“Bipartite life history of marine species and “openness ... · “Bipartite life history of marine species and “openness” of populations ... 1 cm 1000 km100 ... Temporal scale

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“Bipartite life history of marine speciesand “openness” of populations

“Open” Populations“Closed” Populations

SupplyProduction

Little or no exchange among

populations

Significant exchange among

populations

SupplyProduction

Supply ProductionSupplyProduction

2

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:

1) Evidence for long distance dispersal of larvae:a. prolonged pelagic larval duration (PLD)b. Low spatial genetic structure

2) Evidence of shorter distance movement of adults a. Sessile organisms (algae and invertebrates)b. Limited distance of adult movement

Propagule Duration (hr)

Pelagic duration: a proxy for dispersal potential

Dis

per

sal

Dis

tan

ce (

km)

Shanks et al. 2003 Ecological Applications

0.0001

0.001

0.01

0.1

1

10

100

1000

10000

0.01 0.1 1 10 100 1000 10000

invertebrates

fish

Predicted by passive dispersal

3

Western North American Coastal FishTime in Larval Stage

midpoint (range)

Shanks et al. 2003AVERAGE = 94 days

Time in the larval stage (fish)

Pacific Ocean Reef FishesLarval Duration Estimates

10

20

30

40

50

60

70

< 1 1-7 31-90 > 90

Per

cen

t o

f S

pec

ies

Tropical (n= 298 spp)

Temperate (n= 60 spp)

Larval Duration (days)

8-30

Carr and Syms 2006, CA Fishes book

4

Gen

etic

dif

fere

nce

Geographic distance (kilometers)

Geographic distance = genetic difference

Slope measures average dispersal

short

low

long

high

Gen

etic

dif

fere

nce


Geographic distance = genetic difference

Slope measures average dispersal

populations nearby one another

populations further apart

short

low

long

high

5

Gen

etic

dif

fere

nce

Genetic difference

8000 200 400 600

low

hig

h


Copper rockfish

CA snailsRosethorn rockfish

CA corals

Fish: 20-200 kilometers

Palumbi 2003Kinlan and Gaines 2003

Larval dispersalBased on geneticdifference

Inverts: <1-100 kilometers

Num

ber

of s

peci

es

6

Different estimates, similar results

Dispersal distance of fish larvae = 20 - 200 km

Time in larval stage Genetic distance

Time as larvae (hr)

0.0001

0.001

0.01

0.1

1

10

100

1000

10000

0.01 0.1 1 10 100 1000 10000

Dis

per

sal

dis

tance

Dispersal distance of invert larvae = 1 - 100 km

= 30 days

Conclusion: 76% of these species moved less than 0.5 km

Home ranges of 25 west coast rocky habitat fish species

Median max. distance (km)

num

ber

of s

peci

es

0

4

8

12

Limited Adult Movement

Freiwald, J. 2012. Can. J. of Fish. & Aquat. Sci.

7

Some schooling fish

0 – 1 km 1 – 10 km 10 – 100 km 100 – 1000 km > 1000 km

Many rockfish

Other reef fish

Some surfperch

Some surfperch

Some rockfish Some rockfish

Other reef fish

Some flatfish

Few rockfish

Salmon

More flatfish

Some schooling fish

Tunas

Many sharks

Adult Home Range Size Varies Among Species of Fishesas does the “openness” of their populations!

Sources of spatial and temporal variation in recruitment

Larval production

Larval dispersal

Settlement

Post-settlement

8

Seagrass beds

Recruitment is important and fascinating…

but studying it really bites!

1 cm 1000 km100 km

Linear spatial scales

Tem

po

ral s

cale

1 m 10 m 100 m 1 km 10 km1 dm

100 yrs

1 month

1 week

1 day

1 hour

1 min

1 year

1 decade

Turbulent eddies Surfacewaves

Surface tides

Langmuir cells

Planktonmigration

Small-scalefonts, plumes,

runoff

Coastallytrappedwaves

Seasonalupwelling

Mesoscaleeddies

ENSO

10000 km

PDO

Internalwaves

Internaltides

Coastal filaments,Upwelling / relaxation

Seasonalcurrent shifts


Kelpforests

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??

ii) problem of tracking or identifying source of highly dispersive offspring

iii) poor stock-recruitment relationships

i) presumed decoupling

9


1) Reproduction by adults -very little work has been done on this.

Poor stock-recruitment relationships from fisheries statistics:

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

dispersal distance

(% settlers produced locally)


10


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) seasonality (influences dispersal patterns)

vii) location (influences dispersal patterns)

Offspring production: climatic variability

- 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

Power plant impingement of fish larvae:Ocean climate change

1) Love et al. 1998 Fishery Bulletinincluded commercial species

2) Brooks et al. 2002 Mar. Freshwater Res.no commercial spp.

11

Offspring production: climatic variability

Perch recruitment

Population responses:4 surfperch

species

Benthic productivity

Surfperch production

Holbrook et al. 1997 Ecological Applications

Ocean climate change



Not just numbers but larval quality as well:

A) Spatial variation in environmental quality (productivity) and larval quality MacFarlane and Norton 1998 Fishery Bulletin

0

20

40

60

80

100

protein totallipids

esters triacyl-glycerol

s

cholesterol polarlipids

BodegaPioneerAscension

Larval condition of Sebastes jordani (shortbelly rockfish) among three submarine canyons:

12


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


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…

Result: older females spawn earlier, when larvae experience higher survival and recruitment

Number of females

spawning

time

Larval survival

3-4 yr5-6 yr7-8 yr9-10 yr

13

Seagrass beds

Recruitment is important and fascinating…

but studying it really bites!

1 cm 1000 km100 km

Linear spatial scales

Tem

po

ral s

cale

1 m 10 m 100 m 1 km 10 km1 dm

100 yrs

1 month

1 week

1 day

1 hour

1 min

1 year

1 decade

Turbulent eddies Surfacewaves

Surface tides

Langmuir cells

Planktonmigration

Coastallytrappedwaves

Seasonalupwelling

Mesoscaleeddies

ENSO

10000 km

PDO

Internalwaves

Seasonalcurrent shifts

Kelpforests

Larval production

Small-scalefronts, plumes,

runoff


Sources of spatial and temporal variation in recruitment

Larval dispersal

Larval production

14


B) Determinants of larval delivery

What influences the fate of propagule production?

1) Survivalthey 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



a) currents —e.g., California Current

2) Dispersal (advection)what determines patterns of transport?

i) Large-scale (biogeographic) processes

15


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


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)

El Nino

Hypothesis: Change in current patterns influences spatial patterns of sheephead recruitment

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b) currents — e.g., California current - El Nino



Large scale patterns of temporal (episodic) variability

Normal year (La Nada)

El Nino


Specifically, northward El Nino currents would increase recruitment in northern portion of sheephead range.

California sheepheadSemicossyphus pulcher

17



Test: Use annual otolith increments and settlement mark to back-calculate what year individuals settled… Use this to construct strength of year-class recruitment

San Nicolas Is.

Is. Guadalupe

Cabo Thurloe

0 0 0

0ND

ND ND

0ND

10

20

10

20

20

20

10

10

30

40

Is. San Benito

75 77 79 81 83

Year



a) Windward and leeward patterns around islands


ii) Small-scale (localized) processes

- local retention? Implications for “openness” of marine populations

- microchemical signatures in otoliths -“flight recorders”

- two cool examples:Swearer et al. 1999 Nature -St. Croix, CaribbeanJones et al. 1999 Nature -Lizard Island, Australia

18

Pattern: Spatial variation in recruitment of blue head wrasse, Thalassoma bifasciatum, around St. Croix Island

Caselle & Warner, 1996

Mon

thly

rec

ruit

densi

ty (

fish

/m2)

Mon

thly recru

itden

sity (fish/m

2)

0

1.0

2.0

1.5

0.5

N

1.0

2.0

1.5

0.50

5 km

current

wind

Larval dispersal withpatch depletion

Larval retention within island wake

Hypothesis: Patterns of larval transport (delivery and retention) causes spatial pattern of recruitment

Sources of chemical signatures:

Christiansted

Cruzan Rum, Hess Oil, Vialco

Salt River Canyon

Groundwater

Swearer et al. St. Croix, Caribbean

19

Monthly relative recruitment intensity

Mea

n c

anon

ical

fac

tor

1

Dis

per

sal R

eten

tion

Butler Bay (Leeward) Jack's Bay (Windward)

-1.5

-1.0

-0.5

0.0

0.5

1.0

1.5

2.5 3.0

Northstar (Leeward)

1.0 2.0 2.5 0.0 1.0 2.00.5 1.51.52.25 3.25

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

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”)



b) Internal waves

2) Dispersal (advection) what determines patterns of transport?


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!

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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

c) Physical processes and larval behavior



“Structure - schooling”Long larval duration(3 - 4 months)

“Benthic - solitary”Short larval duration(1-2 months)

Black-&-yellow rockfish

Gopher rockfish

Kelp rockfish

Black rockfish

Yellowtail rockfish

Olive rockfish

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(1998) (1999) (2000)El Nino La Nina La Nada

Mid-water complexLong larval duration(3 - 4 months)

Benthic complexShort larval duration(1-2 months)

Kelp, Black-&-yellow, and Gopher rockfish

Olive, Yellowtail and Black rockfish

1007550250

255075

100

Rel

ativ

e A

bu

nd

ance

1986 1992

1.0

0.5

0.5

1.0

0

Pro

po

rtio

n

Lenarz et al. 1995 CalCOFI

Carr and Syms 2006

Pattern: Interannual variation in rockfish recruitment - midwater vs. benthic species

Num

ber

of

fish

per

tra

nse

ct Midwater complex

Benthic Complex

0

10

20

30

40

50

60

70

80

1999 2000 2001 2002 2003 2004 2005

Year

-1

0

1

2

3

Cumulativeupwelling

indexanomaly

(thru June)

22

e.g., Larson et al. 1994, Lenarz et al. 1995, CalCOFI Rpt.s

d) shifts in vertical distribution with ontogeny -- upwelling

- vertical distribution of early and late larval rockfishes

0 0.2 0.4 0.6

13

37

87-117

proportion

depth (m)

“structure - schooling” spp.

“benthic - solitary” spp.

offshore

onshore

depthearly

larvae

latelarvae

pelagicjuveniles

kelp bed



Mid-water complexLong larval duration(3 - 4 months)

Upwelling

10

30

50

70

(1998) (1999) (2000)

El NiñoLa Niña

La Nada

Fis

h p

er 2

40 m

3

Olive, Yellowtail and Black rockfish

Black rockfish

Olive rockfish

Yellowtail rockfish

23

Benthic complexShort larval duration(1-2 months)

Relaxation

(1998) (1999) (2000)

El NiñoLa Niña

La Nada

Fis

h p

er 2

40 m

3

0

4

8

12

16

20 Kelp, Black-&-yellow, and Gopher rockfish

Black-&-yellow rockfish

Gopher rockfish

Kelp rockfish

May June July August

Nu

mb

er o

f fi

shp

er s

amp

lin

g u

nit Benthic complex

n = 363 0.5

0.3

0.4

0.2

0.1

0.0

Mid-water complexn = 227

Nu

mb

er o

f fi

shp

er s

amp

lin

g u

nit

0.5

0.3

0.4

0.2

0.1

0.0

Tem

per

atu

re (

°C)

May June July August

Year 2000

9

10

11

12

13

14

(3) Smaller-scale, more frequent events(Ammann unpublished)

24

Feb Apr Jun Aug Oct

0 0.2 0.4-0.2-0.4

Feb Apr Jun Aug OctFeb Apr Jun Aug Oct

33.3N

37.0N

40.7N

44.4N

2xCO2 – 1xCO2 2xCO2Veg – 2xCO2 2xCO2Veg – 1xCO2

Not looking good… or differences in replenishment may increase!(Diffenbaugh et al., PNAS, 2004)

Wind Stress Curl Anomalies (x 10-7 N/m3) = Upwelling

Application: predicting ecological consequences of regional climate change

Pattern: Interannual variation in rockfish recruitment - midwater vs. benthic species

Num

ber

of

fish

per

tra

nse

ct Midwater complex

Benthic Complex

0

10

20

30

40

50

60

70

80

1999 2000 2001 2002 2003 2004 2005

Year

-1

0

1

2

3

Cumulativeupwelling

indexanomaly

(thru June)

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“Bipartite life history of marine species and “openness ... · “Bipartite life history of marine species and “openness” of populations ... 1 cm 1000 km100 ... Temporal scale