Effects of boat traffic and mooring infrastructure on aquatic vegetation: A systematic review and meta-analysis.

Sustainable management of coastal and inland water areas requires knowledge of how tourism and recreation affects the ecosystems. Here, we present the first systematic review and meta-analysis to quantify to what extent recreational boat traffic and infrastructure for mooring affect the abundance of submerged vegetation on soft bottoms. Our systematic search yielded 25 studies containing data on effects of boat traffic, docks and mooring buoys on vegetation abundance. The abundance below docks was on average 18% of that in controls, and areas with boat traffic had on average 42% of the abundance in control areas. Mooring buoys often created scour areas without vegetation. However, the effects were variable and there were too few studies to test the reasons for this variability. We conclude that boating can cause significant declines in submerged vegetation but that informed management of boat traffic and improved design of docks and buoys can reduce negative impacts.

Cover of coastal vegetation as an indicator of eutrophication along environmental gradients.

Coastal vegetation communities are important for primary production, biodiversity, coastal protection, carbon and nutrient cycling which, in combination with their sensitivity to eutrophication, render them potential indicators of environmental status for environmental policies like the EU Water and Marine Strategy Framework Directives. We evaluated one potential indicator for coastal vegetation, the cumulative cover at depths where the vegetation is light limited, by investigating its response to eutrophication along gradients in natural conditions. We used a large data set covering the Swedish coastline, spanning broad gradients in nutrient level, water clarity, seabed substrate, physical exposure and climate in addition to a salinity gradient from 0.5 to 30.5. Macroalgal cover increased significantly along gradients of declining nutrient concentration and increasing water clarity when we had accounted for diver effects, spatio-temporal sampling variability, salinity gradients, wave exposure and latitude. The developed empirical model explained 79% of the variation in algal cover across 130 areas. Based on this, we identified macroalgal cover as a promising indicator across the Baltic Sea, Kattegat and Skagerrak. A parallel analysis of soft-substrate macrophytes similarly identified significant increases in cover with decreasing concentrations of total nitrogen and increasing salinity, but the resulting empirical model explained only 52% of the variation in cover, probably due to the spatially more variable nature of soft-substrate vegetation. The identified general responses of vegetation cover to gradients of eutrophication across wide ranges in environmental settings may be useful for monitoring and management of marine vegetation in areas with strong environmental gradients.

Divergent ecological strategies determine different impacts on community production by two successful non-native seaweeds.

The consequences of plant introductions into ecosystems are frequently reported from terrestrial environments, but little is known about the effects on ecosystem functioning caused by non-native primary producers in marine systems. In this study we explored the effects of the invasion by the two filamentous red algae Heterosiphonia japonica and Bonnemaisonia hamifera on the primary production of seaweed communities by using single and mixed cultures of non-native and native red algae. The experiments were conducted both in the presence and absence of herbivores. Biomass production of the invaded community increased more than four times in mixed cultures with H. japonica, while introduction by B. hamifera had no significant effect. The different impact on community production could be explained by differences in life history strategies between the invaders; H. japonica grew considerably faster than the native seaweeds which directly increased the community production, while B. hamifera showed a relatively slow growth rate and therefore had no effect. From previous studies it is known that B. hamifera produces a highly deterrent, but also costly, chemical defence. The assessment of survival and growth of a native generalist herbivore further corroborated that the biomass produced by B. hamifera constitutes a very low-quality food, whereas the performance of herbivores on a diet of H. japonica was comparable to that on native algal diets. In summary, this study demonstrates that successful invaders belonging to the same functional group (filamentous red algae) may have distinctly different impacts on productivity in the recipient community, depending on their specific life history traits.

Invasion by mobile aquatic consumers enhances secondary production and increases top-down control of lower trophic levels.

Increased biological diversity due to invasion by non-indigenous species (NIS) is a global phenomenon with potential effects on trophic interactions and ecosystem processes in the invaded habitat. We assessed the effects of resource availability and invasion of three non-indigenous invertebrate grazers (two crustaceans and a snail) on secondary production, relative dominance of NIS grazers and resource depletion in experimental freshwater mesocosms. The relative dominance of NIS grazers increased with increasing initial resource availability, although the effect was largest for one of the three species. The effect was due to the fact that all the included non-indigenous grazers were able to expand their populations quickly in response to resource addition. For the most dominating species, the increased grazer diversity due to invasion in turn resulted in higher production of grazer biomass and a more efficient depletion of the periphyton resource. The effect was largest at high initial resource availability, where NIS dominance was most pronounced. Our results show that an invasion-induced increase in species diversity can increase resource depletion and consequently production, but that the effect depends on identity of the introduced species. The results also suggest that properties of the recipient system, such as resource availability, can modulate ecosystem effects of NIS by affecting invader success and dominance.

Stress ecology in fucus: abiotic, biotic and genetic interactions.

Stress regimes defined as the synchronous or sequential action of abiotic and biotic stresses determine the performance and distribution of species. The natural patterns of stress to which species are more or less well adapted have recently started to shift and alter under the influence of global change. This was the motivation to review our knowledge on the stress ecology of a benthic key player, the macroalgal genus Fucus. We first provide a comprehensive review of the genus as an ecological model including what is currently known about the major lineages of Fucus species with respect to hybridization, ecotypic differentiation and speciation; as well as life history, population structure and geographic distribution. We then review our current understanding of both extrinsic (abiotic/biotic) and intrinsic (genetic) stress(es) on Fucus species and how they interact with each other. It is concluded that (i) interactive stress effects appear to be equally distributed over additive, antagonistic and synergistic categories at the level of single experiments, but are predominantly additive when averaged over all studies in a meta-analysis of 41 experiments; (ii) juvenile and adult responses to stress frequently differ and (iii) several species or particular populations of Fucus may be relatively unaffected by climate change as a consequence of pre-adapted ecotypes that collectively express wide physiological tolerences. Future research on Fucus should (i) include additional species, (ii) include marginal populations as models for responses to environmental stress; (iii) assess a wider range of stress combinations, including their temporal fluctuations; (iv) better differentiate between stress sensitivity of juvenile versus adult stages; (v) include a functional genomic component in order to better integrate Fucus' ecological and evolutionary responses to stress regimes and (vi) utilize a multivariate modelling approach in order to develop and understand interaction networks.

Higher resistance to herbivory in introduced compared to native populations of a seaweed.

Non-indigenous species (NIS) are important components of global change, and in order to manage such species it is important to understand which factors affect their success. Interactions with enemies in the new range have been shown to be important for the outcome of introductions, but thus far most studies on NIS-enemy interactions have considered only specialist herbivores in terrestrial systems. Here we present the results from the first biogeographic study that compares herbivore resistance between populations in the native and new region of a non-indigenous seaweed. We show that low consumption of the non-indigenous seaweed by a generalist herbivore is caused by higher chemical defence levels and herbivore resistance in the new range-and not by the failure of the herbivore to recognise the non-indigenous seaweed as a suitable host. Since most seaweed-herbivore interactions are dominated by generalist herbivores, this pattern could be common in marine communities. Our results also reveal that traits used to predict the invasive potential of species, such as their resistance to enemies, can change during the invasion process, but not always in the way predicted by dominant theories.

Fouling mediates grazing: intertwining of resistances to multiple enemies in the brown alga Fucus vesiculosus.

Macroalgae have to cope with multiple natural enemies, such as herbivores and epibionts. As these are harmful for the host, the host is expected to show resistance to them. Evolution of resistance is complicated by the interactions among the enemies and the genetic correlations among resistances to different enemies. Here, we explored genetic variation in resistance to epibiosis and herbivory in the brown alga Fucus vesiculosus, both under conditions where the enemies coexisted and where they were isolated. F. vesiculosus showed substantial genetic variation in the resistance to both epibiosis and grazing. Grazing pressure on the alga was generally lower in the presence than in the absence of epibiota. Furthermore, epibiosis modified the susceptibility of different algal genotypes to grazing. Resistances to epibiosis and grazing were independent when measured separately for both enemies but positively correlated when both these enemies coexisted. Thus, when the enemies coexisted, the fate of genotypes with respect to these enemies was intertwined. Genotypic correlation between phlorotannins, brown-algal phenolic secondary metabolites, and the amount of epibiota was negative, indicating that these compounds contribute to resistance to epibiosis. In addition, phlorotannins correlated also with the resistance to grazing, but this correlation disappeared when grazing occurred in the absence of epibiota. This indicates that the patterns of selection for the type of the resistance as well as for the resistance traits vary with the occurrence patterns of the enemies.

Increased chemical resistance explains low herbivore colonization of introduced seaweed.

The success of introduced species is often attributed to release from co-evolved enemies in the new range and a subsequent decreased allocation to defense (EICA), but these hypotheses have rarely been evaluated for systems with low host-specificity of enemies. Here, we compare herbivore utilization of the brown seaweed, Fucus evanescens, and its coexisting competitors both in its native and new ranges, to test certain predictions derived from these hypotheses in a system dominated by generalist herbivores. While F. evanescens was shown to be a preferred host in its native range, invading populations supported a less diverse herbivore fauna and it was less preferred in laboratory choice experiments with important herbivores, when compared to co-occurring seaweeds. These results are consistent with the enemy release hypothesis, despite the fact that the herbivore communities in both regions were mainly composed of generalist species. However, in contrast to the prediction of EICA, analysis of anti-grazing compounds indicated a higher allocation to defense in introduced compared to native F. evanescens. The results suggest that the invader is subjected to less intense enemy control in the new range, but that this is due to an increased allocation to defense rather than release from specialized herbivores. This indicates that increased resistance to herbivory might be an important strategy for invasion success in systems dominated by generalist herbivores.

Chemical settlement inhibition versus post-settlement mortality as an explanation for differential fouling of two congeneric seaweeds.

It has been proposed that seaweed secondary metabolites, e.g. brown algal phlorotannins, may have an ecologically important function as a chemical defence against epiphytes, by acting against colonisation of epiphytic organisms. We tested whether the low epiphytic abundance on the invasive brown seaweed Fucus evanescens, compared to the congeneric F. vesiculosus, is due to a more effective chemical defence against epiphyte colonisation. A field survey of the distribution of the common fouling organism Balanus improvisus (Cirripedia) showed that the abundance was consistently lower on F. evanescens than on F. vesiculosus. However, contrary to expectations, results from experimental studies indicated that F. vesiculosus has a more effective anti-settlement defence than F. evanescens. In settlement experiments with intact fronds of the two Fucus species, both species deterred settlement by barnacle larvae, but settlement was lower on F. vesiculosus both in choice and no-choice experiments. Phlorotannins from F. vesiculosus also had a stronger negative effect on larval settlement and were active at a lower concentration than those from F. evanescens. The results show that Fucus phlorotannins have the potential to inhibit settlement of invertebrate larvae, but that settlement inhibition cannot explain the lower abundance of the barnacle Balanus improvisus on F. evanescens compared to F. vesiculosus. Assessment of barnacle survival in the laboratory and in the field showed that this pattern could instead be attributed to a higher mortality of newly settled barnacles. Observation suggests that the increased mortality was due to detachment of young barnacles from the seaweed surface. This shows that the antifouling mechanism of F. evanescens acts on post-settlement stages of B. improvisus.