The ecology of restoration: historical links, emerging issues and unexplored realms

Restoration ecology is a young academic field, but one with enough history to judge it against past and current expectations of the science's potential. The practice of ecological restoration has been identified as providing ideal experimental settings for tests of ecological theory; restoration was to be the 'acid test' of our ecological understanding. Over the past decade, restoration science has gained a strong academic foothold, addressing problems faced by restoration practitioners, bringing new focus to existing ecological theory and fostering a handful of novel ecological ideas. In particular, recent advances in plant community ecology have been strongly linked with issues in ecological restoration. Evolving models of succession, assembly and state-transition are at the heart of both community ecology and ecological restoration. Recent research on seed and recruitment limitation, soil processes, and diversity–function relationships also share strong links to restoration. Further opportunities may lie ahead in the ecology of plant ontogeny, and on the effects of contingency, such as year effects and priority effects. Ecology may inform current restoration practice, but there is considerable room for greater integration between academic scientists and restoration practitioners.     

Exotic Plant Species as Problems and Solutions in Ecological Restoration: A Synthesis

Exotic species have become increasingly significant management problems in parks and reserves and frequently complicate restoration projects. At the same time there may be circumstances in which their removal may have unforeseen negative consequences or their use in restoration is desirable. We review the types of effects exotic species may have that are important during restoration and suggest research that could increase our ability to set realistic management goals. Their control and use may be controversial; therefore we advocate consideration of exotic species in the greater context of community structure and succession and emphasize areas where ecological research could bring insight to management dilemmas surrounding exotic species and restoration. For example, an understanding of the potential transience of exotics in a site and the role particular exotics might play in changing processes that influence the course of succession is essential to setting removal priorities and realistic management goals. Likewise, a greater understanding of the ecological role of introduced species might help to reduce controversy surrounding their purposeful use in restoration. Here we link generalizations emerging from the invasion ecology literature with practical restoration concerns, including circumstances when it is practical to use exotic species in restoration.     

Trends in Ecological Research during the Last Three Decades – A Systematic Review

It is thought that the science of ecology has experienced conceptual shifts in recent decades, chiefly from viewing nature as static and balanced to a conception of constantly changing, unpredictable, complex ecosystems. Here, we ask if these changes are reflected in actual ecological research over the last 30 years. We surveyed 750 articles from the entire pool of ecological literature and 750 articles from eight leading journals. Each article was characterized according to its type, ecological domain, and applicability, and major topics. We found that, in contrast to its common image, ecology is still mostly a study of single species (70% of the studies); while ecosystem and community studies together comprise only a quarter of ecological research. Ecological science is somewhat conservative in its topics of research (about a third of all topics changed significantly through time), as well as in its basic methodologies and approaches. However, the growing proportion of problem-solving studies (from 9% in the 1980s to 20% in the 2000 s) may represent a major transition in ecological science in the long run.     

Science, Art, or Application—the "Karma" of Restoration Ecology

The present state of restoration ecology is far away from Bradshaw’s “acid test for ecology.” The conclusions drawn from the series of papers in this issue and from the Jena workshop suggest some directions in which the field may progress. More attention must be paid to the degraded state, which should be evaluated by its specific features and carefully analyzed before any restoration plan is laid down. Restoration goals have to be realistic, which includes the appreciation of globally changing conditions, resulting in a paradigm‐shift toward “forward‐restoration.” Basically, the transition from the degraded state conditions to the target state is a kind of succession that is manipulated by the application of goal‐orientated and system‐specific disturbances. Whenever possible, restorationists should step back and make use of naturally occurring succession, which requires a sophisticated restoration strategy, involving flexible management responses, multiple alternative target states, robust measurements for the restoration progress, and careful long‐term monitoring. The unique feature of restoration ecology is the involvement of socioeconomic decisions, and conceptual frameworks for ecological restoration have to implement the specific links to natural succession. To bridge the gap between ecological theory and on the ground restoration, it is essential that restoration practice is translated into the vocabulary and thinking of basic ecology. If all these aspects are integrated, ecological restoration as an application—and restoration ecology as an applied science—may develop into an acid test for our understanding of interactions between people and their environment, rather than for pure ecology.     

A call for an end to calls for the end of invasion biology

Calls for the end of invasion biology are misguided. There is no evidence that modern invasion biology has progressed slowly in its short life. Although some aspects of biological invasions fit comfortably in the framework of ecological succession, many others do not. Some native species, particularly in the wake of various anthropogenic impacts, behave like invasive non‐native species, but the probability and degree of harmful impact are greater for non‐native than for native species. Neither native nor non‐native species suffer lack of attention and research by virtue of the fact that invasion biology focuses on the latter. Basing management solely on current observed impact is highly risky because impacts may be subtle but nonetheless important, and impacts often change, as they are contingent on the physical or biotic environment. The known harmful impacts of many non‐native species suggest that recent introductions warrant attention even if impacts are not evident. Neither is the focus of modern invasion biology on non‐native species motivated by xenophobia. Rather, it reflects the recognition of their likelihood of harmful impact. A related call for the end of traditional restoration ecology shares many features with calls to terminate invasion biology, not least because management of invasive non‐native species is a key component of restoration ecology. Such species are a dominant element in generating the ‘novel ecosystems’ that are said to render traditional restoration ecology obsolete. The argument that both invasion management and traditional restoration are largely futile endeavors is contradicted by substantial and growing successes in both fields.     

Four opportunities for studies of ecological succession

Lessons learned from the study of ecological succession have much to offer contemporary environmental problem solving but these lessons are being underutilized. As anthropogenic disturbances increase, succession is more relevant than ever. In this review, we suggest that succession is particularly suitable to address concerns about biodiversity loss, climate change, invasive species, and ecological restoration. By incorporating modern experimental techniques and linking results across environmental gradients with meta-analyses, studies of succession can substantially improve our understanding of other ecological phenomena. Succession can help predict changes in biodiversity and ecosystem services impacted by invasive species and climate change and guide manipulative responses to these disruptions by informing restoration efforts. Succession is still a critical, integrative concept that is central to ecology.     

On the Status of Restoration Science: Obstacles and Opportunities

Terrestrial restoration ecology is not as well developed as aquatic and wetland restoration. There are several key obstacles to progress in restoration ecology, but these obstacles may also be viewed as opportunities to exploit. One obstacle is demonstration science, or an overreliance on simplistic experiments with few treatment factors and few levels of those factors. Complex, multivariate experiments yield greater insights, especially when teamed with sophisticated methods of data analysis. A second key obstacle is myopic scholarship that has led to little synthesis and weak conceptual theory. A greater awareness of and explicit references to ecological principles will help develop the conceptual basis of restoration science. Where should restoration ecology be headed? We should consider forming partnerships with developers, landscape artists, and industry to do complex, large‐scale experiments and make restoration a more common part of everyday life.     

采石场废弃地的生态重建研究进展

采石场的开采严重破坏了植被和土壤,形成了大量的裸露岩石斜坡,造成宏观景观支离破碎和极端的环境条件,限制了植物的生长。由于自然恢复所需时间长久,人工恢复被广泛应用于采石场废弃地的生态重建。自然演替过程是采石场生态重建的理论基础,自然演替理论可以为人工恢复措施提供指导。植物群落演替的早期阶段,非生物因素起主要作用,随着演替的推移,生物因素的重要性增强。邻近自然植被的土壤和繁殖体通过外力的扩散,对恢复起重要作用。除了非生物和其他的限制,先到达恢复地的物种竞争能力的变化能决定了演替过程。演替过程中的干扰因素往往成为演替重要的驱动力。裸露岩石斜坡的物理稳定性对植被恢复有重要影响,有机废物的使用和施肥可以影响恢复演替的方向和生物多样性。播种一定的植物能够改变恢复演替方向,加速演替过程。乡土物种适应了当地气候,能够促进演替。随着修复时间的延长,土壤有机质含量,植被覆盖度和物种丰富度不断增加,土壤微生物生物量随之增加。开展不同地区采石场植物种类的选育、研究乡土物种的功能特性、土壤微生物群落和酶的变化、植被演替过程的定位研究、植物种间的竞争关系、自然演替和人工恢复的比较研究、探索经济高效的采石场生态重建方法是未来的研究方向。     

Changing assembly processes during a primary succession of plant communities on Mediterranean roadcuts

Aims Studying plant ecological succession provides insights into the spatiotemporal processes underlying community assembly and is of primary importance for restoration ecology. We investigate here colonization events and local community assembly over an original primary succession occurring on roadcuts after roadwork. For this, we addressed both the changes in species presence-absence (incidence data) to highlight pre-establishment filters and in species relative abundances to further assess the influence of local biotic processes.Methods We studied 43 limestone roadcuts in Mediterranean France, covering five age classes up to an age of 80 years, along with 13 natural cliffs as a reference, and we counted 14322 plant individuals on these sites. We applied a constrained nonsymmetric correspondence analysis of both the incidence (presence-absence) and abundance data to assess the variation of these data along the chronosequence.Important findings Along the first 30 years, the initially abundant short-lived species declined both in terms of incidence and abundance and were replaced by longer lived herbaceous and woody species. This first phase was characterized by species that are widespread in the surrounding scrublands and was comparable to an early secondary succession there. After 30 years, there were continuing changes in incidence data with age, but no more significant change in species' abundances. This second phase was marked by the late colonization of specialists that did not become dominant. Although colonization and establishment limitation was thereby apparent for specialist species, a slow convergence of community composition toward the situation of natural cliffs could be detected in the older stages of the chronosequence. These findings convey insights into the natural dynamics of man-made outcrop plant communities and may be useful for the ecological management and restoration of such contexts. It also illustrates the interest of comparing incidence and abundance data to investigate the relative influence of ecological determinants on the assembly of plant communities.     

A Striking Profile: Soil Ecological Knowledge in Restoration Management and Science

Available evidence suggests that research in terrestrial restoration ecology has been dominated by the engineering and botanical sciences. Because restoration science is a relatively young discipline in ecology, the theoretical framework for this discipline is under development and new theoretical offerings appear regularly in the literature. In reviewing this literature, we observed an absence of in‐depth discussion of how soils, and in particular the ecology of soils, can be integrated into the developing theory of restoration science. These observations prompted us to assess the current role of soil ecological knowledge in restoration research and restoration practice. Although soils are universally regarded as critical to restoration success, and much research has included manipulations of soil variables, we found that better integration of soil ecological principles could still contribute much to the practice of ecosystem restoration. Here we offer four potential points of departure for increased dialog between restoration ecologists and soil ecologists. We hope to encourage the view that soil is a complex, heterogeneous, and vital entity and that adoption of this point of view can positively affect restoration efforts worldwide.     

Biodiversity Resources for Restoration Ecology

Biological resources can be more usefully incorporated into many aspects of restoration ecology. During the planning and design stage, the wide genotypic variation in natural plant populations must be recognized and exploited. This will ensure that genotypes used on a site are best adapted to local conditions and have a greater probability of survivorship than arbitrarily chosen material. Also, certain unusual genotypes can be located using the principles of evolutionary ecology and can be installed in areas with extreme conditions, such as soils contaminated with heavy metals, in areas where rapid colonizing ability (high seed set and/or clonal growth) is particularly advantageous, or where soils are of poor quality. Similarly, where high herbivore pressure is a threat to restoration, genotypes that are well defended, chemically or mechanically, against animal enemies should be selected to initiate the restoration process. The nursery industry can be encouraged to supply an ecologically wider selection of material for restoration, originating from local biological reserves and natural habitats. During the management phase of a restoration, local natural habitats are critical as reservoirs of biological control agents, seed sources for plant species, and members of higher trophic levels and additional plant species needed during succession. Mutualists such as pollinators, seed dispersers, and mycorrhizal fungi are vital to the success of a restoration project, and these must invade from nearby natural habitats or must be deliberately introduced. During the evaluation phase of restoration, local natural areas should be used as templates of community composition and structure from which one measures success. A functioning restoration project will interact biologically with surrounding areas, the exchange of species and genes being particularly important. Analysis of the microbial and invertebrate communities that have invaded the installed plant community may be useful and accurate determinants of ecological function. For these latter stages of the restoration process, the value of preserving local habitat remnants is high and complements their usefulness as a source of ecologically precise material for installation.     

Floristic patterns and plant traits of Mediterranean communities in fragmented habitats

Aim To contrast floristic spatial patterns and the importance of habitat fragmentation in two plant communities (grassland and scrubland) in the context of ecological succession. We ask whether plant assemblages are affected by habitat fragmentation and, if so, at what spatial scale? Does the relative importance of the niche differentiation and dispersal‐limitation mechanisms change throughout secondary succession? Is the dispersal‐limitation mechanism related to plant functional traits? Location A Mediterranean region, the massif of Albera (Spain). Methods Using a SPOT satellite image to describe the landscape, we tested the effect of habitat fragmentation on species composition, determining the spatial scale of the assemblage response. We then assessed the relative importance of dispersal‐related factors (habitat fragmentation and geographical distance) and environmental constraints (climate‐related variables) influencing species similarity. We tested the association between dispersal‐related factors and plant traits (dispersal mode and life form). Results In both community types, plant composition was partially affected by the surrounding vegetation. In scrublands, animal‐dispersed and woody plants were abundant in landscapes dominated by closed forests, whereas wind‐dispersed annual herbs were poorly represented in those landscapes. Scrubby assemblages were more dependent on geographical distance, habitat fragmentation and climate conditions (temperature, rainfall and solar radiation); grasslands were described only by habitat fragmentation and rainfall. Plant traits did not explain variation in spatial structuring of assemblages. Main conclusions Plant establishment in early Mediterranean communities may be driven primarily by migration from neighbouring established communities, whereas the importance of habitat specialization and community drift increases over time. Plant life forms and dispersal modes did not explain the spatial variation of species distribution, but species richness within the community with differing plant traits was affected by habitat patchiness.     

基于功能性状的毛乌素沙地不同演替阶段适生植物筛选

沙地生态系统修复是恢复生态学研究的热点问题,适生植物筛选是修复的关键。植物功能性状反映了植物在不同环境中的生存策略,探究沙地植物功能性状及其与环境之间的关系,有助于筛选用于植被恢复的物种,为保护沙地生态系统提供理论依据。以毛乌素沙地为研究区,分析了1983-2015年间沙地典型飞播样地群落演替特征及其对环境因子的响应,建立基于10个植物功能性状的毛乌素沙地潜在种库,进一步筛选飞播恢复下沙地不同演替阶段的适生植物。研究表明:(1)飞播恢复下的毛乌素沙地植物群落分为三个演替阶段:固沙先锋物种群落、沙生植物为主的杂类草群落、中生植物为主的杂类草群落。(2)土壤因子是群落演替的主要驱动力,其中土壤全氮、土壤总有机碳、土壤硝态氮是影响群落演替的关键因素。(3)基于功能性状筛选出29种适生物种用于植被恢复,演替第一阶段可用雾冰藜、猪毛菜等,演替第二阶段可用拂子茅、无芒隐子草等,演替第三阶段可用草地风毛菊、猪毛蒿等。通过物种功能性状特征可以快速选择适合沙地退化生态系统修复的候选物种,为植被恢复提供了一定的理论支持。     

Neutral theory and the species abundance distribution: recent developments and prospects for unifying niche and neutral perspectives

Published in 2001, The Unified Neutral Theory of Biodiversity and Biogeography (UNTB) emphasizes the importance of stochastic processes in ecological community structure, and has challenged the traditional niche‐based view of ecology. While neutral models have since been applied to a broad range of ecological and macroecological phenomena, the majority of research relating to neutral theory has focused exclusively on the species abundance distribution (SAD). Here, we synthesize the large body of work on neutral theory in the context of the species abundance distribution, with a particular focus on integrating ideas from neutral theory with traditional niche theory. First, we summarize the basic tenets of neutral theory; both in general and in the context of SADs. Second, we explore the issues associated with neutral theory and the SAD, such as complications with fitting and model comparison, the underlying assumptions of neutral models, and the difficultly of linking pattern to process. Third, we highlight the advances in understanding of SADs that have resulted from neutral theory and models. Finally, we focus consideration on recent developments aimed at unifying neutral‐ and niche‐based approaches to ecology, with a particular emphasis on what this means for SAD theory, embracing, for instance, ideas of emergent neutrality and stochastic niche theory. We put forward the argument that the prospect of the unification of niche and neutral perspectives represents one of the most promising future avenues of neutral theory research.     

Policy Language in Restoration Ecology

Relating restoration ecology to policy is one of the aims of the Society for Ecological Restoration and its journal Restoration Ecology. As an interdisciplinary team of researchers in both ecological science and political science, we have struggled with how policy‐relevant language is and could be deployed in restoration ecology. Using language in scientific publications that resonates with overarching policy questions may facilitate linkages between researcher investigations and decision‐makers' concerns on all levels. Climate change is the most important environmental problem of our time and to provide policymakers with new relevant knowledge on this problem is of outmost importance. To determine whether or not policy‐specific language was being included in restoration ecology science, we surveyed the field of restoration ecology from 2008 to 2010, identifying 1,029 articles, which we further examined for the inclusion of climate change as a key element of the research. We found that of the 58 articles with “climate change” or “global warming” in the abstract, only 3 identified specific policies relevant to the research results. We believe that restoration ecologists are failing to include themselves in policy formation and implementation of issues such as climate change within journals focused on restoration ecology. We suggest that more explicit reference to policies and terminology recognizable to policymakers might enhance the impact of restoration ecology on decision‐making processes.     

Is there a general theory of community ecology?

Community ecology entered the 1970s with the belief that niche theory would supply a general theory of community structure. The lack of wide-spread empirical support for niche theory led to a focus on models specific to classes of communities such as lakes, intertidal communities, and forests. Today, the needs of conservation biology for metrics of “ecological health” that can be applied across types of communities prompts a renewed interest in the possibility of general theory for community ecology. Disputes about the existence of general patterns in community structure trace at least to the 1920s and continue today almost unchanged in concept, although now expressed through mathematical modeling. Yet, a new framework emerged in the 1980s from findings that community composition and structure depend as much on the processes that bring species to the boundaries of a community as by processes internal to a community, such as species interactions and co-evolution. This perspective, termed “supply-side ecology”, argued that community ecology was to be viewed as an “organic earth science” more than as a biological science. The absence of a general theory of the earth would then imply a corresponding absence of any general theory for the communities on the earth, and imply that the logical structure of theoretical community ecology would consist of an atlas of models special to place and geologic time. Nonetheless, a general theory of community ecology is possible similar in form to the general theory for evolution if the processes that bring species to the boundary of a community are analogized to mutation, and the processes that act on the species that arrive at a community are analogized to selection. All communities then share some version of this common narrative, permitting general theorems to be developed pertaining to all ecological communities. Still, the desirability of a general theory of community ecology is debatable because the existence of a general theory suppresses diversity of thought even as it allows generalizations to be derived. The pros and cons of a general theory need further discussion.     

Ecology after 100 years: Progress and pseudo-progress

Has the science of ecology fulfilled the promises made by the originators of ecological science at the start of the last century? What should ecology achieve? Have good policies for environmental management flowed out of ecological science? These important questions are rarely discussed by ecologists working on detailed studies of individual systems. Until we decide what we wish to achieve as ecologists we cannot define progress toward those goals. Ecologists desire to achieve an understanding of how the natural world operates, how humans have modified the natural world, and how to alleviate problems arising from human actions. Ecologists have made impressive gains over the past century in achieving these goals, but this progress has been uneven. Some sub-disciplines of ecology are well developed empirically and theoretically, while others languish for reasons that are not always clear. Fundamental problems can be lost to view as ecologists fiddle with unimportant pseudo-problems. Bandwagons develop and disappear with limited success in addressing problems. The public demands progress from all the sciences, and as time moves along and problems get worse, more rapid progress is demanded. The result for ecology has too often been poor, short-term science and poor management decisions. But since the science is rarely repeated and the management results may be a generation or two down the line, it is difficult for the public or for scientists to decide how good or bad the scientific advice has been. In ecology over the past 100 years we have made solid achievements in behavioural ecology, population dynamics, and ecological methods, we have made some progress in understanding community and ecosystem dynamics, but we have made less useful progress in developing theoretical ecology, landscape ecology, and natural resource management. The key to increasing progress is to adopt a systems approach with explicit hypotheses, theoretical models, and field experiments on a scale defined by the problem. With continuous feedback between problems, possible solutions, relevant theory and experimental data we can achieve our scientific goals.     

Insights from a Cross‐Disciplinary Seminar: 10 Pivotal Papers for Ecological Restoration

Restoration ecology is a deepening and diversifying field with current research incorporating multiple disciplines and infusing long‐standing ideas with fresh perspectives. We present a list of 10 recent pivotal papers exemplifying new directions in ecological restoration that were selected by students in a cross‐disciplinary graduate seminar at the University of California, Berkeley. We highlight research that applies ecological theory to improve restoration practice in the context of global change (e.g. climate modeling, evaluation of novel ecosystems) and discuss remaining knowledge gaps. We also discuss papers that recognize the social context of restoration and the coupled nature of social and ecological systems, ranging from the incorporation of cultural values and Traditional Ecological Knowledge into restoration, to the consideration of the broader impacts of markets on restoration practices. In addition, we include perspectives that focus on improving communication between social and natural scientists as well as between scientists and practitioners, developing effective ecological monitoring, and applying more integrated, whole‐landscape approaches to restoration. We conclude with insights on recurrent themes in the papers regarding planning restoration in human‐modified landscapes, application of ecological theory, improvements to restoration practice, and the social contexts of restoration. We share lessons from our cross‐disciplinary endeavor, and invite further discussion on the future directions of restoration ecology through contributions to our seminar blog site http://restecology.blogspot.com .     

Predicting phenology by integrating ecology,evolution and climate science

Forecasting how species and ecosystems will respond to climate change has been a major aim of ecology in recent years. Much of this research has focused on phenology – the timing of life‐history events. Phenology has well‐demonstrated links to climate, from genetic to landscape scales; yet our ability to explain and predict variation in phenology across species, habitats and time remains poor. Here, we outline how merging approaches from ecology, climate science and evolutionary biology can advance research on phenological responses to climate variability. Using insight into seasonal and interannual climate variability combined with niche theory and community phylogenetics, we develop a predictive approach for species’ reponses to changing climate. Our approach predicts that species occupying higher latitudes or the early growing season should be most sensitive to climate and have the most phylogenetically conserved phenologies. We further predict that temperate species will respond to climate change by shifting in time, while tropical species will respond by shifting space, or by evolving. Although we focus here on plant phenology, our approach is broadly applicable to ecological research of plant responses to climate variability.     

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.