1. “Bipartite life history of marine species and “openness” of populations1 1

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

3. 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 = d, mean realized dispersal distance= km) .
Oligocottus maculosus -- observations of dispersing larvae and their distribution (duration = 30 d, mean realized dispersal distance= < 1 km)

4. 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 = d, mean realized dispersal distance= 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 Ecological Applications 4

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

6. 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

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

8. 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)

9. 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)

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

11. 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 = km
Dispersal distance of fish larvae = km

12. 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 Can. J. of Fish. & Aquat. Sci. 12 12

13. 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

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

15. 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

16. 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

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

18. 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

19. 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)

20. Offspring production: climatic variability
Ocean climate change
Power plant impingement of fish larvae:
1) Love et al Fishery Bulletin
included commercial species
2) Brooks et al 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. ( )
- attributed to reduced production (but maybe larval survival)
- reflecting large-scale decline in productivity

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

22. 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

23. 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

24. 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

25. 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

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

27. 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

28. 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

29. 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

30. 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.

31. Semicossyphus pulcher
California sheephead
Semicossyphus pulcher

32. Example: Cowen 1985 Jour. Mar Research40
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

33. 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 Nature -St. Croix, Caribbean
Jones et al Nature -Lizard Island, Australia

34. 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

35. 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

36. 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

37. START HERE

38. 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!

39. 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

40. “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

41. Mid-water complex Long larval duration Benthic complex
Lenarz et al 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)

42. 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

43. 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 , 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

44. 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)

45. 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)

46. (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

47. 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)

48. 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