Non-native species in urban environments: patterns,processes, impacts and challenges

Although urban ecosystems are hotspots for biological invasions, the field of invasion science has given scant attention to invasion dynamics and the challenges facing managers in towns and cities. This paper provides an introduction to the growing challenges of understanding and managing invasive species in urban systems, and the context for a special issue of Biological Invasions, comprising 17 papers, that arose from a workshop on “Non-native species in urban environments: patterns, processes, impacts and challenges” held in Stellenbosch, South Africa, in November 2016. Contributions explore the following key questions: Are patterns and processes of urban invasions different from invasions in other contexts? Why is it important to manage non-native species in urban ecosystems? What are the special management needs in an urban context? How can we bridge the gaps between science, management, and policy with regards to biological invasions in urban ecosystems? The papers in this special issue show that patterns and processes of urban invasions differ in many ways from invasions in other contexts, and that managing invasive species in cities poses unique and increasingly complex challenges. Progress in urban invasion science requires further work to: (1) address key limitations that hinder our understanding of invasion dynamics in cities; (2) clarify whether fundamental concepts in the field of invasion science are appropriate for urban ecosystems; (3) integrate insights from invasion science with those from the burgeoning literature on the “Anthropocene biosphere”, novel ecosystems, social–ecological systems, human–wildlife conflicts, urban green infrastructure, urban planning and design, and ecosystem services/disservices.     

Ranking of invasive spread through urban green areas in the world’s 100 most populous cities

Urban landscapes are highly fragmented (leading to the extinction of native species) as well as transformed and disturbed (creating novel environments). Such conditions provide non-native species with opportunities to establish and spread through “urban green areas” (UGAs). UGAs can serve as stepping stones for many alien species to recruit and may become sources of propagules to launch invasions in adjoining natural ecosystems. There is great diversity in the spatial structures of UGAs worldwide; these are determined by the city’s level of development, human density, urban planning policy, and history. We explore the invasion risks of, and the potential of invasive spread in, UGAs in the world’s 100 most populous cities (in 40 countries). Based on maps of enhanced vegetation index at 250 m resolution over the extent of 25 by 25 km for each city centre, we simulate the invasion and spread of a reference species (a virtual ruderal invasive species) from the city centre into surrounding urban or rural areas. Doing so allowed us to provide an objective baseline for comparing urban susceptibility to such invasions across diverse cultures, histories and societies. We derive the global ranking of invasive spread potential for each city based on the rate of spread of the reference species, and the ranking of 40 countries, based on the average rate of spread in their cities. We explore correlates of spread rates after 100 time steps (years) by examining the roles of climate (mean annual temperature and rainfall), human demography (city population size and growth rate), and socio-economic indicators [human footprint, human development index and gross domestic product (GDP) per capita]. Small city population size and high GDP per capita are the only significant predictors of high potential for invasive spread. Among the G20 countries, Canada, South Korea, South Africa, France, USA and Brazil all feature in the top-10 countries, and Atlanta, Washington, D.C. and Dallas in the USA, Chittagong in Bangladesh, Toronto in Canada and Brasilia in Brazil are listed among the top 10 cities overall. Our results can serve as a global baseline assessment of invasive spread risks through UGAs, and call for improved protocols for monitoring, planning and management of UGAs.     

基于网络分析的城市生态空间结构优化——以常州市为例

经济社会的高速发展带来城市的快速扩张,造成城市生态空间的萎缩和生态功能的下降,城市生态安全受到严重威胁。系统研究城市生态空间结构,提出针对性保护和优化措施,对于城市的可持续发展具有重大意义。本研究以常州市为研究区,考虑城市生态空间的自然生态功能和社会服务功能两方面,构建基于自然生态的“源地-廊道”与基于人文生态的“供给-需求”两类生态网络,对于源地廊道生态网络,主要从节点重要性、网络连通性与稳定性进行定量分析,对于供给需求生态网络,主要从节点重要性、供需均衡与稳定性进行定量分析。结果表明: 常州市主城区源地廊道生态网络的连通性与稳定性水平不高,供给需求生态网络的稳定性水平一般且存在服务供给与需求的空间错位。从连通性与稳定性提升角度,提出新增12个源地节点与57条廊道的源地廊道生态网络优化方案;从供需均衡与稳定性提升角度,提出新增22个供给节点的供给需求生态网络优化方案。对比初始源地廊道生态网络,优化网络连通性水平提升10%,稳定性提升0.05;对比初始供给需求生态网络,优化网络的服务水平提升4%,网络稳定性提升0.10。最后,综合两类生态网络,分别针对现状保护斑块与新增节点两类对象,提出了保护与管理实施方案。     

Ecological genetics of invasive alien species

There is growing realisation that integrating genetics and ecology is critical in the context of biological invasions, since the two are explicitly linked. So far, the focus of ecological genetics of invasive alien species (IAS) has been on determining the sources and routes of invasions, and the genetic make-up of founding populations, which is critical for defining and testing ecological and evolutionary hypotheses. However an ecological genetics approach can be extended to investigate questions about invasion success and impacts on native, recipient species. Here, we discuss recent progress in the field, provide overviews of recent methodological advances, and highlight areas that we believe are of particular interest for future research. First, we discuss the main insights from studies that have inferred source populations and invasion routes using molecular genetic data, with particular focus on the role of genetic diversity, adaptation and admixture in invasion success. Second, we consider how genetic tools can lead to a better understanding of patterns of dispersal, which is critical to predicting the spread of invasive species, and how studying invasions can shed light on the evolution of dispersal. Finally, we explore the potential for combining molecular genetic data and ecological network modelling to investigate community interactions such as those between predator and prey, and host and parasite. We conclude that invasions are excellent model systems for understanding the role of natural selection in shaping phenotypes and that an ecological genetics approach offers great potential for addressing fundamental questions in invasion biology.     

Unlocking the potential of Google Earth as a tool in invasion science

Distribution data are central to many invasion science applications. The shortage of good information on the distribution of alien species and their spatial dynamics is largely attributable to the cost, effort and expertise required to monitor these species over large areas. Virtual globes, particularly Google Earth, are free and user-friendly software which provide high-resolution aerial imagery for the entire globe. We suggest this has enormous potential for invasion science. We provide suggestions and tools for gathering data on the distribution and abundance of invasive alien trees using visual interpretation of Google Earth imagery, and propose how these data may be used for a number of purposes, including calculating useful metrics of invasions, prioritising species or areas for management and predicting potential distributions of species. We also suggest various practical uses of Google Earth, such as providing a tool for early detection of emerging invasions, monitoring invasions over time, and to help researchers and managers identify suitable field study sites. Virtual globes such as Google Earth are not without limitations and we provide guidance on how some of these can be overcome, or when imagery from Google Earth may not be fit for invasion science purposes. Because of Google Earth’s huge popularity and ease of use, we also highlight possibilities for awareness-raising and information sharing that it provides. Finally, we provide the foundations and guidelines for a virtual global network of sentinel sites for early detection, monitoring and data gathering of invasive alien trees, which we propose should be developed as part of a “citizen science” effort. There has been limited use of virtual globes by invasion scientists and managers; it is our hope that this paper will stimulate their greater use, both within the field of invasion science and within ecology generally.     

Using spatially explicit commodity flow and truck activity data to map urban material flows

To analyze and promote resource efficiency in urban areas, it is important to characterize urban metabolism and particularly, material flows. Material flow analysis (MFA) offers a means to capture the dynamism of cities and their activities. Urban‐scale MFAs have been conducted in many cities, usually employing variants of the Eurostat methodology. However, current methodologies generally reduce the study area into a “black box,” masking details of the complex processes within the city's metabolism. Therefore, besides the aggregated stocks and flows of materials, the movement of materials—often embedded in goods or commodities—should also be highlighted. Understanding the movement and dispersion of goods and commodities can allow for more detailed analysis of material flows. We highlight the potential benefits of using high‐resolution urban commodity flows in the context of understanding material resource use and opportunities for conservation. Through the use of geographic information systems and visualizations, we analyze two spatially explicit datasets: (1) commodity flow data in the United States, and (2) Global Positioning System‐based commercial vehicle (truck) driver activity data in Singapore. In the age of “big data,” we bring advancements in freight data collection to the field of urban metabolism, uncovering the secondary sourcing of materials that would otherwise have been masked in typical MFA studies. This brings us closer to a consumption‐based, finer‐resolution approach to MFA, which more effectively captures human activities and its impact on urban environments.     

Secondary invasion: When invasion success is contingent on other invaders altering the properties of recipient ecosystems

Positive interactions between exotic species may increase ecosystem‐level impacts and potentially facilitate the entry and spread of other exotic species. Invader‐facilitated invasion success—”secondary invasion”—is a key conceptual aspect of the well‐known invasional meltdown hypothesis, but remains poorly defined and empirically underexplored. Drawing from heuristic models and published empirical studies, we explore this form of “secondary invasion” and discuss the phenomenon within the recognized conceptual framework of the determinants of invasion success. The term “secondary invasion” has been used haphazardly in the literature to refer to multiple invasion phenomena, most of which have other more accepted titles. Our usage of the term secondary invasion is akin to “invader‐facilitated invasion,” which we define as the phenomenon in which the invasion success of one exotic species is contingent on the presence, influence, and impacts of one or more other exotic species. We present case studies of secondary invasion whereby primary invaders facilitate the entry or establishment of exotic species into communities where they were previously excluded from becoming invasive. Our synthesis, discussion, and conceptual framework of this type of secondary invasion provides a useful reference to better explain how invasive species can alter key properties of recipient ecosystems that can ultimately determine the invasion success of other species. This study increases our appreciation for complex interactions following invasion and highlights the impacts of invasive species themselves as possible determinants of invasion success. We anticipate that highlighting “secondary invasion” in this way will enable studies reporting similar phenomena to be identified and linked through consistent terminology.     

Model invasions and the development of national concerns over invasive introduced trees: insights from South African history

This article examines how invasions within a discrete geographic, cultural, and ecological context disproportionately shaped awareness of invasions in other places. Such “model invasions” have been valuable for catalyzing national and international interest in biological invasion since the 1980s. Specifically, this article traces how scientific and public perspectives of invasive introduced trees evolved in South Africa during the twentieth century. It argues that concerns about the impact of invasive introduced trees first developed in the Mediterranean-climate region of the southwestern Cape Province during the 1940s and 1950s before emerging elsewhere in South Africa during the 1970s and early 1980s. Though there has been a nation-wide convergence in scientific and public views of invasive trees since the 1980s, there are still stark geographic and cultural knowledge divergences that hinder the effectiveness of contemporary invasive tree management efforts. This study suggests that geographical knowledge imbalances between regions should not be overlooked when historicizing or planning environmental management schemes at national scales.     

Are humans disturbing conditions in ecology?

In this paper I argue, first, that ecologists have routinely treated humans—or more specifically, anthropogenic causal factors—as disturbing conditions. I define disturbing conditions as exogenous variables, variables “outside” a model, that when present in a target system, inhibit the applicability or accuracy of the model. This treatment is surprising given that (1) humans play a dominant role in many ecosystems and (2) definitions of ecology contain no fundamental distinction between human and natural. Second, I argue that the treatment of humans as disturbing conditions is an idealization: since it is, and has long been, known that humans are pervasive, this treatment amounts to an intentionally introduced theoretical distortion. Finally, characterizing this treatment as idealization forces us to confront the question of its justification, and so, drawing on three different kinds of idealization, I evaluate how this treatment may be justified.     

The importance of biological inertia in plant community resistance to invasion

Abstract. Insights into the ecology of historic invasions by introduced species can be gained by studying long‐term patterns of invasions by native species. In this paper, we review literature in palaeo‐ecology, forest‐stand simulation modelling, and historical studies of plant species invasions to illustrate the relevance of biological inertia in plant communities to invasion ecology. Resistance to invasion occurs in part because of environmental, demographic, and biotic factors influencing the arrival and establishment of invading species. We propose that biological inertia within the resident community is a fourth component of resistance to invasion, because of the lag time inherent in eliminating resident species and perhaps their traces after environmental conditions become suitable for invasion by immigrating species. Whether or not an introduced species invades can be conditioned by the presence of the pre‐existing community (and/or its legacy) in addition to the other biotic and abiotic factors.     

The statistics of urban scaling and their connection to Zipf's law

Urban scaling relations characterizing how diverse properties of cities vary on average with their population size have recently been shown to be a general quantitative property of many urban systems around the world. However, in previous studies the statistics of urban indicators were not analyzed in detail, raising important questions about the full characterization of urban properties and how scaling relations may emerge in these larger contexts. Here, we build a self-consistent statistical framework that characterizes the joint probability distributions of urban indicators and city population sizes across an urban system. To develop this framework empirically we use one of the most granular and stochastic urban indicators available, specifically measuring homicides in cities of Brazil, Colombia and Mexico, three nations with high and fast changing rates of violent crime. We use these data to derive the conditional probability of the number of homicides per year given the population size of a city. To do this we use Bayes' rule together with the estimated conditional probability of city size given their number of homicides and the distribution of total homicides. We then show that scaling laws emerge as expectation values of these conditional statistics. Knowledge of these distributions implies, in turn, a relationship between scaling and population size distribution exponents that can be used to predict Zipf's exponent from urban indicator statistics. Our results also suggest how a general statistical theory of urban indicators may be constructed from the stochastic dynamics of social interaction processes in cities.     

Analysis of the impact of ancient city walls on urban landscape patterns by remote sensing

As the most important construction features of ancient Chinese cities, the city walls nowadays have lost their function of enemy defense and turned to affect the urban structure and development. To clarify the impact of ancient city walls on modern urban development, this work was conducted to measure the differences of landscape types and levels between the inner and outer walls of three typical ancient Chinese cities with the help of geoinformatics materials and landscape ecology indices. The results of this research proved that city walls have great impact on landscape pattern. Specifically, the aggregation, fragmentation, diversity and evenness of landscape were strongly affected by well-preserved ancient city walls. By sorting out and consulting historical documents and China's “City Walls Protection Regulations”, we also found that city walls help the old city to retain its original style and design characteristics. The findings of this quantitative-analysis-based historical study can provide a theoretical basis for the protection of historical heritage and landscape design.

     

The impact of interactions on invasion and colonization resistance in microbial communities

In human microbiota, the prevention or promotion of invasions can be crucial to human health. Invasion outcomes, in turn, are impacted by the composition of resident communities and interactions of resident members with the invader. Here we study how interactions influence invasion outcomes in microbial communities, when interactions are primarily mediated by chemicals that are released into or consumed from the environment. We use a previously developed dynamic model which explicitly includes species abundances and the concentrations of chemicals that mediate species interaction. Using this model, we assessed how species interactions impact invasion by simulating a new species being introduced into an existing resident community. We classified invasion outcomes as resistance, augmentation, displacement, or disruption depending on whether the richness of the resident community was maintained or decreased and whether the invader was maintained in the community or went extinct. We found that as the number of invaders introduced into the resident community increased, disruption rather than augmentation became more prevalent. With more facilitation of the invader by the resident community, resistance outcomes were replaced by displacement and augmentation. By contrast, with more facilitation among residents, displacement outcomes shifted to resistance. When facilitation of the resident community by the invader was eliminated, the majority of augmentation outcomes turned into displacement, while when inhibition of residents by invaders was eliminated, invasion outcomes were largely unaffected. Our results suggest that a better understanding of interactions within resident communities and between residents and invaders is crucial to predicting the success of invasions into microbial communities.     

Impact of the Economic Structure of Cities on Urban Scaling Factors: Implications for Urban Material and Energy Flows in China

We explore the population‐scaling and gross domestic product (GDP)‐scaling relationships of material and energy flow (MEF) parameters in different city types based on economic structure. Using migration‐corrected population data, we classify 233 Chinese city propers (Shiqu) as “highly industrial” (share of secondary GDP exceeds 63.9%), “highly commercial” (share of tertiary GDP exceeds 52.6%), and “mixed‐economy” (the remaining cities). We find that, first, the GDP population‐scaling factors differ in the different city types. Highly commercial and mixed‐economy cities exhibit superlinear GDP population‐scaling factors greater than 1, whereas highly industrial cities are sublinear. Second, GDP scaling better correlates with city‐wide MEF parameters in Chinese cities; these scaling relationships also show differences by city typology. Third, highly commercial cities are significantly different from others in demonstrating greater average per capita household income creation relative to per capita GDP. Further, highly industrial cities show an apparent cap in population. This also translates to lower densities in highly industrial cities compared to other types, showing a size effect on urban population density. Finally, a multiple variable regression of total household electricity showed significant and positive correlation with population, income effect, and urban form effect. With such multivariate modeling, the apparent superlinearity of household electricity use with respect to population is no longer observed. Our study enhances understanding of MEFs associated with Chinese cities and provides new insights into the patterns of scaling observed in different city types by economic structure. Results recommend dual scaling by GDP and by population for MEF parameters and suggest caution in applying universal scaling factors to all cities in a country.     

So what is a species anyway? A primatological perspective

Since Darwin's time, the question “what a species” has provoked fierce disputes and a tremendous number of publications, from short opinion papers to thick volumes. 1 The debates covered fundamental philosophical questions, such as: Do species exist at all independently of a human observer or are they just a construct of the human mind to categorize nature's organismic diversity and serve as a semantic tool in human communication about biodiversity? 2 - 4 or: Are species natural kinds (classes) or individuals that are “born” by speciation, change in course of time, and finally “die” when they go extinct or diverge into new species? 5 - 8 Also included was the problem of species as taxa (taxonomic) versus species as products of the speciation process (evolutionary). 9 More pragmatic issues arose, such as: How can we reliably delineate and delimitate species? 10 , 11 The great interest in what a species is reflects the importance of “species” as fundamental units in most fields of biology, especially evolutionary biology, ecology, and conservation. 2 , 12 - 14     

Deciphering the worldwide invasion of the Asian long‐horned beetle: A recurrent invasion process from the native area together with a bridgehead effect

Retracing introduction routes is crucial for understanding the evolutionary processes involved in an invasion, as well as for highlighting the invasion history of a species at the global scale. The Asian long‐horned beetle (ALB) Anoplophora glabripennis is a xylophagous pest native to Asia and invasive in North America and Europe. It is responsible for severe losses of urban trees, in both its native and invaded ranges. Based on historical and genetic data, several hypotheses have been formulated concerning its invasion history, including the possibility of multiple introductions from the native zone and secondary dispersal within the invaded areas, but none have been formally tested. In this study, we characterized the genetic structure of ALB in both its native and invaded ranges using microsatellites. In order to test different invasion scenarios, we used an approximate Bayesian “random forest” algorithm together with traditional population genetics approaches. The strong population differentiation observed in the native area was not geographically structured, suggesting complex migration events that were probably human‐mediated. Both native and invasive populations had low genetic diversity, but this characteristic did not prevent the success of the ALB invasions. Our results highlight the complexity of invasion pathways for insect pests. Specifically, our findings indicate that invasive species might be repeatedly introduced from their native range, and they emphasize the importance of multiple, human‐mediated introductions in successful invasions. Finally, our results demonstrate that invasive species can spread across continents following a bridgehead path, in which an invasive population may have acted as a source for another invasion.     

Ecology and urban planning

Urban areas harbour diverse nature ranging from semi-natural habitats to wastelands, parks and other highly human-influenced biotopes with their associated species assemblages. Maintenance of this urban biodiversity for the residents and for its intrinsic value in the face of increasing population and expanding cities requires that ecological knowledge should be better integrated into urban planning. To achieve this goal understanding of ecological patterns and processes in urban ecosystems is needed. The first step in the necessary urban ecological research is to find out what kind of nature exists in cities. Second, knowledge about ecological processes important in urban nature is required. Although ecological processes in cities are the same as in rural areas, some of them, such as invasion by alien species, are more prevalent in urban than in rural conditions. Third, based on ecological knowledge, management schemes maintaining the diversity of urban nature should be designed. These procedures should also include protection of urban nature, e.g. in urban national parks. Finally, as ecology alone cannot provide the complex information about human influence on urban ecosystems, interdisciplinary research involving natural and social sciences is imperative for a holistic approach to integrating ecology into the process of urban planning.     

The Nature of Urban Soils and Their Role in Ecological Restoration in Cities

Current and predicted trends indicate that an increasing proportion of the world’s population is living in urban and suburban places. The nature of the urban environment becomes an important factor if we are concerned with the restoration and preservation of biodiversity and ecosystems in and around cities. This article highlights the varied impacts of cities on soils and their implications for restoration planning and expectations of restoration “success.” Urban soils exist in different historical and formational trajectories than their local nonurbanized counterparts due to direct anthropogenic disturbance and indirect environmental impacts from urbanization. Therefore, urban soils often exhibit altered physical, chemical, and biological characteristics in comparison to local nonurbanized soils. Several unique features of urban soils and urban ecosystems pose particular issues for ecological restoration or the improvement of degraded soil conditions in cities. The creation of novel soil types, conditions that promote invasion by non‐natives, the strong influence of past land use on soil properties, and the presence of strong interactions and alternative stable states set up unique difficulties for the restoration of urban soils. Soils in urban restorations are a medium that can be deliberately manipulated to improve site conditions or in the monitoring of soil conditions as indices of ecosystem status. Including an explicit role for strong manipulations of soils in urban ecosystems changes how we approach baselines, management, and reference conditions in urban ecological restoration. With an understanding of urban soil ecological knowledge, we can guide aspects of urban ecological restoration toward successful outcomes.     

我国城市人与自然耦合系统的协调度

城市作为人类生存与发展的主要空间载体,是人与自然耦合系统的典型代表。城市人与自然协调度是认识人与自然耦合机制重要内容,对揭示城市人类活动对生态环境的影响,指导城市建设具有重要意义。从水资源开发强度、土地开发强度、水资源供给能力、环境污染物排放强度与碳排放强度五个方面分析城市人与自然协调度特征,评估城市建设与发展对自然环境的影响程度,并评估了我国146个城市的协调度特征,进一步分析了城市人口规模、城市经济规模、城市社会发展水平、不同生态地理区人与自然耦合协调度的关系。结果表明：2016年城市平均协调度指数为87.90,不同城市的协调度差距很大且协调度与城市人口规模、经济规模、发展水平呈显著负相关,而六大生态地理区中,西南地区协调度最高,协调度指数为92.81,西北地区协调度最低,协调度指数为82.25。研究发现我国城市建设与发展对自然环境影响仍然较大,并表现为（1）我国城市发展总体仍处于高需求高排放的发展阶段;（2）生态环境承载力是影响城市人与自然耦合协调度的重要因素;（3）提高能源利用效率,降低碳排放强度在改善城市人与自然协调度中发挥作用。最后,从优化城市布局、提高水、能源利用效率、控制环境污染等方面提出政策建议,为评估城市生态文明建设与可持续发展提供参考与依据。     

海绵城市研究进展综述:从水文过程到生态恢复

在我国快速城市化进程中,不合理的规划建设加剧了城市洪涝、城市缺水、生态功能降低等诸多问题,受国外低影响开发理念启发,国内学者提出了与城市水文过程紧密结合的解决之道——"海绵城市"。在梳理海绵城市概念、内涵和发展现状的基础上,讨论了海绵城市建设与城市水文过程之间的作用关系,指出海绵城市研究涵盖水环境、水资源、水安全和水生态等众多领域且不断融合拓展。归纳了海绵城市的发展趋势,强调应更加遵循因地制宜原则,积极响应多尺度气候变化,科学认知城市水文过程并完善城市生态系统构建。提出了海绵城市建设优化框架,在理论上从单纯地关注水文过程转变到全方位倡导生态水文修复,在技术方法上重点融合生态水文途径改善城市水文空间体系,将生态恢复作为重要建设方式来提升城市应对灾害的弹性。