Ecosystem engineering in space and time

The ecosystem engineering concept focuses on how organisms physically change the abiotic environment and how this feeds back to the biota. While the concept was formally introduced a little more than 10 years ago, the underpinning of the concept can be traced back to more than a century to the early work of Darwin. The formal application of the idea is yielding new insights into the role of species in ecosystems and many other areas of basic and applied ecology. Here we focus on how temporal, spatial and organizational scales usefully inform the roles played by ecosystem engineers and their incorporation into broader ecological contexts. Two particular, distinguishing features of ecosystem engineers are that they affect the physical space in which other species live and their direct effects can last longer than the lifetime of the organism – engineering can in essence outlive the engineer. Together, these factors identify critical considerations that need to be included in models, experimental and observational work. The ecosystem engineering concept holds particular promise in the area of ecological applications, where influence over abiotic variables and their consequent effects on biotic communities may facilitate ecological restoration and counterbalance anthropogenic influences.     

Translocations of digging mammals and their potential for ecosystem restoration: a review of goals and monitoring programmes

1. Globally, translocations are commonly used to improve the conservation status of threatened species. There is increasing recognition that translocations of ecosystem engineers also have the potential to restore ecological processes. Digging mammals are often considered to be ecosystem engineers, as their diggings provide shelter for other species and can significantly alter soil properties, with subsequent changes to vegetation.
2. Using Australian species as a case study, we reviewed published and grey literature on digging mammal translocations to determine how often these translocations are conducted to restore ecosystem processes. We documented ecosystem-level monitoring and research efforts, and assessed whether restoration was perceived to be occurring post-release.
3. At least 208 translocations of 24 digging mammal species have been conducted in Australia, with a further 38 planned for the near future. Prior to 2019, only 3% of translocations included a goal relating to the restoration of ecosystem processes associated with digging activities. Nearly a quarter of pre-2019 translocations have been the subject of some form of ecosystem-level monitoring or research, but long-term ecosystem-level monitoring was very rare. In contrast, 74% of the translocations planned for post-2018 include a goal relating to the restoration of ecological processes and most also include plans to conduct ecosystem-level monitoring.
4. Ecosystem restoration was perceived to be occurring for 26% of the pre-2019 translocations. None of the documents we reviewed indicated that ecological degradation had occurred post-translocation, even when declines in other taxa were recorded.
5. The restoration of ecosystem processes is increasingly being identified as a goal for translocation programmes. Where this is the case, we suggest that translocation practitioners include success criteria for the restoration of ecosystem processes, and commit to long-term monitoring designed to detect ecosystem-level effects of translocations.

     

The role of spontaneous vegetation succession in ecosystem restoration: A perspective

Abstract. The paper summarizes ideas which were discussed during the ‘Spontaneous Succession in Ecosystem Restoration’ conference and elaborated through further discussion among the authors. It seeks to promote the integration of scientific knowledge on spontaneous vegetation succession into restoration programs. A scheme illustrating how knowledge of spontaneous succession may be applied to restoration is presented, and perspectives and possible future research on using spontaneous vegetation succession in ecosystem restoration are proposed. It is concluded that when implementing spontaneous succession for ecological restoration the following points must be considered: setting clear aims; evaluation of environmental site conditions; deciding whether spontaneous succession is an appropriate way to achieve the aims; prediction of successional development; monitoring of the results. The need for interdisciplinary approaches and communication between scientists, engineers and decision‐makers is emphasized.     

Multi‐scale habitat modification by coexisting ecosystem engineers drives spatial separation of macrobenthic functional groups

By changing habitat conditions, ecosystem engineers increase niche diversity and have profound effects on the distribution and abundances of other organisms. Although many ecosystems contain several engineering species, it is still unclear how the coexistence of multiple engineers affects the physical habitat and the structure of the community on a landscape scale. Here, we investigated through a large‐scale field manipulation how three coexisting engineers on intertidal flats (cockles Cerastoderma edule; lugworms Arenicola marina; blue mussels Mytilus edulis) influence the functional composition of the local macrobenthic community and what the consequences are at the landscape level. By using biological trait analysis (BTA), we show that on the local scale biogenic changes in sediment accumulation and organic matter content translated into specific shifts in the distribution of functional traits within the community. At a landscape scale, the co‐occurrence of multiple ecosystem engineers resulted in the spatial separation of different functional groups, i.e. different functional groups dominated unique complementary habitats. Our results emphasize the role of co‐occurring multiple engineers in shaping natural communities, thus contributing to a better knowledge of community assembly rules. This understanding can profitably be used to improve ecosystem‐based management and conservation actions.     

Ecosystem engineering as an energy transfer process: a simple agent-based model

Ecosystem engineers are defined as organisms who modulate the availability of resources for themselves and other organisms by physically changing the environment. Ecosystem engineering is a well-recognised ecological interaction, but there is a limited number of general models due to the recent development of the field. Agent-based models are often used to study how organisms respond to changing environments and are suitable for modelling ecosystem engineering. To our knowledge, agent-based methodology has not yet been used to model ecosystem engineering. In this paper, we develop a simple agent-based population dynamics model of ecosystem engineering as an energy transfer process. We apply energy budget approach to conceptually explain how ecosystem engineers transfer energy to the environment and define various types of energy transfers relative to their effects on the engineers and other organisms. We simulate environments with various levels of resource abundance and compare the results of the model without ecosystem engineering agents to the model with ecosystem engineering agents. We find that in environments with higher levels of resources, the presence of ecosystem engineers increases the average carrying capacity and the strength of population fluctuations, while in environments with lower levels of resources, ecosystem engineering mitigates fluctuations, increases average carrying capacity and makes environments more resilient. Finally, we discuss about the further application of agent-based modelling for the theoretical and experimental development of the ecosystem engineering concept.     

采矿地的生态恢复技术

矿藏开采给生态环境带来严重破坏,矿地与尾矿的自然恢复是相当缓慢的,有些甚至是不大可能自然恢复的,本文总结了在过去20年全球发展起来的各种生态恢复技术措施,包括基质的改良,优良物种的选择等,并总结出一套人工恢复的一般步骤,分析了目前所存在的评判人工生态恢复成功与否的几种评价体系的优劣。     

采矿地的生态恢复技术

矿藏开采给生态环境带来严重破坏.矿地与尾矿的自然恢复是相当缓慢的,有些甚至是不大可能自然恢复的.本文总结了在过去20年全球发展起来的各种生态恢复技术措施,包括基质的改良,优良物种的选择等,并总结出一套人工恢复的一般步骤,分析了目前所存在的评判人工生态恢复成功与否的几种评价体系的优劣.     

The Semiglades: The Collision of Restoration,Social Values,and the Ecosystem Concept

Defining success targets in restoration and how social values affect them are two commonly discussed issues in restoration today. We believe that how success is commonly defined—with vague terms such as “healthy ecosystem” or cited as a return to a previous, historic state—needs to be reevaluated. With the increasing number of novel ecosystems, there is an increasing conflict between the ecosystem concept, social values, and restoration. This arises from the fact that ecosystems are defined by the values of the scientists describing them, necessarily constraining the ecosystem to a generally static concept. It is not directly the concept, but how it is perceived through our filter of social values that represses the creativity and innovation needed in restoration today. Within restoration, we feel that the ecosystem concept does a disservice by ignoring the increasing number of novel systems, and that hinders real progress in a time when hesitation can be costly. To best illustrate this, we offer the example of restoration of the Florida Everglades and how it has become a novel system in pattern and process. We suggest renaming the Everglades “The Semiglades” in hopes of opening a dialog to expose social/ecosystem biases and include novel landscapes in management and planning.     

Restoration Success: How Is It Being Measured?

The criteria of restoration success should be clearly established to evaluate restoration projects. Recently, the Society of Ecological Restoration International (SER) has produced a Primer that includes ecosystem attributes that should be considered when evaluating restoration success. To determine how restoration success has been evaluated in restoration projects, we reviewed articles published in Restoration Ecology (Vols. 1[1]–11[4]). Specifically, we addressed the following questions: (1) what measures of ecosystem attributes are assessed and (2) how are these measures used to determine restoration success. No study has measured all the SER Primer attributes, but most studies did include at least one measure in each of three general categories of the ecosystem attributes: diversity, vegetation structure, and ecological processes. Most of the reviewed studies are using multiple measures to evaluate restoration success, but we would encourage future projects to include: (1) at least two variables within each of the three ecosystem attributes that clearly related to ecosystem functioning and (2) at least two reference sites to capture the variation that exist in ecosystems.     

The Use of Extant Non-Indigenous Tortoises as a Restoration Tool to Replace Extinct Ecosystem Engineers

We argue that the introduction of non-native extant tortoises as ecological replacements for extinct giant tortoises is a realistic restoration management scheme, which is easy to implement. We discuss how the recent extinctions of endemic giant Cylindraspis tortoises on the Mascarene Islands have left a legacy of ecosystem dysfunction threatening the remnants of native biota, focusing on the island of Mauritius because this is where most has been inferred about plant–tortoise interactions. There is a pressing need to restore and preserve several Mauritian habitats and plant communities that suffer from ecosystem dysfunction. We discuss ongoing restoration efforts on the Mauritian offshore Round Island, which provide a case study highlighting how tortoise substitutes are being used in an experimental and hypothesis-driven conservation and restoration project. The immediate conservation concern was to prevent the extinction and further degradation of Round Island's threatened flora and fauna. In the long term, the introduction of tortoises to Round Island will lead to valuable management and restoration insights for subsequent larger-scale mainland restoration projects. This case study further highlights the feasibility, versatility and low-risk nature of using tortoises in restoration programs, with particular reference to their introduction to island ecosystems. Overall, the use of extant tortoises as replacements for extinct ones is a good example of how conservation and restoration biology concepts applied at a smaller scale can be microcosms for more grandiose schemes and addresses more immediate conservation priorities than large-scale ecosystem rewilding projects.     

Rewilding the Scottish Highlands: Do Wild Boar,Sus scrofa,Use a Suitable Foraging Strategy to be Effective Ecosystem Engineers?

Ecosystem engineers are increasingly being reintroduced to restore ecological processes in restoration and rewilding projects. To predict and adaptively manage the impact of such species their behavioral ecology must be understood and quantified. Rooting behavior by wild boar qualifies them as ecosystem engineers due to their impact on vegetation disturbance regimes. The behavioral foraging ecology of wild boar was quantified in a fenced area in the Scottish Highlands in order to provide some of the understanding necessary to predict their ability to affect ecosystem restoration. Five wild boar were monitored within a 125 ha fenced area using Global Positioning System (GPS) collars and behavioral monitoring over a 12‐month period. Their activity budget, the relationship between foraging behavior and vegetation communities, and how these relationships vary between seasons was investigated. The results indicate that wild boar invested approximately four more hours daily to rooting during the autumn and winter than the spring and summer. During the spring and summer, grazing was the dominant foraging behavior (approximately 28% of foraging period) while rooting dominated in autumn and winter (approximately 76% of foraging period). Deep rooting behavior is particularly associated with bracken‐dominated communities. Associations between rooting, vegetation community, and season will have a strong influence on the spatial and temporal distribution of rooting behavior. This variation could have important implications for the impacts of boar on vegetation community dynamics. These results detail some of wild boar's ecosystem engineering behaviors; however, further research is required to consider the wider impacts of a full reintroduction.     

Jump‐starting coastal wetland restoration: a comparison of marsh and mangrove foundation species

During coastal wetland restoration, foundation plant species are critical in creating habitat, modulating ecosystem functions, and supporting ecological communities. Following initial hydrologic restoration, foundation plant species can help stabilize sediments and jump‐start ecosystem development. Different foundation species, however, have different traits and environmental tolerances. To understand how these traits and tolerances impact restoration trajectories, there is a need for comparative studies among foundation species. In subtropical and tropical climates, coastal wetland restoration practitioners can sometimes choose between salt marsh and/or mangrove foundation species. Here, we compared the early life history traits and environmental tolerances of two foundation species: (1) a salt marsh grass (Spartina alterniflora) and (2) a mangrove tree (Avicennia germinans). In an 18‐month study of a recently restored coastal wetland in southeastern Louisiana (USA), we examined growth and survival along an elevation gradient and compared expansion and recruitment rates. We found that the rapid growth, expansion, and recruitment rates of the salt marsh grass make it a better species for quickly establishing ecological structure at suitable elevations. The slower growth, limited expansion, and lower recruitment of the mangrove species show its restricted capacity for immediate structural restoration, especially in areas where it co‐occurs with perennial salt marsh species. Our findings suggest that the structural attributes needed in recently restored areas can be achieved sooner using fast‐growing foundation species. Following salt marsh grass establishment, mangroves can then be used to further assist ecosystem development. This work highlights how appropriate foundation species can help jump‐start ecosystem development to meet restoration objectives.     

The role of wildlife science in wetland ecosystem restoration: Lessons from the Everglades

There has been little discussion of how and when to integrate wildlife science into ecological restoration projects. The recent emergence of wetland ecosystem restoration offers an opportunity to use wildlife science to increase the probability of a project being successful. This paper traces the evolution of wetland ecosystem restoration in North America and proposes three roles for wildlife science in wetland ecosystem restoration: (1) contribute to conceptual ecosystem models, (2) develop quantitative performance measures and restoration targets that track the progress of restoration, and (3) achieve social feasibility by sustaining long-term public support for a project. The extensive knowledge base for many species of wildlife makes them especially useful for contributing to conceptual ecosystem models. Wildlife species are often the subject of long-term monitoring and research because they have commercial value, are conspicuous, or have aesthetic appeal. Wildlife parameters can be good performance measures for large-scale restoration projects because some species integrate information over large spatial scales and are long-lived. Parameters associated with threatened or endangered wildlife species should get special consideration as performance measures because the information will meet multiple needs rather than just those of the conceptual ecosystem model. Finally, wetland ecosystem restoration projects need to sustain funding over decades to ensure the restored system is self-sustaining. Wildlife are a valued resource that can help achieve the social feasibility of a project by providing a way to communicate complex science in terms that society understands and values.     

Ant biodiversity and its relationship to ecosystem functioning: a review

Ants are important components of ecosystems not only because they constitute a great part of the animal biomass but also because they act as ecosystem engineers. Ant biodiversity is incredibly high and these organisms are highly responsive to human impact, which obviously reduces its richness. However, it is not clear how such disturbance damages the maintenance of ant services to the ecosystem. Ants are important in below ground processes through the alteration of the physical and chemical environment and through their effects on plants, microorganisms, and other soil organisms. This review summarizes the information available on ant biodiversity patterns, how it can be quantified, and how biodiversity is affected by human impacts such as land use change, pollution, invasions, and climate change. The role of ants in ecosystems is discussed, mainly from the perspective of the effects of ground-dwelling ants on soil processes and function, emphasizing their role as ecosystem engineers. Some lines of research are suggested after demonstrating the gaps in our current information on ant-soil interactions.     

Optimal control of population recovery – the role of economic restoration threshold

A variety of ecological systems around the world have been damaged in recent years, either by natural factors such as invasive species, storms and global change or by direct human activities such as overfishing and water pollution. Restoration of these systems to provide ecosystem services entails significant economic benefits. Thus, choosing how and when to restore in an optimal fashion is important, but has not been well studied. Here we examine a general model where population growth can be induced or accelerated by investing in active restoration. We show that the most cost‐effective method to restore an ecosystem dictates investment until the population approaches an ‘economic restoration threshold’, a density above which the ecosystem should be left to recover naturally. Therefore, determining this threshold is a key general approach for guiding efficient restoration management, and we demonstrate how to calculate this threshold for both deterministic and stochastic ecosystems.     

Restoration for variability: emergence of the habitat diversity paradigm in terrestrial ecosystem restoration

Ecosystem restoration implies focusing on multiple trophic levels and ecosystem functioning, yet higher trophic levels, that is, animals, are less frequently targeted by restoration than plants. Habitat diversity, the spatial heterogeneity between and within habitat patches in a landscape, is a well‐known driver of species diversity, and offers possible ways to increase species diversity at multiple trophic levels. We argue that habitat diversity is central in whole‐ecosystem restoration as we review its importance, provide a practical definition for its components, and propose ways to target it in restoration. Restoration targeting habitat diversity is used commonly in aquatic ecosystems, mostly to increase the physical diversity of habitats, meant to provide more niches available to a higher number of animal species. To facilitate the uptake of habitat diversity in terrestrial ecosystem restoration, we distinguish between compositional and structural habitat diversity, because different animal groups will respond to different aspects of habitat diversity. We also propose four methods to increase habitat diversity: varying the starting conditions to obtain divergent successional pathways, emulating natural disturbances, establishing keystone structures, and applying ecosystem engineer species. We provide two case studies to illustrate how these components and methods can be incorporated in restoration. We conclude that targeting habitat diversity is a promising way to restore habitats for a multitude of species of animals and plants, and that it should become mainstream in restoration ecology and practice. We encourage the restoration community to consider compositional and structural habitat diversity and to specifically target habitat diversity in ecosystem restoration.     

Restoring grassy ecosystems – Feasible or fiction? An inquisitive Australian's experience in the USA

Many small‐scale projects in Australia suggest that ground‐layer elements of ecosystems can be restored, but scaling up of grassland and grassy understorey restoration has not occurred to date. Paul Gibson‐Roy recently travelled through the USA, where well‐developed markets for restoration have created a large, financially viable native‐herbaceous seed production and restoration sector. Here, he shares his observations, which show how much about the USA situation can be a model and inspiration for Australian grassy ecosystem restoration.     

People as Ecological Participants in Ecological Restoration

In 1987, Bradshaw proposed that ecological restoration is the ultimate “acid test” of our understanding the functioning of ecosystems ( Bradshaw 1987 ). Although this concept is widely supported academically, how it can be applied by restoration practitioners is still unclear. This is an issue not limited to Bradshaw’s acid test, but moreover, reflects a general difficulty associated with the polarization between conceptual restoration (restoration ecology) and practical restoration (ecological restoration), where each has functioned to certain degree in isolation of the other. Outside of the more obvious pragmatic reasons for the relative independence between ecological restoration and restoration ecology, we propose that a more contentious explanation is that the approach taken toward understanding ecosystem development in restoration ecology is tangential to what actually takes place in ecological restoration. Current paradigms assume that the process of ecosystem development in restoration should follow the developmental trajectories suggested by classical ecological succession models. However, unlike these models, ecosystem development in restoration is, at least initially, largely manipulated by people, rather than by abiotic and biotic forces alone. There has been little research undertaken to explore how restoration activities impact upon or add to the extant ecological processes operating within a restoration site. Consequently, ecological restoration may not be so much an acid test of our understanding the functioning of ecosystems, but rather, an acid test of our understanding mutually beneficial interactions between humans and ecosystems.     

Integrating trait‐based empirical and modeling research to improve ecological restoration

A global ecological restoration agenda has led to ambitious programs in environmental policy to mitigate declines in biodiversity and ecosystem services. Current restoration programs can incompletely return desired ecosystem service levels, while resilience of restored ecosystems to future threats is unknown. It is therefore essential to advance understanding and better utilize knowledge from ecological literature in restoration approaches. We identified an incomplete linkage between global change ecology, ecosystem function research, and restoration ecology. This gap impedes a full understanding of the interactive effects of changing environmental factors on the long‐term provision of ecosystem functions and a quantification of trade‐offs and synergies among multiple services. Approaches that account for the effects of multiple changing factors on the composition of plant traits and their direct and indirect impact on the provision of ecosystem functions and services can close this gap. However, studies on this multilayered relationship are currently missing. We therefore propose an integrated restoration agenda complementing trait‐based empirical studies with simulation modeling. We introduce an ongoing case study to demonstrate how this framework could allow systematic assessment of the impacts of interacting environmental factors on long‐term service provisioning. Our proposed agenda will benefit restoration programs by suggesting plant species compositions with specific traits that maximize the supply of multiple ecosystem services in the long term. Once the suggested compositions have been implemented in actual restoration projects, these assemblages should be monitored to assess whether they are resilient as well as to improve model parameterization. Additionally, the integration of empirical and simulation modeling research can improve global outcomes by raising the awareness of which restoration goals can be achieved, due to the quantification of trade‐offs and synergies among ecosystem services under a wide range of environmental conditions.     

Human‐aided admixture may fuel ecosystem transformation during biological invasions: theoretical and experimental evidence

Biological invasions can transform our understanding of how the interplay of historical isolation and contemporary (human‐aided) dispersal affects the structure of intraspecific diversity in functional traits, and in turn, how changes in functional traits affect other scales of biological organization such as communities and ecosystems. Because biological invasions frequently involve the admixture of previously isolated lineages as a result of human‐aided dispersal, studies of invasive populations can reveal how admixture results in novel genotypes and shifts in functional trait variation within populations. Further, because invasive species can be ecosystem engineers within invaded ecosystems, admixture‐induced shifts in the functional traits of invaders can affect the composition of native biodiversity and alter the flow of resources through the system. Thus, invasions represent promising yet under‐investigated examples of how the effects of short‐term evolutionary changes can cascade across biological scales of diversity. Here, we propose a conceptual framework that admixture between divergent source populations during biological invasions can reorganize the genetic variation underlying key functional traits, leading to shifts in the mean and variance of functional traits within invasive populations. Changes in the mean or variance of key traits can initiate new ecological feedback mechanisms that result in a critical transition from a native ecosystem to a novel invasive ecosystem. We illustrate the application of this framework with reference to a well‐studied plant model system in invasion biology and show how a combination of quantitative genetic experiments, functional trait studies, whole ecosystem field studies and modeling can be used to explore the dynamics predicted to trigger these critical transitions.