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Marine heatwaves (MHWs) are increasing in frequency and intensity due to climate change. Several well-documented effects of heatwaves on community structure exist, but examples of their effect on functioning of species, communities or ecosystems remain scarce. We tested the effects of short-term, moderate and strong MHWs on macrofauna bioturbation and associated solute fluxes as examples of ecosystem functioning. We also measured macrofaunal excretion rates to assess effects of temperature on macrofauna metabolism. For this experiment, we used unmanipulated sediment cores with natural animal communities collected from a muddy location at 32 m depth in the northern Baltic Sea. Despite the mechanistic effect of bioturbation remaining unchanged between the treatments, there were significant differences in oxygen consumption, solute fluxes and excretion. Biogeochemical and biological processes were boosted by the moderate heatwave, whereas biogeochemical cycling seemed to decrease under a strong heatwave. A prolonged, moderate heatwave could possibly lead to resource depletion if primary production cannot meet the demands of benthic consumption. By contrast, decreased degradation activities under strong heatwaves could lead to a build-up of organic material and potentially hypoxia. The strong variability and the complexity of the response highlight the context dependency of these processes complicating future predictions.

The paper published in Limnology and Oceanography 67: 784-799 (2022) contains a calculation error. This error concerns the result and discussion sections “Stoichiometric variation in the benthic community”. The benthic community biomass and consequently the community C, N and P content were miscalculated. The corrected benthic community C, N and P content is approximately three times higher than the values presented in the original paper. However, the correction in biomass had no major effect on the community C: N: P ratios. Statistical analysis was performed (two-way PERMANOVA) again for the community stoichiometry, but the results did not change. Importantly, corrections of the community biomass did not affect the main results or the conclusion of the paper. The corrected version of Table 3, Fig. 7 and Supplementary Table 7 are presented below. These corrections change the following: 3 Corrected Table Seasonal variation in the magnitude of the benthic community carbon, nitrogen, and phosphorus pools (g m−2) and elemental ratios. Tot C, N, and P present the values with bivalve shells included, whereas tissue C, N, and P content were calculated without the shells. Molar C:N:P ratios of the community are not accounting for bivalve shells. (Table presented.) 7 Corrected Fig. (Figure presented.) Average community total carbon (C) content g m−2 ± SD over the sampling occasions, and the contribution of the main taxonomic groups to the total biomass. The nitrogen and phosphorus followed the same trend as carbon. 7 Corrected Supplementary Table Results of two-way PERMANOVA show the effect of season and site on community C, N and P content and C:N:P ratios. (Table presented.) Methods: Stoichiometric variation in the benthic community. p. 787: the text should read “The model was based on Euclidean distances, calculated on total C, N, and P content g m-2 and log10(x + 1) transformed C: N: P data (Anderson et al. 2008)”. Results: Stoichiometric variation in the benthic communi, This article corrects the following: Food sources drive temporal variation in elemental stoichiometry of benthic consumers by Saara Mäkelin, Anna Villnäs in Limnology and Oceanography 2022, Volume 67, Issue 4

Benthic consumers profoundly impact nutrient regeneration in coastal marine ecosystems. The concurrent nutrient imbalance and warming of our coastal seas will change the nutritional requirements and metabolic demands of these consumers, which may affect their ability to recycle nitrogen and phosphorous. Here we explore whether nutrient excretion rates of two benthic consumers, the Baltic clam (Macoma balthica) and the invasive spionid polychaete (Marenzelleria spp.) can be quantified with basic biological traits across seasons using allometric and stoichiometric relationships. We found species-specific N and P excretion rates that positively link to allometric traits, i.e., per individual rates increased with body mass and temperature; thus, high mass-specific excretion rates characterized small relative to large macrofaunal individuals. Interestingly, our body size scaling coefficients diverge from predictions by the metabolic theory of ecology (MTE) and the universal model of excretion. Furthermore, stoichiometric traits and stable isotope signatures (delta C-13 and delta N-15) explained a minor additional proportion of variability in excretion rates among species. The excretion rates also varied strongly seasonally, with the highest nutrient recycling rates during summer months, when community NH4-N and PO4-P excretion clearly exceeded net sediment efflux. The seasonal pattern emphasized that changes in temperature and food availability drove metabolic processes and thus excretion rates of the benthic consumers, and indicated that these effects could outweigh the importance of animal biomass. Our results highlight the benefits of using allometric and stoichiometric traits when quantifying species-specific contributions to nutrient recycling in coastal marine environments, and in predicting alteration of function in response to environmental change.

Hypoxia is presently seen as the principal driver behind the decline of the former dominating Eastern Baltic cod stock (EBC; Gadus morhua). It has been proposed that both worsening conditions for reproduction and lower individual growth, condition, and survival are linked to hypoxia. Here, we elucidate the ecological envelope of EBC in terms of salinity stratification, oxygen content, and benthic animal biomasses, and how it has affected EBC productivity over time. The spawning conditions started deteriorating in the Gotland Deep in the 1950s due to oxygen depletion. In contrast, in the Bornholm Basin, hydrographic conditions have remained unchanged over the last 60 years. Indeed, the current extent of both well-oxygenated areas and the frequency of hypoxia events do not differ substantially from periods with high EBC productivity in the 1970s–1980s. Furthermore, oxygenated and therefore potentially suitable feeding areas are abundant in all parts of the Baltic Sea, and our novel analysis provides no evidence of a reduction in benthic food sources for EBC over the last 30 years. We find that while reproduction failure is intricately linked to hydrographic dynamics, a relationship between the spread of hypoxia and the decline in EBC productivity during the last decades cannot be substantiated.

Benthic fauna plays an important role in mediating biogeochemical cycles in coastal areas by storing carbon (C), nitrogen (N), and phosphorus (P) in their body tissues at theoretically homeostatic rates. To maintain homeostasis, the benthic consumers need to be in balance with their resource supply or alter their stoichiometric traits in response to environmental change. However, we lack an understanding regarding the potential variation in the C : N : P content ratios of benthic consumers, especially in marine coastal areas where stoichiometric shifts in macrofauna could have strong effects on sediment carbon and nutrient cycling. By monitoring two sites over a year, we quantified the magnitude and temporal stability of benthic faunal carbon and nutrient pools in coastal soft-sediment habitats. Our results show that benthic fauna is not strictly homeostatic, but instead expresses temporal variation in C : N : P content. These aquatic consumers undergo ontogenetic changes in diet and morphology, which alter their stoichiometric characteristics. In addition, the faunal C : N : P ratios showed strong seasonal variation at both species and community level, and our results suggest that the stoichiometric traits of benthic consumers shift in response to food sources and environmental conditions. The ability to adapt to varying stoichiometric conditions is essential in face of the growing C : N : P imbalance occurring in marine and coastal ecosystems as a consequence of anthropogenic activities. Therefore, it is critically important to identify the stoichiometric plasticity of different species, before environmental change causes a shift in benthic community composition that will alter functions on an ecosystem level.

Explaining the recent decline of eastern Baltic cod (EBC) remains scientifically challenging. Brander proposes in a comment to Svedäng et al.that the observed trend in oxygen in SD 25 supports the idea that juvenile cod are balancing the physiological cost of living under mild hypoxiaby offsetting the risk of being eaten by diving seals and cormorants in shallower water with more oxygen. There are a number of objections tothis conjecture, besides the fact that supporting observations are missing. Hence, it is difficult to reconcile the long-term development of EBCunder varying oxygen conditions with the hypothesis that a small reduction in oxygen content can explain the current strong and uniform declinein growth observed in the entire southern Baltic Sea.

Despite a long history of disturbance-recovery research, we still lack a generalizable understanding of the attributes that drive community recovery potential in seafloor ecosystems. Marine soft-sediment ecosystems encompass a range of heterogeneity from simple low-diversity habitats with limited biogenic structure, to species-rich systems with complex biogenic habitat structure. These differences in biological heterogeneity are a product of natural conditions and disturbance regimes. To search for unifying attributes, we explore whether a set of simple traits can characterize community disturbance-recovery potential using seafloor patch-disturbance experiments conducted in two different soft-sediment landscapes. The two landscapes represent two ends of a spectrum of landscape biotic heterogeneity in order to consider multi-scale disturbance-recovery processes. We consider traits at different levels of biological organization, from the biological traits of individual species, to the traits of species at the landscape scale associated with their occurrence across the landscape and their ability to be dominant. We show that in a biotically heterogeneous landscape (Kawau Bay, New Zealand), seafloor community recovery is stochastic, there is high species turnover, and the landscape-scale traits are good predictors of recovery. In contrast, in a biotically homogeneous landscape (Baltic Sea), the options for recovery are constrained, the recovery pathway is thus more deterministic and the scale of recovery traits important for determining recovery switches to the individual species biological traits within the disturbed patch. Our results imply that these simple, yet sophisticated, traits can be effectively used to characterize community recovery potential and highlight the role of landscapes in providing resilience to patch-scale disturbances.

Macrofauna play a key role in the functioning of mangrove ecosystems. Nevertheless, our understanding of the diversity and functional structure of macrofaunal communities across different habitats in the mangrove forests of the Persian Gulf is limited. In this study, we investigated species diversity and biological trait patterns of macrofauna in different mangrove-associated habitats, i.e., encompassing actual mangrove forests, and adjacent Beaches and Creeks, which exhibit different levels of habitat heterogeneity. Samples were collected from the different habitats in five different locations, over four seasons. A total of 122 macrofauna taxa were identified. The diversity of species was higher in summer than in winter. In the Beach habitats, species diversity showed an increasing trend from land toward the mangrove, whereas in Creek habitats diversity decreased from the Creek toward the mangrove. Multivariate community analysis showed differences in the distribution of abundant species and biological traits across all habitats. Deposit-feeding, crawlers, medium-size, and free-living were the dominant trait modalities in all habitats. The similarities within habitats over the four seasons had the same specific pattern of species and biological trait abundance in the Beach and the Creek, increasing from the non-covered habitat into the mangrove trees. Although many species shared similar traits, the abundance-driven differences in trait expression between habitats showed the importance of habitat filtering. The results of this study will be useful in the conservation of mangrove forests and they give a deeper understanding of the ecological patterns and functions of benthic macrofaunal communities in the Persian Gulf.

Benthic macrofaunal communities have a profound impact on organic matter turnover and nutrient cycling in marine sediments. Their activities are of particular importance in the coastal filter, where nutrients and organic matter from land are transformed and/or retained before reaching the open sea. The benthic fauna modify the coastal filter directly (through consumption, respiration, excretion and biomass production) and indirectly (through bioturbation). It is hard to experimentally quantify faunal contribution to the coastal filter over large spatial and temporal scales that encompass significant environmental and biological heterogeneity. However, estimates can be obtained with biological trait analyses. By using benthic biological traits, we explored how the potential contribution of macrofaunal communities to the coastal filter differ between inner and outer sites in an extensive archipelago area and examine the generality of the observed pattern across contrasting coastal areas of the entire Baltic Sea. Estimates of benthic bioturbation, longevity and size (i.e. 'stability') and total energy and nutrient contents differed between coastal areas and inner versus outer sites. Benthic traits indicative of an enhanced nutrient turnover but a decreased capacity for temporal nutrient retention dominated inner sites, while outer sites were often dominated by larger individuals, exhibiting traits that are likely to enhance nutrient uptake and retention. The overarching similarities in benthic trait expression between more eutrophied inner vs. less affected outer coastal sites across the Baltic Sea suggest that benthic communities might contribute in a similar manner to nutrient recycling and retention in the coastal filter over large geographical scales.

Foundation species host diverse associated communities by ameliorating environmental stress. The strength of this facilitative effect can be highly dependent on the underlying biotic and abiotic context. We investigated community level patterns of macrofauna associated with and adjacent to the marine foundation species eelgrass (Zostera marina) along a hydrodynamic stress gradient. We could demonstrate that the relative importance of this foundation species for its infaunal community increases with environmental variables associated with increasing hydrodynamic stress (depth, sand ripples formation, sediment grain size and organic content). Faunal assemblages in proximity to the Zostera patch edges, however, showed no (infauna) or negative (epifauna) response to hydrodynamic stress. Our study highlights that the facilitative outcome of a foundation species is conditional to the faunal assemblage in question and can be highly variable even between positions within the habitat.

Foundation species host diverse associated communities by ameliorating environmental stress. The strength of this facilitative effect can be highly dependent on the underlying biotic and abiotic context. We investigated community level patterns of macrofauna associated with and adjacent to the marine foundation species eelgrass (Zostera marina) along a hydrodynamic stress gradient. We could demonstrate that the relative importance of this foundation species for its infaunal community increases with environmental variables associated with increasing hydrodynamic stress (depth, sand ripples formation, sediment grain size and organic content). Faunal assemblages in proximity to the Zostera patch edges, however, showed no (infauna) or negative (epifauna) response to hydrodynamic stress. Our study highlights that the facilitative outcome of a foundation species is conditional to the faunal assemblage in question and can be highly variable even between positions within the habitat.

Understanding large-scale spatial variation in ecosystem properties and associated functionality is key for successful conservation of ecosystems. This study provides a template for how to estimate differences in ecosystem functionality over large spatial scales by using groupings of biological traits. We focus on trait groupings that describe three important benthic ecosystem properties, namely bioturbation, community stability, and juvenile dispersal. Recognizing that groups of traits interact and are constrained within an organism, we statistically define important functional trait subgroups that describe each ecosystem property. The sub-groups are scored according to their weighted ecological impact to gain an overall estimation of the cumulative expression of each ecosystem property at individual sites. Furthermore, by assigning each property a value relative to its observed maximum, and by summing up the individual property values, we offer an estimate of benthic ecosystem multi-functionality. Based on a spatially extensive benthic data set, we were able to identify coastal areas with high and low potential for the considered benthic ecosystem properties and the measure of ecosystem multifunctionality. Importantly, we show that a large part of the spatial variation in functional trait sub-groups and in benthic ecosystem multifunctionality was explained by environmental change. Our results indicate that through this simplification it is possible to estimate the functionality of the seafloor. Such information is vital in marine spatial planning efforts striving to balance the utilization with the preservation of natural resources.