How dense is dense? Toward a harmonized approach to characterizing reefs of non-native Pacific oysters – with consideration of native mussels

Abstract

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.