Mya arenaria
0 1000 2000 3000 4000
Mid Intertidal Low Intertidal
Ind. m-2
Exclusion Control Ambient
Macoma balthica
0 500 1000 1500 2000 2500
Mid Intertidal Low Intertidal
Ind. m-2
Exclusion Control Ambient
experimental setup: 2-factorial nested block design
Ecosystem engineering in intertidal sand by the lugworm Arenicola marina
Nils Volkenborn, Lubos Polerecky*, Karsten Reise
AWI, Wadden Sea Station Sylt, *MPI for Marine Microbiology, Bremen
Alfred Wegener Institute
Alfred Wegener Institute forforPolar-Polar-und Marine Research, und Marine Research, WaddenWaddenSeaSeaStation Sylt, contactStation Sylt, contact: : nvolkenborn@awinvolkenborn@awi--bremerhaven.debremerhaven.de Deposit-feeding lugworms are widespread the
dominant large burrowing macrofaunal species and consequently the major source of sediment reworking and bioirrigation of intertidal sediments in the Wadden Sea. In 2002 an ongoing large-scale, long-term lugworm exclusion experiment was started, by meshing the sediment in 10 cm depth on 6 exclusion plots, each with an area of 400m2to reveal significant effects of lugworms on ecosystem functioning.
…biogeochemical processes
Conclusions and perspectives
• Effects of ‘ecosystem engineering’ by lugworms extend beyond the vicinity of individual burrows with considerable implications for sediment properties, biogeochemical processes, and the benthic community.
• Effects are variable in space and time.
• The experiment offers ideal testing ground for further research regarding key species, biodiversity and ecosystem functioning.
• The experiment is a contribution to the European network of Marine Biodiversity and Ecosystem Functioning (MarBEF) and one part of the BMBF project NEBROC (Netherlands Bremen Oceanography).
20 meters
Exclusion plots, December 2002
500 meters Exclusion
(netted) Ambient
(undisturbed)
Control (disturbed, without net)
Low intertidal fine sand Emersion period 3-4 h grain size median ~220 µm
Mid intertidal medium sand emersion period 6-7 h grain size median ~340 µm 20 meters
BLOCK 1 BLOCK 2
BLOCK 3
BLOCK 4 BLOCK 5
BLOCK 6
Lugworms affect...
…intertidal habitat succession
…functional diversity …structural habitat complexity
0 1 2 3 4 5
Exclusion Control Ambient
Ind. in 1.000 m-2
0 2 4 6 8 10
Exclusion Control Ambient 0 1 2 3
ExclusionControl Ambient
1st year 2nd year 3rd year
carnivorous motile worms
subsurface deposit-feeding discretely motile worms surface deposit-feeding discretely motile worms mixed suspension and surface deposit feeding tube worms
M ud Fine
S a nd M ed ium S a nd C o a rse
S a nd H W L
LW L
O2(nmol cm-3)O2(nmol cm-3)
Lugworms maintain sediment permeability and facilitate oxygen penetration with increasing hydrostatic pressures
h = 20
Exclusion site
h = 0
h = 20 h = 30 h = 40 h = 50
h = 10 h = 30 h = 40
h = 0 1 cm
Lugworm site
Berkeleya rutilans spring 2003
Enteromorpha spp. Ulva specc.
summer 2004
Algae fixed to protruding polychaete tubes on lugworm exclusion plots Lugworms facilitate sulphide sensitive
species and inhibit tube building species
Enhanced recruitment of byssus drifting bivalves
-10
0
10
20
30
0 100 200 300 400
µmol L-1
depth (cm
0.0 0.5 1.0 1.5 2.0
Medium Sand Fine Sand
Fine Fraction %
Exclusion Control Ambient
Lugworms inhibit accumulation of fine particles, porewater nutrients, and sulphide
Hypothsized expansion of permeable sandflats at the cost of cohesive mudflats
2003 mid intertidal2004lowintertidal
Sediment fraction < 63 µm Porewater nutirents (NH4)
Lugworms tidal flat and lugworm burrow