Since 1998 the non-indigenous Pacific oyster Crassostrea gigas (Thunberg 1793) has been invading the Wadden Sea of Lower Saxony, southern German Bight. C. gigas settles predominantly on intertidal Mytilus-beds (M. edulis L.) and subsequently create rigid reef-like structures. Both bivalve species are ecosystem engineers in sedimentary tidal flats. They provide hard substrate for sessile species, mobile organisms find refuge within the habitat matrix of dense suspension feeders, and biodeposits enrich the sediments with organic matter. The transformation of Mytilus-beds into Crassostrea-reefs gives rise to the question whether the invader may affect the native community. We investigated two parts of a changing bivalve bed in the backbarrier area of the island of Juist in March 2005. One part was still dominated by M. edulis whereas the other part was already densely colonized by C. gigas. Crassostrea-reefs compensate for the conceivable loss of Mytilus-beds in the intertidal of the Wadden Sea by replacing the ecological function of M. edulis. There was no indication of a suppression of indigenous species. This even applied to M. edulis, which persisted at the site invaded by C. gigas. The associated macrofaunal community showed increased species richness, abundance, biomass, and diversity in the Crassostrea-reef. The latter particularly favored sessile species like anthozoans, hydrozoans, and barnacles. Higher abundance and biomass for vagile epizoic species like the shore crab Carcinus maenas and the periwinkle Littorina littorea also occurred among oysters. Abundance of deposit feeding oligochaetes was enhanced by oysters as well. More opportunistic, facultative filter-feeding polychaetes occurred in the Crassostrea-reef.

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Pacific oysters Crassostrea (Magallana) gigas have been successfully invading ecosystems worldwide. As an ecosystem engineer, they have the potential to substantially impact on other species and on functional processes of invaded ecosystems. Engineering strength depends on oyster density in space and time. Density has not yet been studied on the extent of reef structural dynamics. This study assessed abundance of naturalized Pacific oysters by shell length (SL) of live individuals and post-mortem shells at six sites over six consecutive years during post-establishment. Individual biomass, i.e. live wet mass (LWM), flesh mass (FM) and live shell mass (SM LIVE), were determined from a total of 1.935 live oysters in order to estimate areal biomass. The generic term density attribute was used for SL-related population categories and the biomass variables LWM, FM, SM LIVE and SM. As the oyster invasion modulated resident Mytilus edulis beds, the study was supplemented by contemporaneously assessed data of mussels and corresponding analyses. Interrelations of abundance and areal biomass revealed distinct linkages between specific density attributes. Most importantly, large individuals were identified as intrinsic drivers for the determination of areal biomass. Additionally, allometry of large oysters differed from small oysters by attenuated scaling relations. This effect was enhanced by oyster density as results showed that crowding forced large individuals into an increasing slender shape. The significant relationship between the density attributes large oyster and biomass enabled a classification of reef types by large oyster abundance. Reef type (simple or complex reef) and oyster size (small or large) were considered by implementing a novel concept of weighted twin functions (TF) for the relationship between SL and individual biomass. This study demonstrates that the interplay of scaling parameters (scalar, exponent) is highly sensitive to the estimation of individual biomass (shape) and that putative similar scaling parameters can exceedingly affect the estimation of areal biomass. For the first time, this study documents the crucial relevance of areal reference, i.e. cluster density (CD) or reef density (RD), when comparing density. RD considers reef areas devoid of oysters and results from CD reduced by reef coverage (RC) as the relative reef area occupied by oysters. A compilation of density attributes at simple and complex reefs shall serve as a density guide. Irrespective of areal reference, oyster structural density attributes were significantly higher at complex than at simple reefs. In contrast, areal reference was of vital importance when evaluating the impact of engineering strength at ecosystem-level. While mussel CD was similar at both reef types, RD at complex reefs supported significantly more large mussels and higher mussel biomass than at simple reefs. Although mussels dominated both reef types by abundance of large individuals, oysters were the keystone engineers by dominating biomass. The prominent status of large oysters for both allometric scaling and density, presumably characteristic for Pacific oyster populations worldwide, should be considered when conducting future investigations. The effort of monitoring will substantially be reduced as only large oysters have to be counted for an empirical characterization of Pacific oyster reefs. The large oyster concept is independent of sampling season, assessment method or ecosystem, and is also applicable to old data sets. Harmonization on the proposed density attributes with a clear specification of areal reference will allow trans-regional comparisons of Pacific oyster reefs and will facilitate evaluations of engineering strength, reef performance and invasional impacts at ecosystem-level.

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