Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea. Causes and consequences of a successful invasion.

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HARBASINS Report:
Spread of the Pacific Oyster
Crassostrea gigas in the Wadden Sea
Causes and consequences of a successful invasion
Prepared by: Georg Nehls
Heike Büttger
BioConsult SH
Brinckmannstr. 31
D-25813 Husum, Germany
Web: www.bioconsult-sh.de
On behalf of: The Common Wadden Sea Secretariat, Wilhelmshaven
April 2007
HARBASINS is a project funded under the European Regional
Development Fund INTERREG IIIB North Sea Region Program – A
European Community Initiative concerning Trans National Co-operation
on Spatial Development 2000-2006.

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
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Contents
1 INTRODUCTION AND SCOPE OF WORK.................................................................................3
2 GENERAL ASPECTS.................................................................................................................4
2.1 Distribution and Ecology of Crassostrea gigas.....................................................................4
2.2 History of introduction and geographical spread..................................................................6
3 SPREAD IN THE WADDEN SEA................................................................................................8
3.1 Temporal course.................................................................................................................8
3.2 Development of Pacific oyster density and biomass..........................................................10
3.3 What facilitates the increase of Pacific oysters in the Wadden Sea ?.................................22
3.4 Distribution over different habitats.....................................................................................23
4 ECOLOGICAL CONSEQUENCES...........................................................................................26
4.1 Do Pacific oysters displace blue mussels?........................................................................26
4.2 Blue mussel and oyster beds and their associated species communities...........................30
4.3 Introduction of other species associated with oysters........................................................35
4.4 Consequences for energy flow and top predators (birds)...................................................36
5 OYSTER FISHERIES...............................................................................................................37
6 MANAGEMENT........................................................................................................................38
7 MONITORING/ASSESSMENT.................................................................................................39
8 SUMMARY...............................................................................................................................41
9 LITERATUR.............................................................................................................................43
Annex 1: Conclusions and Recommendations..................................................................................47
Annex 2: Final Workshop Program....................................................................................................52
Annex 3: Workshop Participants.......................................................................................................53

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1 INTRODUCTION AND SCOPE OF WORK
The recent spread of the Pacific oyster Crassostrea gigas in the Wadden Sea is one of the
most spectacular biological invasions in this ecosystem and has attracted the attention of the
public as well as of the scientific community. Pacific oysters have most probably been
brought to European waters since ships are sailing between the continents. They have been
cultivated in various places for decades and were seemingly not adapted to prosper in the
Wadden Sea. Brought out in cultures in France and the Netherlands to substitute the
European Oyster which became extinct in many places, it was reported to spread soon in
warm summers and form stable populations outside the cultures since the seventies which
are increasing until today. Pacific oysters are found in the Wadden Sea since the eighties,
where cultivation has been licensed in the Lister Deep between the island of Sylt and the
mainland since 1986. As commonly reported from other biological invasions, the
development in the Wadden Sea was rather slow over many years and some colonised sites
were abandoned again. However, following a period of rather warm summers, Pacific
Oysters have drastically increased, overgrown many native blue mussel Mytilus edulis beds
and are suddenly an abundant and dominating species on the tidal flats. By forming
extensive and stable reefs on the tidal flats, Pacific oysters create new habitats which may
be suitable for other species living in the mussel bed community. The sudden and fast
spread of this invader has raised concern about the ecological impacts. Will blue mussel
beds be displaced by Pacific oyster reefs, and if so, what will be the consequences for the
benthic communities, how will mussel eating birds be affected? As Pacific oysters mainly
settled into native mussel beds they are also considered as a problem for mussel fisheries.
On the other hand, it is not quite clear how to deal with an invader in the Wadden Sea
ecosystem, a habitat protected in all parts. May fishing or even removal of Pacific oyster
reefs be licensed within the Wadden Sea National Parks?
For the first time a comprehensive data compilation of the entire Wadden Sea was carried
out in the framework of the TMAP to document the spread of the Pacific Oyster with special
emphasis on the recent strong increase. The report aims to provide support for reporting and
assessment under the EU Water Framework Directive and Habitats Directive and to support
necessary updates of the Trilateral Monitoring and Assessment Program.
The help of our colleagues in compiling for the first time data from the entire Wadden Sea to document
the spread of the Pacific oysters is gratefully acknowledged. We would especially like to thank Johan
Craeymeersch, Norbert Dankers, Gerald Millat, Achim Wehrmann, Alexandra Markert, Karsten Reise,
Per Sand Kristensen and Helle Torp Christensen for making unpublished data accessible. We would
further like to thank Harald Marencic and the Common Wadden Sea Secretariat and the Regional
Agency for the Wadden Sea National Park of Schleswig-Holstein for support.

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2 GENERAL ASPECTS
2.1 Distribution and Ecology of Crassostrea gigas
Pacific oysters are cultivated in many countries of the world and they have spread to a
variety of new habitats and successfully established stable populations. The introduction of
this species has been reported for at least 60 countries (Ruesink et al. 2005). However,
although being locally highly important, introduced oyster only make up 6% of the worlds
annual oyster harvest of about 3.3 million tons (Ruesink et al. 2005). The spread of the
species into new habitats is documented in a variety of sources (e.g. www.issg.org,
www.marine.csiro.au, www.nobanis.org) and C. gigas is now globally distributed and occurs
in the wild on all continents (Fig. 1).
Fig. 1: Worldwide distribution of Crassostrea gigas (NIMPIS).
Pacific oysters are currently reported to occur in 24 countries, but only in six of these the
species is considered to be invasive (Tab. 1). Interestingly, until recently, the species has
been considered not to be invasive in most European countries and the Mediterranean.
Tab. 1: List of countries where Crassostrea gigas has established non-native populations (ISSG
2005).
Not invasive invasive not specified
Belgium Australia South Africa
China Canada
Cyprus Chile
Denmark The Netherlands
France New Zealand
French Polynesia USA
Germany
Greece
Korea
Malta
Morocco
Norway
Portugal
Spain
Tunisia
United Kingdom
Vanuatu

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The biology of C. gigas has been recently reviewed by several authors (e.g. Kater 2003,
Nehring 2006) and for detailed reading it is referred to these sources. Looking at the species
from the Wadden Sea point of view, the following aspects are regarded as important:
• The species reproduces in the summer months at water temperatures exceeding 20°
C, but is very tolerant to cold winter temperatures.
• C. gigas may live both in intertidal and subtidal habitats. In the Wadden Sea, it mainly
lives in the intertidal in the same zone as blue mussels.
• Pacific oysters may live for 30 years; reach a shell length of 30 to 40 cm and an
individual weight of more than 1 kg (live wet weight).
• As in other bivalves, Pacific oysters have pelagic larvae spending 3 to 4 weeks in a
free-swimming phase.
• Pacific oyster larvae may settle on all kind of natural and artificial hard substrates as
mollusc shells, living molluscs, wood, stones, concrete and others.
• Pacific oysters are consumed by a variety of marine animals as Asteroid
echinoderms, boring gastropods, boring bivalves, spionid polychaetes. Carcinus
maenas in the intertidal, Cancer irroratus, benthic feeding fish, lobsters in the subtidal
zone, black ducks, eider ducks, and wading birds (see NIMPIS). In the Wadden Sea,
however, predation seems to be very limited.
• As other bivalves, Pacific oysters are filter feeders consuming phytoplancton and
other digestible organic materials from the water column.

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2.2 History of introduction and geographical spread
(from Nehring 2006)
More than once during the 19th century, attempts have been made to revive exploited stocks
of the European oyster (Ostrea edulis) with American oysters (Crassostrea virginica) and
‘Portuguese oysters C. angulata’ at several sites in coastal waters of Northern Europe.
These attempts largely failed (Wolff and Reise 2002). In 1964 Dutch oyster farmers imported
spat of the Pacific oyster (Crassostrea gigas) from British Columbia for aquaculture activities
in the Oosterschelde estuary. In the following years more imports of spat and adult
specimens followed, starting in 1966 also from Japan. In 1975 and 1976 natural spatfalls
occurred during very warm summers and resulted in millions of so-called weed oysters in the
Oosterschelde estuary. Within several years the Pacific oyster has developed explosively
and in the 1980’s other Dutch estuaries started to be colonized (Wolff and Reise 2002). In
1983 first specimens were observed near the island of Texel in the Dutch Wadden Sea,
probably brought there deliberately from the Oosterschelde in the 1970s (Bruins 1983, Wolff
2005). Since the 1990’s this alien species was frequently observed along the entire Dutch
coast and in the Dutch Wadden Sea (Dankers et al. 2004). In 1996 a first settlement of the
Pacific oyster occurred in the western Wadden Sea area of Germany as well, which may
have been dispersed from the Netherlands by natural means (Wehrmann et al. 2000).
In Germany ‘Portuguese oysters Crassostrea angulata’ were imported from Portugal and
Spain and introduced into the Wadden Sea near Norddeich in 1913-14 and in the Jadebusen
and near Sylt in 1954, 1961 and 1964. This did not lead to lasting cultures or establishment
of the species (Meyer-Waarden 1964, Neudecker 1992, Wehrmann et al. 2000). Between
1971 and 1987 spat and larvae of the Pacific oyster were repeatedly imported from Scottish
hatcheries for scientific aquacultural experiments and studies at different sites in the German
Wadden Sea and at the German Baltic Sea coast in the Flensburg Fjord (Meixner and
Gerdener 1976, Seaman 1985, Wehrmann et al. 2000). All these attempts, however, proved
unsuccessful. Since 1986 commercial farming activities began in the northern area of the
German Wadden Sea near the island of Sylt, primarily with spat taken from British and Irish
hatcheries (Reise 1998, Nehring 1999). Pacific oysters are cultivated in plastic mesh bags on
trestles in the intertidal zone. It takes about 2 years until the Pacific oysters reach marketable
size. Shortly after oyster farming had commenced, natural spatfalls occurred and in 1991 the
first Pacific oysters were found outside the culture plot (Reise 1998).
In Denmark Pacific oysters from German aquaculture experiments in the Flensburg Fjord
were planted in the Little Belt of the Baltic Sea around the island Bogø in 1979. During the
last thirty years of the last century large amount of C. gigas (> millions) seed oysters were
imported from England, The Netherlands, and France to different Danish marine waters for
culture in marine aquaculture experiments. The aquaculture took place in the Little Belt, in
the waters south of the island Funen, in Horsens fjord, around the island Samsø in Kattegat

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
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and in Isefjorden (Kristensen, 1986, Kristensen and Hoffmann in press). In Isefjorden
commercial production has taken place and about 100.000 to 300.000 Pacific oysters were
harvested annually between 1986 and 1999. Apparently Pacific oysters were abandoned in
the area, where they survived for several years but did not expand. Pacific oysters imported
by the Limfjord Oyster Company were kept in basins for depuration. They were not cultured -
though there are anecdotal information of finding escapees years after the company ceased
its activities (Kathe Jensen, pers. comm.). At the end of the last century commercial
production has taken place in the Danish Wadden Sea (very limited scale with a few
thousands oysters annually) (Per Dolmer and Per Sand Kristensen, pers. comm.). In 1999
first freeliving specimens were observed, which may have been dispersed from the northern
German Wadden Sea by natural means (Reise et al. 2005).
The main centre of oyster cultivation in Europe is France, where about 150.000 t of C. gigas
are harvested annually. Pacific oysters have spread here in the course of warm summers
and extended their distribution from the culturing areas Arcachon and Marennes to the North.
The spread of C. gigas has not been investigated in France but research projects are now
started (Hily unpubl.).
Fig. 2: First step into the wild: Young oyster attached to a shell of Mya arenaria

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3 SPREAD IN THE WADDEN SEA
3.1 Temporal course
Pacific oysters have been brought on several occasions to the Wadden Sea and apart from
the only existing culture in the Lister Deep, various attempts have been made to cultivate this
species. Tab. 2 lists the documented occasions where Pacific oysters were brought into the
German Wadden Sea.
Tab. 2: Oyster cultures and imports into the German Wadden Sea. Information taken from Wehrmann
et al. (2000) and Neudecker (1984, 1985).
part of the Wadden Sea site year
Nordfriesland
Husum
Nordfriesland
Emmelsbüll
Amrum
Sylt
1880-1957
1930/ 1963
1982/1983
1982/1983
since 1986
culture of Colchester-oysters
import of French and Dutch oysters to
Nordfriesland
field trials for overwintering of spat and halfgrown
C gigas in tanks
field trials for overwintering of spat and halfgrown
C gigas in the harbour
successful culture of Crassostrea gigas
East Frisian
different places
Norddeich
Jadebusen
Neuharlingersiel
Jade
Wangerooge
Norderney
Wilhelmshaven
1911
1913/1914
1964
1974
1976/1982
1982
1987
1982/1983
introduction of Crassostrea virginea
introduction of Ostrea edulis, Crassostrea gigas
and Crassostrea angulata
introduction of Crassostrea angulata
experimental culture plots of Crassostrea gigas
experimental culture plots of Crassostrea gigas
experimental culture plots of Crassostrea gigas
experimental culture plots of Crassostrea gigas
field trials for overwintering of spat and halfgrown
C gigas upwelling ashore, in tanks and
reattachment frame
Cultivation at a larger scale started already in 1964 to the Oosterschelde in the Netherlands.
However, since oysters achieved good spatfall in Dutch waters during some warm summers,
imports were stopped after 1977. This was apparently not considered in a later German
approval of oyster cultures in the Wadden Sea. In 1983 Pacific Oysters were found near
Texel, brought there deliberately or accidentally (Bruins 1983, Dankers et al. 2004). Since
1986 Pacific oysters are in culture on the island of Sylt in the Lister deep (Fig. 3). Though
first wild specimens were found in 1991, it took more than a decade that Pacific oysters
spread all over the Wadden Sea and to establish dense aggregations. First records in 1998

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
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in the East Frisian Wadden Sea were reported by Wehrmann et al. (2000). Reise et al.
(2005) illustrate the development in the entire Wadden Sea (Fig. 4). It demonstrates that the
Pacific oyster most likely originates from two different sources which are the Oosterschelde
and the cultures in the Lister Deep near Sylt (Germany). Most notably since 2003 Pacific
oysters started to increase very strongly in density and biomass (see Fig. 24 and Fig. 25, see
also Dankers et al. 2004, Diederich 2005, Kristensen & Pihl 2006, Nehls & Büttger 2006) and
developed solid reefs (Reise et al. 2005). Overall, the spread of the Pacific oysters follows
the classic development of an alien species which may exist in an ecosystem over a long
period with low abundances before the invasive character becomes obvious. Changing
environmental conditions might support the spread of this species; the fact, that C. gigas
forms reefs and can overgrow existing blue mussel beds, leads to changes in the ecosystem
of the Wadden Sea with unpredictable further consequences.
The development of the C. gigas population in the Wadden Sea was apparently facilitated
from two sources: (1) The Pacific oyster cultures in the Wadden Sea itself (Lister Deep),
where an early and isolated development was documented and (2) a later inflow of Pacific
oysters from the southwest which was most probably enhanced by larvae drift with current
from the western parts of the Wadden Sea, which had most probably been reached by larvae
from the Oosterschelde (Wehrmann et al. 2006). To which extent other sources of
introduction (Tab. 2) have contributed to this development is not clear. The size of the
population in the Oosterschelde and of the culture in the Lister Deep indicate these as the
most important sources of Pacific oyster larvae facilitating the strong spread in the Wadden
Sea.
Fig. 3: Bags with Pacific oysters on the culture in the Lister Deep, Sylt (Germany).

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
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Fig. 4: Pacific oyster in the Wadden Sea. Asterisks indicate sites and years (boxed) of introduction
(Texel, Sylt). Other years indicate first records of settlement by larval dispersal for sites. Circles show
mean abundances in 2003 (from Reise et al. 2005).
3.2 Development of Pacific oyster density and biomass
By 2006 about 61,000 tons of Pacific oysters (live wet weight) were estimated to live on the
tidal flats of the entire Wadden Sea (Fig. 5). This value can be considered as a minimum
because not all scattered occurrences outside dense beds or reefs nor those on sluices,
seawalls and other artificial substrates could be included and little knowledge exists on
Pacific oyster abundance in the subtidal.
As Fig. 6 indicates, Pacific oysters are widely distributed in the Wadden Sea now and they
have overgrown many former mussel beds which have now been turned into dense oyster
reefs. High values are calculated for the western parts along the Eastfriesian islands and the
Dutch Wadden Sea, and again for the northern part north of the island of Sylt. Pacific oyster
population in the central parts of the Wadden Sea, e.g. in the area north and south of the
mouth of the river Elbe (between Jade and Eider) and also in some parts of the Wadden Sea

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
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of Nordfriesland are still comparatively low, but it is not clear, whether this situation will
remain stable.
Fig. 5: Total biomass of Pacific oysters in different parts of Wadden Sea 2006. Biomass is given as life
wet weight (Craeymeersch, Wehrmann & Markert, Kristensen pers. com.).
Fig. 6: Large Pacific oyster reef in the Lister Deep, Schleswig-Holstein.
#
#
#
#
#
643 t
3,289 t
13,838 t
22,747 t
20,510 t

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
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The Netherlands:
Pacific oysters were first recorded in the Dutch Wadden Sea in the early eighties as a result
from direct releases and/or drifted larvae from the Oosterschelde. C. gigas is spreading in
the Oosterschelde since the 1970ies which is the oldest population in the North Sea. It is
most likely that the growing population in the Oosterschelde is the most important source of
larvae facilitating the fast spread of this species in the Wadden Sea. Regarding the fast
development in the Wadden Sea, it is worth to consider that the population increase of C.
gigas in the Oosterschelde might not have reached a ceiling yet. 30 years after this species
has been reported to form wild populations, there is a steady increase in the area of Pacific
oyster reefs (Fig. 7).
Fig. 7: Development of intertidal Pacific oyster reef area in the Oosterschelde (Kater & Baars, 2003,
including RIVO-data, Dankers et al. 2006).
The intertidal area in the Oosterschelde covered by Pacific oysters was repeatedly mapped
using aerial pictures and reached 15 ha in 1980 and 210 ha in 1990. In 2002 640 ha were
registered and a biomass of 89,000 t living oysters was estimated, in 2003 809 ha. For 2005
the biomass was estimated at about 210,000 t living and dead oysters (Dankers et al. 2006,
Geurts van Kessel et al. 2003). In the Oosterschelde, Pacific oysters not only form dense
reefs in the intertidal but also colonize subtidal areas. The percentage of subtidal hard
substrates covered by Pacific oysters strongly increased during the past two decades (Fig.
8). Oyster biomass within the intertidal reefs usually ranges from 30 to 50 kg m-2.
0
200
400
600
800
1000
1200
19701980199020002010
oyster bed area [ha]

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
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Fig. 8: Pacific oyster coverage of subtidal hard substrates in the Oosterschelde (from Dankers et al.
2006).
In 1983, first Pacific oysters were found near the island of Texel, which is the first record for
the Dutch Wadden Sea, and oysters have been spreading since then. Since 2000, C. gigas
occurs in all parts of the Dutch Wadden Sea (Fey et al. 2007). In 2004 the spread of Pacific
oyster increased significantly and therefore oysters occur in many places in massive reefs
where the individuals adopt a vertical position (Fey et al. 2007). After the spread in 2004 the
mean densities on several places (transect of Oudeschild, oyster reef Zeeburg, mussel bed
near Ameland) seem to stabilize, though they still increased on a mussel bed near Ameland
in 2006 (Fey et al. 2007). In 2006 the biomass did not increase further but it is too early to
conclude on a possible stabilization of the development (Fig. 9).
The maximum density reported by Fey et al. (2007) in the Dutch Wadden Sea was found on
the oyster reef near Zeeburg (Texel) with more than 500 indiv./m² in 2003.
Fig. 9. Samples with oysters (n = 1293 samples with 0.4 m²each) and mean biomass in these samples
with standard deviation (from Fey et al. 2007).
Pacific oysters in Dutch Wadden Sea
0
20
40
60
80
100
20022003200420052006
number of samples (n)
0
2000
4000
6000
8000
10000
mean biomass (g)/m2
number of samples with oysters mean biomass/m2 in samples

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
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The area with a clear dominance of Pacific oysters in the Dutch Wadden Sea covered about
430 ha in 2005 and slightly decreased again in 2006 (Fig. 10). Areas with a mixed
community of Pacific oysters and blue mussels increased since 2004, which might be a
result from mussel spatfall. Scattered oysters were found in another 250 ha in 2005 and this
area increased in 2006 up to nearly 400 ha indicating that Pacific oysters start to populate
new areas.
Fig. 10: Development of the area colonised by Pacific oysters in the Dutch Wadden Sea (Goudswaard
et al. in prep.).
Lower Saxony
The immense spread of Pacific oysters is a recent process, mainly of the past five years.
Oysters can be found today on almost all mussel beds in Lower Saxony. In 2004 and 2005
Pacific oyster distribution followed a gradient in west-east direction with high abundances in
the west (Wehrmann et al. 2006), but numbers are increasing in the eastern parts as well
especially after a strong spatfall in 2006 (Markert pers. com.). Pacific oysters increasingly
dominate mussel beds in the Wadden Sea of Lower Saxony. Millat (2006) described some
mussel beds with still very few individuals in 2005 but mentioned that 2006 Pacific oysters
occur on all mussel beds (Fig. 11).
0
100
200
300
400
500
200420052006
Area with oysters dominating Mixed oysters and mussels Scattered oysters
ha

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
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Fig. 11: Proportion of mussel beds with occurrence of Pacific oysters in Lower Saxony (Millat pers.
com.).
Densities of Pacific oysters within the reefs in Lower Saxony cannot be directly compared
with the other surveys in Denmark, Schleswig-Holstein and Netherlands, as Wehrmann et al.
(2006) sampled densities on fixed points which might be situated in an unpopulated area or
where mussel beds vanished. In Lower Saxony maximum densities (highest densities of
single frames) reached 1460 indiv./m² in 2005 (Fig. 13).
Fig. 12: Pacific oysters in the harbour of the island of Langeoog, Lower Saxony (Markert 2006).
0
20
40
60
80
100
199819992000200120022003200420052006
percentage [%]

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
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Fig. 13: Development of Pacific oyster densities in the Wadden Sea of Lower Saxony between 2003
and 2005. Data from Wehrmann et al. (2006). The graphs show mean (above) and maximum
densities.
Wehrmann et al. (2006) also measured the biomass of Pacific oysters as wet weight (WW),
dry weight (DW) and ash free-dry-weight (AFDW) of the flesh. They sampled three beds in
2004 and 2005. The data document the increasing biomass stored in oyster reefs as a result
of individual growth and increasing abundances (Fig. 14).
Fig. 14: Average biomass (wet weight of flesh) of three Pacific oyster beds in Lower Saxony (Data
from Wehrmann et al. 2006).
0
50
100
150
200
250
300
350
Bank 1Bank 2Bank 3Bank 4Bank 5Bank 6Bank 7Bank 8Bank 9Bank 10Bank 11Bank 12Bank 13Bank 14Bank 15
averaged densities/ m²
2003 2004 2005
0
300
600
900
1200
1500
Bank 1Bank 2Bank 3Bank 4Bank 5Bank 6Bank 7Bank 8Bank 9Bank 10Bank 11Bank 12Bank 13Bank 14Bank 15
max. d ensity / m²
2003 2004 2005
0
200
400
600
800
1000
Bank 2Bank 3Banke 12
biomass [g wet weight/ m²/year]
2004 2005

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
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Schleswig-Holstein
In the Wadden Sea of Schleswig-Holstein the spread of the Pacific oyster took place mainly
in the Lister deep and on the tidal flats between the islands of Amrum and Föhr. In the Lister
deep recently nearly all mussel beds have been turned into Pacific oyster reefs (Fig. 16).
This process seems to be in the beginning now between Amrum and Föhr (Fig. 15). The
spread in the Lister deep is well documented since 1991 (see Reise 1998). First surveys (in
1999) resulted in average densities of about 4 indiv./m². In 2003 Pacific oysters reached
densities of > 100 indiv./m2. Because of a good spatfall in 2002, Pacific oyster densities
increased in 2003 and this process still continues. One mussel bed in the Lister Deep
revealed mean densities of 1000 indiv./m2 in 2006. Overall mean densities in the Lister Deep
were 722 (± 267) indiv./m² in 2006. Biomass in Pacific oyster reefs reaches 50 kg/m2 (live
wet weight). This is similar to the highest values in the Oosterschelde and may eventually
represent an upper level. Since 2005 Pacific oysters form massive reefs in the Lister deep.
Pacific oysters settle mainly on former mussel beds but since 2004 some larger areas with
empty shells from former mussel beds were colonized. In the area between Amrum and Föhr
Pacific oysters appeared since 1998 in low densities. In 2004 higher abundances were
recorded. In 2006 a strong spatfall was recorded in many areas of the Wadden Sea of
Schleswig-Holstein.
Fig. 15: Development of the average densities of Pacific oysters in two tidal basins in Schleswig-
Holstein. Lister Deep and Norderaue (between the island of Amrum and Föhr). Data from Diederich et
al. (2005) and Reise (1998).
Different tidal basins in Schleswig-Holstein show different spatial and temporal developments
(Fig. 15, Fig. 24). In the Lister Deep the spread occurred earlier and much stronger than in
other places. This might be caused by the Pacific oyster cultures present in this area and
potentially producing a large number of larvae; here, Pacific oysters are now dominating
most former mussel beds since 2004. In the other tidal basin, the Norderaue, Pacific oyster
distribution occurred at a later time and oysters so far do not cover large areas. This is
0
200
400
600
800
1000
1200
199519992003200420052006
averaged densities/m²
Lister Tief Norderaue

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
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remarkable, as Pacific oysters are present in this region at least since 1995 and in 1998 they
were found already on nearly all mussel beds, though in low densities (< 1 indiv./m2). The
strong spatfall in summer 2003 lead to high abundances in the following years and the
spatfall from 2006 is likely to further increase Pacific oyster numbers in this area.
Going south from these two tidal basins, Pacific oysters are still found in low densities in the
tidal basin south of Amrum and on most mussel beds around the islands of Pellworm,
Hooge, Gröde and Nordstrand, however, with densities lower than 1 indiv./m². Also, the
spatfall in late 2006 appeared to be much less than in the northern area (Fig. 17). The tidal
flats and also the mussel beds are muddier in this part of the Wadden Sea, which is a former
marsh and peat area, as compared to the sandy flats near the islands Sylt, Amrum and Föhr,
which are situated on glacial moraines. Further south, Pacific oysters are still rather rare on
the tidal flats between the Elbe and the Eider mouth, although they are abundant on sluices
and harbours in this area. On the only mussel bed in the Meldorf Bay Pacific oyster density
was still below 1 indiv./m² in 2006. Thus, about 95% of total Pacific oyster biomass in the
Wadden Sea of Schleswig-Holstein is still found in the Lister Deep (13,838 t), whereas oyster
biomass in the tidal basin Norderaue on the tidal flats near Amrum is rather low (643 t) and in
the rest of the area negligible. These differences in the development within the Wadden Sea
of Schleswig-Holstein give first hints about potential differences in habitat suitability,
however, it is too early to draw conclusions, and it cannot be stated yet, whether this reflects
a stable situation or just a delay in the development.
Fig. 16: Dense Pacific oyster reef in the Lister Deep, Schleswig-Holstein.

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 19
Fig. 17: Pacific oyster density (n/m2) on intertidal mussel beds in the Wadden Sea of Schleswig-
Holstein in 2006.
#
##
#
#
#
#
#
#
#
##
#
#
#
##
#
#
#
#
#
#
#
#
#
#
#
#
#
#
Pellworm
Nordstrand
Hooge
Langeness Gröde
061218Kilometer
#1 - 5
#6 - 50
#51 - 250
#251 - 500
#501 - 1000

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 20
Denmark
Pacific oysters are recorded in the Danish Wadden Sea since 1999, but until 2004 oysters
still occurred in low densities with only few individuals per km² (Kristensen pers. com.). In
2004 a strong spatfall of Pacific oysters on former mussel beds in the Danish part of the
Lister deep strongly enhanced the population growth. The biomass was estimated at 1000
tons. Kristensen & Pihl (2006) report biomass values (LWW) of Pacific oysters between 0
and 30.36 kg/m² in 2006. As a result of the growth of the Pacific oysters, biomass increased
to 3,289 tons in the Danish Wadden Sea in 2006 (Fig. 18, Fig. 19).
These biomass values are low in comparison to the other parts of the Wadden Sea but a
further increase is likely to occur.
Fig. 18: Development of Pacific oyster biomass in the Danish Wadden Sea. From Kristensen & Pihl
(2006).
0
500
1000
1500
2000
2500
3000
3500
1995199719992001200320052007
biomass [t]
?
3289

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 21
Fig. 19: Biomass (kg/m²) of blue mussels (Blåmuslinger; above) and Pacific oysters (Stillehavøsters) in
the Danish Wadden Sea in 2006. The question mark indicates an area with no inspections yet. From
Kristensen & Pihl 2006.

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 22
3.3 What facilitates the increase of Pacific oysters in the Wadden Sea ?
The spread of the Pacific oyster in the Wadden Sea follows the classic pattern of biological
invasions with a long phase of stagnancy followed by a fast increase. A small founder
generation has to reach a certain size before a fast growth is possible. However, in the case
of the Pacific oyster, it is likely that the recent spread is facilitated by changing environmental
conditions, especially an increase in summer temperatures (Nehls et al. 2006).
Before Pacific oyster cultivation was officially approved in the Wadden Sea it was argued that
they were not able to proliferate in the Wadden Sea because water temperatures are too low.
The lowest temperature allowing reproduction of C. gigas is about 20° C, a value, which is,
however, frequently reached in the shallow parts of the Wadden Sea in warm summers.
Consequently, already in the warm summers of 1976 and 1982 Pacific oysters revealed good
spatfall in the Dutch Wadden Sea (Drinkwaard 1999).
The increase of Pacific oyster populations in all parts of Wadden Sea since 2003 coincides
with water temperatures above average in summer. In the Lister Deep strong Pacific oyster
spatfall occurred in years with a positive deviation of water temperatures to the long-term
average (Diederich et al. 2005, see Fig. 20). Years of good spatfall are listed in Tab. 3.
Fig. 20: Deviation of mean monthly water temperature in July and August from the long-term mean
(1987 - 2003). Arrows mark years with high C. gigas recruitment (1991, 1994, 1997, 2001, 2002,
2003) (Diederich et al. 2005).
-3
-2
-1
0
1
2
3
4
Year
Temperature (°C)
July
August
'87 '88 '89 '90 '91 '92 '93 '94 '95 '96 '97 '98 '99 '00 '01 '02 '03

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 23
Tab. 3: Years of good spatfall in different regions of the Wadden Sea.
region years with good spatfall literature and comment
The Netherlands
(including
Oosterschelde)
1975, 1976,
1982, 1986, 1987, 1989,
1992
2003
Drinkwaard 1999
Dankers et al. 2006
Lower Saxony 1997 or 1998, 1999,
2004, 2006 Wehrmann et al. 2000 & 2006
Markert pers. com.
Schleswig-Holstein 1991, 1994, 1997,
2001, 2002, 2003, 2004, 2006
Tydeman 1999
Diederich et al. 2005
Nehls & Büttger 2006 & unpubl. obs.
Denmark
Oysters recorded north of Rømø since
1999 and years with spatfall are
assumed to be parallel to Schleswig-
Holstein
• In 2006 we observed a very strong spatfall in most many parts of the Wadden Sea of
Schleswig-Holstein which was confirmed for Lower Saxony too (Markert pers. com.).
More factors enhanced the spread of C. gigas. First, Pacific oysters are highly tolerant
against cold temperatures (Reise 1998, Diederich et al. 2005). Survival of oyster spat in the
first winter depends on temperatures while adult oysters seem not be much affected and
even survive cold winters when ice covers the tidal flats (Diederich 2006). In addition, so far,
no viral diseases are know to impact oyster populations (Reise et al. 2005) and predation
rate is apparently very low (Diederich et al. 2005, Diederich 2006), which facilitates a high
survival even in times, when many potential predators in the Wadden Sea are present.
Finally, they produce very large numbers of larvae and as Pacific oyster larvae prefer to
settle on adult oysters, aggregations grow rapidly and finally form dense reefs (see in
Diederich 2006).
3.4 Distribution over different habitats
Pacific Oysters settle on different hard substrates along the entire coast. They have been
found on blue mussel beds or on conspecifics, rock fills along the coast, sheet pile wall in
harbours, stones, shell beds or single shells and on sand (Dankers et al. 2004). Juvenile
oysters prefer to settle on adult conspecifics (see in Diederich 2006). In the early stage of the
development, Reise (1998) found 85% attached to Mytilus edulis (alive and empty shell) and
8% on other bivalves. In Niedersachsen, Pacific oysters settled primarily on blue mussels,
cockles and attached barnacles in the first years, but with increasing population densities,
Pacific oysters themselves became the most important substrate (Fig. 21).

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 24
Fig. 21: Settling substrates of Pacific oysters in the Wadden Sea of Niedersachsen (Wehrmann et al.
2006).
In general blue mussel beds are the most important location for the establishment of Pacific
oyster reefs so far. This regards not only live blue mussel beds. In Schleswig-Holstein Pacific
oysters also colonised former blue mussel beds which had vanished before the settlement of
Pacific oyster and the oyster larvae used shells of blue mussels and other bivalves as a
substrate (Fig. 22). All areas with a high Pacific oyster density in the Wadden Sea of
Schleswig-Holstein are found in existing or former blue mussel beds. Pacific oysters are also
found on other bivalve shells than mussels or oysters, however, in most places still in low
densities. Even on eroding mudflats with dense layers of dead shells Pacific oysters are
remarkably scarce, so the availability of shell material is apparently not the only factor
required for successful settlement.
Apart from settlement on tidal flats Pacific oysters today form dense layers on sluices,
harbour walls and all kind of solid installations in the Wadden Sea (Fig. 12).
Only limited knowledge exists on the subtidal distribution of Pacific oysters in the Wadden
Sea and they seemed to be restricted to the intertidal during the first years. Outside the
Wadden Sea in a part of the Oosterschelde subtidal occurrences have been surveyed in
2002 by ’side scan sonar’ and about 700 ha are considered by interpolation to be populated
by Pacific oysters (Geurts van Kessel et al. 2003). In the Dutch Wadden Sea subtidal
occurrences are reported by fishermen who fished clusters of Pacific oysters (Dankers et al.
2006). Near Texel subtidal Pacific oysters influence the water flow and in 2004 higher
abundances of fish between oyster beds were mentioned (Dankers et al. 2006).
2003
23%
6% 5% 2%
24%
40%
Mya areneria Littorina littorea
Cerastoderma edule Mytilus edulis
Balanidae Crassostrea gigas
2004
24%
25%
22%
20%
3%
6% 2005
15%
45%
12%
1%
15%
12%

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 25
In Lower Saxony occurrences in the subtidal have been reported by fishermen in southern
Randzel and in the Jade (Wehrmann pers. com.). Damm & Neudecker (2006) refer on
subtidal Pacific oysters near the island of Langeoog.
In Schleswig-Holstein subtidal occurrences are known from the Lister basin. Here, between
1992 and 1996 no subtidal Pacific oysters were found in dredge hauls (Reise 1998), in 1999
single individuals have been found and in 2004 428 individuals were fished in 10 hauls in the
middle of the basin (Diederich et al. 2005). Apart from the Lister basin, subtidal surveys
(dredge hauls) in 2005 and 2006 in different channels (but not in the List basin) in the
Wadden Sea of Schleswig-Holstein did not detect any subtidal Pacific oysters (Nehls &
Büttger 2007). Reise (pers. com.) searched the subtidal parts of the Lister Deep for Pacific
oysters in 2006 and reported a mean density of about 2 oysters per 100 m².
In Denmark subtidal Pacific oysters are reported in the Limfjorden (Kristensen pers. com).
Fig. 22: Scattered Pacific oysters on a former blue mussel bed in the Lister Deep, Schleswig-Holstein.

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 26
4 ECOLOGICAL CONSEQUENCES
4.1 Do Pacific oysters displace blue mussels?
As blue mussel bed area and blue mussel biomass strongly decreased in most places of the
Wadden Sea while Pacific oysters spread out, a burning question about the spread of the
Pacific oyster is whether or not they displace native blue mussels. This will be addressed by
comparing the development of blue mussels and oysters on different scales.
Nehls et al. (2006) and Wehrmann et al. (2006) both compared the densities of blue mussels
and Pacific oysters within existing blue mussel beds and found contrasting results. On a blue
mussel bed near Sylt a positive correlation of blue mussel and Pacific oyster density was
apparent, whereas on a blue mussel bed in Lower Saxony both species were negatively
correlated. The background of these differing results is not quite clear, however, it is possible
that the results from Sylt just reflect the fact that Pacific oysters preferably settled in the
denser parts of the blue mussel bed. Several studies pointed out, that the abundance of blue
mussels apparently remains on a stable level even in dense Pacific oyster reefs. In dense
Pacific oyster reefs near the island of Sylt, blue mussel density until now remained at a level
exceeding 1000 indiv./m2 (Fig. 23), however, densities have decreased and the biomass of
blue mussels in these beds is now much lower than it was before Pacific oysters were
present. This indicates, that Pacific oysters might be able to locally depress abundance and
biomass of blue mussels in dense reefs, however, it is not clear, whether this has an impact
on a larger scale.

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 27
Fig. 23: Mean densities (± standard deviation) of blue mussels (standard sampling method) and
Pacific oysters (counting frame) on two blue mussel beds in the Lister Deep in Schleswig-Holstein.
Comparing the development of blue mussel and Pacific oysters on the scale of whole blue
mussel beds indicates, that the decrease of the blue mussel and the increase of the Pacific
oyster did not occur at the same time and thus are likely to be independent processes. As
shown for two blue mussel beds in Schleswig-Holstein, blue mussels biomass and area
decreased long before the Pacific oysters reached significant densities which could have
affected the blue mussels; the decrease is even apparent on those beds, where oysters still
reach low densities (Fig. 24).
LT01
0
1000
2000
3000
4000
5000
6000
19992000200120022003200420052006
mean densities [n/m²]
Mytilus edulis Crassostrea gigas
LT13
0
1000
2000
3000
4000
5000
6000
19992000200120022003200420052006
mean densities [n/m²]
Mytilus edulis Crassostrea gigas

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 28
Fig. 24: Development of total biomass of Pacific oysters and blue mussels in Schleswig-Holstein on
two mussel beds in the tidal basins Lister Deep and Norderaue. Biomass given as life wet weight.
Fig. 25: Development of Pacific oyster and blue mussel biomass in the Wadden Sea regions between
1998 and 2006. Biomass given as life wet weight (Craeymeersch, Millat, Wehrmann, Markert and
Kristensen pers. com.).
Considering the four regions of the entire Wadden Sea reveals, that blue mussels decreased
in three regions – Lower Saxony, Schleswig-Holstein and Denmark – since at least 1998;
however, a notable increase of the Pacific oyster is only apparent since 2003 (Fig. 25). In all
Wadden Sea countries the total biomass of Pacific oysters is still much lower than the former
blue mussel biomass. There is no indication, that spreading Pacific oysters have caused the
NA12
0
200
400
600
800
1000
1200
1400
1600
1800
2000
199819992000200120022003200420052006
biomass [t]
0
2
4
6
8
10
12
14
bank area [ha]
blue mussel
oyster
bank area [ha]
LT01
0
200
400
600
800
1000
1200
1400
1600
1800
2000
199819992000200120022003200420052006
biomass [t]
0
2
4
6
8
10
12
14
bank area [ha]
blue mussel
oyster
bank area [ha]
0
5000
10000
15000
20000
25000
30000
35000
40000
45000
199819992000200120022003200420052006
biomass [t]
0
200
400
600
800
1000
1200
bank area [ha]
Mytilus edulis
Crassostrea gigas
bank area [ha]
0
20000
40000
60000
80000
100000
120000
1996199719992000200120022003200420052006
biomass [t]
0
500
1000
1500
2000
2500
3000
3500
bank area [ha]
Mytilus edulis
Crassotrea gigas
bank area [ha]
0
10000
20000
30000
40000
50000
60000
70000
199819992000200120022003200420052006
biomass [t]
0
200
400
600
800
1000
1200
bank area [h a]
Mytilus edulis
Crassostrea gigas
bank area [ha]
0
10000
20000
30000
40000
50000
60000
70000
80000
199819992000200120022003200420052006
biomass [t]
0
500
1000
1500
2000
2500
3000
3500
bank area [ha]
Mytilus edulis
Crassostrea gigas
bank area [ha]
The
Netherlands
Lower Saxon
y
Denmark
Schleswig
-
Holstein

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 29
decline of the blue mussels. Interestingly, in the Dutch Wadden Sea blue mussel bed area
and blue mussel biomass recovered from a low level and increased to a total area of 2600 ha
and a biomass of more than 70,000 t in 2004 and remained on a high level as Pacific oysters
spread out.
The contrasting developments of blue mussels and Pacific oysters as well as the different
development of blue mussels in the different regions of the Wadden Sea raises questions
about the causes of this development. Several recent studies highlight the significance of
predation on the recruitment success of bivalves in the Wadden as the most important factor
(e.g. Strasser & Günther 2001, Strasser 2002, Beukema and Dekker 2005). It is likely, that
annual variation in predation rate and recruitment success relates to winter temperatures:
cold winters result in low predation rates because the main predators of bivalve spat such as
shrimps and crabs occur later on the tidal flats; consequently, the bivalve s can successfully
establish new year classes. In turn, warm winters lead to high predation rates and low
recruitment success. These findings explained why recruitment of blue mussels to existing
beds was low and the establishment of new beds was rare during a long period of mild
winters since 1995/96.
Considering the Pacific oyster, it appears that predation plays a minor role in the recruitment
in the Wadden Sea. Thus, the combination of mild winters causing low recruitment rates of
blue mussels and warm summers causing high survival of Pacific Oysters seems to explain
well why blue mussel beds turned into oyster reefs (e.g. Nehls et al. 2006). These findings
also offer a potential perspective of the consequences of climate change in the Wadden Sea:
a warming Wadden Sea will be beneficial for Pacific oysters and lead to further decreases of
blue mussel beds. The development in the Netherlands, however, puts some caution to
these expectations and demonstrates that there is still a lack of knowledge about the ultimate
factors governing these population dynamics; with blue mussel beds increasing in the
Netherlands, questions arise especially about the recruitment process of blue mussels in the
Wadden Sea.
Concerning competition and coexistence of Pacific oysters and blue mussels it will be most
important whether or not blue mussels are able to recruit in greater numbers into oyster reefs
and either coexist or overgrow them in years of good spatfall. The recent development within
the Dutch Wadden Sea offers at present the best opportunities to study the interactions of
these two species.

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 30
4.2 Blue mussel and oyster beds and their associated species communities
The growing Pacific oyster population in the Wadden Sea experienced several strong
spatfalls in the past five years. New Pacific oyster reefs developed mainly on former blue
mussel beds. Blue mussel beds are known as centers of high diversity. Because of their
three-dimensional structure they offer a rich habitat for epibenthic as well as for endobenthic
species and they play an important role in marine food webs (e.g. for birds and other
predators). Several surveys assessed these different relations on blue mussel beds (Asmus
1987, Dittmann 1990, Hertzler 1995, Günther 1996). Now, as Pacific oyster reefs
increasingly replace blue mussel beds in the Wadden Sea, the question arises what happens
to the associated community (Broekhoeven 2005, Görlitz 2005 and Markert 2006).
Broekhoeven (2005) investigated the macrofauna along two transects covering different
Pacific oyster reefs and the adjacent mudflats in the Oosterschelde (NL). The aim was to
investigate the influence of the Pacific oysters on the macrofaunal diversity. In total, 38
species have been found and diversity and abundances were higher on Pacific oyster beds
than on the adjacent mudflats. Species numbers and abundances did not differ significantly
between both sampled beds. However, species composition did, but this might be related to
other factors which have not investigated (e.g. drift direction, elevation above low water line,
compare Saier 2001).
The two other investigations compared the faunal communities of blue mussel beds and
Pacific oyster reefs. Görlitz (2005) compared the associated community of blue-mussel beds
and Pacific oyster reefs in the Lister deep in the Wadden Sea of Schleswig-Holstein in 2004
and also considered beds covered by Fucus vesiculosus forma mytili. It is known that this
algae influences the abundances of different epibenthic species (Albrecht 1990, Albrecht &
Reise 1994). Fucus vesiculosus forma mytili cannot settle on Pacific oyster because this
algae depends on blue mussels which attach them with their byssus threads. Some
epibenthic species are supported by Fucus-cover, however, others occur in lower densities.
Görlitz (2005) found 49 species on Pacific oyster reefs. Species richness did not differ
significantly between blue mussel beds and Pacific oyster reefs, the community of oyster
reefs was more similar to the community of blue mussel beds than to the community of
Fucus-covered beds. Several species had different abundances and therefore it was
concluded that the dominance structure of the associated community will be different
between blue mussel and Pacific oyster beds, which might lead to changing functional
relationships in the ecosystem. Losses of species have not been documented yet. Cirripedia
showed similar densities on blue mussel and Pacific oyster beds, only densities of
Semibalanus balanoides were lower on oysters. Of Littorina littorea, Ralfsia verrucosa and
Polydora ciliata higher densities were registered upon Pacific oyster reefs, whereas only
juvenile crabs (Carcinus maenas) were found in lower densities. The fouling with macroalgae
and sessile invertebrates was higher on Pacific oyster beds. Analysis of sediments did not

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 31
yield significant differences of biodeposits between the different beds (percentage of
sediment fraction < 63µm taken as value for bio deposition).
Markert (2006) surveyed the macrofauna of a blue mussel bed in Lower Saxony near the
island of Juist. This study compared the associated community of Pacific oyster dominated,
blue mussel dominated and mixed beds as well as the adjacent unpopulated mudflats.
Species richness, abundances and biomass were higher on Pacific oyster dominated beds.
45 taxa were found in the Pacific oyster dominated beds. All species found on Pacific oyster
beds have been found on blue mussel beds before (former surveys) and no new species
were detected. Similar to Görlitz (2005), Markert (2005) found higher abundances of
epibenthic species (mobile species of the epibenthic) on Pacific oyster beds. This might be
related to the somewhat larger potential space of the three dimensional structure offered by
oysters positioned in a vertical position. Markert (2005) found a shift in species composition,
from deposit-feeders on blue mussel beds to opportunistic filter-feeders on Pacific oyster
beds However, Markert (2005) assumes impoverishment of endobenthic species below
Pacific oyster layers.
As part of the blue mussel monitoring in the Wadden Sea National Park of Schleswig-
Holstein the associated macrofauna of blue mussel beds has been monitored since 1999
(Nehls & Büttger 2006). The macrobenthic communities of two beds in the Lister Deep near
the island of Sylt have been studied from 1999 to 2005, thus during the period that these
beds turned into in Pacific oyster reef.
Referring to the species list which takes different determination levels into account, between
26 and 48 taxa were found during seven years of monitoring. At all 53 taxa have been found
on both beds (unconsidered steadiness and abundances, only presence/absence-data).
Several taxa occurred just in one year with low abundances, and might represent single
events. For example the species Pagurus bernhardus was observed in several years on
LT01 but never on LT13. The bed LT01 is located close to the low water line adjacent to a
deep subtidal area from where individuals could immigrate. On the other hand Ligia oceanica
was found in some years only on LT13, but always with few individuals only.

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 32
Fig. 26: Number of taxa on two beds between 1999 and 2005 (because of different determination level
some taxa were combined on a higher taxonomic level for comparability).
The number of taxa does not give much information about the community structure and
similarity of the species composition and their abundances between the different years. To
illustrate the relations a non-metric MDS-plot is used (Clark & Warwick 2001). The MDS-Plot
bases on the Bray-Curtis similarity matrix which determines the similarity between each year
to each other. Years which are more similar to each are plotted closer to each other. Cluster
analysis was used to group years which are more similar to each other and clusters formed
at three arbitrary levels are superimposed on the MDS-Plots. Community structure of both
beds was similar with 65% over all years, considering species composition and mean
abundances. The changes in community structure proceeded almost analogue at both beds,
but began before Pacific oysters started to dominate (see Fig. 24 and Fig. 25.
Fig. 27: MDS-Plots (Bray-Curtis-Similarity, fourth root transformation), comparing community
structures of two localities during 1999-2005. Left: considering average densities on bed LT01; right:
considering average densities on bed LT13; Oligochaeta undet. are ignored (see text).
0
10
20
30
40
50
60
1999200020012002200320042005
number of taxa
LT01 (total 64) LT13 (total 61)

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 33
These changes of community structure can be explained with increasing densities and
biomass of the macrozoobenthos. On both beds biomass of macrozoobenthos and the total
number of individuals increased during the study period (Fig. 28) and both beds showed the
same development. Maximum values of biomass were achieved in 2002 and 2003, but
decreased thereafter. Total number of individuals was highest in 2003.
Within the macrozoobentos community, changes were dominated by barnacles. However,
abundances of polychaets, Anthrozoa undet. and Bivalvia increased too, especially from
2003-2005 on both beds (Fig. 29). Abundances of Gastropoda increased, representing
mainly Littorina littorea and Crepidula fornicata, which occurred in higher densities.
Fig. 28: Biomass and number of macrobenthic species on the beds LT01 and LT13 in the Lister Deep
(Oligochaeta undet. not considered; given are average numbers of individuals and standard
deviation).
0
50
100
150
200
250
300
1999200020012002200320042005
AFDW [g/m²]
LT01 LT13
0
10000
20000
30000
40000
50000
60000
70000
1999200020012002200320042005
averaged number of individuals/m²
LT01 LT13

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 34
Fig. 29: Mean denisties of selected classes on the bed LT01 between 1999 and 2005.
Abundances of Oligochaets (only > 1mm due to sieve size) increased during the monitoring
period (Fig. 29). Markert (2006) found highest densities of Oligochaets in oyster beds while
Görlitz (2005) did not find differences between blue mussel and Pacific oyster beds.
These changes form a complex picture probably influenced by three interdependent
processes. First the ageing of the blue mussel beds due to missing spatfall offer a more
diverse physical structure because of dead shells (Tsuchiya & Nishihira 1986) which
facilitates the associated epibenthic community. Secondly on both beds the Fucus-coverage
decreased and it could be assumed that barnacles and some other epibenthic species have
been promoted. Thirdly the increase in three-dimensional structure due to the Pacific oysters
(initially with low densities and since 2003 increasing) enhanced in particular epibenthic
species.
De Kluijver & Dubbeldam (2003) investigated the influence of Pacific oyster densities on the
diversity of subtidal beds in the Oosterschelde (NL). They state that oyster densities up to
50% coverage lead to higher diversity but higher densities caused decreasing diversity. This
aspect might become important also in the Wadden Sea, since the short time period of the
Pacific oyster spread still allows for further developments and increasing impact of this
indigenous species.
In summary, some general statements can be made:
• No species losses were observed yet.
• Blue mussels are able to coexist with Pacific oysters in their reefs.
• Pacific oyster reefs offer species of blue mussel beds an alternative habitat.
0
5000
10000
15000
20000
25000
OligochaetaPolychaetaBalanus sp.GastropodaBivalviaPolyplacophora
AnnelidaArthropodaMolluscaNemertea
denisties [n/m²]
1999 2000 2001 2002 2003 2004 2005

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
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• Changes are mainly caused by increasing abundances of the epibenthic species, but
also abundances of endobenthic species increased.
• Species dominance changed.
• The increase of Pacific oyster densities, biomass and coverage is still an ongoing
process; single surveys and a long-term monitoring is necessary to observe,
document and further analyse and assess this process of changing mussel
communities and structures as well as the changes in the development of the
associated community (species, densities, biomass).
4.3 Introduction of other species associated with oysters
Many new species have been introduced in Europe along with the oyster imports but most
did not establish stable populations (Wolff & Reise 2002). Considering the criteria that a)
species occur in Japan or on the pacific coast of North America, b) their occurrence
coincides with the period of oyster imports, Wolff & Reise (2002) present a list of species
which could have potentially been introduced with oysters (Tab. 4)
Tab. 4: Species which might have been introduced with oysters (Wolff & Reise 2002).
scientific name
Gymnodinium miki
Alexandrium leeii
Fibrocapsa japonica
Chattonella sp.
Thalassiosira punctigera
Coscinodiscus wailesii
Sargassum muticum
Undaria pinnatiiida
Grateloupia doryphora
Dasysiphonia sp.
Anotrichium furcell
Polysiphonia senticulosa
Pileolaria berkeleyana
Hydroides ezoensis
Ammothea hilgendorfi
Reise (1998) mentions species which were introduced (Aplidium nordmanni, Verruca
stroemia, Ascophyllum nodosum) or re-introduced (Pomatoceros triqueter) with oyster
imports to Wadden Sea around the island of Sylt (Germany). Two species, Sargassum

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 36
muticum and Styela clava, are established now but might be imported on other paths than
oyster imports.
Nehring & Leuchs (1999) give a list of species which have been introduced and background
information how they reached the German coast. For the Pacific Oyster they mention five
species, which have been introduced with oyster imports:
Anthrozoa: Diadumene cincta: introduced by oyster imports or as fouling on ships,
distributed only around Helgoland, common.
Haliplanella luciae introduced as fouling on ships, mussel imports as vector
seem to be possible. No recent population along the German coast.
Mollusca: Crepidula fornicata: introduced with mussel imports to GB, further distribution
of larvae by current to the Netherlands and Belgian. Import to Lower Saxony
and Schleswig-Holstein with Ostrea edulis.
Petricola pholadiformes: introduced in GB by oyster import, eastward dispersal
by larvae with current and/or imports of Ostrea edulis
Tunicata: Aplidium nordmanni: first records 1992 on oyster cultures near the island of
Sylt, probably introduced with oyster imports for the culture. Only around Sylt
and rare.
Recently Dekker found the alien species Hemigrapsus penicillatus on oysters near Texel
(Cadée 2007).
Oysters also brought some of their parasites (Mytilicola orientalis and Myicola ostreae) with
them to France and the Netherlands. These parasites were found in other species of shellfish
too but seem to be harmless to them (Wolff & Reise 2002 therein cited His 1977 and Stock
1993). Wolff & Reise (2005) mentioned no strong evidence about introduced shellfish
diseases with oyster imports.
4.4 Consequences for energy flow and top predators (birds)
Bivalve suspension feeders are very important members in intertidal food webs (e.g. Asmus
et al. 1998). In the Wadden Sea, blue mussel beds represent the most productive benthic
communities; even though only about 1% of the tidal flats are covered with blue mussel
beds, they reach a significant proportion of the total energy flow due to their intensive
production per area. In the Lister Deep, blue mussel bed annual production reaches 762 g C
m-2 and thus is 28 times higher than the average of the other communities (Asmus et al.
1998). An important aspect of the high production of blue mussel beds is, that a high share of
this production is available for higher trophic levels especially birds. On blue mussel beds in
the Lister Deep, bird predation by eider duck Somateria mollissima, oystercatcher
Haematopus ostralegus and herring gull Larus argentatus almost matches the annual
production of the blue mussel bed (Nehls et al. 1997). As Pacific oysters are not eaten by
eiders and only rarely by other birds, production and biomass of this species is hardly
available for higher trophic levels and the energy flow will be redirected into the storage
compartment (cf. Asmus 1987) and microbial decomposition.

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
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To which extent this has an effect on the bird population in the Wadden Sea is not clear and
needs further investigation. For mussel eating birds, the change of blue mussel beds to
Pacific oyster reefs is certainly not an advantage. While oystercatchers and herring gulls may
adapt to the new species and learn how to handle at least small individuals of this new prey,
this will certainly not be the case with eider ducks which swallow their prey with shells.
In general, the ecological consequences of the spread of the Pacific oyster and the change
of the blue mussel bed community is far from clear and urgently needs further investigations
because it poses a variety of changes to the entire ecosystem. At present it is not clear, how
the change in the blue mussel bed community will affect biomasses, productivities and
filtration rates and what may be the impacts on the surrounding tidal flats.
5 OYSTER FISHERIES
Although cultured Pacific oysters are sold at higher prices as most other shellfish, wild
oysters have attracted only very limited action of the fisheries. As wild Pacific oysters are
often attached to big clumps, exhibit a variety of forms, are overgrown with barnacles and
other epifauna and reach sizes which would overlap a plate, they do not fulfil the standards
of high-priced food. Consequently, Pacific oysters are collected by some local fishermen and
other people walking on the tidal flats of the Wadden Sea but there is apparently limited
interest in harvesting the new species at a larger scale. In the Oosterschelde large amounts
of Pacific oysters were removed from some sites, but as there was no demand for further use
of the oysters, they were just dumped at some other locations (Baptist 2005, Wijsman et al.
2006). At present, in no part of the Wadden Sea Pacific oysters are commercially fished and
at present there seems to be no interest of the fisheries to do so in the near future.
In Schleswig-Holstein, the company running the culture in the Lister Deep has been licensed
to collect seed oysters from the Wadden Sea in 2005. However, as only single-grown Pacific
oysters of a limited size are useful for the culture, only a small fraction of the wild oyster
population is suitable for their purposes and the demand to collect seed oysters appears to
be limited.
Though the interest of the fisheries in Pacific oysters is apparently low, the example from
Schleswig-Holstein shows, that some fishing activity may still develop and on the other hand,
fisherman may be interested to remove oysters from mussel beds in order to improve the
mussel fisheries, as has been done in the Netherlands (see below).

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
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6 MANAGEMENT
The spread of the Pacific oyster has to be regarded as the consequence of a careless
introduction of an alien species to a sensitive and protected ecosystem. However, no options
are available to prevent the species from further spread. It is important to note, that the
reproductive capacity of the Pacific oyster is too high and that the oyster population has
grown far beyond a point, that it could be removed from the Wadden Sea again. The Pacific
oyster obviously has established a stable population not only in the Wadden Sea but also in
many other European coastal waters and as other invaders, like the Soft-shelled clam Mya
arenaria and the American razor clam Ensis americanus, they have to be regarded as
permanent members of our coastal ecosystems.
In the Oosterschelde some measures have been taken to locally remove oyster reefs
(Wijsman et al. 2006). They showed that Pacific oysters could locally be removed efficiently
even in dense reefs, however, the effort required to do so was considerable (20 days per ha)
and due to a lack of commercial interest in the Pacific oysters, these were just dumped in
deep gullies. The effect of these measures on the ecosystem is not clear yet and subject of
an ongoing study.
Blue mussel fisheries in the Wadden may be affected by the spread of the Pacific oyster in
the future from two reasons. First, Pacific oysters may settle on culture lots and overgrow the
blue mussels. At present, it seems to be unlikely that this will be a major problem for the
fisheries, as Pacific oysters apparently rarely settle on young blue mussels and in general do
not settle in high densities in the subtidal. As blue mussel cultures are stocked with young
seed mussels and are located always in the subtidal it seems at present to be unlikely, that
they might be overgrown by Pacific oysters. Second, oyster may be present on seed mussel
beds and make it impossible to fish purely for blue mussels. This might be the case, if blue
mussel spatfall occurs in Pacific oyster reefs, but it is not clear yet to which extent this may
happen. As long as blue mussel seed is fished in the subtidal this will be of no problem for
the fisheries, because of low Pacific oyster densities in these places. In the intertidal where
Pacific oysters build up dense reefs, they may in fact make these areas unattractive for blue
mussel fisheries and offer some protection for intertidal habitats, however, at present it is not
clear to which extent this may be a problem for the fisheries. From an ecological point of view
any intentions to remove Pacific oysters in order to promote blue mussel fisheries must be
regarded with caution, as this may well lead to more damage then benefit. Any measure
should clearly comply with the guiding principle of the Wadden Sea, which is to achieve, as
far as possible, a natural and sustainable ecosystem in which natural processes proceed in
an undisturbed way. It is recommended that any steps should be based on a thorough
impact assessment according to the habitat directive before approval.

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
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7 MONITORING/ASSESSMENT
The spread of the Pacific oyster poses major changes to the Wadden Sea and there is an
urgent need to monitor the further development also to fulfil the monitoring requirements from
the habitat directive and the water framework directive. Once Pacific oysters had been able
to establish dense populations in the Wadden Sea, they have started to form dense reefs
which harbour a diverse benthic community similar to mussel beds. It must be noted,
however, that these reefs, initiated by an alien species, are situated within a protected area,
the Wadden Sea. Although they form a diverse habitat by itself which might be of some
added value for the resident species ecological as well as formal aspects have to be taken
into account and it has to be assessed in the near future, whether oyster reefs fulfil the
definition of the habitat type 1170 reef of the habitat directive or not.
It is recommended to extend and update the TMAP manual on monitoring (TMAG/CWSS
1997) with respect to the spread of this important new species. Regarding the fast
development during the last years, it is strongly recommended that the development of the
Pacific oyster becomes subject of a detailed monitoring program covering the following
aspects in a yearly rhythm:
• Extension of the area covered by Pacific oysters
• In selected areas: density, size structure and biomass of both, blue mussels and
Pacific oysters, respectively
• Annual recruitment of blue mussels and Pacific oysters
These data will allow to follow the most important aspects of the general development but
more detailed investigations on filtration rate, production and energy flow as well as
investigations into the changes of the associated benthic community are highly
recommended.
Biomass of Pacific oysters should be measured as cooked flesh weight. Live wet weight,
which is often used in the monitoring of blue mussels is useful, however, data from both
species cannot be compared directly as the flesh content of oysters is much lower than that
of blue mussels.
The monitoring of Pacific oyster beds can in many aspects follow the standards of the blue
mussel monitoring implemented in the Wadden Sea (TMAG/CWSS 1997), but it has to be
stressed, that the current monitoring activities on blue mussels will not allow to monitor this
fast development adequately. Like the blue mussel beds, oyster reefs can easily be identified
from aerial photos, however, both communities cannot be distinguished in most cases, thus
there is an added need for ground-based investigations. Similar to blue mussel beds, oyster
reefs can be mapped at the tidal flats with a GPS. Sampling of oyster reefs needs somewhat
larger frames as used for blue mussels because the density is lower. In the monitoring of
blue mussel beds in the Wadden Sea National Park of Schleswig-Holstein a 25 x 25 cm
frame is now used to sample both species and it is recommended to use it as a standard for

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 40
further monitoring. Samples may both be taken either randomly over the whole bed or in the
covered heaps only if coverage of the bed is assessed in the same time.
Fig. 30: Oyster reefs near Sylt – a familiar view in many parts of the Wadden Sea.

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
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8 SUMMARY
For the first time a comprehensive data compilation of the entire Wadden Sea was carried
out in the framework of the TMAP to document the spread of the Pacific Oyster Crassostrea
gigas with special emphasis on the recent strong increase in all parts of the Wadden Sea.
This study describes the successful spread of an alien species following a careless release
which is now leading to remarkable changes in the ecological structure of the Wadden Sea.
Oysters have been brought into the Wadden Sea to stock cultures for more than 30 years
and a permanent culture was licensed near the island of Sylt, Schleswig-Holstein in 1986.
Pacific oysters are today found in all parts of the Wadden Sea. They form dense layers which
have all characteristics of reefs on former beds of the blue mussel Mytilus edulis and settle
on all other kind of hard substrates. Total biomass in the Wadden Sea is calculated at 61,000
live wet weight (LWW) in 2006. Regional figures are as follows: The Netherlands: 20,510 t,
Lower Saxony: 22,747 t, Schleswig-Holstein: 14,481 t, Denmark: 3,289 t. In Schleswig-
Holstein, about 95% of the oysters biomass (LWW) are found in the Lister Deep near the
island of Sylt. Distribution and temporal course of the spread of the Pacific oysters indicate
two main pathways: The oyster culture in the Lister Deep and an inflow of larvae from areas
south of the Wadden Sea, most probably from the Oosterschelde where Pacific oysters are
cultured and spreading since the 1970ies. Within dense reefs oysters reach abundances of
about 1000 indiv./m2 and reach a biomass of up to 50 kg/m2 live wet weight. The spread of
the Pacific oyster is apparently highly facilitated by warm summer temperatures and years
with good spatfall are characterised by water temperatures above average. Climate change –
as it will lead to higher temperatures - will thus further promote the development of this
introduced species.
A comparison of the development of Pacific oyster and of blue mussel provided no indication,
that oysters are the cause of declining mussel stocks. In three regions – Lower Saxony,
Schleswig-Holstein and Denmark – mussel bed area and biomass showed a long-term
decrease which started before oysters were spreading and in The Netherlands, mussel beds
markedly recovered to high values in the same period as oysters spread out. Although
competition of the two species on a local scale cannot be ruled out, the decline of mussel
beds on a larger scale cannot be explained by the spread of the oyster.
Studies of the macrofauna community structure of oyster beds revealed a diverse community
which is in many aspects similar to that associated with of blue mussel beds. So far no
species losses have been detected and oyster reefs seem to serve as a new habitat for all
indigenous species of mussel beds. However, comparison of results indicated differences in
dominance structure and in endobenthic functional groups composition between blue mussel
beds and Pacific oyster beds. Therefore, further spread of oysters might lead to functional

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
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changes in the Wadden Sea and to changes in the whole associated community. However,
this is an ongoing process and further development cannot be predicted.
The consequences of the spread of the oysters for the Wadden Sea ecosystem cannot be
overseen yet and a further rapid extension of the population is expected. Unlike blue mussels
oyster are only consumed by a few species and suffer little predation. Energy flow of an
oyster reef is thus anticipated to be highly different from mussel beds and not directed to
higher trophic levels. Oyster reefs are apparently of little value for mussel eating birds and
especially eider ducks Somateria mollissima cannot make use of the oysters.
Consequences for mussel fisheries are expected to be rather low at present, as the
distribution of Pacific oysters is still mainly restricted to the intertidal parts and there is no
indication yet, that oysters have an impact on mussel spatfall.
No options exist for management to reduce or even stop the spread of the Pacific oyster. The
Pacific oyster obviously has established a stable population not only in the Wadden Sea but
also in a number of other European coastal waters and as other invaders, like the Soft-
shelled clam Mya arenaria and the American razor clam Ensis americanus, the species has
to be regarded a permanent members of our coastal ecosystems. Any intentions to remove
Pacific oysters for whatever reason should be assessed carefully, as this would be a
considerable impact with yet unforeseen consequences and could potentially lead to more
damage then benefit. Any measure should clearly comply with the guiding principle of the
Wadden Sea, which is to achieve, as far as possible, a natural and sustainable ecosystem in
which natural processes proceed in an undisturbed way. It is recommended that any steps
should be based on a thorough impact assessment according to the habitat directive before
approval.
The further spread of the Pacific oyster and possible consequences to the Wadden Sea
ecosystem should be subject of detailed research and monitoring. Monitoring standards
should be harmonized within the entire Wadden Sea and comply with the objectives of the
habitat directive as well as of the water framework directive.

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
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9 LITERATUR
Albrecht, A. 1990. Einfluß der Braunalge Fucus vesiculosus auf die Lebensgemeinschaft von
Miesmuschelbänken im Wattenmeer. Diplomarbeit, Universität Göttingen.
Albrecht, A. & K. Reise 1994. Effects of Fucus vesiculosus covering intertidal mussel beds in
the Wadden Sea. Helgol. Meeresunters. 48: 243-256.
Asmus, H. 1987. Secondary production of an intertidal mussel bed community related to its
storage and turnover compartments. Mar. Ecol. Prog. Ser. 39: 251-266.
Asmus, H., D. Lackschewitz, R. Asmus, G. Scheiffarth & G. Nehls. 1998. Carbon Flow in the
Food Web of Tidal Flats in the Sylt-Römö Bight. In: Gätje, C. & K. Reise (Eds.): The
Wadden Sea Ecosystem. Exchange, Transport and Transformation Processes.
Springer.
Baptist, H. 2005. Habitattoets proef weghalen oesterbanken in de Oosterschelde. Unpubl.
Report, Ecologisch Adviesbureau Henk Baptist
Beukema, J. J. & R. Dekker 2005: Decline of recruitment success in cockles and other
bivalves in the Wadden Sea: possible role of climate change, predation on postlarvae
and fisheries. Mar. Ecol. Prog. Ser. 287: 149–167.
Broekhoeven, W. v. 2005. Macrofaunal diversity on beds of the Pacific oyster (Crassostrea
gigas) in the Oosterschelde estuary. RIVO-report number 05.002.
Bruins, R. W. B. 1983. Crassostrea gigas (Thunberg, 1973) op Texel. Corresp.-blad Ned.
Mal. Ver. 215: 1436-1438.
Cadée, G. C. 2007. Vervabgeb de recente Japanse oesterriffen de vroegere oesterbanken?
De Levende Natuur 108: 64-67..
Clark, K. R. & R. M. Warwick 2001. Change in marine communities: an approach to
statistical analysis and interpretation, 2nd edition. Primer-E, Plymouth.
Damm, U. & T. Neudecker 2006. Zweikampf im Wattenmeer – Wird die heimische
Miesmuschel verdrängt? Inf. Fischereiforsch. 53:46-48.
Dankers, N, E Dijkman, M de Jong, G de Koort & A Meijboom 2004: De verspreiding en
uitbreiding van de Japanse Oester in de Nederlands Waddenzee. Alterra-rapport 909,
Texel.
Dankers, N., A. Meijboom, M. de Jong, E. Dijkman, J. Cremer, F. Fey, A. Smaal, J,
Craermeersch, E. Brummelhuis, J. Steenbergen & D. Baars 2006. De ontwikkeling van
de Japanse oester in Nederland (Waddenzee en Oosterschelde). Wageningen
IMARES Rapport nr. C040/06
Diederich S. 2005. Invasion of Pacific oysters (Crassostrea gigas) in the Wadden Sea:
competitive advantage over native mussels. PhD thesis, University of Kiel.

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 44
Diederich, S. 2006. High survival and growth rates of introduced Pacific oysters may cause
restrictions on habitat use by native mussels in the Wadden Sea. J Exp mar Biol 328:
211-227, doi:10.1016/j.jembe.2005.07.012.
Diederich, S., Nehls, G., Beusekom, J. E. E. van, Reise, K. 2005. Introduced Pacific oysters
(Crassostrea gigas) in the northern Wadden Sea: Invasion accelerated by warm
summers?, Helgol Mar Res 59: 97-106, doi:10.1007/s10152-004-0195-1.
Dittmann, S. 1990. Mussel beds – amensalism or amelioration for intertidal fauna? Helgol.
Meeresunters. 44: 335-352.
Drinkwaard, A. C. 1999. Introductions and developments of oysters in the North Sea area: a
review. Helgol. Meeresunters. 52: 301-308.
Fey, F., N. Dankers, A. Meijboom, M. de Jong, P.-W. van Leeuwen, E. Dijkman & J. Cremer
2007. De ontwikkleing van der Japanse oester in de Nederlandse Waddenzee: Situatie
2006. Wageningen IMARES Interne Rapportage Nummer: 07.003.
Geurts van Kessel, A. J. M., B. J. Kater & T.C. Prins 2003. Verandernde draagtracht van de
Oosterschelde voor kokkels. RIVO rapport C062/03.
Görlitz, S. 2005. Neue Riffe im Wattenmeer: Die Pazifische Auster Crassostrea gigas und
ihre assoziierte Lebensgemeinschaft. Diplomarbeit, Universität Kiel.
Goudswaard, P. C., J. Steenbergen, J. M. D. D. Baars, J. J. Kesteloo, J. Jol & M. R. van
Stralen 2006. Het mosselbestand en het areaal aan mosselbanken op de
droogvallende platen in de Waddenzee in het voorjaar van 2006. Wageningen Rapport
nr. C059/06.
Goudswaard, P. C, J. J. Kesteloo & M. R. van Stralen 2007. Het bestand aan Japanse
Oesters in het litoraal van Nederlands deel van de Waddenzee in 2004 – 2006.
Unpubl. Report, IMARES.
Günther, C. P. 1996. Development of small Mytilus beds and its effects on resident intertidal
macrofauna. Marine ecology publicazioni della stazione zoologica di napoli I 17: 117-
130
Hertzler, I. 1995. Nahrungsökologische Bedeutung von Miesmuschelbänken für Vögel (Laro-
Limikolen) im Nordfriesischen Wattenmeer. Diplomarbeit, Universität Göttingen.
Kater, B. J. 2003. Ecologisch profiel van de Japanse oester. Rivo rapport nr. C032/03
Kater, B. J. & J. M. D. D Baars 2003. Reconstructie van oppervlakten van litorale japanse
oesterbanken in de Oosterschelde in het verleden en een schatting van het huidig
oppervlak. RIVO rapport C017/03.
Kluijver, M. de & M. Dubbeldam 2003. De sublitorale hard-substraat levensgemeenschappen
in de Oosterschelde – Evaluatie van de ontwikkeling in de periode 1985-2002. In
opdracht van: Rijksintituut voor Kust en Zee. Rapportnummer: 1973-2.
Kristensen, P.S. 1989. Mussel and oyster culture in Denmark. In: N. De Pauw, E. Jaspers, H.
Ackerfors, N. Wilkins (eds.) Aquaculture - A biotechnology in progress. European
Aquaculture Society, Bredene, Belgium: 342-350.

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 45
Kristensen, P. S. & N. J. Pihl 2006. Blåmuslinge- og stillehavsøstersbestanden i det danske
Vadehav efteåret 2006. DFU-rapport nr. 167-06
Markert, A. 2006. Untersuchungen der Lebensgemeinschaft eines durch die Pazifische
Auster (Crassostrea gigas) neu entstandenen Austernriffs im Ostfriesischen
Wattenmeer. Diplomarbeit, Freie Universität Berlin.
Meixner, R. & Ch. Gerdener 1976. Untersuchungen über die Möglichkeiten einer
Fortpflanzung der eingeführten Pazifischen Auster Crassostrea gigas in deutschen
Küstengewässern. Arch. Fisch. Wiss. 26: 155-160
Meyer-Waarden, P.F. 1964. Mastversuche mit portugiesischen Austern im deutschen
Wattenmeer. Inf. Fischwirtschaft 2: 77-78
Nehls, G., I. Hertzler & G. Scheiffarth 1997: Stable mussel beds in the Wadden Sea - they
are just for the birds. Helgol. Meeresunters. 51: 361-372
Nehls, G., S. Diederich, D. Thieltges & M. Strasser 2005: Perishing blue mussels and
invading aliens – What are the reasons for ecological turnover in the Wadden Sea ?
Wadden Sea Newsletter 2005-1: 17-20
Nehls, G. & H. Büttger 2006: Miesmuschelmonitoring im Nationalpark Schleswig-
Holsteinisches Wattenmeer 1998 – 2005. Husum.
Nehls, G. & H. Büttger 2007: Miesmuschelmonitoring im Nationalpark Schleswig-
Holsteinisches Wattenmeer 2006. in prep., Husum.
Nehls, G., S. Diederich, D. Thieltges & M. Strasser 2006: Wadden Sea mussel beds invaded
by oysters and slipper limpets: competition or climate control? Helgol Mar Res 2006 60:
135–143, doi:10.1007/s10152-006-0032-9.2005
Nehring, S. 1999. Oyster beds and Sabellaria reefs. – In: De Jong, F., J. F. Bakker, C. J. M.
van Berkel, N.M.J.A. Dankers, K. Dahl, C. Gätje, H. Marencic & P. Potel (Hrsg.),
Wadden Sea Quality Status Report. Common Wadden Sea Secretariat
Wilhelmshaven, Wadden Sea Ecosystem No. 9: 146-147.
Nehring, S. 2006. NOBANIS - Invasive Alien Species Fact Sheet - Crassostrea gigas. –
Online Database of the North European and Baltic Network on Invasive Alien Species -
NOBANIS www.nobanis.org: 13 S.
Nehring, S. & H. Leuchs 1999. Neozoa (Makrozoobenthos) an der deutschen Nordseeküste
– Eine Übersicht. Bundesamt für Gewässerkunde, Koblenz, Bereicht-BfG-1200: 131S.
Neudecker, T. 1992. The history of the former German oyster fishery and mariculture: 400
years of crown law on oysters (4.2.1587). - Dt. hydrogr. Z., Erg.-H. B, 22: 518-525.
Reise, K. 1985: Tidal flat ecology. Springer Verlag.
Reise K 1998: Pacific oysters invade mussel beds in the European Wadden Sea. Senckenb
Marit 28:167–175

Nehls & Büttger: Spread of the Pacific Oyster Crassostrea gigas in the Wadden Sea
page 46
Reise, K., E. Herre & M. Sturm 1989. Historical changes in the benthos of the Wadden Sea
around the island of Sylt in the North Sea. Helgol. Meeresunters. 43: 417-433
Reise K., N. Dankers & K. Essink 2005: Introduced species. In: Essink K., C. Dettmann, H.
Farke, K. Laursen, G. Lüerßen, H. Marencic & W. Wiersinga (eds) Wadden Sea quality
status report 2004. Common Wadden Sea Secretariat, Wilhelmshaven, pp 155–161
Ruesink, J. L., H. S. Lenihan, A. C. Trimble, K. W. Heiman, F. Micheli, J. E. Byers & M. C.
Kay 2005. Introduction of non-native oysters: ecosystem effects and restoration
implication. Ann. Rev. Ecol. Evol. Syst. 36: 643-689
Saier, B. 2001. Ecological comparison of intertidal and subtidal mussel beds (Mytilus edulis
L.) in the Wadden Sea. Dissertation, Universität Hamburg.
Seaman, M.N.L. 1985. Ecophysiological investigations on the oyster, Crassostrea gigas, in
Flensburg fjord. - Veröff. Inst. Küst. Binnenfisch., Hamburg Bd. 89: 71 pp.
Strasser, M 2002: Reduced epibenthic predation on intertidal bivalves after a severe winter in
the European Wadden Sea. Mar. Ecol. Prog. Ser. 241: 113-123
Strasser, M. & C.-P. Günther 2001. Larval supply of predator and prey: temporal mismatch
between crabs and bivalves after a severe winter in the Wadden Sea. J. Sea Res. 46:
57-67.
Tsuchiya, M. & M. Nishihira 1986. Islands of Mytilus edulis as a habitat for small intertidal
animals: effect of Mytilus age structure on the species composition of the associated
fauna and community organization. Ecol. Prog. Ser. 31:171-178.
Tydeman, P. 1999. Japanse Oester in de Eemshaven. Het Zeepaard 59: 58-64.
Wehrmann A, M Herlyn, F Bungenstock, G Hertweck, G Millat 2000: The distribution gap is
closed—first record of naturally settled Pacific oysters (Crassostrea gigas) in the East
Frisian Wadden Sea, North Sea. Senckenb Marit 30:153–160
Wehrmann, A., A. Markert, P. May, P. Schiek & A. Schmidt 2006. Gefährdungspotential der
eulitoralen Miesmuschelbänke im Niedersächsischen Wattenmeer durch die
Bioinvasion der Pazifischen Austern Crassostrea gigas. Abschlussbericht Projekt 7/02
der Niedersächsischen Wattenmeer-Stiftung.
Wijsman, J. W. M., M. van Stalen, M. Dubbeldam, R. Geene, M. de Kluijver, E. van Zanten &
A. C. Smaal 2006. Wegvisproef Japanse oesters in de Oosterschelde. Tussentijdes
rapportage T2. Wageningen IMARES Rapport nr. C077/06.
Wolff, W.J. 2005. Non-indigenous marine and estuarine species in The Netherlands. - Zool.
Med. Leiden 79: 1-116. Wolff, W.J. and Reise, K. 2002. Oyster imports as a vector for
the introduction of alien species into northern and western European waters. - In :
Leppäkoski, E., Gollasch, S. and Olenin, S. (eds), Invasive aquatic species of Europe.
Distribution, impacts and management. Kluwer, Dordrecht: 193-205
Wolff, W.J. & K. Reise 2002.Oyster imports as a vector for the introduction of alien species
into the northern and western European waters, Leppäskoski, E., S. Gollasch & S.
Olenin (eds.) Invasive aquatic species of Europe : distribution, impacts, and
management. Dordrecht [u.a.] : Kluwer, 193-205.

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Annex 1: Conclusions and Recommendations
Conclusions and Recommendations
from the Trilateral Workshop on
Pacific Oyster Invasion in the Wadden Sea
Consequences for Ecology, Monitoring and Management
22 March 2007, Wilhelmshaven
Background
The Pacific Oyster has spread over the entire Wadden Sea. Results from ongoing research
projects have indicated that spreading of the Pacific oyster has increased considerably
during the last 5 years.
In order to obtain an up-date of the status of the Pacific oyster invasion and to assess the
possible consequences for the ecosystem, as well as for monitoring and management, also
with regard to the EC Habitats and Water Framework Directive, a trilateral report with a
comprehensive data analysis was compiled by Georg Nehls and Heike Büttger (BioConsult
SH) on behalf of the Common Wadden Sea Secretariat.
The work was co-funded by the Interreg IIIB project HARBASINS and supported by the
Research Institute Senckenberg (Wilhelmshaven), the Schleswig-Holstein National Park
Agency (Tönning) and the Lower Saxon National Park Administration (Wilhelmshaven).
Additionally, the Institute of Marine Research-IMARES (Texel), the Danish Fisheries
Research Institute and the Alfred Wegener Institute-AWI (List/Sylt) provided recent data on
oyster distribution.
The report was the starting point for a discussion at a trilateral workshop on 22 March 2007
in Wilhelmshaven with scientists and managers from the Wadden Sea and England to
discuss the recent status and possible consequences of the Pacific oyster spreading in the
Wadden Sea and to prepare recommendations regarding follow up-activities in the fields of
research, monitoring and management. The workshop program is in Annex 1 and the list of
participants is in Annex 2.
Conclusions
Sporadic occurrence of the Pacific oyster in some parts of the Wadden Sea is known for over
20 years, but a rapid increase in the entire area has been documented only for the last 5
years. Areas populated by the Pacific oyster (blue mussel beds, hard substrates) increased
significantly in the past years, also including formerly soft sediments. The Pacific oysters
form massive reefs in all parts of the Wadden Sea. The total oyster biomass in the intertidal
area of the entire Wadden Sea has increased to about 61.000 tons fresh weight in 2006 (The
Netherlands: 20.510, Lower Saxony: 22.747, Schleswig-Holstein: 14.481; Denmark: 3289).
In many places Pacific oysters have overgrown former blue mussel beds to form dense and
solid reefs. Abundance in dense reefs is about 1000 oysters m-2 with a mean biomass 30 to
50 kg m-2 live wet weight.

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1. Factors influencing oyster distribution
Abiotic factors:
As a main abiotic factor the substrate availability must be pointed out. Pacific oysters settle
on hard substrates which are of limited occurrence in the Wadden See. They often settle
initially upon barnacles, blue mussels and dead bivalve shells. Further on Pacific oysters
create their own substrate and as oyster larvae prefer conspecifics to settle, they initiate
massive clumps which grow further as more oyster larvae settle on top.
Salinity does not seem to significantly influence the distribution of the Pacific oyster in the
Wadden Sea.
The reproduction of the Pacific Oyster in the Wadden Sea is highly correlated to summer
temperatures. The high summer temperatures in the last 10-12 years have been identified as
a main factor causing the recent increase of the Pacific oyster. A few days in July and August
with water temperatures above 18-20 °C may already trigger the reproduction of the oyster.
On the other hand, the Pacific oyster is able to survive cold winters better than previously
expected.
Mass mortality among Pacific oysters was observed occasionally in shallow bays and
harbours. In late summer after spatfall (when condition of oysters is low).This is probably
caused by local factors such as limited water exchange.
Biotic Factors
The Pacific oyster has almost no natural predators in the Wadden Sea. But predation by
birds like oystercatchers and herring gulls has been observed. So far neither starfish, shore
crabs, birds nor parasites induce high mortality that could stop or reduce the oyster’s
population growth.
The observed population development is considered to have been promoted by an extension
of phytoplankton blooms in late summer which enhance especially successful settling of
oyster larvae.
On the other hand, phytoplankton spring blooms have developed later in the year (due to
increased grazing pressure during mild winters) providing less food for blue mussels, cockles
and Baltic tellins during their reproductive period.
2. Future development
From experiences in the Dutch Delta region (Oosterschelde), where the Pacific oyster
showed a continuous increase over the last 30 years, it can be assumed that the spread in
the Wadden Sea will continue further. Total biomass (life wet weight) of Pacific oysters in
Wadden Sea is still less than half of that known from the Oosterschelde.
Climate change will further enhance the spread of the Pacific oyster. Warmer winters are
assumed to have a negative influence on reproduction of blue mussels and other bivalves,
because predators like shore crabs and shrimps return earlier in spring to the tidal flats
reducing bivalve spat significantly. On the other hand, proliferation of oysters is facilitated by

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page 49
warm summers. The warm summer of 2006 lead to a strong oyster spatfall in the entire
Wadden Sea and further spread and increasing abundances and biomass are to be
expected.
The questions whether or not native blue mussel beds will disappear from the Wadden Sea
due to the invasive nature of the Pacific oyster or whether a coexistence of Pacific oysters
and blue mussels is possible cannot be answered so far. In future, all three types of beds
might occur: blue mussel beds, oyster reefs and mixed beds.
There are several examples of co-existence of blue mussels and oysters in mixed beds.
In the Dutch Wadden Sea, blue mussels have successfully re-established a strong
population in the last years. If sufficient blue mussel recruits manage to settle, new beds may
develop and blue mussels may co-exist with oyster reefs.
To evaluate the further development of the Pacific Oyster in the Wadden Sea annual
monitoring should be carried out, accompanied with investigations on ecology and
development in other coastal waters of our globe.
3. Consequences for the ecosystem
Until now the species composition of the associated fauna of oyster reefs compared to that of
blue mussel beds does not differ significantly. Negative effects on the biodiversity have not
been detected, but the dominance structure of the associated fauna has changed.
Most bird species seem to be able to adapt to the appearance of oyster reefs because they
feed mainly on the associated fauna, such as worms and shore crabs. Birds with food
preference for blue mussels, like Eider and Oystercatcher, are not able to use oysters as
food resource.
In the Oosterschelde (SW Netherlands) a management experiment was carried out
consisting of the removal of 50 ha of oyster reefs in March 2006. Effects on sedimentation
and benthic infauna community at the oysters plots and reference areas were also
investigated. The interim report (Wijsman et al. 2006) stated that the removal is effective.
Oyster beds exhibit muddy sediments with higher organic carbon content which becomes
apparent in the associated community (higher diversity, characterized by polychaetes,
decapods and amphipods). Oyster removal might lead to sandy sediments with a less
diverse community (more molluscs). A negative side effect may occur through suffocation of
the benthic fauna living in areas where the removed oyster bed is dumped.
4. Human use
Negative effects on recreational activities can be expected because oyster shells are sharp
and swimmers and surfers may be hurt; however, this will be restricted to few areas only.
The spread of the Pacific oyster may have some effects on the blue mussel fishery because
mixed beds with blue mussels and oysters can no longer be exploited. At present no specific
data on this issue are available.
At present there is only limited interest in fisheries for Pacific oysters as the wild oysters are
of little commercial value.

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Positive effects of developing oyster reefs on coastal protection issues are not considered to
be of significance.
5. Monitoring
The spread of the Pacific oyster is a major change in the Wadden Sea ecosystem and
should therefore be properly documented to assess future changes. A higher effort is
therefore necessary to gather the required data.
The ongoing national monitoring programs in the TMAP document the location of blue
mussel beds and in this context also the occurrence of Pacific oyster. However, no
information on oysters is available for areas which are not surveyed in the mussel
monitoring.
There is a gap in the monitoring in the Netherlands in 2007 (only few beds are monitored
regularly in connection with blue mussel fishery).
Apart from the monitoring program, more research is urgently needed to investigate and
assess the changes of the Wadden Sea ecosystem which may be induced by the change of
a key ecological community. Such data are also needed against the background of the
Habitat Directive.
Methods of the Trilateral Monitoring must be standardized with respect to the adequate
monitoring of oyster reefs.
6. Management
There are no successful management options available for removal of the Pacific oyster from
the Wadden Sea. Therefore, these oysters have to be considered as permanent members of
the Wadden Sea ecosystem. The ecosystem has been always changing and interference by
severe management measure should be avoided. This would be in line with the guiding
principle that natural processes in the Wadden Sea should proceed as undisturbed as
possible.
In order to control, or reduce the occurrence of the Pacific oyster in the Wadden Sea, no
viruses or diseases should be introduced because of their unforeseeable and uncontrollable
effects.
Local management / removal may be possible as already discussed for the Oosterschelde.
In any case a better scientific basis has to be established to assess the impact of such a
management measure on the ecosystem and its processes.
Recommendations
1. Research
The workshop underlined the urgency of further ecological research in order to better assess
the consequences of the Pacific oyster invasion for the Wadden Sea ecosystem. A good
scientific foundation has to be prepared now and in this phase of oyster spreading, to be able
to better assess future changes.

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In general there is a need for more basic knowledge on ecosystem effects in order to support
management decisions. Experiences from other coastal waters of the world should be used.
Cooperation of research institutes in the Wadden Sea and other countries (F, UK, Asia) and
exchange of knowledge is recommended. The CWSS is asked to coordinate research
activities and future initiatives, and to investigate possible funding from the EU (such as
COST, FP7).
2. Monitoring
At present, the monitoring of Pacific oysters in the Wadden Sea is not done in a harmonized
way. For example, no monitoring takes places in the Dutch Wadden Sea in 2007, whereas
further assessments in Niedersachsen and Schleswig-Holstein are guaranteed until 2008.
Pacific oyster monitoring cannot be simply included in the existing monitoring of blue mussel
beds, and therefore requires extra funding.
The further developments of the Pacific oyster and its effects on the ecosystem should be
part of the harmonized assessment of the data for the entire Wadden Sea. The Trilateral
Monitoring and Assessment Program (TMAP) is the instrument to provide Wadden Sea wide
data in a harmonized and effective way.
The present gaps in monitoring of the Pacific oyster should be filled as soon as possible by
establishing a regular monitoring within the framework of the TMAP, starting already in spring
2007
The CWSS in cooperation with N. Dankers, P. S. Kristensen, G. Millat, G. Nehls and A.
Wehrmann will prepare a proposal for a TMAP manual for Pacific oyster monitoring.
3. Management
The spread of the Pacific oyster in the Wadden Sea has to be regarded as the consequence
of a careless introduction of an alien species to a sensitive and protected ecosystem. This
underlines that the introduction of alien species in an ecosystems has to be avoided as much
as possible.
The judgement on possible removal of Pacific oyster beds as a management option needs
more supporting scientific knowledge and must be based on thorough impact assessment.
The first trial has been done in the Oosterschelde in March 2006 (Wijsman et al. 2006) and
more results have to be awaited.

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Annex 2: Final Workshop Program
Co-funded by ERDF Trilateral Workshop on
Pacific Oyster Invasion in the Wadden Sea
Consequences for Ecology, Monitoring and Management
22 March 2007
Senckenberg Research Institute,
Südstrand 40, Wilhelmshaven
Organized by the Common Wadden Sea Secretariat in the framework of the HARBASINS project
Program
11.00 Welcome and introduction (Chair: Karel Essink)
11:10 Distribution of the Pacific Oyster in the Netherlands, Germany and Denmark:
Temporal and spatial trends (Georg Nehls, BioConsult SH)
11.40 Distribution of the Pacific Oyster in South-west England (Gemma Couzens,
Natural England, UK)
11:50 Distribution of Pacific Oyster in the Danish Limfjord (Helle Torp Christensen,
Danish Technical University)
12:00: Topical presentations
• 12:00 Associated fauna (Heike Büttger, BioConsult SH, Husum)
• 12:10: Filtration experiments (Achim Wehrmann, Research Institute Senckenberg,
Wilhelmshaven)
12:20– 13:15 Lunch
• 13:15 .Field experiments on artificial oyster reefs (Karsten Reise, Alfred-Wegener-
Institute, Sylt)
• 13:30 Larviphagy by oysters, mussels and cockles (Wim Wolff / Karin Troost, Uni
Groningen)
• 13:45 Consequences for birds (Bruno Ens, SOVON, NL / Gregor Scheiffarth, Inst.
Avian Research, Wilhelmshaven)
• 14:00 Oyster monitoring in the TMAP (Georg Nehls, BioConsult SH, Husum)
14:15 – 14:30 Break
14:30 Plenary Discussion
Topics
- Oyster distribution in the Wadden Sea: main factors, future developments;
- Consequences for the ecosystem,
- Consequences for human use (recreation, fisheries, coastal protection);
- Adaptation of monitoring and management,
15:45 Conclusions
• Main findings and recommendations
• Follow up (research projects, monitoring proposals)
16:00 Closing

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Annex 3: Workshop Participants
Name Institute
The Netherlands
Essink Karel Chairman
Dankers Norbert Institute for Marine Resources & Ecosystem Studies
Fey-Hofstede Frouke Institute for Marine Resources & Ecosystem Studies
Wijsman Jeroen Institute for Marine Resources & Ecosystem Studies
Braaksma Sytze Min. Landbouw, Natuur & Voedselkwaliteit
Kouwenhoven Angelo Min. Landbouw, Natuur & Voedselkwaliteit
Schermer Voest Wilbert Min. Landbouw, Natuur & Voedselkwaliteit
Ens Bruno SOVON
de Vlas Jaap RIKZ
Wolff Wim Uni Groningen
Germany
Reise Karsten Alfred-Wegener-Institute
Ruth Maarten ALR-Kiel, Abt. Fischerei
Büttger Heike BioConsult SH
Nehls Georg BioConsult SH
Rolke Manfred Bundesamt für Seeschifffahrt und Hydrographie
Marencic Harald Common Wadden Sea Secretariat
de Jong Folkert Common Wadden Sea Secretariat
Markert Alexandra Forschungsinstitut Senckenberg
Schmidt Andreas Forschungsinstitut Senckenberg
Wehrmann Achim Forschungsinstitut Senckenberg
Zwaka Hanna Forschungsinstitut Senckenberg
Lenz Mark IFM-GEOMAR
Dittmann Tobias Institut für Vogelforschung
Esser Wiebke Institut für Vogelforschung
Exo Michael Institut für Vogelforschung
Scheiffarth Gregor Institut für Vogelforschung
de Leeuw Andries Landesverband Schleswig-Holsteinischer Angler und Fischer e.V.
Oberdoerffer Philipp Landwirtschaftskammer Niedersachsen
Knoke Vera Ministerium für Landwirtschaft, Umwelt und ländliche Räume
Schl.-Holst.
Eskildsen Kai Nationalparkamt Schleswig-Holst.
Borchardt Thomas Nationalparkamt Schleswig-Holst.
Millat Gerald Nationalparkverwaltung Niedersächsiches Wattenmeer
Claußen Alina Nationalparkverwaltung Niedersächsiches Wattenmeer
Stede Michael Nds. Landesamt f. Verbraucherschutz u. Lebensmittelsicherheit
Gubernator Manuela Niedersächsische Muschelfischer GbR

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Name Institute
Herlyn Marc Niedersächsischer Landesbetrieb für Wasserwirtschaft, Küsten-
und Naturschutz
Pogoda Bernadette Universität Bremen, Marine Zoologie
Denmark
Jensen Jan Steinbring Danish Forest and Nature Agency
Christensen Helle Torp Danish Institute for Fisheries Research Technical University of
Denmark
Knudsen Tom Ministry of the Environment Environmental Center Ribe
UK
Couzens Gemma Natural England
Donnelly Conor Natural England