An experimental approach to assessing the impact of ecosystem engineers on biodiversity and ecosystem functions.

<p>Geomorphology has long considered the role of abiotic factors in geomorphic processes, including tectonics, geology, climate and relief, as well as humans impact upon them. Biotic factors however, including not only plants but also bacteria and protists, biofilms, fungi, insects, invertebrates, and animals are increasingly recognized as governing geomorphic processes on many spatial and temporal scales. We argue that if fauna are important as geological agents, then understanding the complex response of geomorphic systems to fauna is necessary to understand the past, present and future of the fluvial environment. It is not surprising that studies of the Late Quaternary evolution of fluvial morphodynamics have largely focused upon changes in the sedimentary soil-sediment sequences that result from climate change; (ii) vegetation change; and/or (iii) human impacts. Reconstruction of vegetation and climate from pollen and other records facilitates these analyses. But if animals are shown to be an important influence on geomorphic processes today, then it is quite possible that they were also important historically. For example, conclusive interpretation of Holocene river changes may be limited because of an incomplete or partial account of the presence and/or absence of data on the role of ecosystem engineers in modifying the riparian and aquatic ecosystems, including hydro-geomorphic processes. DNA found within historical deposits may be used to constrain the role of past ecosystem engineers. Analysis of ancient environmental DNA up to date includes palaeo-environmental DNA from sedimentary deposits (sedaDNA) from disseminated genetic material found within sedimentary archives, including paleo-dietary ancient DNA. Here, we use an analogue study investigating the present hydro-geomorphic and biogeochemical changes that the ecosystem engineer beaver (Castor fiber) creates at four sites in central Europe to better understand and quantify the effects of beaver ecosystem engineering on a seasonal to decadal scale. We utilize these results to interpret the chrono-stratigraphy of two Holocene beaver sites, including macro-fossil and sedaDNA sampling, and test for the first time if sedaDNA can support the investigation of beaver-induced palao-environmental conditions in river floodplains. We find that sedaDNA data and other palaeo-botanical proxies complement each other showing wider diversity of species than if the methods are used separately. However, care must be taken with regards of experimental setup, and further investigation into the effects of transport processes and/or quantitative representativeness is needed.</p>

The problem of assessing the impact of biodiversity on the climate-regulating functions of forests is fundamental. It is of great applied importance for sustainable forest management in the context of global climate change. On the one hand, climate change affects biodiversity; on the other hand, biodiversity underlies the mechanisms of adaptation of forests and society to these changes, because it is a provider of all ecosystem functions. This article aims to discuss scientific issues currently faced by scientists, such as the relationships between biodiversity and climate-regulating functions of forests, and to outline the perspective of the studies. There are numerous studies that describe the influence of certain plant and animal species – ecosystem engineers – on the ecosystem, including climate-regulating functions of forests. However, we lack estimates of the combined effect of the diversity of biota of different trophic levels and groups on the completeness of the implementation of climate-regulating functions of forests of different types/at different succession stages. We emphasise the importance of accounting for such estimates as taxonomic, including genetic, and the functional and structural diversity of forests. We considered various concepts of forest management, taking into account the conservation and restoration of biodiversity. The most important aspect of this problem is estimates and forecasts of interrelationships (trade-offs and synergies) between climate-regulating and other ecosystem functions of forests characterised by different levels of biodiversity, with their natural development and with the combined impact of various natural and anthropogenic factors on forests, including climate change, fires, and forest management regimes. Integration of mathematical models is a promising approach to assess and predict the dynamics of relationships between various ecosystem functions of forests.

Summary Intertidal zones have a key ecological role in estuaries. The area covered by the intertidal mudflats in the Seine estuary (NW France) was divided by 3 over the last centuries. The preservation of these areas is a priority but requires a detailed understanding of their functioning. The latter is complex and notably depends on the presence and activity of benthic macrofauna, which can act as ecosystem engineers, strongly affecting the other organisms. The aim of this study was to assess the influence of the two predominant ecosystem engineers in the Seine Estuary mudflats (Hediste diversicolor and Scrobicularia plana) on the dissolved and sedimentary organic matter (OM) quality. The role of the ecosystem engineers on the quality of the sedimentary OM was evaluated through the analysis of complementary lipid biomarkers. The characteristics of the dissolved OM (DOM) from the corresponding interstitial waters were concomitantly determined by spectroscopic analyses. Even though it was shown that H. diversicolor and S. plana play a major role on the microphytobenthos dynamics, the seasonality was shown to have a more pronounced effect on the sedimentary OM characteristics than benthic organisms. In contrast, the DOM was suggested to be sensitive and reactive to the ecosystem engineer activities.

Abstract. Ecosystem engineers (EEs) are present in every environment and are known to strongly influence ecological processes and thus shape the distribution of species and resources. In this study, we assessed the direct and indirect effect of two EEs (perennial shrubs and ant nests), individually and combined, on the composition and function of arid soil bacterial communities. To that end, topsoil samples were collected in the Negev desert highlands during the dry season from four patch types: (1) barren soil; (2) under shrubs; (3) near ant nests; or (4) near ant nests situated under shrubs. The bacterial community composition and potential functionality were evaluated in the soil samples (14 replicates per patch type) using 16S rRNA gene amplicon sequencing together with physico-chemical measures of the soil. We have found that the EEs affected the community composition differently. Barren patches supported a soil microbiome, dominated by Rubrobacter and Proteobacteria, while in EE patches Deinococcus-Thermus dominated. The presence of the EEs similarly enhanced the abundance of phototrophic, nitrogen cycle, and stress-related genes. In addition, the soil characteristics were altered only when both EEs were combined. Our results suggest that arid landscapes foster unique communities selected by patches created by each EE(s), solo or in combination. Although the communities' composition differs, they support similar potential functions that may have a role in surviving the harsh arid conditions. The combined effect of the EEs on soil microbial communities is a good example of the hard-to-predict non-additive features of arid ecosystems that merit further research.

Effects of plant functional diversity induced by grazing and soil properties on above- and belowground biomass in a semiarid grassland

Biodiversity is crucial for supporting ecosystem functioning, yet some species play a disproportionate role in maintaining complex ecological processes. Ecosystem engineers are species that directly influence ecosystems by modifying biophysical environments, creating novel habitats, altering biogeochemical cycles, increasing biodiversity and/or modulating ecological processes. Although these species may substantially influence ecosystem functioning, their role is often overlooked and difficult to quantify. Understanding the status, dynamics and trends of ecosystem engineers is essential for mitigating biodiversity loss and maintaining healthy ecosystems. This review reveals the common but underappreciated roles that ecosystem engineers play in ecosystem functioning across many different taxa, biomes and ecological processes. We first synthesise how knowledge of ecosystem engineering improves our understanding of species interactions and the ecological processes underlying both ecosystem functioning and BEF relationships. We provide a conceptual model for addressing the effects of ecosystem engineers in BEF research and ecological dynamics. We provide a ‘how to’ analytical framework for monitoring and quantifying changes in ecosystem engineers and their effects on ecosystem functioning. This framework includes (i) what variables to measure, how and at which scale; (ii) experiments involving species exclusion or removal, introduction and comparative designs when experimental manipulation is not feasible and (iii) statistical, data‐driven and theory‐driven models. We discuss how to leverage ecosystem engineering in the context of current global change and ecosystem restoration efforts. Including ecosystem engineers in conservation and restoration programs, when implemented in the appropriate context and supported by an understanding of ecological mechanisms and processes, may be crucial for sustaining biological diversity and functional ecosystems. Read the free Plain Language Summary for this article on the Journal blog.

Plant functional diversity mediates the effects of vegetation and soil properties on community-level plant nitrogen use in the restoration of semiarid sandy grassland

Ecosystem engineers alter, and can be influenced in turn by, the ecosystems they live in. Woodpeckers choose foraging and nesting sites based, in part, on food availability. Once abandoned, these cavities, particularly within areas of high forage, may be crucial to secondary cavity‐nesting birds otherwise limited by cavities formed through decay. Our study examined factors that influence the nesting success of primary cavity nesters and the subsequent impact on secondary cavity‐nesting birds. Using 5 years of point count data, we monitored the outcomes of cavity‐nesting birds in South Texas. We used logistic‐exposure models to predict daily survival rates based on cavity metrics and used woodpecker foraging trends and insect surveys to determine if nesting where woodpeckers actively forage benefits secondary cavity‐nesting birds. Both woodpeckers and secondary cavity nesters shared predictors of daily survival; nests were more successful in cavities with small openings in minimally decayed trees. All secondary cavity nesters had higher probabilities of success when nesting in an abandoned woodpecker cavity, opposed to ones formed by decay. Woodpeckers tended to forage in areas with higher‐than‐average levels of the insect orders Coleoptera, Hymenoptera, and Orthoptera, and secondary cavity nesters had higher rates of success when nesting in these areas. Our results suggest abandoned woodpecker cavities may be constructed in a way that directly benefit secondary cavity nesters. Additionally, we suggest an interplay between these ecosystem engineers, food availability, and secondary cavity nesters: Woodpeckers engineer superior nesting cavities in areas where food is more abundant, and the resultant cavities in areas of high forage may benefit local secondary cavity nesters. Our findings indicate that there is still much to be explored in the role of ecosystem engineers, and how they influence local communities on multiple trophic levels.

An important and highly active research agenda has developed at the interface of fluvial geomorphology and ecology that addresses the capacity for vegetation and animals to act as ecosystem engineers within fluvial systems. This paper briefly introduces this research domain and describes the 15 papers that contribute to the special issue on 'Dynamic riverine landscapes: the role of ecosystem engineers'. The papers illustrate the breadth of research activity at this interface, investigating the influence of a range of ecosystem engineering organisms through a combination of field study, laboratory experiments, numerical simulation and analysis of remotely sensed data. Together, the papers address a series of key themes: conceptual frameworks for feedbacks between aquatic biota, hydraulics, sediment dynamics and nutrient dynamics and their quantification through experimental and field research; the potential contribution of ecosystem engineering species to assist river recovery and restoration; and the contribution of riparian vegetation to bank stability and morphodynamics across a range of spatio‐temporal scales. Copyright © 2015 John Wiley & Sons, Ltd.

Ecosystem engineers are organisms that change the distribution of materials and energy in the abiotic environment, usually creating and maintaining new habitat patches in the landscape. Such changes in habitat conditions have been widely documented to affect the distributions and performances of other species but up to now no studies have addressed how such effects can impact the biotically driven physicochemical processes associated with these landscapes, or ecosystem functions. Based on the widely accepted positive relationship between species diversity and ecosystem functions, we propose that the effects of ecosystem engineers on other species could have an impact on ecosystem functions via two mutually inclusive mechanisms: (1) by adding new species into landscapes, hence increasing species diversity; and (2) by improving the performances of species already present in the landscape. To test these hypotheses, we focused on the effects of a high-Andean ecosystem engineer, the cushion plant Azorella monantha, by comparing the accumulation of plant biomass and nitrogen fixed in plant tissues as species richness increases in landscapes with and without the engineer species. Our results show that both ecosystem functions increased with species richness in both landscape types, but landscapes including A. monantha cushions reached higher outcomes of plant biomass and nitrogen fixed in plant tissues than landscapes without cushions. Moreover, our results indicate that such positive effects on ecosystem functions could be mediated by the two mechanisms proposed above. Then, given the conspicuousness of ecosystem engineering in nature and its strong influence on species diversity, and given the well-known relationship between species diversity and ecosystem function, we suggest that the application of the conceptual framework proposed herein to other ecosystems would help to advance our understanding of the forces driving ecosystem functioning.

Cascading effects of long‐term land‐use changes on plant traits and ecosystem functioning

Introduced ecosystem engineers can severely modify the functioning on invaded systems. Species-level effects on ecosystem functioning (EF) are context dependent, but the effects of introduced ecosystem engineers are frequently assessed through single-location studies. The present work aimed to identify sources of context-dependence that can regulate the impacts of invasive ecosystem engineers on ecosystem functioning. As model systems, four locations where the bivalve Ruditapes philippinarum (Adams and Reeve) has been introduced were investigated, providing variability in habitat characteristics and community composition. As a measure of ecosystem engineering, the relative contribution of this species to community bioturbation potential was quantified at each site. The relevance of bioturbation to the local establishment of the mixing depth of marine sediments (used as a proxy for EF) was quantified in order to determine the potential for impact of the introduced species at each site. We found that R. philippinarum is one of the most important bioturbators within analysed communities, but the relative importance of this contribution at the community level depended on local species composition. The net contribution of bioturbation to the establishment of sediment mixing depths varied across sites depending on the presence of structuring vegetation, sediment granulometry and compaction. The effects of vegetation on sediment mixing were previously unreported. These findings indicate that the species composition of invaded communities, and the habitat characteristics of invaded systems, are important modulators of the impacts of introduced species on ecosystem functioning. A framework that encompasses these aspects for the prediction of the functional impacts of invasive ecosystem engineers is suggested, supporting a multi-site approach to invasive ecology studies concerned with ecosystem functioning.

The positive relationship between biodiversity and ecosystem functioning is mainly derived from studies concerning primary producers, whereas a generalization of this relationship for higher trophic levels is more difficult. Furthermore, most evidence of the biodiversity–ecosystem functioning relationship is derived from experiments manipulating only one trophic level and, as a consequence, interactive diversity effects at multiple trophic levels have mostly been ignored. Here, we performed a mesocosm experiment in which we manipulated functional group diversity at two trophic levels (primary and secondary consumers) applying a full‐factorial design. More specifically, we asked whether 1) predator functional diversity affects prey mortality rates, 2) prey functional diversity affects prey mortality rates, 3) whether there are interactive effects of simultaneous diversity changes at both trophic levels. For each trophic level we used two functional groups, i.e. organisms belonging to two different habitat domains: at the higher trophic position 1) a ground foraging spider species and 2) a spider species foraging in the vegetation canopy and at the lower trophic position 3) a ground living cricket species and 4) leafhoppers living in the vegetation canopy. Increasing predator functional group diversity increased prey mortality by 53%, and increasing prey functional group diversity increased prey mortality by 24%. Further, prey mortality was highest at the uppermost level of functional group diversity (142% increase in prey mortality compared to single prey and predator functional diversity), most likely due to resource partitioning between the predators. This finding demonstrates that a multi‐trophic perspective is necessary, and that previous studies focusing on only one trophic level have most likely underestimated the strength of the relationship between biodiversity and ecosystem functioning.

Organisms as cooperative ecosystem engineers in intertidal flats

Ecosystem engineers are organisms whose presence or activity alters their physical surroundings or changes the flow of resources, thereby creating or modifying habitats. Because ecosystem engineers affect communities through environmentally mediated interactions, their impact and importance are likely to shift across environmental stress gradients. We hypothesize that in extreme physical environments, ecosystem engineers that ameliorate physical stress are essential for ecosystem function, whereas in physically benign environments where competitor and consumer pressure is typically high, engineers support ecosystem processes by providing competitor- or predator-free space. Important ecosystem engineers alleviate limiting abiotic and biotic stresses, expanding distributional limits for numerous species, and often form the foundation for community development. Because managing important engineers can protect numerous associated species and functions, we advocate using these organisms as conservation targets, harnessing the benefits of ecosystem engineers in various environments. Developing a predictive understanding of engineering across environmental gradients is important for furthering our conceptual understanding of ecosystem structure and function, and could aid in directing limited management resources to critical ecosystem engineers.