Fossil insects and ecosystem dynamics in wetlands: implications for biodiversity and conservation

We review the uses of fossil insects, particularly Coleoptera (beetles) and Chironomidae (non-biting midges) from ancient deposits to inform the study of wetland ecosystems and their ecological and restoration processes. In particular, we focus on two contrasting ecosystems, drawing upon research undertaken by us on British raised mire peats and shallow lake systems, one an essentially terrestrial ecosystem, the other aquatic, but in which wetland insects play an important and integral part. The study of raised mire peats suggests that faunal stability is a characteristic of these wetland systems, over what appear to be extensive periods of time (up to several millennia), whilst studies of shallow lake ecosystems over recent timescales indicates that faunal instability appears to be more common, usually driven by increasing eutrophication. Drawing upon a series of fossil Coleoptera records spanning several thousand years from Hatfield Moors, south Yorkshire, we reconstruct in some detail the mire’s ontogeny and fluctuations in site hydrology and vegetation cover, illustrating the intimate association between substrate, topography and peat development. A comparison between fossil and modern beetle populations indicates that the faunal characteristics of this mire and its adjacent neighbour, Thorne Moors, become established during the early phases of peat development, including its rare endemics, and that the faunal biodiversity on the sites today is dictated by complex site histories. The over-riding characteristic of these faunas is of stability over several thousand years, which has important implications for the restoration of degraded sites, especially those where refugial areas are limited. In contrast, analyses of fossil Chironomidae from shallow lakes allow researchers to track changes in limnological status and while attempts have been made to reconstruct changes in nutrient levels quantitatively, the chironomids respond indirectly to such changes, typically mediated through complex ecosystem dynamics such as changes in fish and/or macrophyte communities. These changes are illustrated via historic chironomid stratigraphies and diversity indices from a range of shallow lakes located across Britain: Slapton Ley, Frensham Great Pond, Fleet Pond, Kyre Pool and Barnes Loch. These sites have shown varying degrees of eutrophication over recent timescales which tends to be associated with a decline in chironomid diversity. While complex functional processes exist within these ecosystems, our evidence suggests that one of the key drivers in the loss of shallow lake chironomid diversity appears to be the loss of aquatic macrophytes. Overall, while chironomids do show a clear response to altered nutrient regimes, multi-proxy reconstructions are recommended for a clear interpretation of past change. We conclude that if we are to have a better understanding of biota at the ecosystem level we need to know more of the complex interactions between different insect groups as well as with other animal and plant communities. A palaeoecological approach is thus crucial in order to assess the role of insect groups in ecosystem processes, both in the recent past and over long time scales, and is essential for wetland managers and conservation organisations involved in long term management and restoration of wetland systems     

What is a healthy ecosystem?

Rapid deterioration of the world's major ecosystems has intensified the need for effective environmental monitoring and the development of operational indicators of ecosystem health. Ecosystem health represents a desired endpoint of environmental management, but it requires adaptive, ongoing definition and assessment. We propose that a healthy ecosystem is one that is sustainable – that is, it has the ability to maintain its structure (organization) and function (vigor) over time in the face of external stress (resilience). Various methods to quantify these three ecosystem attributes (vigor, organization, and resilience) are discussed. These attributes are then folded into a comprehensive assessment of ecosystem health. A network analysis based ecosystem health assessment is developed and tested using trophic exchange networks representing several different aquatic ecosystems. Results indicate the potential of such an ecosystem health assessment for evaluating the relative health of similar ecosystems, and quantifying the effects of natural or anthropogenic stress on the health of a particular ecosystem over time.     

Back-to-the-future: a fresh policy initiative for fisheries and a restoration ecology for ocean ecosystems

'Back-to-the-future' (BTF) is an integrative approach to a restoration ecology of the oceans that attempts to solve the fisheries crisis. To this end, it harnesses the latest understanding of ecosystem processes developments in whole ecosystem simulation modelling, and insight into the human dimension of fisheries management. BTF includes new methods for describing past ecosystems, designing fisheries that meet criteria for sustainability and responsibility, and evaluating the costs and benefits of fisheries in restored ecosystems. Evaluation of alternative policy choices, involving trade-offs between conservation and economic values, employs a range of economic, social and ecological measures. Automated searches maximize values of objective functions, and the methodology includes analyses of model parameter uncertainty. Participatory workshops attempt to maximize compliance by fostering a sense of ownership among all stakeholders. Some challenges that have still to be met include improving methods for quantitatively describing the past, reducing uncertainty in ecosystem simulation techniques and in making policy choices robust against climate change. Critical issues include whether past ecosystems make viable policy goals, and whether desirable goals may be reached from today's ecosystem. Examples from case studies in British Columbia, Newfoundland and elsewhere are presented.     

生态系统红色名录——一种新的生物多样性保护工具

生物多样性为人类提供了生存所必需的一系列生态系统服务和功能。然而,由于人为活动的加剧,生物多样性不断丧失。传统的生物多样性保护主要关注物种多样性,存在着对生物多样性的代表性不足,不能及时反应生态系统多样性的变化等缺点。近年来,生态系统层次上的多样性保护成为研究热点,一些国家和组织相继开展了大尺度的生态系统评估工作。文章回顾了已有的生态系统评估方案,发现当前生态系统评估多采用IUCN物种红色名录的分级标准体系,主要评估生态系统的濒危程度,评估标准主要是分布范围和功能的变化,不同评估方案采用的指标和阈值有差异,需要建立统一的生态系统分类体系和评价方案。同时,结合国内生态系统评价的现状,提出了在我国开展生态系统红色名录研究的若干可行建议。     

When is restoration not?: Incorporating landscape-scale processes to restore self-sustaining ecosystems in coastal wetland restoration

With increasing restoration initiatives for coastal wetlands, the question of ‘What are we restoring to?’ becomes more pressing. The goal of this paper is to explore restoration concepts, examples, and challenges from the Pacific and Gulf coasts. One of the fundamental concepts explored is change over time – either in the controlling processes or the restoration structure – and how such changes can be meshed with the goals of various restoration efforts. We subsequently review the concepts of ecosystem trajectories, alternative restoration approaches, and the ideal attributes of functional self-sustaining restoration in the context of realities of restoration planning, design, and implementation. These realities include the dynamics of the ecosystems being restored, very real constraints that are imposed by the contemporary physical and human landscape, and the need to plan for the long term development of restoration sites recognizing that both project performance and expectations may change over time.     

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.     

云贵高原生态恢复空间分异及其影响因素

全面深入评估生态恢复效益并明晰其自然—社会经济驱动机制,对维护区域生态安全、实现可持续发展目标具有重要意义。然而,在区域尺度量化分析时间序列生态恢复程度及其影响因素的案例研究较少。以云贵高原为研究对象,基于多源遥感数据,运用模型模拟、时间序列趋势分析等方法,从生态系统宏观结构、质量和服务3个层次评估了2000—2019年云贵高原地区的生态恢复效益,使用随机森林方法探究了生态恢复的自然-社会经济影响因素。结果表明：(1)近20年云贵高原地区生态系统宏观结构变化明显,主要表现为森林、草地和聚落生态系统面积的增加,以及农田生态系统面积的减少;(2)生态系统质量和服务恢复态势总体稳定向好,部分区域转差;(3)云贵高原地区生态系统的恢复程度整体较高。其中,恢复程度中等及以上的地区占总面积的35.84%;生态恢复程度有所转差及以下的面积最少,占总面积的7.54%;(4)地形、气候和土壤等生态本底特征是云贵高原地区生态恢复最重要的影响因素;在局地尺度,造林面积、人口密度年变化率等人类活动因子对生态恢复具有关键的作用。未来生态系统管理和生态恢复工程的实施应综合考虑区域生态背景和尺度效应,因地制宜采取恢...     

不同生态恢复方式下生态系统服务与生物多样性恢复效果的整合分析

全球范围内关键生态系统服务的减少使人类社会面临巨大的威胁,生物多样性是生态系统提供各种产品和服务的基础。生态恢复工程对退化的生态系统服务和生物多样性进行修复,对于缓解人类环境压力具有非常重要的意义。长期的理论和实践工作形成了多种生态恢复措施:(1)单纯基于生态系统自我设计的自然恢复方式,(2)人为设计对环境条件进行干预,反馈影响生态系统的自我设计,(3)人为设计对目标种群和生态系统进行直接干预和重建。这3类恢复方式可以在不同程度上定向的影响生态系统的恢复进程,反映了人类对生态系统的低度、中度和高度介入。哪种恢复方式和介入程度能够实现更好的恢复效果,是生态恢复学中的一个关键问题,但到目前为止,虽广有争议,却无定量的分析和结论。针对这个空白,通过对ISI Web of Knowledge数据库中生态恢复相关文献的整合分析,基于数学统计的方法定量比较在不同条件下低度介入(自然恢复)、中度介入(环境干预)和高度介入(直接干预)3种恢复方式对生态系统服务与生物多样性的恢复效果。论文从4个方面展开研究:(1)低度、中度、高度介入生态恢复方式的划分,(2)比较3大类介入方式对生态系统服务和生物多样性恢复效果的差异,(3)不同气候条件、生态系统类型和恢复时间等背景因素的影响,(4)生物多样性恢复和生态系统服务恢复之间的关系。研究结果揭示了不同生态恢复方式的适用条件,以及对生物多样性和生态系统恢复相互关系的作用,对生态恢复实践中恢复方式的选择有指导作用。对未来的研究也有启示意义,如针对特定生态系统服务或具体研究问题进一步探索低度、中度和高度介入生态恢复方式的作用规律和机制;将地区的社会经济水平、生态系统的受损程度等因素纳入生态恢复方式的考察,以最优化生态恢复成本-效率等。     

Grieving for the Past and Hoping for the Future: Balancing Polarizing Perspectives in Conservation and Restoration

I consider the possibility that people engaged in conservation and management of species and ecosystems are experiencing grief related to ongoing loss of species, assemblages, ecosystem integrity, and so on. In human psychology, five stages of the grieving process have been identified, namely denial, anger, bargaining, depression, and acceptance. Although presented as a series of stages, it is recognized that the progression through them is not linear and people can move quickly among them or experience more than one at any one time. I then consider whether current polarized debates in conservation and restoration, for instance in relation to non‐native species and novel ecosystems, result in part from people operating from different places in the grief spectrum. Throughout the grieving process, hope remains a constant feature, and it is important to recognize the place of hope in motivating and sustaining people engaged in conservation and restoration. Although restoration provides great hope that losses can be minimized and, in some cases, reversed, this hope needs to be grounded in a realistic assessment of what is possible in a rapidly changing world.     

A conceptual framework for urban ecological restoration and rehabilitation

Urban greenspace has gained considerable attention during the last decades because of its relevance to wildlife conservation, human welfare, and climate change adaptation. Biodiversity loss and ecosystem degradation worldwide require the formation of new concepts of ecological restoration and rehabilitation aimed at improving ecosystem functions, services, and biodiversity conservation in cities. Although relict sites of natural and semi-natural ecosystems can be found in urban areas, environmental conditions and species composition of most urban ecosystems are highly modified, inducing the development of novel and hybrid ecosystems. A consequence of this ecological novelty is the lack of (semi-) natural reference systems available for defining restoration targets and assessing restoration success in urban areas. This hampers the implementation of ecological restoration in cities. In consideration of these challenges, we present a new conceptual framework that provides guidance and support for urban ecological restoration and rehabilitation by formulating restoration targets for different levels of ecological novelty (i.e., historic, hybrid, and novel ecosystems). To facilitate the restoration and rehabilitation of novel urban ecosystems, we recommend using established species-rich and well-functioning urban ecosystems as reference. Such urban reference systems are likely to be present in many cities. Highlighting their value in comparison to degraded ecosystems can stimulate and guide restoration initiatives. As urban restoration approaches must consider local history and site conditions, as well as citizens’ needs, it may also be advisable to focus the restoration of strongly altered urban ecosystems on selected ecosystem functions, services and/or biodiversity values. Ecosystem restoration and rehabilitation in cities can be either relatively inexpensive or costly, but even expensive measures can pay off when they effectively improve ecosystem services such as climate change mitigation or recreation. Successful re‐shaping and re-thinking of urban greenspace by involving citizens and other stakeholders will help to make our cities more sustainable in the future.     

Restoration for multiple use

Management of restored ecosystems for multiple use is a modern necessity given a growing human population and dwindling supplies of ecosystem goods and services. Multiple use management refers to managing resources simultaneously for sustainable output of many goods and services. Within any restoration, thoughtful planning and early stakeholder engagement can help harmonize seemingly competing multiple uses. Although the field of ecological restoration is young and there are few long‐term lessons to draw from, we can infer from ecological theory that maximization of native biodiversity can impart resilience in the restored ecosystem and can buffer against the stress of multiple use management. Restoration for multiple use should be accompanied with an acknowledgment that humility is required and monitoring is needed to keep the restored ecosystem on an acceptable trajectory. The field of ecological restoration was founded upon the notion that ecosystems would be restored for ethical reasons, but modern realities have necessitated a more utilitarian approach to restoration that requires restoring ecosystems for multiple uses. This reality represents a grand challenge for the next generation of restoration ecologists.     

典型脆弱生态区生态技术评价方法及应用专题导读

全球经济发展和日益增强的人类活动给业已脆弱的生态系统带来了巨大挑战,寻求尊重自然规律、环境友好的生态治理与恢复技术成为实现联合国可持续发展目标的重要组成部分。近年来,国内外科学家和生态治理机构研发出了一系列技术体系和技术模式,对脆弱生态区退化生态系统展开了全面的治理和恢复。然而,缺乏对这些治理技术的评价和优选方法的研发和应用在很大程度上限制了优良技术的筛选和推广应用,造成了资金和人力的浪费和损失。针对这些问题,科技部在2016年启动了国家重点研发计划项目"生态技术评价方法、指标体系与全球生态治理技术评价",其中重要的任务之一是对生态技术进行梳理、评价和优选。本文旨在对已有的评价方法和指标体系的研究进展进行系统梳理,对生态技术的特征、生态技术评价的基本步骤和原则、生态技术的三阶段评估方法等进行了界定和深入分析,对本专题收录的14篇学术文章所涵盖的评价方法和模型研发、案例应用等进行了介绍,以期为筛选优良技术、提高生态技术应用效果、促进优良技术的输出和引进提供参考。     

Stretch Goals and Backcasting: Approaches for Overcoming Barriers to Large-Scale Ecological Restoration

The destruction and transformation of ecosystems by humans threatens biodiversity, ecosystem function, and vital ecosystem services. Ecological repair of ecosystems will be a major challenge over the next century and beyond. Restoration efforts to date have frequently been ad hoc, and site or situation specific. Although such small‐scale efforts are vitally important, without large‐scale visions and coordination, it is unlikely that large functioning ecosystems will ever be constructed by chance through the cumulative effects of small‐scale projects. Although the problems of human‐induced environmental degradation and the need for a solution are widely recognized, these issues have rarely been addressed on a sufficiently large‐scale basis. There are numerous barriers that prevent large‐scale ecological restoration projects from being proposed, initiated, or carried through. Common barriers include the “shifting baseline syndrome,” the scale and complexity of restoration, the long‐term and open‐ended nature of restoration, funding challenges, and preemptive constraint of vision. Two potentially useful approaches that could help overcome these barriers are stretch goals and backcasting. Stretch goals are ambitious long‐term goals used to inspire creativity and innovation to achieve outcomes that currently seem impossible. Backcasting is a technique where a desired end point is visualized, and then a pathway to that end point is worked out retrospectively. A case study from the Scottish Highlands is used to illustrate how stretch goals and backcasting could facilitate large‐scale restoration. The combination of these approaches offers ways to evaluate and shape options for the future of ecosystems, rather than accepting that future ecosystems are victims of past and present political realities.     

The importance of ecological memory for trophic rewilding as an ecosystem restoration approach

Increasing human pressure on strongly defaunated ecosystems is characteristic of the Anthropocene and calls for proactive restoration approaches that promote self‐sustaining, functioning ecosystems. However, the suitability of novel restoration concepts such as trophic rewilding is still under discussion given fragmentary empirical data and limited theory development. Here, we develop a theoretical framework that integrates the concept of ‘ecological memory’ into trophic rewilding. The ecological memory of an ecosystem is defined as an ecosystem's accumulated abiotic and biotic material and information legacies from past dynamics. By summarising existing knowledge about the ecological effects of megafauna extinction and rewilding across a large range of spatial and temporal scales, we identify two key drivers of ecosystem responses to trophic rewilding: (i) impact potential of (re)introduced megafauna, and (ii) ecological memory characterising the focal ecosystem. The impact potential of (re)introduced megafauna species can be estimated from species properties such as lifetime per capita engineering capacity, population density, home range size and niche overlap with resident species. The importance of ecological memory characterising the focal ecosystem depends on (i) the absolute time since megafauna loss, (ii) the speed of abiotic and biotic turnover, (iii) the strength of species interactions characterising the focal ecosystem, and (iv) the compensatory capacity of surrounding source ecosystems. These properties related to the focal and surrounding ecosystems mediate material and information legacies (its ecological memory) and modulate the net ecosystem impact of (re)introduced megafauna species. We provide practical advice about how to quantify all these properties while highlighting the strong link between ecological memory and historically contingent ecosystem trajectories. With this newly established ecological memory–rewilding framework, we hope to guide future empirical studies that investigate the ecological effects of trophic rewilding and other ecosystem‐restoration approaches. The proposed integrated conceptual framework should also assist managers and decision makers to anticipate the possible trajectories of ecosystem dynamics after restoration actions and to weigh plausible alternatives. This will help practitioners to develop adaptive management strategies for trophic rewilding that could facilitate sustainable management of functioning ecosystems in an increasingly human‐dominated world.     

Setting Effective and Realistic Restoration Goals: Key Directions for Research

Restoration ecology has made significant advances in the past few decades and stands to make significant contributions both to the practical repair of damaged ecosystems and the development of broader ecological ideas. I highlighted four main areas where progress in research can assist with this. First, we need to enhance the translation of recent advances in our understanding of ecosystem and landscape dynamics into the conceptual and practical frameworks for restoration. Second, we need to promote the development of an ability to correctly diagnose ecosystem damage, identify restoration thresholds, and develop corrective methodologies that aim to overcome such thresholds. This involves understanding which system characteristics are important in determining ecosystem recovery in a range of ecosystem types, and to what extent restoration measures need to overcome threshold and hysteresis effects. A third key requirement is to determine what realistic goals for restoration are based on the ecological realities of today and how these will change in the future, given ongoing changes in climate and land use. Finally, there is a need for a synthetic approach which draws together the ecological and social aspects of the issues surrounding restoration and the setting of restoration goals.     

Restoring Kangaroo Grass (Themeda triandra) to grassland and woodland understoreys: a review of establishment requirements and restoration exercises in south-east Australia

Summary Temperate grassy ecosystems are amongst Australia's most endangered ecosystems. Most remnants are small, fragmented and highly degraded. Practical methods for restoring native understorey species are urgently required. Dominant native grasses such as Kangaroo Grass (Themeda triandra Forssk.) and Tussock‐grasses (Poa species) have been eliminated from many remnants by heavy grazing in the past. The reintroduction of these grasses is a critical step for understorey restoration. This paper (i) reviews the literature on Themeda seed biology and seedling establishment; (ii) summarizes the lessons learnt from three major attempts to establish Themeda stands in south‐east Australia; and (iii) identifies the research needed to enhance Themeda restoration. Considerable information is available on Themeda seed and establishment biology, and restoration exercises have shown that Themeda stands can be readily established by surface‐spreading awned seeds in seed‐bearing hay. However, many practical challenges remain, including the need to identify optimal sowing periods, create better seedbed conditions, develop practical mulching techniques, and improve weed control. The use of seed‐bearing hay has constrained restoration to relatively small areas in the past. Future trials may benefit by using more concentrated seed products such as seed‐bearing florets and pure seeds which permit larger areas to be restored at one time.     

Swamp rabbits as indicators of wildlife habitat quality in bottomland hardwood forest ecosystems

Reforestation of bottomland hardwood (BLH) forests has occurred within the Lower Mississippi Alluvial Valley (LMAV), USA, to support a wide range of ecosystem services, but especially wildlife habitat enhancement. As ecosystem restoration efforts proceed in BLH ecosystems, managers and policymakers are seeking criteria to evaluate wildlife habitat enhancement goals. Specialist wildlife that evolved within forest ecosystems can be sensitive to the composition, structure, and function of an ecosystem in relation to the system's natural or historical range of variation and thereby serve as indicators of habitat quality. The swamp rabbit (Sylvilagus aquaticus) is a specialist species of BLH forests throughout the LMAV and therefore may be an appropriate indicator species for this ecosystem. To address this, we reviewed peer-reviewed literature to evaluate the utility of swamp rabbits as an indicator species according to three commonly-used criteria: habitat factors defining swamp rabbit relationships to BLH forests, the importance of swamp rabbit habitat to other wildlife, and the efficiency of swamp rabbit monitoring. We conclude that the swamp rabbit is a suitable indicator of wildlife habitat quality in BLH ecosystems in the LMAV because they evolved and remain endemic to the ecosystem, use habitat that integrates desirable characteristics that positively influence wildlife biodiversity, and are easy to monitor routinely.     

Evaluating Ecosystem Services Provided by Non-Native Species: An Experimental Test in California Grasslands

The concept of ecosystem services – the benefits that nature provides to human''s society – has gained increasing attention over the past decade. Increasing global abiotic and biotic change, including species invasions, is threatening the secure delivery of these ecosystem services. Efficient evaluation methods of ecosystem services are urgently needed to improve our ability to determine management strategies and restoration goals in face of these new emerging ecosystems. Considering a range of multiple ecosystem functions may be a useful way to determine such strategies. We tested this framework experimentally in California grasslands, where large shifts in species composition have occurred since the late 1700''s. We compared a suite of ecosystem functions within one historic native and two non-native species assemblages under different grazing intensities to address how different species assemblages vary in provisioning, regulatory and supporting ecosystem services. Forage production was reduced in one non-native assemblage (medusahead). Cultural ecosystem services, such as native species diversity, were inherently lower in both non-native assemblages, whereas most other services were maintained across grazing intensities. All systems provided similar ecosystem services under the highest grazing intensity treatment, which simulated unsustainable grazing intensity. We suggest that applying a more comprehensive ecosystem framework that considers multiple ecosystem services to evaluate new emerging ecosystems is a valuable tool to determine management goals and how to intervene in a changing ecosystem.     

基于生态系统演变机理的生态系统脆弱性、适应性与突变理论

随着气候变化影响广度与深度的增加,生态系统脆弱性、适应性与突变理论逐渐被广泛应用到生态学研究领域中,探讨和评估各类生态系统对气候变化的敏感性、脆弱性和适应性,可谋求更好的方式来应对气候变化对区域生态系统带来的深远影响,服务于国家生态系统可持续管理及生态安全建设.虽然相关研究已获取许多进展,区分了气候敏感区和某些生态系统...     

Restoration Ecology to the Future: A Call for New Paradigm

The discipline of restoration ecology has grown remarkably in the past decades, providing new ideas and opportunities for conserving biological diversity, managing ecosystems, and testing ecological theories. On the other side, its past‐oriented, static, and idealistic approach has been criticized for subjectivity in determining restoration goals, inapplicability to dynamic ecosystems, and inability for restoring certain irreversible losses. Moreover, unpredictable sustainability of the restored ecosystems, which were modeled after its historical fidelity, adds our skepticism under the changing environment. This paper calls for a new paradigm of ecological restoration to the future. A future‐oriented restoration should (1) establish the ecosystems that are able to sustain in the future, not the past, environment; (2) have multiple alternative goals and trajectories for unpredictable endpoints; (3) focus on rehabilitation of ecosystem functions rather than recomposition of species or cosmetics of landscape surface; and (4) acknowledge its identity as a “value‐laden” applied science within economically and socially acceptable framework. Applicability of ecological theories to restoration practice is also discussed in this paper.