Johan S. Eklöf

Confronting Feedbacks of Degraded Marine Ecosystems

In many coastal areas, marine ecosystems have shifted into contrasting states having reduced ecosystem services (hereafter called degraded). Such degraded ecosystems may be slow to revert to their original state due to new ecological feedbacks that reinforce the degraded state. A better understanding of the way human actions influence the strength and direction of feedbacks, how different feedbacks could interact, and at what scales they operate, may be necessary in some cases for successful management of marine ecosystems. Here we synthesize interactions of critical feedbacks of the degraded states from six globally distinct biomes: coral reefs, kelp forests, seagrass beds, shallow soft sediments, oyster reefs, and coastal pelagic food webs. We explore to what extent current management captures these feedbacks and propose strategies for how and when (that is, windows of opportunity) to influence feedbacks in ways to break the resilience of the degraded ecosystem states. We conclude by proposing some challenges for future research that could improve our understanding of these issues and emphasize that management of degraded marine states will require a broad social–ecological approach to succeed.

Consumers mediate the effects of experimental ocean acidification and warming on primary producers

It is well known that ocean acidification can have profound impacts on marine organisms. However, we know little about the direct and indirect effects of ocean acidification and also how these effects interact with other features of environmental change such as warming and declining consumer pressure. In this study, we tested whether the presence of consumers (invertebrate mesograzers) influenced the interactive effects of ocean acidification and warming on benthic microalgae in a seagrass community mesocosm experiment. Net effects of acidification and warming on benthic microalgal biomass and production, as assessed by analysis of variance, were relatively weak regardless of grazer presence. However, partitioning these net effects into direct and indirect effects using structural equation modeling revealed several strong relationships. In the absence of grazers, benthic microalgae were negatively and indirectly affected by sediment-associated microalgal grazers and macroalgal shading, but directly and positively affected by acidification and warming. Combining indirect and direct effects yielded no or weak net effects. In the presence of grazers, almost all direct and indirect climate effects were nonsignificant. Our analyses highlight that (i) indirect effects of climate change may be at least as strong as direct effects, (ii) grazers are crucial in mediating these effects, and (iii) effects of ocean acidification may be apparent only through indirect effects and in combination with other variables (e.g., warming). These findings highlight the importance of experimental designs and statistical analyses that allow us to separate and quantify the direct and indirect effects of multiple climate variables on natural communities.

Female egg investment in relation to male sexual traits and the potential for transgenerational effects in sexual selection

Life-history theory predicts that individuals should increase their reproductive effort when the fitness return from reproduction is high. Females mated with high-quality males are therefore expected to have higher investment than females mated with low-quality males, which could bias estimates of paternal effects. Investigating the traits females use in their allocation decisions and the aspects of reproduction that are altered is essential for understanding how sexual selection is affected. We studied the potential for differential female allocation in a captive population of a precocial bird, the Chinese quail, Coturnix chinensis. Females paired with males with large sexual ornaments laid larger, but not more, eggs than females paired with males with small sexual ornaments. Furthermore, female egg mass was also significantly positively affected by male testis size, probably via some unknown effect of testis size on male phenotype. Testis size and ornament size were not correlated. Thus, both primary and secondary male sexual traits could be important components of female allocation decisions. Experimental manipulation of hormone levels during embryonic development showed that both male and female traits influencing female egg size were sensitive to early hormone exposure. Differences in prenatal hormone exposure as a result of maternal steroid allocation to eggs may explain some of the variation in reproductive success among individuals, with important implications for non-genetic transgenerational effects in sexual selection.

Biodiversity mediates top–down control in eelgrass ecosystems: a global comparative‐experimental approach

Nutrient pollution and reduced grazing each can stimulate algal blooms as shown by numerous experiments. But because experiments rarely incorporate natural variation in environmental factors and biodiversity, conditions determining the relative strength of bottom-up and top-down forcing remain unresolved. We factorially added nutrients and reduced grazing at 15 sites across the range of the marine foundation species eelgrass (Zostera marina) to quantify how top-down and bottom-up control interact with natural gradients in biodiversity and environmental forcing. Experiments confirmed modest top-down control of algae, whereas fertilisation had no general effect. Unexpectedly, grazer and algal biomass were better predicted by cross-site variation in grazer and eelgrass diversity than by global environmental gradients. Moreover, these large-scale patterns corresponded strikingly with prior small-scale experiments. Our results link global and local evidence that biodiversity and top-down control strongly influence functioning of threatened seagrass ecosystems, and suggest that biodiversity is comparably important to global change stressors.

Effects of Altered Offshore Food Webs on Coastal Ecosystems Emphasize the Need for Cross-Ecosystem Management

By mainly targeting larger predatory fish, commercial fisheries have indirectly promoted rapid increases in densities of their prey; smaller predatory fish like sprat, stickleback and gobies. This process, known as mesopredator release, has effectively transformed many marine offshore basins into mesopredator-dominated ecosystems. In this article, we discuss recent indications of trophic cascades on the Atlantic and Baltic coasts of Sweden, where increased abundances of mesopredatory fish are linked to increased nearshore production and biomass of ephemeral algae. Based on synthesis of monitoring data, we suggest that offshore exploitation of larger predatory fish has contributed to the increase in mesopredator fish also along the coasts, with indirect negative effects on important benthic habitats and coastal water quality. The results emphasize the need to rebuild offshore and coastal populations of larger predatory fish to levels where they regain their control over lower trophic levels and important links between offshore and coastal systems are restored.

Experimental climate change weakens the insurance effect of biodiversity

Ecosystems are simultaneously affected by biodiversity loss and climate change, but we know little about how these factors interact. We predicted that climate warming and CO (2) -enrichment should strengthen trophic cascades by reducing the relative efficiency of predation-resistant herbivores, if herbivore consumption rate trades off with predation resistance. This weakens the insurance effect of herbivore diversity. We tested this prediction using experimental ocean warming and acidification in seagrass mesocosms. Meta-analyses of published experiments first indicated that consumption rate trades off with predation resistance. The experiment then showed that three common herbivores together controlled macroalgae and facilitated seagrass dominance, regardless of climate change. When the predation-vulnerable herbivore was excluded in normal conditions, the two resistant herbivores maintained top-down control. Under warming, however, increased algal growth outstripped control by herbivores and the system became algal-dominated. Consequently, climate change can reduce the relative efficiency of resistant herbivores and weaken the insurance effect of biodiversity.

Use of vision in prey detection by brown long-eared bats Plecotus auritus

Abstract We investigated the ability of brown long-eared bats to make use of visual cues when searching for food. By using petri dishes containing mealworms that were subjected to different levels of illumination, we presented four bats with different sensory cues: visual, sonar and a combination of these. The bats preferred situations where both sonar cues and visual cues were available, and the visual information was more important than the sonar cues. The bats did, however, emit echolocation calls throughout the experiments. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Spatially extended habitat modification by intertidal reef-building bivalves has implications for consumer-resource interactions

Ecosystem engineers can strongly modify habitat structure and resource availability across space. In theory, this should alter the spatial distributions of trophically interacting species. In this article, we empirically investigated the importance of spatially extended habitat modification by reef-building bivalves in explaining the distribution of four avian predators and their benthic prey in the Wadden Sea—one of the world’s largest intertidal soft-sediment ecosystems. We applied Structural Equation Modeling to identify important direct and indirect interactions between the different components of the system. We found strong spatial gradients in sediment properties into the surrounding area of mixed blue mussel (Mytilus edulis) and Pacific oyster (Crassostrea gigas) reefs, indicating large-scale (100s of m) engineering effects. The benthic community was significantly affected by these gradients, with the abundance of several important invertebrate prey species increasing with sediment organic matter and decreasing with distance to the reefs. Distance from the reef, sediment properties, and benthic food abundance simultaneously explained significant parts of the distribution of oystercatchers (Haematopus ostralegus), Eurasian curlews (Numenius arquata), and bar-tailed godwits (Limosa lapponica). The distribution of black-headed gulls (Chroicocephalus ridibundus)—a versatile species with many diet options—appeared unaffected by the reefs. These results suggest that intertidal reef builders can affect consumer-resource dynamics far beyond their own boundaries, emphasizing their importance in intertidal soft-bottom ecosystems like the Wadden Sea.

The fundamental role of ecological feedback mechanisms for the adaptive management of seagrass ecosystems - a review

Seagrass meadows are vital ecosystems in coastal zones worldwide, but are also under global threat. One of the major hurdles restricting the success of seagrass conservation and restoration is our limited understanding of ecological feedback mechanisms. In these ecosystems, multiple, self‐reinforcing feedbacks can undermine conservation efforts by masking environmental impacts until the decline is precipitous, or alternatively they can inhibit seagrass recovery in spite of restoration efforts. However, no clear framework yet exists for identifying or dealing with feedbacks to improve the management of seagrass ecosystems. Here we review the causes and consequences of multiple feedbacks between seagrass and biotic and/or abiotic processes. We demonstrate how feedbacks have the potential to impose or reinforce regimes of either seagrass dominance or unvegetated substrate, and how the strength and importance of these feedbacks vary across environmental gradients. Although a myriad of feedbacks have now been identified, the co‐occurrence and likely interaction among feedbacks has largely been overlooked to date due to difficulties in analysis and detection. Here we take a fundamental step forward by modelling the interactions among two distinct above‐ and belowground feedbacks to demonstrate that interacting feedbacks are likely to be important for ecosystem resilience. On this basis, we propose a five‐step adaptive management plan to address feedback dynamics for effective conservation and restoration strategies. The management plan provides guidance to aid in the identification and prioritisation of likely feedbacks in different seagrass ecosystems.

No lunar phobia in swarming insectivorous bats (family Vespertilionidae)

The purpose of this study was to evaluate the impact of moonlight on the swarming activity of bats at an abandoned mine in southern Sweden. The mine serves as a hibernaculum for six species of insectivorous bats. Swarming activity at one of the mine entrances was measured on 12 nights between 1 August and 8 September 2000, by electronically counting the number of bats passing through an opening (about 30×40 cm) leading to a mine tunnel. The number of bat passes outside the mine was also counted, using a bat detector. Most bats were males, as confirmed by mist-netting outside the mine. Low frequency vocalizations, indicating territorial interactions, were heard frequently. The number of bats entering the mine was closely correlated with the number of bats flying outside, and neither was affected by moonlight. We conclude that the insectivorous bats at high latitudes may not have been exposed to significant nocturnal predator pressure, leading to the evolution of lunar phobia, as have many tropical bats.