“Bipartite life history of marine species and “openness” of populations 1 1

“Closed” Populations “Open” Populations Production Supply Production Supply Little or no exchange among populations Significant exchange among populations Production Supply Supply Production

So, how open are marine populations? Depends on how far their larvae travel, especially, relative to how far adults travel Lines of evidence of population openness: Evidence for long distance dispersal of larvae: a. prolonged pelagic larval duration (PLD) b. Low spatial genetic structure Evidence of shorter distance movement of adults a. Sessile organisms (algae and invertebrates) b. Limited distance of adult movement Lutjanus kasmira -- spread subsequent to introduction (duration = 25-47 d, mean realized dispersal distance= 33-130 km) . Oligocottus maculosus -- observations of dispersing larvae and their distribution (duration = 30 d, mean realized dispersal distance= < 1 km)

Pelagic duration: a proxy for dispersal potential 0.0001 0.001 0.01 0.1 1 10 100 1000 10000 invertebrates fish Predicted by passive dispersal r2= 0.61, P= 0.001 Dispersal Distance (km) Lutjanus kasmira -- spread subsequent to introduction (duration = 25-47 d, mean realized dispersal distance= 33-130 km) . Oligocottus maculosus -- observations of dispersing larvae and their distribution (duration = 30 d, mean realized dispersal distance= < 1 km) Propagule Duration (hr) Shanks et al. 2003 Ecological Applications 4

Time in the larval stage (fish) Western North American Coastal Fish Time in Larval Stage midpoint (range) AVERAGE = 94 days Shanks et al. 2003

Pacific Ocean Reef Fishes Larval Duration Estimates Tropical (n= 298 spp) Temperate (n= 60 spp) 70 60 50 Percent of Species 40 30 20 10 < 1 1 - 7 8 - 30 31 - 90 > 90 Larval Duration (days) Carr and Syms 2006 in: Allen et al. The Ecology of Marine Fishes

Geographic distance = genetic difference “Isolation by Distance” high Genetic difference Slope measures average dispersal low short long Geographic distance (kilometers)

Geographic distance = genetic difference “Isolation by Distance” populations nearby one another populations further apart high Genetic difference Slope measures average dispersal low short long Geographic distance (kilometers)

Genetic difference Copper rockfish CA snails High dispersal Rosethorn rockfish Intermediate dispersal CA corals Low dispersal high Genetic difference low 200 400 600 800 Geographic distance (kilometers)

Larval dispersal Based on genetic difference Inverts: <1-100 kilometers Number of species Fish: 20-200 kilometers Palumbi 2003 Kinlan and Gaines 2003

Different estimates, similar results Time in larval stage Genetic distance 10000 1000 100 = 30 days 10 Dispersal distance 1 0.1 0.01 0.001 0.0001 0.01 0.1 1 10 100 1000 10000 Time as larvae (hr) Dispersal distance of invert larvae = 1 - 100 km Dispersal distance of fish larvae = 20 - 200 km

Limited Adult Movement Home ranges of 25 west coast rocky habitat fish species 12 Conclusion: 76% of these species moved less than 0.5 km 8 number of species 4 But at a minimum… reserves of appropriate size must consider how far adult fish move and here is an example of how we draw from the literature on fish movement to provide guidance on MPA size. This graph shows the number of west coast rocky reef fishes whose home range, the distance they move in their lifetime, is less than half a kilometer, up to 30 kilomters, note the scale is logarithmic, doubling from one interval to the next! Roughly ¾ of these species have home ranges less than ½ a kilometer, far fewer move on the order of ten’s of kilometers. So reserves on the order of 5-10 km can encompass the home range and portect many species reef associated species of fishes. 0.5 1 5 10 20 30 Median max. distance (km) Freiwald, J. 2012. Can. J. of Fish. & Aquat. Sci. 12 12

Adult Home Range Size Varies Among Species of Fishes as does the “openness” of their populations! 0 – 1 km 1 – 10 km 10 – 100 km 100 – 1000 km > 1000 km Many rockfish Some rockfish Some rockfish Few rockfish Some schooling fish Some schooling fish Tunas Some surfperch Other reef fish Other reef fish Salmon Because species vary greatly in how far they move, changes in reserve size will determine how many species are protected by reserves of a given size. For example, reserves on the order of 10 km on a side will protect the many species whose adults move limited distances. Expanding that area will protect more. Some species move so much that only very large protected areas would encompass them but they can benefit otherwise… by protecting habitats where they feed and breed. (refer to Lindholm on giant seabass) Many sharks Some flatfish More flatfish Some surfperch 13 13

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

but studying it really bites! Recruitment is important and fascinating… 100 yrs Seasonal current shifts PDO 1 decade Seagrass beds Kelp forests ENSO 1 year Seasonal upwelling Mesoscale eddies 1 month Temporal scale Coastally trapped waves 1 week Small-scale fonts, plumes, runoff 1 day Plankton migration Surface tides 1 hour Langmuir cells Internal waves Coastal filaments, Upwelling / relaxation Turbulent eddies 1 min Internal tides Surface waves 1 cm 1 dm 1 m 10 m 100 m 1 km 10 km 100 km 1000 km 10000 km Linear spatial scales but studying it really bites! Carr and Syms 2006, CA Fishes book

V) Factors affecting recruitment assume complex life history, and focus on scenarios where settlement is potentially limiting to the level where it affects adult populations and communities A) Production and availability of propagules (spores, eggs, larvae) Determinants: 1) Reproduction by adults - very little work has been done on this - why?? i) presumed decoupling ii) problem of tracking or identifying source of highly dispersive offspring iii) poor stock-recruitment relationships

V) Factors affecting recruitment 1) Reproduction by adults - very little work has been done on this. Poor stock-recruitment relationships from fisheries statistics:

V) Factors affecting recruitment A) Presence and abundance of propagules arriving at a site 2) Reproduction by adults - very little work has been done on this - problem of open populations and following offspring, but it should look like this: importance of local production to local recruitment (% settlers produced locally) dispersal distance

V) Factors affecting recruitment 3) Determinants of larval production (sources of variation): i) population size ii) size /age distribution iii) density (mate availability, Allee effect) iv) sex ratio (mate availability) iv) condition (food / energy availability - benthic, oceanographic) v) resource availability (spawning sites) vi) spawning seasonality (influences dispersal patterns) vii) spawning location (influences dispersal patterns)

Offspring production: climatic variability Ocean climate change Power plant impingement of fish larvae: 1) Love et al. 1998 Fishery Bulletin included commercial species 2) Brooks et al. 2002 Mar. Freshwater Res. no commercial spp. Roemmich, D. and J. McGowan 1995, Science - 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

Holbrook et al. 1997 Ecological Applications Population responses: Offspring production: climatic variability Holbrook et al. 1997 Ecological Applications Ocean climate change Surfperch production Population responses: 4 surfperch species Benthic productivity Perch recruitment

V) Factors affecting recruitment 3) Determinants of larval production (sources of variation): Not just numbers but larval quality as well: Spatial variation in environmental quality (productivity) and larval quality MacFarlane and Norton 1998 Fishery Bulletin Larval condition of Sebastes jordani (shortbelly rockfish) among three submarine canyons: 20 40 60 80 100 protein total lipids esters triacyl- glycerols cholesterol polar Bodega Pioneer Ascension

3) Determinants of larval production (sources of variation): Not just numbers but larval quality as well: B) Larvae produced by older females grow faster and survive better Steve Berkeley 2004 Ecology; 2004 Fisheries black rockfish, Sebastes melanops similar relationships between larval condition /performance and size of oil globule  energy stores suggests age-based energy allocation by females

Number of females spawning 3) Determinants of larval production (sources of variation): Not just numbers, or quality, but timing as well: B) Females of different ages / sizes, spawn at different times over the spawning season Bobko, S. & S. Berkeley. 2004, Fishery Bulletin - examined maturity, ovarian cycle, fecundity, and age-specific parturition of black rockfish (Sebastes melanops). Conceptually… Larval survival Number of females spawning 9-10 yr 7-8 yr 5-6 yr 3-4 yr time Result: older females spawn earlier, when larvae experience higher survival and recruitment

Larval production Recruitment is important and fascinating… 100 yrs Seasonal current shifts PDO 1 decade Seagrass beds Kelp forests ENSO 1 year Seasonal upwelling Mesoscale eddies 1 month Temporal scale Coastally trapped waves 1 week Small-scale fronts, plumes, runoff 1 day Plankton migration Surface tides 1 hour Langmuir cells Internal waves Turbulent eddies 1 min Surface waves 1 cm 1 dm 1 m 10 m 100 m 1 km 10 km 100 km 1000 km 10000 km Linear spatial scales but studying it really bites! Carr and Syms 2006, CA Fishes book

Larval dispersal Larval production Sources of spatial and temporal variation in recruitment Larval dispersal Larval production

V) Factors affecting recruitment B) Determinants of larval delivery What influences the fate of propagule production? 1) Survival they can’t get there if they don’t survive (poorly understood, topic of hypotheses re: fisheries management: match/miss-match) 2) Dispersal (advection) what determines patterns of transport (small and large-scale processes) 3) Depletion (by settlement) fewer available as they settle elsewhere

Pattern: northern boundary of many species ranges at Pt Conception V) Factors affecting recruitment B) Determinants of larval delivery 2) Dispersal (advection) what determines patterns of transport? i) Large-scale (biogeographic) processes a) currents —e.g., California Current Pattern: northern boundary of many species ranges at Pt Conception

2) Dispersal (advection) what determines patterns of transport? i) Large-scale (biogeographic) processes a) currents —e.g., California Current Example: Doyle 1984, Gaines 1997 Gen. Hypothesis: larval supply limits biogeographic ranges Specific Hypothesis: if barnacle larvae transported above Pt. Conception, they would survive Test: Transplanted recently settled juveniles above Pt. Conception Result: They survived! Conclusion: Currents around Pt. Conception limited northern boundary of barnacle range

2) Dispersal (advection) what determines patterns of transport? i) Large-scale (biogeographic) processes b) currents — e.g., California current - El Nino Example: Cowen 1985 Jour. Mar Research Large scale patterns of temporal (episodic) variability Normal year (La Nada) Hypothesis: Change in current patterns influences spatial patterns of sheephead recruitment El Nino Specifically, northward El Nino currents would increase recruitment in northern portion of sheephead range.

Semicossyphus pulcher California sheephead Semicossyphus pulcher

Example: Cowen 1985 Jour. Mar Research 40 Hypothesis: Change in current patterns influences spatial patterns of sheephead recruitment 30 20 10 Test: Use annual otolith increments and settlement mark to back-calculate what year individuals settled… Use this to construct strength of year-class recruitment 20 10 ND San Nicolas Is. 20 Is. San Benito 10 ND ND Is. Guadalupe 20 Cabo Thurloe 10 ND 75 77 79 81 83 Year

V) Factors affecting recruitment B) Determinants of larval delivery 2) Dispersal (advection) what determines patterns of transport? ii) Small-scale (localized) processes a) Windward and leeward patterns around islands - local retention? Implications for “openness” of marine populations - microchemical signatures in otoliths -“flight recorders” - two cool examples: Swearer et al. 1999 Nature -St. Croix, Caribbean Jones et al. 1999 Nature -Lizard Island, Australia

Pattern: Spatial variation in recruitment of blue head wrasse, Thalassoma bifasciatum, around St. Croix Island 1 . 2 5 Caselle & Warner, 1996 Monthly recruit density (fish/m2) N 1 . 2 5 density (fish/m2) Monthly recruit

Hypothesis: Patterns of larval transport (delivery and retention) causes spatial pattern of recruitment Larval retention within island wake 5 km Swearer et al. St. Croix, Caribbean current wind Larval dispersal with patch depletion Sources of chemical signatures: Salt River Canyon Groundwater Christiansted Cruzan Rum, Hess Oil, Vialco

Result: fish that had recruited on leeward sides mostly had retention signatures, whereas fish that recruited on windward side mostly had “dispersal” signatures (“blue water”) Monthly relative recruitment intensity Mean canonical factor 1 Dispersal Retention B u t l e r a y ( L w d ) J c k ' s W i n - 1 . 5 2 3 N o h Multivariate measure of relative abundance of elements in otoliths Conclusion: recruitment on windward side from elsewhere, recruitment on leeward side from retention of locally produced larvae

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V) Factors affecting recruitment B) Determinants of larval delivery 2) Dispersal (advection) what determines patterns of transport? ii) “Smaller-scale” (more localized) processes b) Internal waves Alan Shanks 1983, MEPS ONSHORE TRANSPORT - onshore currents caused by tidal action - form at interface of salinity, temperature (= density) strata - occur on 14-day frequency - form surface slicks above trough with warm water Shanks sampled larvae as waves passed by and detected concentrations above troughs, identified intertidal waves as mechanism for onshore transport of larvae!

B) Determinants of larval delivery ii) Smaller-scale (more localized) processes c) Physical processes and larval behavior Example: Norris 1963, Ecological Monographs - Opal eye (Girella nigricans) - Pattern: recruitment related to tide pool temp. - lab experiments: thermal preference - Pattern: coast-wide, recruitment inversely related to upwelling - hypothesized mechanisms of larval delivery: interaction among… - internal waves - thermal / structural cues - upwelling

“Structure - schooling” Long larval duration 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

Mid-water complex Long larval duration Benthic complex Lenarz et al. 1995 CalCOFI 1.0 Mid-water complex Long larval duration (3 - 4 months) 0.5 Proportion 0.5 Olive, Yellowtail and Black rockfish 1.0 1986 1992 Carr and Syms 2006 100 Kelp, Black-&-yellow, and Gopher rockfish 75 50 25 Benthic complex Short larval duration (1-2 months) Relative Abundance 25 50 75 100 El Nino La Nina La Nada (1998) (1999) (2000)

Cumulative upwelling index anomaly (thru June) Pattern: Interannual variation in rockfish recruitment - midwater vs. benthic species 80 Midwater complex 70 Benthic Complex 60 -1 1 2 3 50 Cumulative upwelling index anomaly (thru June) Number of fish per transect 40 30 20 10 1999 2000 2001 2002 2003 2004 2005 Year

B) Determinants of larval delivery ii) Small-scale (localized) processes d) 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 “structure - schooling” spp. proportion “benthic - solitary” spp. 0.2 0.4 0.6 depth 13 kelp bed late larvae early larvae depth (m) 37 onshore pelagic juveniles sea floor 87-117 offshore

Mid-water complex Upwelling Long larval duration Fish per 240 m3 (3 - 4 months) Upwelling 70 Olive, Yellowtail and Black rockfish Olive rockfish 50 Fish per 240 m3 30 Yellowtail rockfish 10 El Niño La Nada Black rockfish La Niña (1998) (1999) (2000)

Relaxation Fish per 240 m3 Benthic complex Short larval duration Kelp rockfish Relaxation Gopher rockfish Black-&-yellow rockfish Kelp, Black-&-yellow, and Gopher rockfish 20 16 Fish per 240 m3 12 Benthic complex Short larval duration (1-2 months) 8 4 El Niño La Nada La Niña (1998) (1999) (2000)

(3) Smaller-scale, more frequent events (Ammann unpublished) Temperature (°C) May June July August Year 2000 9 10 11 12 13 14 Mid-water complex n = 227 per sampling unit Number of fish 0.5 0.3 0.4 0.2 0.1 0.0 May June July August per sampling unit Number of fish Benthic complex n = 363 0.5 0.3 0.4 0.2 0.1 0.0

Application: predicting ecological consequences of regional climate change Wind Stress Curl Anomalies (x 10-7 N/m3) = Upwelling decreased upwelling increased upwelling Not looking good… or differences in replenishment may increase! (Diffenbaugh et al., PNAS, 2004)

Cumulative upwelling index anomaly (thru June) Pattern: Interannual variation in rockfish recruitment - midwater vs. benthic species 80 Midwater complex 70 Benthic Complex 60 -1 1 2 3 50 Cumulative upwelling index anomaly (thru June) Number of fish per transect 40 30 20 10 1999 2000 2001 2002 2003 2004 2005 Year