west side story the context, causes, and consequences of the pacific oyster introduction to...
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West Side Story the context, causes, and consequences of the Pacific oyster introduction to Washington State
Jennifer Ruesink
University of Washington
• Context– Ruesink, Lenihan, Trimble, Heiman, Micheli, Byers, Kay. 2005.
Introduction of non-native oysters: ecosystem effects and restoration implications. Annu Rev Ecol Evol Syst 36:643-689
• Causes– White, Ruesink, Trimble. History and management of native oysters
(Ostrea conchaphila) in Washington State. In preparation for special issue of J Shellfish Res
• Consequences– Ruesink, Feist, Harvey, Hong, Trimble, Wisehart. 2006. Changes in
productivity associated with four introduced species: Ecosystem transformation of a “pristine” estuary. Mar Ecol Prog Ser 311:203-215
– Trimble, Ruesink, Dumbauld. Factors preventing recovery of a historically overexploited shellfish species. In preparation.
Outline
Context
• Oyster introductions have occurred frequently
• High establishment rate
• Vector for numerous other species
• Nearly complete replacement of native oyster production
Compilation of oyster introductions
• Introduction = movement of species from one country/ region to another where it was not previously present
• Sources = UN FAO, several earlier reviews (Eldredge 1994, NRC 2004), published papers, personal communications
• Number of introductions = _____• Number of recipient locations = ____• Number of species = ____
Compilation of oyster introductions
• Introduction = movement of species from one country/ region to another where it was not previously present
• Sources = UN FAO, several earlier reviews (Eldredge 1994, NRC 2004), published papers, personal communications
• Number of introductions = _164_• Number of recipient locations = _73_• Number of species = _16_
When did oyster introductions occur?
0
20
40
60
80
100
120
1850 1870 1890 1910 1930 1950 1970 1990 2010
Cumulative number of transfers
Charles Elton fingered oyster introductions as the “greatest agency of all that spreads marine animals to new corners of the world”
Where have oysters been introduced?
18
61
6
19
9
6
41
3
4
Which oyster species have been introduced?
O’Foighil & Taylor 2000 Molecular Phylogenetics & Evolution 15:301
C. virginica = 16C. rhizophorae = 3
C. gigas = 64 + 6C. ariakensis = 3S. commercialis = 6S. cucullata = 3
O. edulis = 11
T. chilensis = 3C. densalamellosa = 1
O. conchaphila = 1O. puelchana = 1
C. echinata = 5C. sikamea = 3
C. iredalei = 2C. belcheri = 1
C. cortezensis = 1
How well did they do?
0
10
20
30
40
50
60
70
C. gigasC. virginica
O. edulisC. angulata
S. commercialis
C. echinataC. ariakensisC. rhizophoraeC. sikameaS. cucullataT. chilensis
Frequency of locations
Unknown
Failed
Established
Why were new oysters introduced?
• Replace native species
• Begin new product (Pacific islands)
• Research (12)
• Range expansion (4)
• By-product (3)
Introduced oysters have replaced native oyster production
Africa W USA E USA CanadaS AmericaAustralia/NZEurope
Asia
10-year production 1993-2002(metric tons shucked)
0
1e+6
2e+6
3e+7
Introduced Native Uncertain
Introduced oysters have
vectored many other
species
Context
• Oyster introductions have occurred frequently• High establishment rate• Vector for numerous other species• Nearly complete replacement of native oyster production
However, ecological impacts of introduced oysters are poorly studied:•To what extent do introduced oysters replace “ecosystem function”?•Is recovery of native oysters improved or impaired?
Causes
• Why were Pacific oysters (Crassostrea gigas) introduced to Washington State?– Harvesting native oysters was no longer
economical… but therein lies a story
VANCOUVER
SEATTLE
PORTLAND
Willapa Bay
Puget Sound
Native oyster
Baker 1995
Ostreola conchaphila (Carpenter, 1857)
•Historic Range: Sitka, AK to Mexico
•Maximum 6 cm
•Protandrous Hermaphrodite
•Brooding Females: ~250,000 larvae/adult
•Spawning above 12C for 3+ months/yr.
•Subtidal accumulations of shell
The way it used to be? Willapa Bay at low water, late 1800s
Willapa Bay – current population between 0 and -2’ MLLW is sparse in eelgrass
North Bay, Puget Sound – ~25 million oysters between 0 and -4’, newly recovered
Washington Ostreola conchaphila Harvest
0.0E+00
2.0E+04
4.0E+04
6.0E+04
8.0E+04
1.0E+05
1.2E+05
1.4E+05
1850 1860 1870 1880 1890 1900 1910 1920 1930 1940 1950 1960
Sacks
Willapa Bay
Puget Sound
Trimble, unpubl.
1840 1850 1860 1870 1880 1890 1900 1910
White population exploiting oysters0
50
100
150
200
250
Price $/basket
0
5
10
15
20
25
30
35
White fishers Price/basket
Willapa Bay timeline
1840 1850 1860 1870 1880 1890 1900 1910
White population exploiting oysters0
50
100
150
200
250
Price $/basket
0
5
10
15
20
25
30
35
White fishers Price/basket
First harvest restrictions
1840 1850 1860 1870 1880 1890 1900 1910
White population exploiting oysters0
50
100
150
200
250
Price $/basket
0
5
10
15
20
25
30
35
White fishers Price/basket
Native oysters described scientifically
1840 1850 1860 1870 1880 1890 1900 1910
White population exploiting oysters0
50
100
150
200
250
Price $/basket
0
5
10
15
20
25
30
35
White fishers Price/basket
Tideflat privatization
1840 1850 1860 1870 1880 1890 1900 1910
White population exploiting oysters0
50
100
150
200
250
Price $/basket
0
5
10
15
20
25
30
35
White fishers Price/basket
Marine Reserves
Collins, 1888
Oyster Reserves: •Legislated ~1900•Almost perfect overlap with original native oyster beds
1840 1850 1860 1870 1880 1890 1900 1910
White population exploiting oysters0
50
100
150
200
250
Price $/basket
0
5
10
15
20
25
30
35
White fishers Price/basket
C. virginica introductionSubsequent introductions:
C. gigas - establishedO. edulisC. sikameaC. ariakensis
Commercial interest shifted to Crassostrea gigas: Imports of spat to the west coast
1920 1930 1940 1950 1960 1970 1980
Cases
0
20000
40000
60000
80000
100000
120000
White, Ruesink, Trimble, unpubl.
Secondary production in Willapa Bay shifted from native to non-native shellfish (filtration too)
Ruesink, Feist, Harvey, Hong, Trimble, Wisehart. 2006. Mar Ecol Prog Ser
Research efforts also shifted
• Number of holdings in the University of Washington library referring to each species
• Scientific literature includes <15 modern papers on O. conchaphila – but this will soon change!
0
5
10
15
20
25
30
35
<1930 1930-60 1960-80
Publications
Native
Introduced
Has C. gigas functionally replaced O. conchaphila?
• Native oyster– Brooding– Slow growth– Subtidal– Loose shell
• New oyster– Broadcasting– Rapid growth– Intertidal– Reef-building
Consequences
• What are the ecological impacts of Pacific oysters (Crassostrea gigas)?– Interactions with native oysters – directly and
indirectly– Effects of shellfish and aquaculture practices
on sediment properties, eelgrass, epibiota, fish and crabs are the focus of targeted research funded in part by the shellfish industry
Potential Factors Limiting Population Recovery
- Reproductive Failure (no)
-----------------------
- Settlement Habitat Change (yes)
- Pollution and Sedimentation (yes)
- Competition (space- yes)
- Predation (yes)
- Disease (limited)
Middle Sands Reserve
Long Island Reserve
Spatfall onCinder Block Anchors
(-10m)
Native oyster recruitment
remains high
Quantitative recruitment time series
• Weekly records of spatfall from 1947-1987, 2002-2006
• Native oyster usually > Pacific oyster
0
10
20
30
40
50
60
70
80
90
100
1945 1965 1985 2005
Spat Per Shellface
1947-2006 Spatfall: O. conchaphila and C. gigas
O.conchaphilaC. gigas
But where do those larvae now settle?
What native oyster beds used to be… maybe… perhaps?
Settlement Habitat Change
C. gigas intertidal reefs
Settlement Habitat Change
Increased sedimentation from logging in watershed
Quantitative assessment of recruitment across habitats
Recruitment rate across habitatsOstreola conchaphila
Habitat
Shell Bare Eelgrass
Recruits per 10 shellfaces
0
50
100
150
200
250
300
Above MLLW Below MLLW
Mos
t av
aila
ble
habi
tat
2-factor experiment: 3 elevations, +/- competitors
Tiles on mooring
Day 0 (1 month old) Day 63 Day 304
MLLW
(- 30cm)
(+ 30cm)“HIGH”
“LOW”
“MOORING” (-1 m)
F8,112 = 2.8, P = 0.007
Survival declines when not submerged
Impacts of competitors
• Ectopleura crocea• Botryllus spp.• >45 introduced species
reported in Willapa Bay
Wonham & Carlton 2005 Biol Inv
F1,112 = 42.4, P < 0.001
Competitors Reduce Survival
F4,61 = 5.5, P = 0.001
Competitors Reduce Growth
Interactions between Washington’s native and non-native oysters
Recruitment “sink”:Intertidal aquacultureNaturalized intertidal reefs
Historical vector of non-native predators and competitors
Recruitment, but poor intertidal survival
-
Overgrowth?
High sustained recruitment indicates presence of core reproductive population
Bare Gravel Pacific shell
Crushed Pacific shell Native shell Live natives
May 2004
Recruitment improved at
lower elevations
and on natives
A
Elevation (m MLLW)
-0.6 -0.4 -0.2 0.0 0.2 0.4 0.6
Recruit density (number per 0.0125 m
2)
0
2
4
6
8
10
12
14
16
18
Total October live April live
B
Substrate type
O.c. live O.c. shell C.g. crushed C.g. shell Gravel Bare
Recruit density (number per 0.0125 m
2)
0
5
10
15
20
25
A
Rosette Bag Ground Shell
Count per shell
0
2
4
6
8
10
12
14
16
18
20
B
Density: Low High Low High Stable Unstable
Rosette Bag Ground Shell
Shell length (mm)
25
30
35
40
RogersNemahLong IslandMill Channel
Unstable treatments washed away at most sites
Stable treatments were heavily fouled at most sites
Rosettes – natives grew and survived
Rosettes – buried at other sites
Shell on ground – not a functional replacement for natives