400 0 0 40 80 120 2 Macrocystis Density (stipes / 30 m )

Sources of spatial and temporal variation in recruitment Settlement: - larval cues - habitat structure - priority effects conspecific cues predation competition Not a lot of information!

Sources of spatial and temporal variation in recruitment Early post-settlement: - survival - growth - movement predation competition

Early post-settlement: predation 1.0 per-capita mortality kelp perch 1.0 0.8 0.6 0.4 0.2 0.0 Anderson 2002 Ecology black eyed goby 0 5 10 15 Initial density predators present predators absent 1.0 Steele 1997 Oecologia kelp rockfish Johnson unpublished 0 20 40 60 0 20 40 60 80 100

Early post-settlement: competition Conspecific and interspecific resident effects e.g., Steele 1997a, Ecology - black-eyed and blue-banded gobies in So. California - manipulated presence of adults of both - settlement of black-eyed (-) influenced in presence of adult conspecifics - settlement of black-eyed not influenced by presence of adult blue-banned - settlement of blue-banded (+) influenced in presence of adult conspecifics - settlement of blue-banded not influenced in presence of adult black-eyed

Sources of spatial and temporal variation in recruitment - survival - growth - movement competition predation Late post-settlement: adult and juvenile interactions

Role of fishes in kelp forest communities (1) Keystone predator -- trophic cascades e.g., Cowen 1983, Oecologia - sheephead (Semicossyphus pulcher) in So. Calif. - manipulated local presence of sheephead and observed red sea urchin behavior - urchins more exposed and mobile in absence of sheephead

Cascading Effects of Predator Removal Southern California Central California lobster sheephead sea otters sea urchins rockfishes barrens kelp forest

Another example from down under: New Zealand snapper Sea urchin banded wrasse barrens Ecklonia

Role of fishes in kelp forest communities (2) Enhanced nutrient availability and productivity e.g., Bray et al. 1981, Science - planktivorous blacksmith (Chromis punctipinnis) hole up in crevices at night - monitored nutrient availability and macroalgal production in crevices with and without blacksmith - greater nutrient availability and macroalgal production in crevices with blacksmith - example of planktivorous fishes directing planktonic production to benthos

Role of fishes in kelp forest communities (3) Planktivorous fishes reducing larval supply e.g., Gaines and Roughgarden 1987, Science - reduced recruitment of intertidal barnacles in years with thick Macrocystis forest at Hopkins - high recruitment of planktivorous juvenile rockfishes in years with thick Macrocystis forest - could be hydrodynamic influence of kelp or predation by planktivorous juvenile rockfishes - used barnacle molts to decouple potential causes - barnacle molts not reduced as they passed through forest, concluded reduction due to planktivory

Percentages Average = ~ 76 High Score = 94 Low Score = 53 Midterm Results Points Total possible points 83 35 pts for scantron + 48 points for written