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EcologicalInteractions
James L. Bodkin and M. Tim Tinker
PreyDirectEffects
IndirectEffects
Urchins large, & abundant, Kelps rare
Sea Otters Rare or Absent
Urchins small & Kelps abundant
Sea Otters Abundant
MACROPHYTES and ECOSYSTEM STRUCTURE & FUNCTION
• Structural complexity
• Habitat diversity• Species diversity• Refugia• Complementary
production• Food web
subsidies• Hydrodynamics• Carbon
sequestration• Primary
production
vonBiela et al. 2015Most carbon in muscle of these fish derived from macro algae, in some locations up to 90%
Similar work demonstrates importance of kelp derived carbon in other nearshore taxa (invertebrates, fishes, birds, mammals) as well as export of kelp‐derived carbon to offshore food webs
Dunton et al. 1987, Duggins et al. 1989 Bustamante and Branch 1996, Fredrickson et al. 2003, Estes et al. 2004, Reisewitz et al. 2006 ..…….
Indirect effects on fish
Ecological Interactions: Nearshore Mixed Sediment Habitats
• Sheltered Shores• Macrophyte production (understory kelps and eel grass)
• Sediment excavations/disturbance• Altered abundance & size classes of infaunal bivalves
• Modified filtration rates
• Exposed Shores• Perhaps the most under‐studied of the sea otter habitats
0
50
100
150
200
250
300
350
HighMediumLowNone
Edible Biomass (g wet wt./
0.25
m2 )
Sea Otter Density
Post Pre
Biomass of Clam Prey Species at Glacier Bay Sites With Increasing Sea Otter Density
Summary Ecological Interactions, SEAK• Current status of sea otters and nearshore marine ecosystems
are an artifact of human activities over 250 years• Direct effects of otter predation can include maintenance of
abundance, sizes, and biomass of prey• Indirect effects of reduced herbivory documented across many
habitats following the removal of sea urchins• Restoration of kelp forests & other
macrophytes have ecosystem level effects related to habitat and increased primary production
• Our understanding of long term direct & indirect effects of sea otterrecovery are still incomplete
Sea Otters and Ecological Interactions,California
San Nicolas Sub-tidal sites
Spatial variation in predators and prey
San Nicolas, North Side (low otter effect)Otter effect Otter effectFrequent crashes
of urchin pop’n.s…most likelydisease driven
No relationship between otters and urchins
San Nicolas West End (high otter effect)
Strongylocentrotus franciscanus Strongylocentrotus putpuratus
West end site 1 West end site 1
Otter effect Otter effect
Red urchins start to decline by mid 1990s
Purple urchinsabundant until early 2000s
Kenner, M.C. and M.T. Tinker 2018. Stability and change in kelp forest habitats at San Nicolas Island. Western Naturalist. 78(4): 1-11
What about other coastal habitats, such as estuaries?
Another Opportunistic Experiment:Elkhorn Slough
©Keith Ellenbogen
Sea otter colonization of Elkhorn Slough
0
20
40
60
80
100
120
140
1980 1985 1990 1995 2000 2005 2010 2015 2020
Annu
al Spring Co
unt o
f Sea
Otters
Sea otters in Elkhorn Slough
Trophic Impacts to Estuarine food webs: early stages (1990s – 2010)
0
0.05
0.1
0.15
0.2
0.25
0.3
0.35
0.4
0.45
0.5
Cancercrab
clam small un-idcrab
mussel worm smallcrustacean
shore crab star sand crab snail kelp crab sand dollar
Pro
porti
on o
f tot
al b
iom
ass
cons
umed
Diet Composition 1999-2010
Top-down effects: Sea otters reduced the overall biomass and size of crabs
Hughes, B. B., R. Eby, E. Van Dyke, M. T. Tinker, C. I. Marks, K. S. Johnson, and K. Wasson. 2013. Recovery of a top predator mediates negative eutrophic effects on seagrass. Proceedings of the National Academy of Sciences, doi:10.1073/pnas.1302805110
Trophic Cascade in Sea Grass Habitat
Hughes, B. B., R. Eby, E. Van Dyke, M. T. Tinker, C. I. Marks, K. S. Johnson, and K. Wasson. 2013. Recovery of a top predator mediates negative eutrophic effects on seagrass. Proceedings of the National Academy of Sciences, doi:10.1073/pnas.1302805110
600% Increase in Seagrass Biomass!!
Trophic Impacts to Estuarine food webs: later stages (2012– 2015)
Sea otter diets in upper estuary
Otter Exclusion Procedural Control
0.0
0.5
1.0
1.5
2.0
Otter Exclusion Control
Cra
b de
nsity
cha
nge
*tra
p-1
P < 0.05
0
50
100
150
200
250
300
350
Otter Exclusion Control
Abo
vegr
ound
bio
mas
s (g
DW
) *m
-2
P < 0.05
Reduction in burrowing healthier marsh
Conclusions: Sea otter ecosystem effects vary between coastal habitatsKelp forests• primary productivity, species diversity,
enhanced nursery habitat for fish, shoreline stability, carbon fixation, etc.
Seagrass:• Sea otters reduce crab populations,
facilitating grazers in seagrass beds• Positive effects for eelgrass resilience.Salt marsh:• Sea otters reduce burrowing crabs,
healthier pickleweed• Appear to reduce bank erosion &
marsh loss (ms. in prep)
Ginny L. Eckert Juneau Center, College of Fisheries and Ocean Sciences,
University of Alaska Fairbanks
S E A O T T E R S A N D F I S H E R I E S ,S O U T H E A S T
A L A S K A
Photo credit: Deborah Mercy
Geoduck clam density
Hoyt 2015Before After
Without Sea otters
With Sea otters
Photo credit: Phillip Collaç
California sea cucumber,Parastichopus californicus
Dungeness crab,Cancer magister
Geoduck clam,Panopea generosa
Red sea urchin, Strongylocentrotus
franciscanus
Pinto abaloneHaliotus kamtchatKanaWhat are
sea otters eating?
DU
NG
ENES
S C
RA
B
clam
snai
l
shrim
p
star
othe
r
Prey type (grouped)
cont
ribut
ion
to d
iet (
biom
ass)
0.0
0.1
0.2
0.3
0.4
RED
SEA
UR
CH
IN
GEO
DU
CK
CLA
M
CA
SEA
CU
CU
MB
ER
PIN
TO A
BA
LON
E
RED
KIN
G C
RA
B
fish
mus
sel
scal
lop
othe
r mol
lusk
urch
ins
(gre
en)
crab
othe
r cuc
umbe
r
wor
m
Sea otter diets
n= 699 bouts from 6,117 dives
46% of diet commercially important
Hoyt 2015
NW Dall Island
cont
ribut
ion
to d
iet
0.0
0.2
0.4
0.6
0.8
1.0
RED
UR
CH
INS
GEO
DU
CK
CLA
MS
CA
SEA
CU
CU
MB
ER
S
RED
KIN
G C
RAB
clam
s
mus
sels
clam
s
snai
l
shrim
p
Oth
er c
ucum
bers
Oth
er
wor
m
ABAL
ON
E
Oth
er m
ollu
sks
crab
s
Urc
hin
(gre
en)
star
fish
DU
NG
EN
ES
S C
RA
B
Coronation Island
cont
ribut
ion
to d
iet
0.0
0.2
0.4
0.6
0.8
1.0
Geographic differences in diet
Hoyt 2015
Hoyt 2015
0.
0.25
0.5
0.75
1.
0. 7.5 15. 22.5 30.
Commercial con
tribution to diet
Maximum observed density (otters km‐2)
Commercial contribution as a log‐linear function of density
NW Dall Island (99% red sea urchins)
Photo Credit: Enrique R. Aguirre Aves
Sea Otter by Victor Tumuluk (Hydaburg)
Photo Credit: Enrique R. Aguirre Aves
Ibarra in progress
H U M A N S - S E A O T T E R S - B U T T E R
C L A M S
0%
10%
20%
30%
40%
1‐10 21‐30 41‐50 61‐70 81‐90 101‐110 121‐130
Freq
uency (%
)
Length (mm)
No Sea Otters or Humans
Clam size distribution without sea otters
Ibarra in progress
Clam size distribution with sea otters
Only Sea Otters
0%
10%
20%
30%
40%
1‐10 21‐30 41‐50 61‐70 81‐90 101‐110 121‐130
Freq
uency (%
)
Length (mm)
Ibarra in progress
0%
10%
20%
30%
40%
1‐10 21‐30 41‐50 61‐70 81‐90 101‐110 121‐130
Freq
uency (%
)
Length (mm)
Sea Otters & Humans
Clam size distribution with sea otters & humans
Ibarra in progress
Sea otters and humans• Commercial species in diet greatest early after colonization: long‐established populations supported by clams
• Effect on subsistence clam populations, both abundance and size structure
• Coexistence will be challenging: must account for spatial structure of sea otter populations, and dynamic nature of diet and interactions
T H A N K S
Photo credit: Deborah Mercy
Zac HoytSean LarsonSonia N. IbarraWendel W. RaymondStephen J. Langdon Sunny RiceVerena GillTim Tinker
Alaska Sea Grant, North Pacific Research BoardSoutheast Alaska Regional Dive Fishery AssociationNational Science Foundation